Veterinary Nutrition and Dietetics PDF

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Veterinary Nutrition and Dietetics
First Edition
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VETERINARY
NUTRITION AND DIETETICS
First Edition
Edited by:
Sándor Gy. Fekete, DVM, PhD, DSc

Professor of Veterinary Nutrition and Laboratory Animal Science
Formerly Head of the Department of Animal Nutrition
Faculty of Veterinary Science Budapest,
Szent István University Gödöllő, Hungary
With 31 Co-Authors
“Pro Scientia Veterinaria Hungarica“
Budapest
2008
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“Pro Scientia Veterinaria Hungarica“ Fooundation
© The Foundation (as a whole) and The Authors (their sections) 2008. All rights reserved
No part of this manual may be reproduced or used in any form or by any means, electron-
ic or mechanical, including photocopying or by any information storage and retrievel sys-
tem, without the written permission of the Authors and Publisher
First Paperback Editon 2008

ISBN: 978-963-06-5166-0
Reviewers of the Hungarian Version

-Emese Andrásofszky, Head of Laboratory
-Prof. Dr. Lajos Fekete, DSc
-Habil. Prof. Dr. István Hullár, CSc
-Prof. Dr. Gyula Huszenicza, DSc
-Prof. Dr. Miklós Mézes, DSc
Reviewers of the English Version (selected chapters):

-Assoc. Prof. Dan L. Brown, PhD (Conell University, NY, USA)
-Prof. Richard O. Kellems, PhD (BYU University, UT, USA)
-James Sales, PhD (University of South Czech Reoublic)
-Prof. Dr. János Vetter, DSc (SZIU Faculty of Veterinary Science)
Language Reviewers
-Dr. habil. Sándor Fekete, PhD (SZIU Gödöllő)
-Dr. András Székely
-Katalin Tátrai
Special Note: Readers are advised to check the most current information before applying
any included data. To the fullest extent of the law, neither the authors nor the publisher
assume any liability for any injury or damage.
The Publisher: “Pro Scientia Veterinaria Hungarica“ Foundation

H-1077 Budapest, Rottenbiller u. 50. Hungary
Picture on the cover page: Delacroix. Moroccan and his horse; Madas-Hamilton: Cock,
pigeions and guinea pig © Museum of Fine Arts Budapest, Hungary
Printing office: OOK-Press Kft, Veszprém, Hungary.

Responsible manager: Attila Szathmáry
PRINTED IN HUNGARY
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PREFACE
his English-language textbook is intended for veterinary students, vet-
T erinary practitioners and other professionals working in the field of

veterinary medicine, including veterinary surgeons and nurses.
Although numerous good textbooks on nutrition are available in the agri-
cultural literature, the last manual using a veterinary approach was pub-
lished in 1961. The Hungarian version of the present manual was published
in 2003. Sensing the need for an international version, a wide range of lead-
ing specialists have been gained as contributors. The value of the material
is proved by more than 32 years of teaching practice and the success
achieved at several international competitions, including three first “Best of
Class” (BOC) prizes won in Europe, as well as a second and a third BOC
prize.
In the present publication practically all of the important topics are
represented. The scope of the book will be veterinary nutrition ranging from
pets to cattle, and everything in between, including fish and laboratory ani-
mal nutrition. Feeding and nutrition of the species are classified according
to the evolutionary status of the digestive system: omnivores, carnivores,
herbivores and ruminants are presented separately and the last chapters
are devoted to the nutrition of small and exotic pets and laboratory animals.
The textbook encompasses the full range of the curriculum of veterinary
schools and future diplomates of the European College of Veterinary and
Comparative Nutrition (ECVCN). Needless to say that such a book is unique
and gap-filling, and is expected to become an international standard in vet-
erinary nutrition. A list of self-evaluating questions (SEQs) and a collection
of abbreviations facilitate the students’ learning process. The entire text has
undergone thorough review. Reviewers are highly recognised specialists of
the field. Prof. Bernard Paragon (Alfort) and Prof. Patrick Nguyen (Nantes)
provided great help also by their valuable suggestions. I am obliged to my
Faculty of Veterinary Science Budapest and to the Hungarian-American
Entreprise Scholarship Fund (HAESF) for supporting my sabbatical in 2006
at Brigham Young University (Provo, UT, USA) and Cornell University (New
York, NY, USA).
The authors intended to serve the needs of veterinary practitioners
and veterinary students. Therefore, the topics are treated from a clinical
approach, and special attention is given to deficiency syndromes and meta-
bolic disorders. The authors have continuously kept in view the mission
statement of the European Society of Veterinary and Comparative Nutrition.
I would like to emphasize my appreciation for the understanding and help
to my wife, Mária Siklódi, who provided with her ideas and patience the
spiritual background of this work.
Budapest-Zamárdi-Zebegény, 2008
Sándor György Fekete
www.dietvet.hu
V
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CONTRIBUTORS
Prof. Dr. habil. László BABINSZKY, PhD (Wageningen)

Chairman of the Editorial Board of Animal Feeding & Nutrition
(“Takarmányozás”), Director of Institue for Animal Physiology and
Nutrition, Head of the Department of Animal Nutrition at the
University of Kaposvár, H-7401 Kaposvár, POB 16. (Hungary),
babin@u-kaposvar.hu
Assoc. Prof. Dr. András BERSÉNYI, DVM, PhD (Budapest

Institute of Animal Breeding, Nutrition and Laboratory Animal
Science, Szent István University, Faculty of Veterinary Science
Budapest, H-1400 Budapest, POB 2. (Hungary),
Bersenyi.Andras@aotk.szie.hu
Assoc. Prof. Dr. Géraldine BLANCHARD, DVM, PhD (Alfort)

Dipl. ESVCN, Alfort National Veterinary School, Nutrition Unit, F-
94704 Maison Alfort cedex, (France), gblanchard@vet-alfort.fr
Assoc. Prof. Dan L. BROWN, PhD (Cornell, NY)

New York College of Agriculture & Life Sciences, A statutory College
of the State University, Cornell University, Department of Animal
Science, Morrison Hall, Ithaca, N.Y. 14853 (USA),dlb20@cornell.edu
Prof. Dr. Tony C. A. BUFFINGTON, DVM, PhD (Davis, CA)

Dipl. ACVN, The Ohio State University, Veterinary Teaching
Hospital, 601 Tharp Street, Columbus, OH 43210-1089 (USA),buff-
ington.1@osu.edu
Prof. Dr. habil. Burghard JILGE, DVM, PhD

Director of the Animal Research Cenre of the University of Ulm
(Germany)
Prof. D. Carlos CASTRILLO Gonzales, PhD (Leon)

Chairman of the Department of Animal Production and Food
Science. Veterinary Faculty. Miguel Servet 177. 50013 Zaragoza
(Spain), ccastri@posta.unizar.es
Prof. Dr. habil. Éva CENKVÁRI, MSc, CSc

Institute of Animal Breeding, Nutrition and Laboratory Animal
Science, Szent István University, Faculty of Veterinary Science
Budapest, H-1400 Budapest, POB 2. (Hungary),
Cenkvari.Eva@aotk.szie.hu
VI
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Prof. André CHWALIBOG, DSc (University of Copenhagen)

Leader of the Centre of Experimental Animal Nutrition and
Physiology, Leader of the Research School in Animal Nutrition and
Physiology, Head of Division of Nutrition at the University of
Copenhagen, Faculty of Life Sciences, Department of Basic Animal
and Veterinary Sciences, Grønnegårdsvej 3, 1870 Frederiksberg C
(Denmark), ac@life.ku.dk
Prof. Dr. habil. János CSAPÓ, MSc, DSc (Hungarian Academy of Sciences)
Director of the Chemical Institute and Head of the Department of
Biochemistry and Food Chemistry at the University of Kaposvár, H-
7401 Kaposvár, POB 16. (Hungary), Csapo@mail.atk.kaposvar.hu
Prof. Dr. Sándor György FEKETE, DVM, MSc, PhD (Moscow-Budapest),

DSc (Hungarian Academy of Sciences), Dipl. ECVCN, Chairman of
the HLASA, Natl. Repr. Of ESLAV, Institute of Animal Breeding,
Nutrition and Laboratory Animal Science, Szent István University,
Faculty of Veterinary Science Budapest, H-1400 Budapest, POB 2.
(Hungary), dietvet@yahoo.com; www.dietvet.hu
Prof. Dr. Jose Antonio GUADA Vallepuga, DVM, PhD

Department of Animal Production and Food Science. Veterinary
Faculty. Miguel Servet 177. 50013 Zaragoza (Spain),
jguada@unizar.es
Prof. Dr. habil. János GUNDEL, MSc, CSc (Hungarian Academy of Sciences)
Editor-in-chief of the Journal of Animal Production (“Állattenyésztés
és takarmányozás”), Vice Generaldirector of Research Institute for
Animal Breeding and Nutrition, H-2053 Herceghalom (Hungary),
jgundel@atk.hu
Prof. Dr. Gyula HUSZENICZA, DVM, PhD (Budapest),

DSc (Hungarian Academy of Sciences) Dipl. of ECAR
Szent István University, Faculty of Veterinary Science Budapest,
Department for Obstetrics and Reproduction, H-1078 Budapest,
Istvan u. 2. (Hungary), Huszenicza.Gyula@aotk.szie.hu
Prof. Dr. Christine IBEN, DVM, PhD (Vienna)

Professor at the Institute of Nutrition, Department for Farm Animals
and Veterinary Public Health, University of Veterinary Medicine
Vienna, Veterinärplatz 1, A-1210 Wien (Austria), Christine.Iben@vu-
wien.ac.at
VII
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Prof. Dr. hab.dr h.c., dr h.c. Dorota JAMROZ

Chairman of the Polish Husbandry Committee of Polish Academy of
Science, Head of the Department of Animal Nutrition and Feed
Quality, Wroclaw University of Environmental and Life Sciences, ul.
Chelmonskiego 38C, 51-630 Wroclav (Poland),
djamroz@zoo.ar.up.wroc..pl
Prof. Geert P. J. JANSSENS, PhD (Ghent)

Professor in Animal Nutrition at the Faculty of Veterinary Medicine
of Ghent University, Head of the Laboratory of Animal Nutrition.
Heidestraat 19 B-9820 Merelbeke (Belgium),
Geert.Janssens@UGent.be
Prof. Dr. Ǻshild KROGDAHL, PhD (Oslo)

Professor in Animal Nutrition, Leader of the Gut and Health Group of
the Aquaculture Protein Centre (CoE), Department of Basic Sciences
and Aquatic Medicine, Norwegian School of Veterinary Science, P.O
Box 8146 Dep., N-0033 Oslo (Norway). ashild.krogdahl@veths.no
M.Sc. eng. Janusz KUBIZNA

Department of Animal Nutrition and Feed Quality, Wroclaw
University of Environmental and Life Sciences, ul. Chelmonskiego
38C, 51-630 Wroclav (Poland), janusz.kubizna@up.wroc.pl
Prof. Dr. Dr. h.c. mult. Josef LEIBETSEDER, DVM

General of the Thematic Priority “Food Quality and Safety” Advisory
Group of the Research Directorate, Co-founder of European Society
of Veterinary and Comparative Nutrition, h.c. Diplomate of
European College of Veterinary and Comparative Nutrition,
President of “Animals for Therapy” Society. Institute of Nutrition,
University of Veterinary Medicine Vienna, Veterinärplatz 1, A-1210
Wien (Austria), josef.leibetseder@vu-wien.ac.at
Dr. Richard C. NAP, DVM, PhD (Utrecht)

Consultant to the Animal Health community. Director Vetstart
International Ltd. UK. Former Associate-Professor Orthopedic
Medicine and former Associate-Director External Relations, Procter
& Gamble. Diplomate European College of Veterinary Surgeons and
Diplomate College of Veterinary and Comparative Nutrition.
Uppertunity Consultants. E-mail: richard@uppertunity.com
VIII
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Dr. Fatma Karakas OGÜZ, DVM, PhD (Burdur)

Assoc. Prof. at the Department of Animal Nutrition and Animal
Diseases of Burdur Veterinary Faculty, Mehmet Akif Ersoy
University, Burdur (Turkey), fatmakarakas@yahoo.com
Prof. Dr. Merel RITSKES-HOITINGA, PhD, Dipl ECLAM

In 2005 and 2006: president of FELASA (Federation of European
Laboratory Animal Science Associations). Managing director of the
Central Animal Facility of the Radboud University Medical Center in
Nijmegen, the Netherlands (The Netherlands), Professor in
Laboratory Animal Science, Founder of the 3RRC
(www.umcn.nl/3rrc) M.ritskes@cdl.umcn.nl
…Prof. Dr. Peter RUDAS, DVM, PhD (Budapest), DSc (Hungarian Academy
of Sciences), President of International Society of Farm Animal
Endocrinology, Vice President of the World Organisation of
Hungarian Veterinarians, Head of the Department of Physiology and
Biochemistry, SZIU Faculty of Veterinary Science Budapest, H-1400
Budapest, POB 2.
James SALES, PhD (Agric.), PhD (Fish.Sci.)
Member of the Steering Committee of World Poultry Science
Association (Ph.D. Agric., Ph.D. Fish. Sci.), Research associate at the
University of South Bohemia in Ceske Budejovice, Research
Institute of Fish Culture and Hydrobiology at Vodnany (USB RIFCH)
Zatisi 728/II, 389 25 Vodnany, (Czech),
James_Sales_1@hotmail.com
Bart SAVENIJE, PhD (Groningen)

coordinator of the 3R Research Centre at the Central Animal Facility
of the Radboud University Nijmegen Medical Centre (The
Netherlans), B.Savenije@cdl.umcn.nl
Prof. Ewa SAWOSZ, Dr. habil.

Head of the Department of Animal Nutrition and Feed Science,
Warsaw University of Life Sciences, Ciszewskiego 8, 02-786 Warsaw
(Poland) ewa_sawosz@sggw.pl
Prof. Anne-Helene TAUSON, PhD

Professor of Clinical Animal Nutrition, Department of Basic Animal
and Veterinary Sciences, Faculty of Life Sciences, University of
Copenhagen, Grønnegårdsvej 3, DK-1870 Frederiksberg C,
(Denmark), aht@life.ku.dk
IX
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Dr. Angela TROCINO, PhD (Padova)

Contact Researcher, Member of the World Rabbit Science Association
(WRSA),Department of Animal Science, Faculty of Agriculture,
University of Padova, (Italy) angela.trocino@unipd.it
Prof. Gerolamo XICCATO, PhD (Padova)

President of the Italian Rabbit Science Association, President of World
Rabbit Science Association., Full professor in Poultry and Rabbit
Science at the Department of Animal Science, Faculty of Agriculture,
University of Padova, (Italy) gerolamo.xiccato@unipd.it
Prof. Dr. Annette ZEYNER, Habil. (Göttingen)

Professor for Nutritional Physiology and Animal Nutrition, Institute of
Farm Animal Sciences and Technology, Faculty of Agricultural and
Environmental Sciences, University of Rostock, Justus-von-Liebig-Weg
8, D-18059 Rostock, (Germany), annette.zeyner@uni-rostock.de
X
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TABLE OF CONTENTS
Page
Preface
Contributors
Table of Content HELYE !!!!!!
A) GENERAL PART
Introduction
I. Importance of animal nutrition. Basic notions about feedstuffs.
Role of water in animal organism (Sandor Gy. FEKETE)
II. Animal nutrition, health and food safety (Josef LEIBETSEDER)
III. Composition of the plant and animal body; measuring methods
(Sandor Gy. FEKETE)
IV. Chemical analysis and metabolic importance of feed ingredients
(Janos CSAPO)
V. Basics of material, energy and water balance
(Andre CHWALIBOG and Ewa SAWOSZ
VI. Regulation of energy metabolism and voluntary feed intake
6.1. Overview of endocrine control system of energy balance
(†Péter RUDAS)
6.2. Practical aspects of feed intake regulation
(Sandor Gy. FEKETE)
VII. Digestibility of nutrients
7.1. General considerations regarding digestibility and its determi
nation (Sandor Gy. FEKETE)
7.2. Factors affecting digestibility (Sandor Gy. FEKETE)
7.3. Ileal digestibility of amino acids in pigs and poultry feeds and
their use in diet formulation (Laszlo BABINSZKY)
VIII. Biologically active secondary plant products
8.1. Antinutritíonal factors and antimetabolites. (Sandor Gy. FEKETE)
8.2. Plant poisonings and selected phytotoxins (Dan L. BROWN)
IX. Energetic evaluation of feeds (Sandor Gy. FEKETE)
Poultry Section: Fatma KARAKAS OGUZ
X. Protein evaluation for monogastrics and ruminants
(Carlos CASTRILLO Gonzales, Eva CENKVARI and
Sandor Gy. FEKETE)
10.1. Monogastrics
10.2. Ruminants
10.2.1. Physiological basics
10.2.2. Overview of countries
XI
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XI. Minerals in the animal nutrition (Sandor Gy. FEKETE)

11.1. Mineral composition of the animal body and feedstuffs miner-
als. General part.
11.2. Nutritional importance of minerals. Special part.
XII. Vitamins in the animal nutrition (Sandor GY. FEKETE)
12.1. Generally abouts vitamins
12.2. Nutritional importance of vitamins. The individual vitamins.
XIII. The ruminal and intestinal flora (Sandor GY. FEKETE)
13.1. Intestinal flora and fauna of monogastric animals
13.2. Ruminal flora and fauna
XIV. Deterioration of feeds, their microbiology and mycology.
(Sandor Gy. FEKETE)
14.1. Deterioration of feeds. Warranty questions
14.2.Microbiological deterioration and mycotoxins contamination of
feeds
XV. Feed additives and nutriceuticals (Sandor Gy. FEKETE )
15.1. Generally about growth promoters
15.2. Direct effect on metabolism
15.3. Influencing the ruminal and intestinal flora
15.4. Nutriceuticals efforts
15.5. Stress – Immune status – Performance enhancers
XVI. Nutrition-reproduction interaction
16.1. Effect of nutrition on the reproductive function
(Sandor Gy. FEKETE
16.2. Endocrine and metabolic factors of reproductive functions of
dairy cows (Gyula HUSZENICZA)
XVII. Interrelationships of feeding with immunity, parasitic infection and
genomics (Sandor Gy. FEKETE)
17.1. The nutrition and the immune functions; the acute phase
17.2. Nutrition and immunodeficiency
17.3. Influence of nutrient deficiency on the immunity
17.4. Feed intolerance, allergy and aversion
17.5. Nutrition and parasitic infection
17.6. Nutrigenomics
XVIII. Feed law and relating regulation in the EU
(Josef LEIBETSEDER)
B) APPLIED ANIMAL NUTRITION
XIX. Evolution of digestion (Sandor Gy. FEKETE)

XX. Feeding and nutrition of pig
Digestive physiology of pig (Sandor Gy. FEKETE)
Feeding of piglets, weaning systems (Sandor Gy. FEKETE)
Theoretical basics of pig growth (“fattening”) (Sandor Gy. FEKETE)
Applied nutrition of fattening (growing-finishing) pig (Janos GUNDEL)
XII
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Feeding and nutrition of breeding sow and boar (Sandor Gy. FEKETE)
Feed related diseases, herd health aspects (Sandor Gy. FEKETE)
XXI. Feeding and dietetics of dog and cat
(Sandor Gy. FEKETE, if it is not otherwise indicated)
21.1. .Feeding and dietetics of healthy dog and cat
21.2. Theoretical bases of dog and cat dietetics
-Feline Liver Diseases (Geraldine BLANCHARD)
-Nutritional Therapy of Stones in the Urinary Bladder
of Cats and Dogs (Tony C.A. BUFFINGTON)
-Nutrition and the skeletal system in dogs
(Richard C. NAP)
21.3. Practical aspects of dog and cat dietetics
XXII.Feeding and nutrition of poultry
22.1. Digestive characteristics of birds (Sandor Gy. FEKETE)
22.2. Feeding and nutrition of healthy poultry
(Dorota JAMROZ and Janusz KUBIZNA)
-Ostrich nutrition (Sandor Gy. FEKETE)
22.3. The most important digestive and metabolic diseases of poultr
(Sandor Gy. FEKETE)
XXIII.Feeding and nutrition of the rabbit
23.1. Digestive physiological characteristics (Sandor Gy. FEKETE)
23.2. Nutrition and feeding of breeding does and bucks
(Gerolamo XICCATO and Angela TROCINO)
23.3. Practical rabbit nutrition (Sandor Gy. FEKETE)
23.4. Digestive troubles and nutritional toxicosis in rabbits
(Sandor Gy. FEKETE)
XXIV. Horse feeding and nutrition
24.1. Digestive characteristics of horses (Sandor Gy. FEKETE)
24.2. Nutrient requirements of horses (Annette ZEYNER)
24.3. Practical nutrition of the healthy horse
(Annette ZEYNER and Sandor Gy. FEKETE
24. 4. Realisation of the practical nutrition of the healthy horse
(Sandor Gy. FEKETE)
24.4. Horse dietetics. (Sandor Gy. FEKETE)
XXV. Cattle feedings and dietetics (Sandor Gy. FEKETE)
25.1. Dairy heifer and cow
25.2. Beef heifer and cow
25.3. Beef fattening
25.4. Herd health
XXVI. Sheep feeding and metabolic troubles:
26.1. Breeding sheep (Jose Antonio GUADA Vallepuga)
26.2. Growing-fattening lamb (Sandor Gy. FEKETE)
26.3. Deficiency symptoms and metabolic troubles
(Sandor Gy. FEKETE)
XXVII. Goat feeding and metabolic troubles
(Sandor Gy. FEKETE and Eva CENKVARI)
XIII
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27.1. ”Standard” European goat

27.2. Biological adaptations in goats
27.3. Production and reproduction cycles of dairy goats
27.4. Nutrient requirements
27.5. Feeding the kids.
27.6. Feeding of growing goats from 7 to 12 month of age
27.7. Feeding of dairy goat
27.8. Nutrition and feeding of breeding bocks
27.9. Feeding strategies in goats
27.10. Frequent feeding systems of dairy goats
27.11. Metabolic disorders and nutrition related diseases
XXVIII. Production fish nutrition and feeding (Ǻshild KROGDAHL)
28.1. Physiological characteristics of importance for
nutrient requirement and feeding
28.2. Fish and the environment
28.3. Nutrient supply
28.4. Nutrition and feeding of fish larvae
28.5. Nutrition-related disorders
XXIX. Keeping and feeding of exotic animals (Sandor Gy. FEKETE)
29.1. Environment
29.2. Feeding and nutrition
XXX. Feeding of ornamental fish, amphibia and reptiles
(Sandor Gy. FEKETE and James SALES)
30.1. Feeding of ornamental fish
30.2. Feeding of amphibians
XXXI. Feeding and housing of small mammals (Christine IBEN)
31.1. Ration design and feed composition
31.2. Special needs of the individual species
31.3. Feeding of neonates
XXXII. Cage bird and pigeon feeding and nutrition
(Geert JANSSENS and James SALES)
32.1. Cage birds
32.2. Pigeons
XXXIII. Feeding and nutrition of laboratory animals
33.1. Zoological features, nutrient requirements (Sandor Gy. FEKETE
33.2. New FELASA recommendation: transport, harmfull material,
environmental enrichment, circadian rhythm.
(Merel RITKES-HUIZINGA, Bart SAVENIJE and Jilge BUGHART)
XXXIV. Furbearer animal feeding and nutrition
34.1. Nutrient and energy utilization in fur animal production
(Anne-Helen TAUSON)
34.2. Deficiency symptoms and metabolic troubles
(Andras BERSENYI)
LIST OF ABBREVIATION
Self Evaluating Questions (SEQs)
XIV
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INTRODUCTION
The evolution of veterinary nutrition and clinical dietetics into

an independent discipline had started in the distant past. The main
reason for this is that the majority of the world’s human population
are not vegetarians but consume more or less products of animal
origin as well. Thus the health status and reproductive potential of
animals are closely related to the well-being of human societies,
since “a bite to eat every day is the most fundamental and essen-
tial need of humans. Food is also a source of pleasure, and this
dual role makes it a focus of tension and complex emotions strong-
ly affected by power relations and social inequalities” (MASSIMO
MONTANARI: Hunger and abundance. The European cultural history
of nutrition. Atlantisz, Budapest, 1996). Further links were repre-
sented by the role of dogs, a species domesticated very early, in
hunting, house-watching and guarding grazing animals, not to
speak of the role played by horses in warfare.
I will only highlight the main milestones in the unique devel-
opment of this field of science. Although already the book of MÁRTON
TSEH (Martin Böhme) first published in Hungarian in 1656 (“New
Booklet on the Proven Medicines of Horses”) had contained dietetic
references (e.g. “how to treat a small foal if one wishes to accustom
it to fodder”), it still contained a mixture of medieval superstitions
and rational empiricism. In GYÖRGY FEKÉSHÁZY’s book published in
Pest in the year 1794 (“On the cattle cadaver, its original causes
and own medicines”) separate chapters were devoted to nutrition
(“Hunger and Bad Food”, “Feed”, “Bad Water, Bad Drink”). When
veterinary education was introduced in Hungary in 1787, nutrition
became part of its regular curriculum by the name “dietetics”,
under the professorship SÁNDOR TOLNAI (1787–1818). During the
time when BÉLA TORMAY was professor of animal breeding
(1873–1888), the subjects taught by his department, including ani-
mal nutrition, reached the contemporary world standard. TORMAY
already distinguished and considered essential the veterinarian’s
tasks related to nutrition. “Through their studies and their dual
qualifications gained through the former, veterinarians should be
qualified to participate in animal breeding, animal nutrition corre-
1
sponding to the recommendations, proper animal care, the ration-

al use and utilisation of domestic animals and the most advanta-
geous use of animal products” – writes TORMAY in his work entitled
“The veterinarian’s tasks in public breeding” (Hungarian Veterinary
Society, Budapest, 1902, Veterinary Manuals, Volume V, page 11).
The first handbook that contained the words “veterinary
dietetics” already in its title, was published in Cambridge in 1921
(R. G. LINTON: Animal Nutrition and Veterinary Dietetics). In
Hungary, during the period when Professor ZOLTÁN CSUKÁS acted as
head of the Department of Animal Breeding (1946–1957), from
1951 the veterinary students studied animal nutrition as a sepa-
rate subject in the second semester of the second year of studies,
in three hours of theoretical and two hours of practical training per
week (total: 75 hours). This semester was closed by an independent
final examination. After the second year of studies, veterinary stu-
dents had to complete a one-month practice in a state farm where
animal breeding was practised. The conscious veterinary attitude of
Professor Csukás to the discipline of animal nutrition is well reflect-
ed in his book written in 1952, from the preface to which I would
like to quote the following sentence: “Since the increasing intensity
of production results in a faster reproduction rate and increasing
production level of our animals and partially also in the decreasing
conformity of animal feeds with the natural feed qualities, it is obvi-
ous that we have to increase and update our knowledge of the
metabolic aspects of reproductive phenomena (oestrus, gestation,
milk production, growth) and the biological evaluation of feeds so
that we can serve the interests of live veterinary practice.” He also
states that he intends his book for his students and colleagues. His
successors at the Department, Professors JÓZSEF MÁRKUS, SÁNDOR
SZENTMIHÁLYI, KÁROLY BAINTNER and JÓZSEF BOKORI, preserved the
attitude of Professor Csukás, improving and maintaining the sub-
ject of animal nutrition on the contemporary world standard.
The manual published in Edinburgh in 1961 (J. T. ABRAMS:
Animal Nutrition and Veterinary Dietetics. W. Green & Son, Ltd.)
already devotes a separate chapter to defining the criteria of veteri-
nary dietetics as opposed to the earlier used term, “feeding”; name-
ly, it dwells at length on interactions of the host animal with the
rumen and intestinal microflora, the effects of feed on animal prod-
ucts, mineral and vitamin deficiencies, and malnutrition. In the
years that have elapsed since that time, clinical dietetics, which
had previously been taught in the framework of clinical subjects,
2
has developed into an independent discipline separately for all

species of domestic animals. A very important milestone in this
process was the establishment of the “European Society of
Veterinary and Comparative Nutrition” in 1996, which then creat-
ed its professional college (European College of Veterinary and
Comparative Nutrition) in 1997. The “European Society of
Veterinary Internal Medicine (ESVIM)” adopted the latter organisa-
tion as its member in 1999, paving the way for EU-recognised inter-
national veterinary specialist training in this special field.
Previously such veterinary specialist training was possible only in
the special fields of cardiology, oncology, nephrology and urology,
gastroenterology and hepatology, clinical pathology and dermatol-
ogy. (“Mission statement: ESVCN was founded to increase interac-
tion of European veterinary nutritionists, compare and harmonize
veterinary nutrition education from basic to specialist level in
Europe, advance applied and scientific veterinary nutrition”).
The current manual was preceded by a Hungarian-language
textbook published in 2003, edited by SÁNDOR FEKETE. Many chap-
ters of that book are included in the present updated and expand-
ed English-language version, written by a international collective of
authors of high reputation. With a view to the considerations
detailed above, this textbook and manual lays special emphasis on
topics in the processing of which the veterinary approach and atti-
tude can facilitate problem-solving. For those interested in more in-
depth details of classical nutritional and feed analytical knowledge
we recommend (among others) the works of M. BECKER and K.
NEHRING (1969): Handbuch der Futtermittel. Paul Parey, Hamburg-
Berlin; T. KAKUK and J. SCHMIDT (1988): Takarmányozástan.
Mezőgazdasági Kiadó. Budapest; H. JEROCH, G. FLACHOWSKY and F.
WEISSBACH (1993): Futtermittel-kunde. Gustav Fischer Verlag,
Jena–Stuttgart, R.O. KELLEMS and D.C. CHURCH (2002): Livestock
feeds and feeding. Fifth Ed. Upper Saddle River, NJ and D.A. TISCH
(2006): Animal feeds, feeding and nutrition, and ration evaluation
with CD–ROM. Thomson-Delmar Learning, Clifton Park, NJ.
3
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JAV_1_VND_FSGy1:MINTA J.qxd 2008.09.03. 14:26 Page 5
Chapter I
IMPORTANCE OF ANIMAL NUTRITION.
BASIC NOTIONS.
ROLE OF WATER IN ANIMAL ORGANISM
© Sandor Gy. Fekete
1.1. Definition of feed

1.2. Main features of feedstuffs (pyramid-conception )
1.2.l. Energy value.
1.2.2. Quantity and quality of proteins (exclusivity!)
1.2.3. Mineral and vitamin content.
1.2.4. Dietetic effect
1.2.5. Preservation
1.2.6. Transportability
1.2.7. Specific effects:
1.2.8. Taste, flavour and smell.
1.2.9. Price (on site, on market)
1.2.10. Concentration, bulkiness
1.3. Importance of protein
1.4.Water, electrolyte and acid-base balance
1.4.1.Role of water in the animal organism
1.4.2. Water losses from the animal body
1.4.3. Water supply
1.4.4. Water requirement
5
CHAPTER I: BASIC NOTIONS - IMPORTAQNCE-ROLE OF WATER
nimal nutrition has a key role in the food supply of the humanity. The
A main source of the problem is that while the growth of food produc-
tion is only linear, that of the population is exponential. According to
the recent predictions a world population of 6.5 milliard may reach 8-9 mil-
liards until 2020. To aggravate the situation, the population increase is
uneven distributed geographically, being faster in the underdeveloped
regions.
Consequently, agriculture strives to use both the classical and the
biotechnological ways of development. An increase in crop production is not
enough to solve this problem. To support this statement, let us compare the
macro and micronutrients’ requirements of leguminous and cereal plants
with the mono gastric and ruminant animals. (Abbreviations: AA=amino
acid; NPN=non protein nitrogen; B=boron; Ca=calcium)
ALFALFA PIG
N need need for proteins
do not requires AA AA requirement
no vitamin needs need for vitamins
CORN RUMINANTS
higher N need to a production level)
(no N-fixation) no need for AA only for NPN
more B need less need for vitamins
less Ca requirement (ruminal synthesis)
In the light of competition for certain feedstuffs between man and ani-
mals, a re-evaluation of animal production has to be made on the basis of
that proportion of the diet which also be used by man. Enormous advantage
lies in the fact that material unfit for human consumption can be convert-
ed into high-quality animal protein although the conversion efficiency is
generally low. First the basic notions should be explained.
6
1.1.Definition of feed
Classically, feed has to meet the following two criteria:
1. it should contain nutrient for the animal,
2. it may be fed without health alteration.
A material in question can only be treated as feedstuff if it satisfies
the two criteria simultaneously. For example the saw dust or sand may be
fed without health damage, but they have no nutrient; the seed of castor oil-
plant (Ricinus communis) contains proteins and oil, but cannot be consid-
ered as feedstuff because it is highly toxic. The classification, of course, is
not rigid, because the corn stalk is a feedstuff for ruminant, but not for a
one-day chicken. Different processing may influence whether a certain
material can be used as feedstuff or not (e.g. extrusion of corn starch for
carnivores).
1.2. Main features of feedstuffs (pyramid-conception)

1.2.l. Energy value.
1.2.2. Quantity and quality of proteins (exclusivity!)
Monogastrics are able to synthesize protein exclusively from proteins
(amino acids). Protein synthesis requires energy.
1.2.3. Mineral and vitamin content.
1.2.4. Dietetic effect shows the effect of physico-chemical characteristics
of feedstuff on the taste sensation, function of the gastro-intestinal system,
the peristalsis, the consistency of digesta/faeces and the well-being of the
animal. Dietetic effect may be:
a) good, for example the meadow grass for ruminants, barley
grain for pig, and oats for horse;
b) laxative as the water melon for pig;
c) constipating like leguminous seeds (e. g. bean, pea) and the
tannic acid containing feeds (sorghum) for monogastrics and
d) causing bloat or flatulence, for example the wet alfalfa for
grazing cattle; leguminous seeds with Non Starch Polysaccharides (NSP)
content, like soybean, lentil etc.
Dietetic effect may be modified by the:
- hygienic state of the feed
- fermentation, moulding, mustiness
- total germ count.
1.2.5. Preservation e.g. the roots and tubers like pumpkin, carrot have
good dietetic effects, but they cannot be kept for a long period of time.
1.2.6. Transportability: e.g. dried sugar beet pulp is favourable than the
wet one having less water content; dry versus canned pet food etc.
1.2.7. Specific effects: those features that cannot be explained with the
presence of major chemical nutrients (e.g. feeding of coconut oil increases
milk fat, whereas beef tallow doesn’t. Some meadows, oats, rye stimulate
sexual functions having phytooestrogen content.
7
1.2.8. Taste, flavour and smell.

1.2.9. Price (on site, on market)
1.2.10. Concentration, bulkiness
CONCENTRATION: energy content in the dry matter.
BULKINESS: ratio of dry matter to volume (in natural form, see alfalfa
hay versus cube).
Examples: concentrated, no bulky: corn meal (cf. oats grain),
concentrated, bulky: potatoes, milk,
no concentrated, no bulky: does not exist in the practice
no concentrated, bulky: straw.
1.3. Importance of protein

In feeding humanity the foods of animal origin are not completely
Table I-1: Contribution of foods of animal origin to the supply of human nutrients’, miner-
al and vitamin requirements, %
*pig, cattle, poultry and fish **except butter
replaceable by the vegetable ones. On average, in the nutrition of the world

population 70% of the protein, 80% of the calcium and practically 100% of
the vitamin B12 derives from food of animal origin (Table I-1).
Conversion of feed protein: if the efficiency of production is expressed

in terms of human edible protein return as the percentage of total digestible
protein input, pork, poultry and milk production have efficiencies of 30-40%
and beef production of only 5%. However, the picture is entirely different if
human edible protein return is expressed as a percentage of input of
digestible protein edible also by man. The efficiencies are 170-180% for
dairy cattle, 110% for beef cattle, 80-90% for pork and 70-80% for poultry
production. To express the nutrient density, the Index of Nutritional Quality
(INQ) describes the quality of food for human nutrition with regard to its
components. It shows the ratio of protein (amino acid=AA), mineral or vita-
min to the energy content. The daily requirement is assumed as “1.0”.
INQ = proportion of protein (AA), mineral or vitamin in the food to the
daily need/ratio of the energy amount of the same food to the daily require-
ment or one serving, in percentage.
8
For example, let’s calculate the INQ of milk for calcium. The daily
energy requirement of an adult human involved in light physical work is 10
MJ ME and the daily minimum calcium allowance is 1.2 gram. The energy
value of 1 litre cow’s milk (2.8% fat) is 2.52 MJ and it contains 1.2 gram cal-
cium. Accordingly, by drinking 1 litre milk one can cover 25% of the daily
energy and 100% of the daily calcium needs. Thus
INQ of milk for calcium= 100%/25% = 4.0
It means that if the daily energy requirement were covered exclusively with
milk, one would get 4 times more calcium than the requirement. The greater
is this number, the more valuable is the food for the (micro)nutrient in ques-
tion. Knowledge of the INQ help to select the food in order to complement
each others. Table I-2 presents the INQ of the most important foods of ani-
mal origin.
Table I-2: Index of nutritional quality of the most important foods of animal origin
Based on these data one can conclude that – on population level –

relying exclusively only on foods of plant origin the complete nutrient, min-
eral and vitamin requirement of human cannot be assured. This is espe-
cially true for children, pregnant women and elderly people (FIGURE-I-1)
The situation is different in case of lacto-, or lacto-ovo-vegetarians, who con-
sumes milk or milk and egg also.
1.4. Water, electrolyte and acid-base balance
uge area of the Earth’s surface (363000 km2) is covered by water.
H Without water, there would be no life on Earth. Approx. 70% of the

mammalian body is water. Life probably began in water. Reactions
in living cells take place in solutions. Water, having a very high specific heat
capacity, can hold large amounts of heat without a substantial rise in tem-
perature. As a consequence, water does not only supply the matrix in which
all living processes occur, but it also participates in those processes. One of
the most important properties of water in nutrition is its remarkable ability
9
to dissolve substances. This property is due to its high dielectric constant,

which promotes ionization of electrolytes and allows oppositely charged ions
to move independently of each other. This facilitates cell reactions and
makes water a remarkable solvent. As a solvent, water performs the vital
functions of transporting nutrients and metabolites throughout the body to
satisfy the needs of cells and excrete waste products. Biochemical reactions
in the body do not only take place in solutions, but water actually takes part
in reactions.
1.4.1. Role of water in the animal organism

Some of the important reactions in digestion and intermediary metabolism
involve the chemical addition or release of water, and water catalyses
numerous other reactions. It is doubtful whether any chemical reaction in
the body proceeds in its absence. To be able to accomplish the control of
influx and efflux of materials, most cells are freely permeable to water and
water continuously enters and leaves cells diffusing readily along its con-
centration gradient throughout the tissues. Changes in osmotic pressure
cause immediate water movement and the selective permeability of mem-
branes to various ions operate to regulate cell water.
Water helps in maintaining the shape of body cells; it lubricates and
cushions joints and organs in the body cavity. Most of the waste compounds
are eliminated through the urine and faeces. The high specific heat capaci-
ty and high heat of vaporization of water are due to its hydrogen bonds. A
given amount of water requires more heat to raise its temperature and to
convert it from liquid to vapour than almost any other substance. These
properties make it possible for the large amounts of heat produced by
metabolism to be dissipated with very little change in body temperature.
1.4.2. Water losses from the animal body

Water losses occur in urine, from the skin, with expired gases and in the
faeces. Lactating animals also lose large amounts of water in milk. End-
products of protein catabolism (urea in mammals, uric acids in reptiles and
birds) are excreted by the kidneys. Thus, urine comprises many end-prod-
ucts of his protein catabolism (urea in mammals, uric acids in reptiles and
birds) and minerals, too. Concentrated water solution of urea is toxic for
cells and tissues; in the urine it is diluted to an innocuous concentration
and eliminated. Water requirement of birds related to protein intake is
smaller than in mammals, because the protein to uric acid pathway pro-
duces more metabolic water, than the protein to urea pathway does and the
dry matter content of the bird urine is lower.
Water losses in faeces, in comparison with other species are the high-
est in ruminants where it equals approximately the urinary water losses. (In
humans the faecal water losses are only 7-10% of the amount lost in urine
water.) Faecal water content of high fibre forage eating cattle may reach 60-
10
70%. The majority of faeces water in ruminants derives from the saliva and
digestive juices. Diarrhoea or use of purgatives may significantly increase
faecal water losses.
The water cycle in animal is presented through the example of an
adult horse. The daily water consumption is 14-20 litres. By means of sali-
va 40, gastric juice 20 and the intestinal juice 20 litres liquid enters the gas-
tro-intestinal tract. From the hind gut 80-90 litres are reabsorbed. By exha-
lation and perspiration 6-16, by faeces 4-5 and by urine 4-8 litres of water
leaves the body. It is clearly shown that although a huge amount of water
participates in the daily cycle, owing to the intestinal reabsorption
(enterosystemic cycle) the real water requirement is relatively small.
1.4.3. Water supply

Water is gained by ingestion or as an end product of cellular metabolism.
While much of water loss from the body is continuous, the ingestion of
water is periodical, due to habit or to a daily rhythm of eating and drinking.
(see Chapter XXXIII) Water deprivation causes both the sensation of thirst
and an associated behavioural drive to drink water. Thirst is characterized
by a dryness of the throat and mouth due to a decrease in salivary secre-
tion. Besides, signals arise from the alimentary tract which informs the cen-
tral nervous system of the approximate amount of water being ingested. The
neuronal system controlling thirst and drinking behaviour is located in the
hypothalamic region of the brain.
It is well known that animals are less resistant against water defi-
ciency than against feed deprivation. First sign of a slight water restriction
is a reduced feed intake and productivity. Severe water deprivation leads to
weight loss and dehydration. Dehydration also means an increased excre-
tion of nitrogen and electrolytes such as sodium and potassium. Ten % loss
of the total body water is already critical and a loss of 20% may cause death.
In contrast, adult animals in feed deprivation can survive 40% decrease in
live weight.

Drinking water consumption is affected by many factors. Environmental
temperature and humidity is largely involved, at heat stress increase water
consumption occurs. At moderate temperature water consumption is direct-
ly correlated with the dry matter intake and with the faecal water losses.
Thus, eating fibrous roughage increases the water requirement.
Water content in or on feed is highly variable; grains may range from
less than 8% to over 30% water. Forages may range from less than 5% in
dry hay to over 90% water in a lush young grass. Precipitation or dew on
feed may decrease drinking water consumption. The metabolic water, which
results from the oxidation of organic nutrients in the tissues may account
for 5-10% of the total water intake.
11
CHAPTER I: BASIC NOTIONS- IMPORTAQNCE-ROLE OF WATER
High water content of the feed reduces the need for drinking, but high
salt or protein concentration of diet increases it. The use of animals may
modify water intake, because milk production needs a lot of water. The type
of urinary system and the excretory capacity are also important, and great
differences are found when comparing mammalian and avian urinary sys-
tems. Water quality should also considered: good water should have less
than 2500 mg/litre of dissolved solids; water containing over 1 g/litre sul-
phates may cause diarrhoea and levels of 100-200 mg/litre nitrates are
potentially toxic.
Owing to the variables mentioned above, the water requirements are
not fixed. However, in normal conditions, water requirements can be gener-
ally related to feed intake. Some approximations of the needs for water in
livestock vary from twice the amount of dry feed for rabbit, 2.5 times for
pigs, 3-4 to 1 for lactating sows, 5.5-6.5 to 1 for calves and increasing with
temperature from 3.5-5.5 to 1 for cattle. Sheep and poultry need less except
for egg laying.
With increasing temperature the respiratory (vaporization from the
lungs) and insensible (dissipation through the skin) losses increase.
Animals adapt by decreasing dry matter intake and increasing water con-
sumption.
FOR FURTHER READING
Garrow, J.S., James, W.P.T. and Ralph, A. (1993). Human nutrition an dietetics. Churchill
Livingstone. Edinburgh, London, Madrid, Melbourne, New York, Tokyo
Janick, J., Noller, C.H. and Rhykerd, C.L. (1976): The cycles of plant and animal nutrition.
Sci. American 235(3), 75-86.
Lofgreen, M. and Speckmann, P.A. (1979): Importance of Animal products in the human
diet. Dairy Sci. 63, 1019-1025.
National Academy Press (1988): Recommended Dietary Allowances. (10th ed.)
Whashington, D.C.
Shils, M.E. and Young, V.R. (1988): Modern nutrition in health and disease. Lea & Feer.
Philadelphia.
Wildman, R.E.C. and Medeiros, D.M. (2000): Advanced human nutrition. CRC Press.Boca
Raton, London, New York, Washington D.C.
12
FIGURE I-1: The feed-food pyramid (© FEKETE, S.GY.)
13
JAV_2_VND_FSGy:MINTA J.qxd 2008.09.03. 14:27 Page 15
Chapter II
NUTRITION, ANIMAL HEALTH AND FOOD SAFETY

© Josef Leibetseder
2.1. Deficiency
2.1.1.Energy eficiency
2.1.2. Nutrient deficiency
2.2. Unbalanced diet
2.3. Undesirable substances
2.4. Contaminants
2.4.1. Ergotism
2.4.2. Mycotoxins
2.4.3. Prions
2.4.4. Elements
2.4.5. Agrotechnical substances
2.4.6. Environmental pollutants
15
CHAPTER II: ANIMAL HEALTH FOOD SAFETY
dequate nutrition is a necessary prerequisite for health. Adequate
A nutrition means not only the intake of a well balanced diet in suffi-
cient amounts, but also the intake of a diet containing no undesirable
substances or concentrations which are not considered to be harmful. First
of all, inadequate feeding causes health problems in the animals, in case of
food producing animals it might also have negative impacts on food safety
and/or food quality. Because feeding is the most expensive factor in animal
husbandry and inadequate nutrition leads to economic losses: the interest
in proper nutrition of the livestock was always strong. It is, therefore, not
surprising that the scientific work in animal nutrition started about 200
years ago and that the farmers were prepared to introduce the results into
practice. Innumerable studies were performed to determine energy and
nutrient requirements, the digestibility and availability of nutrients, to clear
up the metabolism of substances and to evaluate the nutritive value of
thousands of feeding stuffs. The results of these scientific studies are rec-
ommendations on energy and nutrient supply of animals at different phys-
iological states like maintenance and all kind of performance and extensive
tables on the energy and nutrient content of feedingstuffs. Numerous unde-
sirable substances in animals have also been intensively studied and per-
missible levels in feedingstuffs were established in order to avoid risks for
animal health or non acceptable residues in food of animal origin. At pres-
ent the consequences of inadequate nutrition and the intake of undesirable
substances are well known and can be correctly diagnosed in such cases.
Recently the impact of animal feeding on the environment has also to be
considered. Not only for economic reasons but also for animal welfare, the
influence on food safety and the protection of the environment, farmers are
obliged to adhere to the legal regulations on animal husbandry and nutri-
tion.
2.1. Deficiency
Deficiency can concern the insufficient intake either of energy or of specific
essential nutrients. Insufficient intake may result from low concentration of
energy or nutrients in the diet, from reduced intake of adequate feed or from
the combination of both.
16
2.1.1. Energy deficiency leads in all species to general signs like reduction
of body weight and impaired performance (growth, production of meat, milk
or eggs), in severe cases to cachexia and negative effects on other systems,
i.e. reproductive and immune system.
In ruminants energy deficiency causes metabolic disturbance called
KETOSIS. In high yielding cows the feed intake is not sufficient to meet the
energy requirements in the transient (fresh) period, especially if these cows
were overfed in the dry period and contain therefore higher amounts of body
fat. During the deficient period fat is mobilised by lipolysis and the fatty
acids produce acetyl-CoA by ?-oxidation. Because of the reduced feed intake
precursors of glucose (propionic acid, glucogenetic amino acids) are not
available in sufficient amounts to form enough glucose as source of pyru-
vate. Pyruvate reacts with CO2-forming oxalacetic acid, the starting com-
pound of the Krebs (TCA) cycle which binds the acetyl-CoA. If oxalacetate is
not present in sufficient amounts because of lack of pyruvate, the acetyl-
CoA reacts with itself forming keton bodies (acetoacetic acid, ?-hydroxybu-
tyric acid and acetone). The consequences of ketosis in dairy cows are a
strong decrease of milk production and in severe cases liver damage. Oral
sodium propionate and glucose infusion is the adequate treatment of sick
animals. Preventive measures are to assure the correct body condition at
the end of pregnancy by appropriate energy supply (requirement for main-
tenance plus only energy for pregnancy corresponding to the requirement
for about 5 kg milk production/day), intensive feeding post partum and
supplementation of propionate or propylene glycol.
Ketosis may also occur in young overfed heifers if the feed intake is
reduced for several reasons, i.e. transportation for some days if heifers are
sold for breeding purposes and are not fed properly. Pregnancy toxicosis in
ewes is caused by ketosis seen especially in ewes with twins or triplets (for
details see Chapter XXVI).
2.1.2. Nutrient deficiency

About 50 substances (essential amino acids, essential fatty acids, vitamins,
minerals and trace elements) are recognized as essential. The deficiency of
an essential nutrient leads to general deficiency symptoms and/or to more
or less specific signs.
A lack of essential amino acids reduces the protein synthesis.
Therefore, growth in young animals is decreased, adult animal lose body
weight. In addition, the formation of antibodies may be reduced and the
immune response may be impaired. Animals deficient in essential fatty
acids don’t grow and show skin and kidney lesions. There is also a change
in the synthesis of specific prostaglandins and leukotrienes.
The most frequently seen mineral deficiencies are calcium and mag-
nesium deficiency in dairy cows. Ca deficiency (HYPOCALCAEMIA, MILK FEVER)
is a regulatory problem post partum in high yielding cows. The excretion of
high amounts of calcium can neither be compensated by the absorption of
adequate amounts of calcium from the intestine nor by the sufficient mobil-
17
isation of calcium from the skeleton (bones). Those cows show a severe
paresis and are not able to stand up. Generally overfed cows and cows with
high Ca intake before parturition are particularly at risk, as are cows with
insufficient Mg supply. Intravenous Ca administration, sometimes in com-
bination with Mg is effective. Nutritional prevention can be done by reduc-
ing the Ca intake before birth, avoiding overfeeding and appropriate Mg
supply. Application of vitamin D (especially 25-dihydro-cholecalciferol) and
highly available Ca compounds (calcium chloride gel) is also indicated.
In the Northern part of Europe, cows may suffer from cramps when
they are put out to pasture because of hypomagnesaemia (“GRASS TETANY”).
The aetiology of hypomagnesaemia can be the low content of Mg in young,
fast growing grass (low Mg concentration of the soil, intensive fertilisation
with K and N) and low Mg absorption in the rumen because of high rumen
pH (high ammonia concentration owing to protein intake), high K and/or
insufficient Na intake. Preventive measures are Mg fertilisation, careful
transient feeding from stable to pasture (addition of roughage) and Mg sup-
plementation. In lactating bitch of large litter size the high Ca and Mg loss-
es through the milk may cause puerperal eclampsia in weeks 2-4 after
whelping.
Nowadays deficiencies of minerals and trace elements are rare
because of supplementation of all trace elements and vitamins in sufficient
amounts. In former times when requirements were not well known and sup-
plementation was not usual, specific signs of deficiencies were frequently
observed in all species. Iron deficiency (ANAEMIA) in suckling animals (espe-
cially in piglets) was common because of the low iron concentration in the
milk when animals did not have access to soil or the dam’s faeces, which
are good source of iron. Also iodine deficiency was often seen especially in
piglets, if the pregnant sow’s supply was insufficient. In some deficient
areas a lack of Se was observed in different species (heifer, lamb, pig).
The situation of vitamins is similar. Under normal conditions defi-
ciencies can hardly be seen in practice. Only in ruminants, mainly in inten-
sively concentrate fed fattening bulls; vitamin B1 might become deficient
due to increased thiaminase activity in the rumen because of disturbed fer-
mentation or thiaminase formation by fungi. Another vitamin deficiency in
ruminants is caused by inadequate intake of beta-carotene. In cattle beta
carotene is not only a provitamin (precursor of vitamin A) but also an essen-
tial vitamin by itself. Since corn silage widely used in dairy cows is low in
beta-carotene it might become deficient without supplementation. The main
symptom of deficiency is INFERTILITY.
2.2. Unbalanced diet

A well-balanced diet is the prerequisite of normal digestion. Especially her-
bivores developed a complicated gastrointestinal tract and are therefore
more sensitive to unbalanced diets. The intake of higher amounts of easily
fermented carbohydrates in the rumen (sugar, starch) leads to the forma-
tion of volatile fatty acids (mostly propionic acid) and lactic acid in greater
18
amounts with a decrease of rumen pH (below 6.0). RUMEN ACIDOSIS causes

reduction of feed intake, lesions of the rumen mucosa, pododermatitis in
acute cases, in chronic cases parakeratosis of the rumen mucosa and
necrotic lesions in liver and kidneys. Dietetic measures are balancing the
diet (reduction of sugar and starch, higher amounts of roughage), adminis-
tration of sodium bicarbonate alone or in combination with magnesium
oxide. In acute cases the rumen content should be removed.
Diets high in protein and low in carbohydrates can cause rumen alkalosis
(pH above 7.0). Consequences are again reduction of feed intake, liver prob-
lems and in severe cases ammonia intoxication.
The intake of higher amounts of lactose may cause problems in poultry and
adult carnivores because of lack of lactase in the intestinal tract. Lactose is,
therefore, intensively fermented by the flora in the large intestine which
leads to OSMOTIC DIARRHOEA.
2.3. Undesirable substances

Undesirable substance shall mean any substance or product, with the
exception of pathogenic agents, which is present in and/or on the product
intended for animal feed and which presents a potential danger to animal
or human health or to the environment or could adversely affect livestock
production. Products intended for animal feed may be used only if they are
sound, genuine and of merchantable quality and therefore when correctly
used do not represent any danger mentioned above. Feedstuffs therefore
shall not contain undesirable substances or only in such amounts which
are not dangerous. Investigations shall be carried out to identify the sources
of undesirable substances. Maximum contents of undesirable substances of
feedingstuffs shall be set by legal regulations. In cases of increased levels it
may be necessary to set action thresholds.
There are different categories of undesirable substances in feedstuffs.
Feedstuffs as such may contain naturally undesirable substances (i.e. poi-
sonous plants, trypsin inhibitor in soybeans, cyanogenic glycosides,
saponins, coumarin etc.), or feedingstuffs may contain contaminants (alive
or unanimated).
Naturally occurring undesirable substances in plants. Recently the
incidence of intoxication by poisonous plants has increased for several rea-
sons. Mainly cattle and horses are endangered grazing on pastures with poi-
sonous plants or by intake of poisonous ornamental plants in parks. In con-
trast, the quality of traded commodities used as ingredients in feed and, as
a consequence, the quality of feedingstuffs has substantially improved over
the last decades. Nonetheless, feed ingredients are inevitably contaminated
with a variety of materials of botanical origin. Depending on the feed or feed
ingredient, these may derive from weeds, growing within the crop or at field
margins at the time of harvest or from adventitious contamination during
storage and/or transport. The vast majority of such contamination is
innocuous and only rarely is noxious or potentially toxic. Some plants or
plant products contain besides valuable nutrients different categories of
19
undesirable substances (Table II-1).
Table II- 1: Examples of naturally occurring undesirable substances in

feedingstuffs (KAMPHUES et al. 1999, modified)
20
Table II-1: continued
2.4. Contaminants
Contaminants can occur naturally (bacteria, fungi, viruses, prions, para-
sites) or anthropogenically (toxic elements, pesticides, dioxin etc.). The most
harmful bacteria in food of animal origin are Salmonella spp., Listeria,
Campylobacter and Clostridium botulinum causing infectious diseases or
intoxication by specific toxins of the bacteria (botulismus toxin).
2.4.1. Ergotism
Fungi can inhibit the development of plants and reduce the nutritive value.
Some of the fungi produce secondary metabolites
(mycotoxins) which are highly toxic for men and
animals. For a long time ergotism has been well
known in animals as well as in humans caused
by the ergot alkaloids of the fungus Claviceps pur-
purea which changes the normal kernels of rye,
triticale, wheat and the seed of different grasses
into the ergot (Secale cornutum). Sings of ergotism
are disorders of circulation, convulsions and
damage to the central nervous system. Nowdays
ERGOT INTOXICATION occurs more often, again
because in some cases the size of ergot is not as
big as it used to be earlier and can not be sepa-
rated completely by the milling process.
FIGURE II-1: Ergot (Secale cornutum)
21
2.4.2. Mycotoxins
Other mycotoxins has become important since the outbreak of Turkey X
disease in England in the early sixties of the last century. In the meantime
it has been established that mycotoxins are responsible for a variety of ani-
mal and human diseases and even death. Although mycotoxins have caused
some dramatic epidemics in humans and animals, such outbreaks are very
rare. Mycotoxicosis is essentially a chronic problem caused by an underly-
ing contamination of crops, particularly cereals, with toxigenic fungi.
Fungal toxins are estimated to affect as much as 25% of the world’s crops
each year. However, the variable production of mycotoxins together with ill-
defined symptoms makes it difficult to estimate the real incidence of myco-
toxicosis. The biological effects are numerous. They can be acutely and/or
chronically toxic, depending on their chemical structure and concentration,
the extent of exposure of animal consuming contaminated feed and the
health status. In animals, targets for acute effects include liver, bone mar-
row, kidney, central nervous system, skin, reproductive and immune sys-
tem. Some mycotoxins are carcinogenic.
Mycotoxin contamination of forages and cereals frequently occurs in
the field following infection of plants with particular pathogenic fungi or
with symbiotic endophytes. Production of mycotoxins can also occur during
processing and storage of harvested feed materials when environmental
conditions such as moisture and ambient temperature appropriate for
development of spoilage fungi are met. It is conventional to subdivides tox-
igenic fungi into “FIELD” OR PLANT PATHOGENIC AND “STORAGE” or saprophyt-
ic/spoilage organisms. Fusarium spp. are representatives of field fungi,
while strains of Asperillus spp. producing aflatoxins and Penicillium spp. are
common storage fungi. Mycotoxigenic species may be further distinguished
on the basis of geographical prevalence due to the specific environmental
requirements for growth and secondary metabolism: Aspergillus spp. prolif-
erate under warm, humid conditions, while Penicillium spp. develop under
temperate climate. Consequently aflatoxins predominate in plant products
emanating from the tropics and other warm regions. Interactions of several
factors operating simultaneously are usually more important than any sin-
gle factor controlling mycotoxin production. Visible fungal growth on the
grains does not necessarily mean that they are contaminated with myco-
toxins, and vice versa. Although fungal growth may not be evident on the
kernels, for example due to drying or to use of fungicides, high concentra-
tions of mycotoxins may still be found. It is important to recognise that two
or more mycotoxins can be produced by the same species of fungus and
that some mycotoxins are produced by more than one fungal species.
Analysis of a single commodity often shows the presence of several myco-
toxins.
A large number of fungal secondary metabolites have been identified,
many of which have been shown to be toxic for animals and humans. Novel
metabolites are constantly being identified and therefore this field needs to
be regularly reviewed. Among these aflatoxins (B1, B2, G1 and G2) are the
22
most important toxins because they are potent carcinogens. The metabolite
of aflatoxin B1 appears in milk and milk products as a direct result of intake
of contaminated feed. Ochratoxins (A and B) are formed by Penicillium and
Aspergillus strains, predominately by Penicillium verucosum and Aspergillus
ochraceaus and Aspergillus niger. Ochratoxins are partially absorbed from
the gastrointestinal tract in monogastrics. Consequently ochratoxins have
been found in edible tissues and products of monogastric animals. In rumi-
nants, ochratoxin A is mainly metabolised by the rumen microbiota to
ochratoxin B before absorption. This major metabolite appears less toxic
than ochratoxin A. Field cases of ochratoxicosis in farm animals (pigs, poul-
try) have been reported, the primary manifestation being chronic nephropa-
thy. Ochratoxin A is considered genotoxic, nephrotoxic and teratogenic and
may cause immunotoxic effects. There is currently inadequate evidence in
humans for the carcinogenicity.
Fusarium species produce a variety of mycotoxins. Of particular inter-
est are zearalenone, the trichothecenes, the fumonisins and moniliformin.
Zearalenone is a (pseudo)estrogenic compound. It induces estrogenic effects
in mammals, including early maturity of mammary glands and reproductive
organs and increases their size. At higher doses it interferes with concep-
tion, ovulation, implantation, foetal development and viability of newborn
animals. Pigs appear to be the most sensitive species. Due to its adverse
effects on mammals, zearalenone is one of the most important mycotoxins
from the point of view of animal health. Amongst the trichothecenes,
deoxynivalenol (DON, vomitoxin) has been shown to have the greatest
adverse impact on animals. Pigs are the most sensitive species. Chronic
exposure causes decreased body weight gain, feed refusal, liver damage,
decreases humoral and cell-mediated immunity and reduces host resist-
ance. Recent findings indicate some genotoxic effects in human cell lines.
Poultry and to a greater extent ruminants are more resistant, whereas fish
have been found susceptible. T–2 toxin was one of the first trichothecenes
to be identified and is known to be amongst the most potent mycotoxins. It
has been associated with the major outbreak of Alimentary Toxic Aleukia in
humans in Russia in 1944. In animals it has been reported to have extreme-
ly toxic effects on skin and mucous surfaces. One of the significant effects
of T–2 toxin is its immunosuppressive activity (lymphocyte depletion in
spleen, liver, bone marrow). Non-ruminants seem to be more sensitive than
ruminants, coprophag and caecotroph animals (rabbit, guinea pig) are the
most sensitive.
At least 12 fumonisin analogues are known, the most important being
the B series (B1, B2 and B3). The most significant crop, in which fumon-
isins occur, is corn, particularly that grown in warmer regions. However,
sorghum and rice are also occasionally affected. In animals fumonisins (par-
ticularly B1) are known to cause a wide range of different illnesses, such as
equine leuko-encephalomalacia and porcine pulmonary oedema. Fumonisin
B1 has been shown to be carcinogenic in rodents causing both liver and kid-
ney tumours. Moniliformin has been detected predominantly in corn, but
23
also in other cereals. Moniliformin is toxic to animals with effects that

include haemorrhages in the gastrointestinal tract and damage to liver and
heart. Reduced body weight gain was also recorded.
Recently two fungal metabolites have been shown to cause toxic
effects in animals. Mycophenolic acid and cyclopiazonic acid are produced
by species of different fungal genera. Mycophenolic acid blocks the biosyn-
thesis of purine and therefore the proliferative response of T and B-lym-
phocytes and inhibits both antibody formation and the production of cyto-
toxic T cells. The toxic effects of cyclopiazonic acids in poultry, pigs and
sheep are well documented. They include body weight loss and diarrhoea,
alimentary tract hyperaemia, haemorrhage and focal ulceration. It also has
the ability to chelate metal ions and this may be an important mechanism
of its toxicity.
2.4.3. Prions
The transmissible spongiform encephalopathy in sheep (scrapie) was known
for more than 250 years. The incidence of scrapie was low and, therefore,
the economic loss unimportant. In 1986 the first case of bovine spongiform
encephalopathy (BSE) was observed in England (WILESMITH et al. 1988) and
this was the beginning of a disastrous outbreak of the disease with tremen-
dous economic losses. A certain protein (proteinaceous infectious particles,
prions) was identified as the pathogen agent which can not be inactivated
by heat and UV or radioactive irradiation, only a 20-min long, 133 oC heat
treatment (maximum particle size 50 mm) under 3 bar pressure may inac-
tivate it. In order to avoid the transmission of the disease the use of meat
and bone meal as feeding material is prohibited in the EU member states,
which cause difficulties in the protein supply of meat producing animals.
2.4.4. Elements. Probably all elements mentioned below are essential, but
the requirements are very low (ultra-trace elements) and deficiencies have
never been observed in practice because these elements are ubiquitous and
present to a sufficient extent in feeding materials. Like all trace elements,
these elements are also tolerated only up to a certain limit by animals and
humans. Above that limit, their toxic potential leads to detrimental effects.
In most cases the toxicity depends to a great extent upon the chemical form
(for instance organic or inorganic). For the evaluation of the toxicity this
aspect needs particular considerations. For that reason the toxic aspect of
these elements is more important than meeting the requirements. Being
either from geologic origin or anthropogenic (air, soil contamination), ele-
ments have an uneven distribution. As a consequence, their occurrence in
feed materials is variable and may exceed tolerable levels. For instance, con-
comitant presence of noxious elements (F, As) in phosphates obtained from
mining is well known. These elements naturally present or brought by
anthropogenic contamination are considered as undesirable not only
because of toxic effects in animals, but also because of a possible increase
of human exposure due to residues in food of animal origin.
24
LEAD is widely used for technical purposes in both organic and inor-
ganic forms. This has led to its widespread distribution in the biosphere.
Consecutive to the ban of the use of organic lead in petrol the contamina-
tion of the environment is decreasing. Sources of lead contamination are
use of contaminated sludge and wastes as fertilizers and industrial emis-
sions. Higher body burden of lead in domestic animals is mostly caused by
airborne deposition of lead on the surface of plants and on soil. Lead is also
often present in mineral feed material, like phosphates, and can contribute
significantly to the diet contamination. Lead is a chronic and cumulative
poison. It affects enzymes, provokes anaemia, renal toxicity, carcinogenici-
ty, and has cardiovascular and neurological/behavioural impact and nega-
tive consequences on the reproductive system.
MERCURY in the natural environment is found in both inorganic and
organic forms. The inorganic forms are less toxic. The most toxic compound
is methyl-mercury. Mercury is widely used for industrial purposes and is
released from industrial sites into the environment, especially into rivers
and sees. In former times mercury compounds were used as fungicides in
seeds causing some cases of intoxication in animals. Because of the low
concentration of mercury in pastures and crops the mercury content of edi-
ble tissues and products from farm animals is rather low. High concentra-
tions can be found in long-living marine predators. Methyl-mercury in fish
and fish products represents up to 85 % of the mercury in total intake from
the diet in humans. Clinical symptoms in animals fed higher levels are inap-
petence, anorexia, ataxia, abnormal behaviour, fatty liver, enlargement of
lymph nodes, necrosis of the buccal cavity and colon, nephrosis and
reduced fertility. Mercury is known to be teratogenic and carcinogenic in
mammals.
CADMIUM. Industrially produced contamination and use of fertilisers
has increased the background levels of cadmium in soil, water, and organ-
isms. In contrast to other elements, cadmium is rather mobile and can be
absorbed by plants via roots. Cadmium impurities are often present also in
mineral feed additives. Experiments with cadmium in different animal
species showed impaired growth, anaemia, hypertension, impaired renal,
reproductive and haematopoietic functions, depressed immune response.
Additionally, congenital defects and abortion were observed in cattle and
sheep. In humans, toxic effects have been observed after long-term expo-
sure in kidneys. Cadmium excess in food has been associated with a severe
bone disease. Cadmium has been shown to be carcinogenic in rat and ter-
atogenic in ruminants.
ARSENIC is present in all types of soil. Apart from the geological origin,
it also comes from emissions from coal fired power plants, smelters, use in
wood preservation and the now discontinued use of arsenical pesticides.
Some organic arsenic compounds have been used as growth promoters in
swine and poultry. Their use has been abandoned in Europe but they are
still in use in third countries such as USA. The toxicity is depending on
chemical form and valency. Trivalent arsenic and inorganic forms are much
25
more toxic than pentavalent arsenic compounds and organic arsenic,

respectively. It is considered that the available evidence on inorganic arsenic
indicated that arsenic is genotoxic.
Some other elements like FLUORINE, CHROMIUM, and ALUMINIUM are con-
sidered to be of minor importance in regard to intoxication of animals and
humans, but collection of data on the presence of these elements in feed
and food should still be continued.
2.4.5. Agrotechnical substances. In the conventional kind of production in

agriculture different categories of substances are used as pesticides, feed
additives and drugs. In very rare cases these substances are harmful to the
animals because of overdosing. The more important aspect is the formation
of residues in edible tissues of animals, but the disturbed reproduction is
also worth mentioning (e.g. 2,4-dichlor-phenoxy acetic acid). The use of
these substances is strongly regulated by law, but nevertheless, the regular
monitoring of residues is necessary to avoid misuse, or to detect failure in
the application.
2.4.6. Environmental pollutants

Dioxins and dioxin like compounds are created by manufacture of chlorine
and such chlorinated compounds as chlorinated phenols (PCBs), phenoxy
herbicides, chlorinated benzenes, chlorinated aliphatic compounds, chlori-
nated catalysts and halogenated biphenyl ethers. The most toxic compound
is 2,3,7,8-tetrachlorodibenzo-p-dioxin (TCDD), therefore the toxicity of other
dioxins, and chemicals like PCBs that act like dioxin, is measured in rela-
tion to TCDD (TEQ=average toxic equivalent). Dioxins are widely distributed
in the environment, and have the tendency to accumulate in the fat tissue,
egg and the milk. Dioxins are highly toxic, and besides cancer, even very
small amount can cause damage of liver and nervous system. They may also
cause abortion, development and birth defects. The greatest exposure to
dioxin is from the food.
CONCLUSION
Although different kinds of undesirable substances exist their risk for ani-
mal and human health is low because good knowledge about occurrence
and effects of these substances is available. For most substances the risk
can be assessed and therefore the maximum tolerable amount (MTA) in feed
material as well as in complete diets can be regulated. In consequence feed
and food in the EU show a high standard of safety and quality.
26
FOR FURTHER READING
Ferrando, R. (1979): Pollution, alimentation et santé publique. Cah. Méd.Vét.

pp 48; 35-45. and 86-89.
Kamphues, J., Schneider, D. and Leibetseder, J. (1999): Supplemente zu

Vorlesungen und Übungen in der Tierernährung. 9th edition, M.&H. Schaper,
Alfeld-Hannover, pp 110-111.
Lemming, E. (1992): The report of the expert group on animal feedingstuffs.

HMSO. London.
Spongiform Encephalopathy Advisory Committee: Transmissible spongiform
encephalopathies. A summary of present knowledge and research. HMSO. London
Wilesmith, J. W., Wells, G.A.H., Cranwell, M.P. and Ryan, J.B.M. (1988):
Bovine spongiform encephalopathy: Epidemiological studies. Vet. Rec. 123, 638-
644.
Wilgus, H.S. (1980): Disease, Nutrition-Interaction. Poult. Sci. 59, 772-781.
27
Chapter III
BODY COMPOSITION OF PLANTS, ANIMALS AND
METHODS FOR MEASUREMENT
©Sandor Gy. Fekete
3.1. Comparison of the body composition of plants

and animals
3.2. Factors influencing the body composition
3.3. Details about the methods
29
CHAPTER III: BODY COMPOSITION
lthough studies in connection with total body composition have more,
A than a hundred years past (VON BEZOLD, 1857), in the last few years
– as a consequence of the new genetic, endocrinological and biotech-
nological results – they got again into the centre of interest. Because of the
interventions the body composition of the animals is changing (or might
change), so it is indispensable to study these modifications. The clinical
importance of the body composition is also constantly growing (obesity,
osteoporosis, and anaesthesia). The true feed conversion can be calculated
from the amount of the ingested feed and the total body composition (data
of retention), which makes the real estimation of the nutrient requirement
possible.
3.1. Comparison of the body composition of plants and animals

One of the main differences between the body composition of plants and
animals is that the supporting tissue of plants is made of fibre, while the
animal body - except the digesta - does not contain any fibre. Plants consist
of considerable amount of carbohydrates (starch and sugar); the carbohy-
drate content of the animal body is only 0.5-1.5%. With the exception of the
oil seeds, plants contain less protein and fat, whereas the main part of the
dry matter content of the animal body is protein. There is more potassium
(K) in the plants, while in the animals the sodium (Na) has a major share.
3.2. Factors influencing the body composition

The body composition is influenced by age, sex, physiological status (preg-
nancy, lactation), genotype (e.g. lean swine hybrids), and nutrition. The
water- and protein content of the body is decreasing with age, whilst the
proportion of fat is increasing. On the basis of this MOULTON (1923) intro-
duced the concept of the CHEMICAL MATURITY, which refers to the stability of
the ratio of fat free dry matter to water as protein and ash content within
the fat-free dry matter. The influence of the sex is manifested by the higher
fat content of the female body.
The composition can be expressed in different ways; the base of com-
parison is generally the original dry matter content and the fat-free dry mat-
30
ter content. The average adult mammalian and avian body contain an aver-
age of 55-60% water, 15-20% protein, 18-25% fat and 3-4.5% ash (minerals).
The fat free body contains 72-75% water, 20-23% protein and 4-5% ash.
After reaching the so-called chemical maturity, the ratio of the main com-
ponents of the fat free body (protein, ash, water) is quite constant. Animals
reach chemical maturity at 3-4% of total life span.
The water content of the different tissues is characteristically diverse.
The water content of the blood and body fluids is above 95%, the muscle
has 70-80%, the bone 40%, the dental enamel 5% and the fat cells
(adipocytes) practically 0% water content. 70-80% of the protein can be
found in the muscle tissue.
The fat can be found around the kidneys, in the abdominal fat tissue
(birds), in the epididymal fat pad (rat) in the liver and muscles. (From chem-
ical and bio-energetic point of view, the marbled meat and the abdominal fat
“cost” the same feed energy, but it is not indifferent regarding the meat
quality.) The ratio of ash compared to protein is rather stable. The water is
also closely related to fat, but in a reverse proportion, the fat always “fills”
the place of water. Starvation increases body water and decreases body pro-
tein and fat, but except the fat, changes are trifle. It is also known that main-
tenance and sub-maintenance feeding compared to the ad libitum, result in
an increase in muscle water content at growing animals.
Based on the total body water, the fat – and with less accuracy – the
protein content can be calculated. This is favourable because only certain
measurements must be performed, and then the other parameters can be
calculated by regression.
The possibilities of the methods of body composition measurements are
demonstrated in Table III-1.
31
Table III-1: Methods for measurement/assessment of the body composition
3.3. Details about the methods

WEIGHING. Its disadvantage is that its value may not be changing while the
composition does (e.g.: water incorporation instead of fat). The result is
highly influenced by the quantity of the stomach and intestinal content,
especially in case of ruminants (For them the mean value of 3-day-weighing
must be considered).
BODY CONDITION SCORING (its unit is the BCS=body condition score).
Dairy cattle: five-point scale Scottish system, scoring by the amount of fat
covering the rump and tailhead area. Beef cattle: nine-point scale Nebraska
system, scores determined by the palpation of the ribs. The condition scor-
ing of the horses is similar. In case of sheep, scoring is based on feeling the
layer of muscles and fat deposition over and around the backbone vertebrae
in the loin region. (FIGURE III-1)
32
FIGURE III-1: BCS system in sheep
Dog and cat: scoring based on the thickness of the subcutaneous

connective tissue of the ribs and the base of the tail, as well as the shape
of waist and abdomen.
PROTEIN AND ENERGY BALANCE. Based on these values one can tell
whether it is positive or negative, but this does not give any information
about the body composition, only about its changes.
DIRECT CHEMICAL ANALYSIS. This is the most accurate method, but it is
labour intensive and expensive. Its other disadvantage is that it cannot be
applied in living animals.
DENSITOMETRY. Based on Archimedes’ Law. (The apparent weight loss
of a body immersed in a fluid is equal to the weight of the displaced fluid.)
The density=specific gravity of the fat is less than 1, that of the protein is
higher than 1. The density of the body can be calculated by weighing the
carcasses in the air and when submerged below water. The density is in
strong correlation with the fat content of the body. This method is accurate
when applied on carcasses, since in case of living animals the air in the
lungs influences the result (in humans it is possible to use because people
can co-operate during the examination).
TOTAL BODY WATER DETERMINATION. Its principle is that if the quantity
33
of the ingoing and outgoing water is not changing during a period of time,
the total body water can be considered constant (in case of dairy animals
this one-pool model is not right, or when the environmental temperature is
very variable, and the water losses of the expiration and sweating increas-
es). The changes of the concentration can be calculated from the dilution of
the indicator, which can be extrapolated to minute zero (the moment of the
administration). With full knowledge of the concentration and the adminis-
tered quantity of the indicator, the volume (total body water) can be calcu-
lated (FIGURE III-2). This method works on condition that the administered
indicator must be spread evenly in the body water. Knowing the water vol-
ume, the other major chemical components can be assessed, based on
regression. For this method D2O (deuterium) or TOH (tritium) is used (the
later is easier to measure because of the radiation, but it is more dangerous
for that very reason), the other possibilities are ethyl alcohol and – until the
beginning of the urinary excretion – the urea.
FIGURE III-2: Example of urea disappearance curve in the nursing foal

(urea = concentration minus basal concentration)
DETERMINATION OF TOTAL BODY POTASSIUM. 0.3% of the total potassium

content of the body is radiating 40K isotope. If the 40K emission is detected,
the total potassium content can be calculated, and from this data we can
assess the other chemical components (mainly the protein), because their
ratio to the K is constant.
X-RAY BASED DENSITOGRAPHY OR COMPUTERISED TOMOGRAPHY (CT).
Computed tomography is based on the X-ray principal: as X-rays pass
through the body they are absorbed or attenuated at differing levels by the
tissues creating a matrix or profile of X-ray beams of different strength. This
X-ray profile of pixels is registered on a film or detector, creating an image.
34
CT produces cross-sectional scans of the body. An X-ray tube sends a beam

of photons toward a detector. As the beam rotates around the animal, data
is collected, stored, and applied to complex algorithms to build images that
calculate body components. The X-ray absorption capacity of the body tis-
sues is different, it is expressed by the Houndsfield unit (HU) between -1024
and +1024 (FIGURE III-3).
FIGURE III-3: Typical CT-graph (HOUNSFIELD, 1980)
Each pixel is being assigned for a value on a scale on which air is -

1000, water and electrolytes are 0 and compact bone is between +600 and
+1000, muscle is between +30 and +200, lungs filled with air are between -
200 and -400, fat is between -20 and -200 (to compare: value for lead is
+5000 HU). The scale is chosen according to the X-ray attenuation of the
tissues to be examined. CT images axial layers, its picture elements, the pix-
els indicate the density of the body “slices”, based on this - via regression
correlations – it is possible to assess the body composition (FIGURE III-4).
35
FIGURE III-4: Frequencies of CT values from a fat (- - - ) and a lean (____) goat of approxi
mately the same live weight. The peak around -100 HU represents adipose tissue and the
peak around 50 HU represents muscle tissue (after SØRENSEN, 1992)
DUAL ENERGY X-RAY ABSORPTIMETRY, OR D(E)XA. DEXA uses a whole

body scanner that has two different X-rays sources that read bone and soft
tissue mass simultaneously. It is a relatively new technology that is very
accurate and precise; DEXA is based on a three-compartment model that
divides the body into total body mineral, fat-free soft (lean) mass and fat tis-
sue. This method is used in humans to study osteoporosis, and in control-
ling the level of obesity in the small animal praxis.
ADIPOCYTE MORPHOMETRY. The adipocytes taken from the predilection
depot (e.g. epidydimal fat pad of rats) are stained by osmium-tetraoxide. Cell
size (diameter, surface) is in correlation with the total fat content of the
body.
VLDL MEASUREMENT OF THE BLOOD PLASMA. The VLDL (very low-densi-
ty lipoprotein) level is higher if the body contains more fat. This level is influ-
enced by the time passed since the last feed intake, thus it is very impor-
tant to determine accurately the time of sampling (at least 2 hours later
than the last feeding). In poultry, this method is applied rather to assess the
quantity of abdominal fat, and it is considered in selection respects.
TOTAL BODY ELECTRICAL CONDUCTIVITY (TOBEC). This method is based
on lean tissue being a better conductor of electricity than fat, i.e. on the dif-
ferent electrical conductivity of the tissues (FIGURE III-5). The electrical
conductivity is decreasing in the following direction: muscles>organs>bone
marrow>fat.
36
FIGURE III-5: The basic principle of the TOBEC-method (frequency dependency of the
resistivity for several tissues)
The latest biotechnological methods (growth hormone injection, transgenic

production of growth hormone), the genetic work, the repartioning agents
like β-2 antagonists, L-carnitine, trivalent chromium may change the body
composition, thus experimental and practical importance of the above-
described methods is increasing. Hereby only, the most important methods
have been mentioned; there is a much wider and constantly increasing
spectrum of the available methods and equipments.
FOR FURTHER READING
Fekete, S. and Brown, D.L. (1993): The major chemical components of the rabbit whole
body measured by direct chemical analysis, deuterium oxide dilution and total
body electrical conductivity. J. Vet. Nutr. 2, 23-29.
Forbes, G.B. (1987): Human body composition. Springer Verlag. Nerlin - Heidelberg - New
York
Lister, D. (1984): In vivo measurement of body composition in meat animals. Elsevier
Applied Science Publishers. London - New York
Speakman, J.R. (2001): Body composition analysis of animals. A handbook of non-destruc
tive methods. Cambridge University Press.
Susenbeth, A. (1984): Berechnung der Körperzusammensetzung von Schweinen aus dem
Hilfe von D2O bestimmten Körperwasser. Dissertation. Stuttgart
37
Chapter IV
METABOLIC SIGNIFICANCE AND ANALYSIS OF

FEED INGREDIENTS
© Janos Csapo
4.1.Metabolic significance and chemical analysis

of feed components
4.1.1. Sample collection and preparation
4.1.2. Determination of dry matter content
4.1.3. Determination of (crude) ash content
4.1.4. Determination of mineral content
4.1.5. Determination of the fat content
4.1.6. Determination of the iodine value, acid number and
peroxide number in feed mixtures and in their components
4.1.7. Determination of the fatty acid composition
4.1.8. Determination of protein content
4.1.9. Determination of the amino acid composition
4.1.10. Determination of the carbohydrate content
4.1.11. Determination of the (crude) fibre content
4.1.13. Determination of the nitrogen free extract
4.1.14. Determination of some antinutritive components
4.1.15. Determination of the vitamins
4.2. Heat damage of proteins
4.2.1. The aims of the heat treatment of the fodder
4.2.2. Types of heat treatment
4.2.3. The effect of the heat treatment on the biological value
of proteins
4.2.4. Types of protein damage
4.2.5. Evaluation of protein damage by heat treatment
39
4.1. Metabolic significance and chemical analysis of feed

components
The different compounds in the living organisms have different roles. They
participate in the construction and maintenance of the animal organisms.
They construct the structure of the body, the bones, the muscles, the skin,
the different organs, the feather, the hair, the horn and the others. During
the life of the organisms, the different compounds have to be resynthetised
and substituted, because of the degeneration, wear and decomposition.
Proteins, fats, the minerals and the water are the most important compo-
nents in this respect.
The most important source of energy are carbohydrates, the fats and
proteins. Molecules in these component groups have also other important
roles, they regulate the biochemical and physiological processes in the body
as enzymes, hormones, regulators, vitamins, minerals, essential amino
acids and fatty acids, and they also have basic role in the production of milk
and egg.
Food of animal or vegetable origin sometimes shows significant fluc-
tuations regarding valuable constituents. Quality control is aimed at a rou-
tine inspection of production processes to achieve a standardised, defined
product. The food manufacturer wants to know not only the composition of
the food produced, but also the variations in the concentrations of the com-
pounds in the raw materials. In order to adjust the final product to the
desired quality reliable and actual composition data should be available. In
the case of animal production the knowledge of the exact composition of
feeds is essential in order to provide animals with the required nutrients. In
this respect the most important nutrients are starch and sugars, fat, pro-
tein and non starch polysaccharides, which can be used to estimate the
energy content of the feed, and the knowledge of the amino acids composi-
tion is also very important.
The aim of this chemical analysis is to provide accurate and repro-
ducible data for the producer or the consumer. There are only a few analyt-
ical methods, which are accurate, reliable, rapid and cheap. The methods
are continuously being modified, and new methods are being developed, but
40
CHAPTER IV: CHEMICAL ANALYSES
there are many factors reducing the accuracy of the methods, some of them
are very expensive and time consuming to elucidate. The matter of choice is
depending on the requirements with respect to accuracy, precision, and
rapidity and also on the financial possibilities.
4.1.1. Sample collection and preparation

One of the most important parts of an analytical method is the sample
collection and the sample preparation for analysis. If the sample collection
and sample preparation are inadequate, the results will be questionable
even if the best method is used. In the case of the feed samples the matrix
is very variable. The first requisite for successful sampling is to obtain a
representative sample. Food samples are extremely complex, containing
diverse compounds. The compounds are often heterogeneously distributed
in the sample, thus making sampling very difficult. For the representative
sampling large sample sizes or greater numbers of sub samples have to be
taken.
Sample preparation involves modifying the sample in some way that
renders it suitable for analysis. Sample drying is often required prior to
analysis. When compounds of interest are heat sensitive, the use of freeze-
drying may be necessary. The dry samples have to be ground to reduce par-
ticle size to a maximum of 1 mm. There are several types of grinders: some
of them force the sample through a mesh, in some cases the sample is
crashed by a large ball, others have jacket surrounding the chamber that
can be filled with liquid nitrogen. Sometimes samples should be ground
manually with a mortar and pestle.
The ideal case is, when the sample is analysed immediately after its
collection, because all compounds undergo chemical or structural changes
over time. For example the conformation of the enzymes changes and there-
fore they loose their biological activity within a few hours at higher temper-
ature. Most vitamins can rapidly be deteriorated if samples are stored at
light and in the presence of air. There are many factors that can accelerate
the degradation of some constituents, including heat, ultraviolet light, pH,
temperature, and the presence of reactants. For example ascorbic acid is
susceptible to oxidation; therefore it needs to be stored under oxygen free
conditions. Sample weighing can also be a major source of analytical error.
It is very important that the analytical balance has to be calibrated. To min-
imise moisture uptake, samples should be allowed to equilibrate at room
temperature or be kept in desiccators before the sample containers are
opened.
Some of the rapid methods use Infra-red Spectroscopy (IR), Near
infra-Red (NIR) and Medium Infra-red (MIR) light absorption. Infra-red light
is an electromagnetic radiation in the visible spectral range towards the
longer wavelength. In general, infra-red light has a wavelength of 760 nm to
0.5 mm. The wavelength at near infra-red is 760 nm – 2.5 mm, at medium
41
infra-red 2.5 mm to 25 mm, at far infra-red 25 mm to 500 mm. Infra-red

radiation causes heat development because infra-red light excites molecu-
lar bounds causing vibration. Part of the infra-red radiation energy is con-
sumed by vibration excitation and therefore, absorbed. If only specific wave-
lengths are allowed to pass through a filter, only defined molecular bounds,
characteristic for certain constituents, are excited. The absorption as a
result, can be measured by using different (NIR-Reflection-, Near
Transmission Measurements) techniques.
4.1.2. Determination of dry matter content

The reference methods are the gravimetric determinations after thermal
drying in a drying chamber. The dry matter content of the samples is deter-
mined by drying to constant weigh at 105oC.
4.1.3. Determination of (crude) ash content

The samples are incinerated at 550oC in a furnace until the organic mate-
rials are totally oxidised and disappeared from the sample. The ash of the
sample contains metal oxides originating from the organic material. If the
food or feed sample is contaminated with soil, the ash content is higher
because most of the soil components stay in the pot. During the combus-
tion some of the volatile elements (for example iodine, selenium, and mer-
cury) disappear from the ash.
4.1.4. Determination of mineral content

Metal content even in low concentration can be determined successfully by
means of atomic absorption spectrophotometry or atomic emission spec-
trophotometry with inductively coupled plasma. The atomic absorption pho-
tometric method has high selectivity, low detection limit and can be applied
in case of several matrixes. The atomic absorption spectrometry (AAS) tech-
niques can be divided into the following groups: flame-, furnace-, cold
vapour- and hydride AAS. In practice the flame and the furnace AAS are the
most widely used techniques. When flame AAS is used metal solution is
sprayed into the flame for atomising; the aerosol formation is done with a
pneumatic nebulizer. With the graphite furnace technique due to the longer
retention period of the atoms in the furnace, the detection limit for the met-
als is 100 times lower than that of at the flame technique.
When emission spectroscopy is used, the excitation is carried out with
the use of inductively coupled plasma in plasma emission spectrometry
(ICP). The favourable characteristics of the ICP source allow a suitable com-
promise of conditions for the simultaneous excitation of almost all the ele-
ments. The high temperature of the plasma allows hardly any interference,
and allows rapid determination by optical emission spectrometry with pho-
tomultipliers as highly sensitive detectors. In the ICP-AS sample solutions
are analysed and the sample preparation techniques are used before AAS
42
can usually be applied prior to analysis with ICP. The ICP instrument can
be used as sequential or simultaneous arrangement of spectrophotometers.
In the ICP-MS systems the ideal excitation concept of the ICP has been com-
bined with the very sensitive mass spectrometric detection.
4.1.5. Determination of the fat content

The reference method is a gravimetric determination after extraction with a
suitable solvent, carried out with or without previous acid- or alkaline
decomposition, depending on the sample material. These methods are
known from the name of their developers (Stoldt-Weibull, Röse-Gottlieb), and
there are many different rapid methods for fat determination based on
refractometry, volumerty, radiometry, NIR/MIR and NMR. Generally the fat
content is determined by the Stoldt method. 2.5 g of the sample to the near-
est 1 mg is weighed into a 400 cm3 beaker and 100 cm3 3M hydrochloric
acid is added to it. The beaker is covered with a watch glass and the mix-
ture is boiled gently over a hot plate for one hour. The product is not to be
allowed to stick to the sides of the container. After cooling, the material is
filtrated to prevent any loss of fat during filtration. The residue is washed
with cold water until a neutral filtrate is obtained. The residue is placed on
a watch glass and dried for one and a half hours in an oven at 1003oC. The
filter paper containing the dry residue is placed in an extraction thimble and
covered with a fat-free cotton wool. The thimble is placed in an extractor
and the fat content is determined by the Soxhlet method.
4.1.6. Determination of the iodine value, acid number and peroxide

number in feed mixtures and in their components
Unsaturated fat readily unites with oxygen and other compounds, for exam-
ple iodine. The double bond in the alkyl chain of the unsaturated fatty acid
can be saturated with oxygen or iodine. The iodine number, the number
derived from the grams of iodine absorbed by a hundred grams of fat, is an
excellent criterion for the evaluation of the degree of unsaturation. The
unsaturated bound is very sensitive to oxidation, resulting in off-flavours
and odours in fats and oils. This process occurs slowly and spontaneously,
and light, heat, and heavy metals accelerate it. The oxidative deterioration
of the fats is resulted in the formation of peroxides and free fatty acids. The
acid number and peroxide number give information about the quality of the
fat. The first step of the determination is the cold extraction of the fat from
the feedstuff. The peroxide number is determined by iodometric titration,
namely the peroxide in the rancid fat converts the sodium iodide to iodine,
which can be measured by thiosulphate solution.
Fats and the fatty acids that comprise them are described as saturat-
ed and unsaturated. These terms refer to the status of hydrogen in the fatty
acid. The iodine value is appropriate to measure the degree of unsaturation
of fat. Each double bonds takes up two atoms of iodine, therefore unsatu-
43
ration is quantified as the number of mg of iodine absorbed per 100 gram

of fat. The definition of the peroxide number or value (PV): the volume of 1
mole of sodium thiosulphate solutions in cm3 of the titration solution,
which is equal to the iodine that reacting with the peroxide in 1 kg of fat
(ether extract). The acid number is determined by titration with alcoholic
potassium hydroxide solution. The definition of the acid number or level of
acidity: the mass of potassium hydroxide in mg, which is necessary to neu-
tralise the acids present in 1 g of fat. Both numbers refer to the mass of fat,
and not that of the feedstuff. The anisidine value (AV) provides an estima-
tion of the presence of fatty aldehydes arising from peroxide decomposition.
In case of oils “totox” values are often quoted and are defined as
totox=2PV+AV. The maximum totox value in refined oil is 10. For veterinary
aspects see Chapter XIV.
4.1.7. Determination of the fatty acid composition

The fatty acid composition of samples is determined by gas chromatography
usually equipped by a flame ionisation detector. For quantitative evaluation,
the weight percentage proportions of the methyl esters are regarded as
equal to the proportions of the corresponding peaks in the chromatogram.
The fatty acid composition for unknown samples is calculated as a function
of the comparative mass percentage of fatty acid methyl-esters.
The following fatty acids are determined during gas chromatographic
analysis: Acetic acid (C2), Butyric acid (C4), Valeric acid (C5), Caproic acid
(C6), Heptanoic acid (C7), Caprylic acid (C8), Pellargonic acid (C9), Capric
acid (C10), Undekanoic acid (C11), Lauric acid (C12), Myristic acid (C14),
Myristoleic acid (C14:1), Pentadecanoic acid (C15), Palmitic acid (C16),
Palmitoleic acid (C16:1), Margaric acid (C17), Stearic acid (C18), Oleic acid
(C18:1c), Elaidinic acid (C18:1t), Nonadekanoic acid (C19), Linoleic acid
(C18:2), Arachic acid (C20), Gadoleic acid (C20:1), Eicosadienoic acid
(C20:2), Linolenic acid (C18:3), Eicosatrienoic acid (C20:3), Arachidonic acid
(C20:4), Eicosapentaenoic acid (C20:5), Behenic acid (C22), Eruic acid
(C22:1), Docosahexanoic acid (C22:6), Lignoceric acid (C24), Selacholic acid
(C24:1), Cerotic acid (C26), Montanic acid (C28). In the brackets there are
the numbers of the carbon atoms in the molecule, followed by the numbers
that of the unsaturated bounds. These are the major fatty acids in feed-
stuffs. Little is known about the nutritional importance of the many minor
fatty acids which generally occur in proportion less than 1%; with the
exception of polyunsaturated minor fatty acids from which linoleic acid and
arachidonic acid are essential. Linolenic acid is however a semiessential
fatty acid for the most sorts of animals.
4.1.8. Determination of protein content

The accurate and precise quantitation of protein and specific amino acids is
essential for feed evaluation and feed analysis. Laboratories have adapted a
44
wide range of techniques. Proteins are made up of amino acids that are
chemically quite diverse. Some of these amino acids are chemically reactive,
therefore can be targeted for determining protein content. Methods such as
direct measurement of absorbance at 280 nm and a variety of dye binding
methods are based on target key characteristics of certain amino acids. In
order to determine the protein content, the nitrogen content is measured by
several methods, but not only the whole amount of the amino acids in meas-
ured. The Dumas and Kjeldahl methods are based on the fact that the pro-
tein is quantitatively the most significant nitrogen containing compounds
present in feedstuffs. The Dumas methods involve the combustion of the
sample and the volume of the nitrogen gas released is measured. The
Kjeldahl method involves the transformation of the nitrogen containing
materials of the sample into ammonia that is titrated against a weak acid.
Proteins contain ~16% nitrogen, and consequently a conversion factor of
6.25 is used to convert nitrogen content to crude protein. This conversion
factor may be unsuitable in case of proteins containing large amount of
tryptophan and basic amino acids (histidine, lysine and arginine) with high-
er nitrogen content. In some cases different conversion factors are used: for
example grain millers often use 5.7 and the dairy industry often uses 6.38.
In most countries the reference method is the volumetric determina-
tion of the released ammonia after sulphuric acid decomposition according
to Kjeldahl. Protein can also be determined by direct distillation, colour
binding method, NIR/MIR or NMR. The rapid and very accurate determina-
tion of the protein contents can be done by automated Kjeldahl method
(Kjel-Foss or Kjel Tec Automatic). Total protein contents of the samples is
measured by Kjel-Foss 16200 typ. nitrogen analyser (protein content = N%
× 6.25). The sample is digested by sulphuric acid in the presence of potas-
sium-sulphate, mercury-oxide and hydrogen-peroxide. The acidic solution
is alkalised with sodium-hydroxide solution. The ammonia is distilled and
collected and its quantity is measured, and ammonia is titrated with a stan-
dard solution of sulphuric acid. After introduction of the sample into the
equipment, the whole classical Kjeldahl method is fully automated. The
equipment is arranged as a carousel, and the individual processing steps
are divided into six interlocking automatic steps: sample digestion,
hydrolysate cools down and diluted with water, sample alkalisation and dis-
tillation of the free ammonia into a simultaneously running titration device
in which the colour change is determined photometrically, the Kjeldahl flask
is emptied automatically, the catalyst used precipitates as a sulphide and
finally the flask is ready for receiving another sample. Each step take about
three minutes and the result is available in 12 minutes.
Although the Kjeldahl method is the most popular method for protein
determination, the near infrared spectroscopy has received considerable
interest in recent years. This technique is very rapid and uses the near
infrared absorption spectra of feedstuffs with a set of calibration samples for
45
which protein content has been determined by conventional methods.
4.1.9. Determination of the amino acid composition

The amino acid compositions of the samples are measured by automatic
amino acid analyser. Hydrolysis of the samples is performed in reusable
Pyrex hydrolysis tubes.
Amino acids can be analysed with an automatic amino acid analyser
based on ion exchange chromatography (IEC). The amino acid composition
can also be determined by high performance liquid chromatography (HPLC).
The tryptophan content of the samples is determined after barium hydrox-
ide hydrolysis. Mostly the following amino acids and amino acid derivatives
are determined during ion exchange chromatography: the amino acids and
the ninhydrin positive compounds in the order of appearance on the chro-
matogram are:
1. cysteic acid, 2. taurine, 3. phosphoserine, 4. urea, 5. aspartic acid
(Asp), 6. hydroxyproline, 7. threonine (Thr), 8. serine (Ser), 9. asparagine
(Asn), 10. glutamic acid (Glu), 11. glutamine (Gln), 12. a-aminoadipic acid,
13. proline (Pro), 14. glycine (Gly), 15. alanine (Ala), 16. citrulline, 17. a-
aminobutyric acid, 18. valine (Val), 19. cystine (Cys), 20. methionine (Met),
21. cystathione, 22. isoleucine (Ile), 23. leucine (Leu), 24. tyrosine (Tyr), 25.
phenylalanine (Phe), 26. b-alanine, 27. b-aminoisobutyric acid, 28. g-
aminobutyric acid, 29. chlorophenylalanine, 30. ethanolamine, 31. ammo-
nia, 32. ornithine, 33. lysine (Lys), 34. histidine (His), 35. 1-methylhistidine,
36. 3-methylhistidine, 37. arginine (Arg). (In the brackets there are the
abbreviations of the protein building amino acids.)
4.1.10. Determination of the carbohydrate content

Reference methods are reduction methods according to Luff-Schorl, polari-
metric methods and enzymatic methods.
4.1.11. Determination of the (crude) fibre content

Fibre designates the complex of carbohydrates and other substances that
are present mainly in the cell walls of the plants. Fibre includes cellulose,
hemicellulose, gums, mucilages, pectin substances and lignin. This sub-
stance is poorly digested by monogastrics, but the ruminants can utilise
this fraction well. During the determination of the crude fibre, the sample is
boiled in 1.25% sulphuric acid. After neutralisation with distilled water it is
boiled again in 1.25% sodium hydroxide, than the sample is filtered,
washed, dried and measured. Then the ash content of the fibre is deter-
mined, and it is subtracted from the weigh of the fibre that resulted in the
amount of the true fibre.
4.1.12. Determination of fibre fractions by Van Soest analysis

Van Soest developed a new method of fibrous fodder analysis that is able to
46
separate the cell content with different fodder value and the fibre fractions.
As a first step the sample is cooked in a neutral detergent solution (3% Na-
lauril-sulphate, buffered for pH 7) for 60 minutes than it is filtered. The
neutral detergent soluble part (NDS) equals to the cell content, which is
built up of lipids, sugars, starch and protein, and their digestibility is near-
ly 100%. The insoluble part is the NDF (neutral detergent fibre) which con-
tains all of the cell wall constituents (CWC= cell wall constituents) consist-
ing of mainly cellulose, lignin, silica, hemicellulose and some protein. (In the
human medicine apart from NDF the “dietary fibre” expression is used.)
The NDF determined by the above mentioned method comprises prac-
tically all the lignin and hemicellulose, but in the case of the Weende analy-
sis some parts of them – differing in proportion in the different feedstuffs –
appear in the nitrogen free extract and not in the raw fibre. That is why the
NDF content of every fodder is higher than their raw fibre content. The com-
pounds of the NDF can hardly be digested and only bacteria can do it. The
lignin and the silica are indigestible. Lignin affects negatively the digestion
of the hemicellulose and that of cellulose as well.
A further differentiation within the NDF content is done with the use
of acidic detergent solution (sulphuric acid cetyl trimetyl ammonium bro-
mid). The NDF fraction is boiled in it for 60 min following filtration. The
insoluble remaining part is the acid detergent fibre (ADF= acid detergent
fibre) which is comprised of cellulose, lignin and silica (quartz). The differ-
ence between the NDF and the ADF content gives the amount of hemicellu-
lose.
For determining the lignin the ADF is bathed for 3 hours in 72% sul-
phuric acid and then filtered. The insoluble part is the lignin and the silica.
The latter can be measured by burning (as leftover after burning). The lignin
determined by this method is called ADL (acid detergent lignin) as opposed
to permanganate lignin in acetic acid agent oxidisable by potassium per-
manganate. This latter in some cases can give smaller values than the ADL,
because it does not measure cutin, while the sulphuric acid method does
so. The MAILLARD-complex (e.g. brown hay) appears in the lignin fraction and
the insoluble organic materials (e.g. some plastics) are mixed to the feed-
stuff as well.
On the basis of the results obtained we can calculate the digestibility
coefficient of the dry material of the roughages and forages:
DCorganic material = 0.98 NDS+(1.472-0.789 log10ADL) × NDF,
where DC= digestibility coefficient, NDS and NDF is the dry material
appearing as the % of the ADL and ADF.
4.1.13. Determination of the nitrogen free extract

The nitrogen free extract (NFE) of the sample is not measured, but calcu-
lated from the other results of theanalysis. During the calculation the crude
protein, crude fibre, crude fat and crude ash content is subtracted from the
47
dry material content; so the nitrogen free extract includes the sum of sug-
ars, starch, the soluble part of cellulose and hemicellulose, and other mate-
rials which was not determined from the sample. Recently in the dairy cat-
tle nutrition (NRC, 2001) the nitrogen-free extract fraction from proximate
analysis has been replaced with the fractions referred to as nonfibre carbo-
hydrates (NFC=100% of DM-%NDF-%CP-%-%ash-%ether extract) and non-
structural carbohydrates (NSC, determined by enzymatic analysis. These
latter two fractions do differ, especially for ensiled forages, because of the
concentration of organic acids. The NFC fraction analysis can be refined by
determining the nitrogen in the NDF fraction to provide for NDFN-free.
4.1.14. Determination of some antinutritive components

4.1.14.1. Determination of the urease enzyme activity from soybean. The
method can be applied to the determination of the urease enzyme activity of
foodstuffs, and the determination of the degree of heat treatment of soy-
bean. The soybean samples are put into two buffer solutions with the pH of
7.5. The first solution contains urea, whereas the second solution does not
contain any. After 30-minute incubation at 35oC, the pH of the two differ-
ent solutions is measured, and the pH difference is calculated.
The soybean sample is milled as fine as flour (the size of the particles
should be less than 0.3 mm). From this fine material 2x1 g is measured into
two 100 cm3 beakers. Buffer solution containing urea is added to the first
sample and only buffer solution (without urea) is added to the second sam-
ple. The content of the beakers is mixed thoroughly with a glass mixer, and
the bakers are put into a thermostat at 35oC for half an hour. While the
samples are in the thermostat, they are mixed up at every five minutes.
After 30 minutes the samples are filtered through a fine filter paper and the
pH of the clear solution is determined by a pH meter. If the pH difference
between the two samples is (sample in buffer, and sample in buffer con-
taining urea):
1.7-2.5: this is a crude soybean, the sample was not treated by heat, or the
heat treatment was not sufficient;
0.2-1.7: the sample was heat treated only for a few minutes, insufficient
heat treatment;
0-0.2: good heat treated sample, this sample is sufficient for animal or
human consumption.
4.1.14.2. Determination of the trypsin inhibitor activity in soybean
The trypsin cleaves off a yellow colour compound (p-nitro aniline) from the
artificial substrate of N-a-benzoil-DL-arginin-p-nitroanilid-hydrochlorid
(DL-BAPA or BAPA). In the presence of trypsin inhibitor the colour intensi-
ty will be lower, and this way the enzyme inhibition can be measured pho-
tometrically. The air dry samples are defatted by cold extraction with petro-
leum ether. The defatted sample is milled as fine as flour (the size of the par-
ticle should be less than 0.1 mm). From this fine material 2 g is measured
48
into a 100 cm3 baker, it is suspended by 60-70 cm3 distilled water, the pH
of this solution is set by 1M sodium hydroxide to 9.8, and the total materi-
al is diluted with distilled water up to the volume of 100 cm3. This solution
is mixed by magnetic stirrer for 3 h.
From the solution 0.6, 1.0, 1.4 and 1.8 cm3 are measured into a test
tube, and distilled water is added to them until the volume of each solution
is 2 cm3. 2 cm3 trypsin solution is added to every test tube, and the test
tubes are put into 37oC water thermostat for 5 minutes. After this 5 cm3
BAPA is added to each test tube, and mixed. They are immersed into the
thermostat again for 10 minutes at 37oC, and then 1 cm3 30% acetic acid
is added to it at room temperature. The samples are filtered into dry test
tubes, and after 30 minutes the colour intensity is measured at 410 nm.
For the investigations the following solution must be made: reagent
blank solution that contains all the reagents except for sample (“0-solution”
which does not contain soybean). Soybean blind solution: that contains only
soybean and BAPA, but does not contain trypsin. This value has to be sub-
stracted from the colour intensity of the solution that containing trypsin.
The determination of the trypsin inhibitor unit (TIU): based on the
determination of the colour intensity of the soybean solution that contains
trypsin minus the colour intensity of the 0-solution. The TIU can be calcu-
lated from the calibration curve, (which was measured by different measur-
ing). If the colour intensity is plotted against the different volumes, the TIU
can be measured where the calibration curve will reach the y-axis. The
result of the trypsin inhibitor activity determination is given as TIU/1 g fat
containing soybean sample.
4.1.15. Determination of the vitamins

Vitamins are organic substances that are essential in small amounts to
health, growth, reproduction, and maintenance of one or more animal
species. They must be included in the diet since they cannot be synthesized,
or their synthesis is limited and therefore insufficient related to the needs
of the organisms. The vitamins have very different structures and charac-
teristics, and the sensitivity of the vitamins for oxygen, heat treatment, irra-
diation and storage is also very diverse. The potency of vitamin content of
feeds can be determined by biological, microbiological, chemical and physi-
cal methods. At biological assay the laboratory animals are first deprived of
the particular vitamin being studied by being fed a diet lacking it. Then the
different groups of animals are provided with different levels of the known
vitamin, and the growth response of the animal is measured and recorded
in a standard response curve. During the microbiological assay the ability
of vitamin to promote growth/multiplication of microorganisms is meas-
ured. Nowadays vitamins can be analysed by different chemical procedures,
and the vitamin content is expressed in units of weight mg or mg. Chemical
assays are much faster than biological or microbiological assays, but the
49
results must be compared to bioassays in order to exclude the possibility

that with the use of the chemical investigation biological inactive form of the
vitamin is detected and treated as if it was active form.
4.2. Heat damage of proteins
4.2.1. The aims of the heat treatment of the fodder

During the production of feed components and complete feeds, heat treat-
ment is frequently used. The aim of this could be:
-Ensuring the proper microbiological quality (for instance sterilization of
slaughterhouse by-products)
-Improving digestibility (for instance treating materials containing keratin
etc.),
-Improving the shelf-life of the product (for instance drying corn grains, milk
powder, yeast, animal protein flour, green flours),
-Inactivating antinutritive factors (for instance elimination of the trypsin
obstructive effect of raw legumes etc.),
-Terminating the non-favourable enzyme activity (for instance inactivating
thiaminase in fish debris, eliminating the enzyme activity of yeast milk etc.),
-Extracting the (dis)solvent during any isolation technology (for instance in
the vegetable oil industry from the extracted breeze),
-Processing (“opening”) starch in the feed of carnivore animals (for instance
extruding dog and cat feeds),
-Flavour improvement (for instance terminating the raw flavour in case of
potato).
Furthermore, heat is developed during every grounding and granulat-
ing operation and also during some non-favourable fermentation (for
instance warming up of silage hay) and in chemical reactions (for instance
autooxidation, rancidity of oil content of fish meal) as well. During heat
treatment the chemical reactions lead to the decomposition of nutrient
materials when the kinetics of the reactions fasten up, but the enzyme reac-
tions due to denaturation of proteins are eliminated.
4.2.2. Types of heat treatment

The effects of heat treatment vary greatly altering depending on whether
heat is overtaking the feeds in dry or humid environment. The difference
cannot be explained by the fact that the humid heat treatment is oxygen
free or that it creates an environment lacking oxygen but there is a lot of dif-
ference in the special (possibilities of changing the configuration) behaviour
of proteins as well.
50
Of the above mentioned heat treatment procedures the infra red and
microwave heat treatment is apparently appropriate for the treatment of
small amount of material.
4.2.3. The effect of the heat treatment on the biological value

of proteins
During heat treatment the configuration of the protein molecule changes
first. In the native protein the polypeptide chain which is built up by amino
acids connected with peptide bonds create a three dimensional configura-
tion, which is stabilised by chemical bonds (for example disulphide bridges)
and other, supplementary bindings (for example hydrogen bridges) between
the side chains of the amino acids.
Due to the heat treatment in wet medium the heat movement of mol-
ecules is intensified and so heating proteins results in the bounds being
broken, which in turn causes the denaturation of the protein (for example
this is taking place while cooking the eggs). During the denaturation of pro-
tein the quaternary, tertiary and secondary configuration of the protein are
altered but the primary structure (peptide chain) is not broken.
DENATURATION means an irreversible process going hand in hand with
51
the cessation of the biological activity connected to the configuration of the

protein. So enzymes, immunoglobulines, lysozyme and lactotransferrin
transferring the iron in the mother’s milk etc. could be inactivated. From the
point of view of feeding though denaturation is favourable as the digestibil-
ity of the protein is improving (boiled meal) and as well the organoleptic fea-
tures. The other favourable effect of denaturation is that the activity of some
heat sensitive antinutiritive materials is decreased or ceased.
When not only the configuration but the side chains of the amino
acids creating the primary structure of the protein is altered due to heat
treatment this change leads to the deterioration of protein, when biological
value is diminished.
These sorts of changes can be:
a) the reaction of the free e-amino group on the side chain of lysine
with the carboxyl group of reducing sugars,
b) or with the side chain of other amino acids (for example glutamine
or asparagine).
4.2.4. Types of protein damage

During heat treatment the side-chains of the amino acids can react with
each other within the polypeptide chain, or between the polypeptide chains
of different proteins, and they can be reacted as well with other biomole-
cules e.g. carbohydrates, and lipids, too.
4.2.4.1. Protein–protein type interactions

When proteins are exposed to heat (this is typically happens during the pro-
duction of animal protein flours) in the absence of carbohydrates the fur-
ther important reactions occur:
intramolecular crosslink-formation between the free e-amino group of the
lysine and other side chains of amino acids;
during the decomposition of the amino acid side-chain dezamination
(ammonia release) and desulphuration (release of hydrogen sulphide and
mercapthane) can occur and
oxidation procedures can take place.
CROSSLINK FORMATION has an importance regarding the nutritional
value of proteins, e.g. the crosslink between the e-amino group of lysine and
the acidamid group of glutamine can lead to a decrease in the digestibility
of protein. Ammonia release can be significant during strong heat treatment
at the production of feather meal, which on the one hand decreases the raw
protein content of the final product, but on the other, it causes pollution of
the environment. On the other hand, during the harmless production of ani-
mal meals containing protein the hydrogen sulphide released from sulphur
containing amino acids causes on the one hand corrosion of iron made sur-
faces of facilities, but on the other, in the case of the mercapthans, a strong
smell formation occurs. Of course it is accompanied by the decrease of the
52
amount of the sulphur containing amino acids as well.

Heating at elevated temperatures exerts an effect on the disulphide
bridge of the cystine in the feather keratin. Cystine is decomposed and
turns into inactive lanthionine in terms of feeding. The structure of lan-
thionine is similar to that of cystine, but instead of a disulphide bridge it
contains only one sulphur atom. Oxidation effects are accelerated due to
heat treatment. Methionine is particularly susceptible to these processes.
On account of a more gentle oxidation methionine sulphoxide (Met-SO) is
formed, but in a reductive environment, the methionine can be regenerat-
ed. The biological activity of methionine sulphoxide is about half than that
of methionine. In the case of a more intensive oxidation effect the methion-
ine sulphoxide turns into methionine sulphone (Met-SO2). This reaction is
irreversible, and since the methionine sulphone has no biological activity, it
cannot be used as a source of methionine.
Due to severe oxidation effect cysteine irreversibly transforms into
stable cysteine acid which is extracted during the quantitative measure-
ment of cysteine.
4.2.4.2. Protein–carbohydrate type interactions

The most well-known protein impairment is the reaction of proteins with
reductive sugars, which is called non-enzymatic browning or Maillard-reac-
tion. The Maillard reaction involves a condensation reaction between reduc-
ing sugars and protein. In the Maillard-reaction at first the free e-amino
group on the side chain of the lysine and the carbonyl-group of the reduc-
tive sugars take part. The subsequent degradation and modification of
Amadori compounds (1-amino-1-deoxy-2-ketose) result in the formation of
brown pigments and flavours ranging from burnt, hay-like caramel. The
reaction sequence is a complex and dependent on temperature and pH.
Thus the reaction creates a brownish product due to some elaborate proce-
dures. The Maillard-reaction leads to the decreasing of the utilisable
amount of lysine since the substituted lysine (deoxifructosil-lysine) on the
e-amino group can not be used for protein synthesis and is emptied with the
urine at a constant form.
Some intermediers of the Maillard-reaction (furosine, piridosine) can
be measured analytically and their amount is characteristic to the heat
impairment of the protein. In the case of Maillardised feedstuff protein (for
example browned milk powders, overdried corns, or green flours, brown
hay) we have to consider whether the bioavailability of lysine is decreased,
which can lead to the lower weight gain in animals. A small amount of
Maillardisation can have a taste improving effects (for example the crust of
the bred). The flavours arising from the Maillard reaction, plus a controlled
proteolysis of poultry viscera, which in turn, can lead to the production of
low molecular weight peptides with low flavour thresholds gives the taste
and preference of so-called DIGEST.
53
4.2.4.3. Protein–lipid type interactions

The easily disintegrating adducts formed between the proteins and lipids
were claimed to have no importance with respect to feeding. On the other
hand when lipid oxidation occurs, the oxo-groups of the oxidation products
can act with the side chains of the amino acids like reducing sugars in the
Maillard-reaction, thus through these processes, lipid oxidation can also be
led to the decrease of the bioavailabitity of lysine.
4.2.5. Evaluation of protein damage by heat treatment

The degree of heat treatment and that of protein impairment are indicated
by numerous phenomena:
54
FOR FURTHER READING
Allen, J.C. and Hamilton, R.J. (ed.) (1994): Rancidity in foods. Third Edition. Blackie
Academic & Professional. London, Glasgow, Weinheim, New York, Tokyo,
Melbourne, Madras
A.O.A C. (1975): Official methods of analysis. (12th Ed). Association of Official Analytical
Chemists. Washington, D.C.
El-Khoury, A.E. (Ed.) (1999): Methods for Investigation of Amino Acid and Protein
Metabolism. CRC Press. Boca Raton-London-New York–Washington, D.C.
van Es, A.J.H. and van der Meer, J.M. (1980): Methods of analysis for predicting the ener
gy and protein value of feeds for farm animals. Lelystad
Moughan, P.J., Verstegen, M.W.A. and Visser-Reyneveld, M.I. (2000): Feed evaluation.
Principles and practice. Wageningen Pers, Wageningen
Pellet, I.P. and Young, V.R. (1980): Nutritional evaluation of proteins in food. The United
Nation University, Tokyo
55
Chapter V
ENERGY AND PROTEIN METABOLISM

© Andre Chwalibog and Ewa Sawosz
5.1. Principles of energy metabolism
5.2. Energy-yielding processes (catabolism)

5.2.2. Oxidation of fat
5.2.3. Oxidation of proteins
5.3. Energy-requiring processes (anabolism)
5.4. Methods for the quantitative determination of

heat production
5.5. Partition of energy in the body
5.6. Energy and protein utilization and requirements

5.6.1. Requirements for maintenance
5.6.2. Requirements for growth
57
Chapter V: ENERGY AND PROTEIN METABOLISM
Ife of animals depends to a large extent on energy supply. Energy is
L needed feed ingestion, for nutrient absorption, endocrine and nervous

regulation, absorbed substrates utilisation, maintenance and synthe-
sis, and for physical activities such as work, defence, etc. Homoeothermic
animals, moreover, use energy to maintain the constant body temperature.
5.1. Principles of energy metabolism
The word energy originates from Greek and means “in work” (en ergon).
Energy is the capacity to do work and can be measured only in its trans-
formation from one form to another. Originally, all measurements of energy
transactions made by animal nutritionists were expressed in terms of kilo-
calories. The calorie (cal) is defined as the amount of energy required to
raise temperature of 1 g of water by 1oC, measured from 14.5 to 15.5 oC.
Since this unit is too small for most convenient use in nutrition, it has been
replaced by the kilocalorie (kcal). Although the calorie is the basic unit of
heat energy, the joule was adopted in the International System of Units as
the preferred unit for expression of electrical, mechanical, and chemical
energy (Table V-1). The joule is defined in terms of the International Metric
System as one kg-m2/sec2 or 107 erg. Because all measurements of ener-
gy are defined in terms of the fundamental metric units of mass (kg), dis-
tance (m), and time (sec), joules are readily converted to calories and vice
versa. Measured in terms of joule 1 kcal = 4.185 kilojoules (kJ). The heat-
technical calculations are based on the unit of watt with the use of a heat
production unit (hpu) defined as 1 hpu = 1000 W. Since 1 W = 0.860
kcal/hour or 3.6 kJ/hour, 1 hpu = 3600 kJ/hour.
58
Table V-1: Energy units used in biological and technical calculations
There are two categories of energy: potential and kinetic. Potential

energy is the energy of position or bound energy which can be released to
produce kinetic energy or to do work. Kinetic energy is the energy of move-
ment. In the living organism the energy of random molecular motion is heat
energy. Energy supplied to animals in chemical form can be transformed to
heat when utilized in support of life processes, stored in chemical form as
in growth, transferred in chemical form to a second animal as in pregnan-
cy and lactation, or transferred to surroundings as in work. Bioenergetics of
animals and man is primarily based on the two laws of thermodynamics.
The first law of thermodynamics is the law of conservation of energy and

holds that energy can be transferred or transformed but neither can be cre-
ated nor destroyed.
The second law of thermodynamics states that all forms of energy are
quantitatively convertible to heat, that heat is the lowest energy form and
that the driving force of all energy transactions is the tendency to reach the
lowest energy form, i.e. heat.
The second law can also be stated as the tendency for energy with a high
degree of orderliness to be converted to energy with a lower degree of order-
liness. The degree in which the total energy of a system is uniformly dis-
tributed (randomness) and thus unavailable to do work is expressed by the
term entropy. In any isolated system, entropy tends to increase because
randomness is more probable than orderliness. The more disordered or ran-
dom a system becomes the more entropy it has. Some molecules have a
greater order than others and so they have lower entropy. For example pro-
teins are highly ordered but upon denaturation they change to a much more
random structure, hence the increase in entropy during denaturation is
considerable. Generally solids are more ordered than liquids which are more
ordered than gases. According to the second law, any change in the total
energy content of a system (e.g. heat of combustion in a biological oxidation)
results in a change in both free energy and the entropy of the system. Since
only the former can be utilized to perform work of any kind, energy-yielding
59
reactions invariably have an efficiency of less than 100%.

It can also be stated that in accordance with the first law the total ener-
gy of the universe remains constant, and in accordance with the 2nd law
the entropy of universe always increases. Both laws of thermodynamics
emphasis that all forms of energy (in living organism) can be measured by
heat. Therefore determination of heat production is theoretically a sound
measure of metabolism; it is also a convenient one.
The total energy content of an isolated system, for example that of an
animal, cannot be measured, but it is possible to measure the energy
changes - transfer and/or transformation - which occur within the system.
The difference between the energy content of the substrates and that of the
products is called the heat of reaction - enthalpy of change (ΔH). If heat is
gained from the environment ΔH is positive (exothermic reaction); if it is lost
the enthalpy is negative (endothermic reaction). For example if glucose is
oxidized in a bomb calorimeter, 2.8 MJ of heat are liberated per mole of glu-
cose oxidized, the ΔH is then -2.8 MJ. Provided one has a set of heats of for-
mation of compounds, then the enthalpy of any reaction, whether exother-
mic or endothermic, can be calculated as the difference between the heats
of formation of products and reactants. It is most important to note that the
value of ΔH is independent of the chemical pathway followed by the reac-
tion, that is, it is independent of the mechanism of the reaction. Whether
glucose is oxidized by burning in a calorimetric bomb in the laboratory or
by the action of glycolysis in the body, ΔH is the same. Although the
enthalpy changes from a specified substrate to the known product is the
same whatever is the mechanism of conversion, the amount of work done -
amount of useful energy - will depend on the mechanism.
The amount of useful energy- free energy - ‘orderly energy’ i.e. capable
of doing work which is liberated in a chemical reaction, cannot be deter-
mined from the heat of reaction. Another factor must be calculated; the
change in free energy (ΔG) which is a difference between free energy of reac-
tants minus free energy of products. This change for a reaction occurring
reversibly at constant temperature and pressure is defined as: ΔG = ΔH -
TΔS, where T is the absolute temperature, ΔS is the change in entropy. If ΔG
is negative, the reaction can occur spontaneously, if ΔG is positive, it can-
not.
Potential energy derived from the metabolism of foods is stored in liv-
ing organisms as “high energy compounds”. The breakdown of these com-
pounds liberates free energy, which can accomplish work together with heat
energy, which cannot do work, and thus a measure it is of the energetic
inefficiency of the reaction. The energy liberated by chemical reactions in
the cell is used to drive other reactions. These require some methods for
coupling the reactions that liberate energy to those that use it. The first step
involves storing the energy derived from combustion of foodstuffs in certain
high energy compounds. The major system in cells that exploits energy from
60
oxidation of fuels for ATP synthesis is the oxidative phosphorylation. Fuel

oxidation generates reducing equivalents, which funnel into mitochondrial
electron transport chain, which pumps protons into the mitochondrial
inter-membrane space. The inner mitochondrial membrane is impermeable
to protons, and the energy of the proton gradient is recaptured by ATP syn-
thetase in the inner membrane. This enzyme allows protons to flow back
down their concentration gradient across the membrane, and the process
uses the energy of the gradient to drive ATP synthesis. Oxidative phospho-
rylation, however, comes with an additional cost, the production of reactive
oxygen species, potentially harmful for organism (HARPER et al. 2004 and
SAWOSZ et al., 2005).
The most important compound of the high energy compounds is
adenosine triphosphate (ATP). In a simple way, we may say that a cell is
supplied with energy stored in ATP, while the energy derived from oxidation
is used to re-synthesize ATP. The free energy released in the hydrolysis of
ATP to ADP (adenosine diphosphate) under standard conditions (standard
free energy measured when 1 mole of reactant is converted to 1 mole of
product at 37oC and pH=7) is generally accepted to be about -30 kJ per
mole. The standard ΔG of the reaction ADPΔ AMP (adenosine monophos-
phate) or AMPΔ phosphate is slightly lower, about 27 kJ/mole. However,
under the conditions of pH and temperature in the body, and because most
of the ATP and ADP in the intact cells is presented as Mg++ complex, which
has the effect of shifting the equilibrium of ATP hydrolysis, the standard free
energy change of ATP hydrolysis is much higher. On average the amount of
energy that is available from the phosphate bond of ATP is 52 kJ/mole.
Directly measured standard free energy of hydrolysis of phosphate com-
pounds is demonstrated in Table V-2.
Table V-2: Standard free energy of hydrolysis
61
Although the standard free energy of ATP hydrolysis occupies an inter-

mediate position, the role of ATP is unique. In comparison with other phos-
phates, ATP is the only phosphate which serves as a carrier of chemical
energy between donors and low-energy phosphate acceptors. ATP is a com-
mon intermediate in both energy-delivering and energy-requiring reactions
of the cell, and it is the only form of chemical energy that can be converted
into all others form of energy used by living organisms. It therefore fills a
role analogous to that of money -“energy currency of the cell”. The main
weakness of the analogy is that money, unlike ATP, can be accumulated for
later use.
5.2. Energy-yielding processes (catabolism)
Animals require energy requirement for maintenance and production (meat,

milk, foetus, eggs, wool, and work). The energy is provided by degradation
of the nutrients absorbed by the animal. Under normal conditions the ener-
gy is produced by oxidation processes in the three main groups of nutrients:
carbohydrates, fat and protein according to the following formula:
Nutrient + O2 → 6 CO2 + H2O + energy
↑
Catalysts
(intercellular enzyme systems)
The oxidation proceeds through a series of processes, in which the
degradation from one stage to the next is catalyzed by intercellular enzyme
systems, such as the dehydrogenases, which liberate hydrogen, and the
oxydases, which activate oxygen. Part of the energy resulting from oxidation
of the organic substances in the body is transferred to energy-rich phos-
phorus compounds (mostly ATP) (FIGURE V-1) from which they are later
mobilized and utilized for syntheses and other life processes (work, intake
and transport of food, absorption of nutrients etc.)
FIGURE V-1: Model of ATP synthesis and hydrolysis
62
Oxidation of a typical carbohydrate, fat and protein can be summarized

in the following stoichiometric equations:
Glucose:
1 g glucose + 0.747 litre O2 →0.747 litre CO2 + 0.60g H2O - 15.65 kJ
1 mole C6H12O6 + 6 O2 → 6 CO2 + 6 H2O - Energy
180 g 134.5 litre 134.5 litre 108 g 2817 kJ
Tripalmitate:
1 g trip. + 2.011 litre O2 →1.416 litre CO2 + 1.09 g H2O - 39.8 kJ
1 mole C51H98O6 + 72.5 O2 → 51 CO2 + 49 H2O - Energy
807 g 1625 litre 1143 litre 883 g 32 MJ
Meat protein:
for mammals:
1 g protein + 0.992 l O2 →0.848 l CO2 + 0.38 g H2O +0.332 g urea-18.4 kJ
for birds:
1 g protein+0.893 l O2→0.649 l CO2 +0.49 g H2O+0.498 g uric acid-17.8 kJ
5.2.1. Oxidation of carbohydrates

Carbohydrates are broken down in all cells, especially in the liver cells,
where glucose is synthesised to glycogen which is stored in the liver and the
muscles. Under catabolic conditions the glycogen is hydrolysed to CO2 and
H2O while taking up O2. The complete oxidation of glucose in the body can
be described as follows:
C6H12O6 + 6 O2 = 6 CO2 + 6 H2O - 38 mol ATP
In the combustion of 1 mole of glucose, 6 moles O2 are used, 6 moles CO2
are produced, and 38 moles ATP (adenosine triphosphate) are liberated,
which then act as energy donors for the anabolic; energy-consuming
processes. Assuming that each mole of ATP contains 52 kJ, 1 mole of glu-
cose will produce 38×52=1976 kJ in the body. Since the total energy con-
tent determined by combustion in the bomb calorimeter, i.e. outside the
body, is 2817 kJ/mole, this will be equivalent to an efficiency of 70%, (1976
kJ/2817 kJ). This shows that only 70% of the total energy of the glucose is
available for the anabolic processes, while the rest is lost as heat. The
release of energy in utilizable form of glycogen that necessitates its break-
down to glucose which is then oxidized as previously described. The net
effect of glycogen breakdown is the production of glucose-1-phosphate plus
a little glucose. Glucose-1-phosphate and the residual glucose are convert-
ed to glucose-6-phosphate which enters the pentose phosphate pathway.
The production of glucose-6-phosphate does not involve expenditure of ATP
so that ATP provided from glycogen is slightly higher that it is from glucose
(Table V-3).
Not all tissues obtain energy from aerobic processes. Erythrocytes,
white blood cells, kidney, tissue of the eye and cancer cells gain most of
their energy from the anaerobic conversion of glucose to lactate. According
to the generally accepted view, metabolism is aerobic at rest and during
63
moderate work. At some level of work, aerobic metabolism becomes inade-

quate to supply energy, and energy deficit is supplied from anaerobic
metabolism. Under anaerobic conditions the formation of 2 moles of lactic
acid from 1 mole of glucose is associated with the net formation of only two
moles of ATP. The anaerobic breakdown of 1 molecule of glucose to 2 mole-
cules of lactic acid is called anaerobic glycolysis. Two molecules of ATP are
utilized in the early stages of glycolysis - in the phosphorylation of glucose
and fructose 6-phosphate. At later stages 4 moles of ATP are formed, so that
there is a net yield of 2 moles of ATP for each mole of glucose utilized:
Glucose + 2 P + 2 ADP → 2 lactic acid + 2 ATP.
Anaerobic glycolysis can be reversed when oxygen becomes available.
If the oxygen supply is restored and the lactic acid disappears, a portion is
oxidized to CO2 and water and the reminder reconverted to glycogen.
Table V-3. Yield of ATP from the catabolism of carbohydrates
5.2.2. Oxidation of fat
The majority of body cells are able to oxidize and convert lipids and to use
fat as a source of energy. The major part of the energy derived from fat is
provided by fatty acids. Fatty acids are made to be available from dietary
lipids and from triacylglyceroles stored in the body when glucose is unable
to provide sufficient energy. The breakdown of lipids starts by the hydrolyt-
ic processes in the intestine, where the fatty acids are split off from their
esters with the glycerol part. The absorbed glycerol is oxidized like a carbo-
hydrate via glycolysis and the tricarboxylic acid cycle, in which process 1
mole of glycerol yields 21 moles of ATP. The fatty acids are broken down to
acetyl-CoA, and then to CO2 and H2O. Taking tripalmitin as a model; in the
cell 408 moles of ATP are formed from each mole of tripalmitin. The energy
stored per mole of ATP is 52 kJ, corresponding to 21216 kJ/mole tri-
palmitin oxidised in the body. The energy yield of palmitic acid in the bomb
calorimeter is 9981 kJ/mole, 1 mole glycerol yields 1655 kJ, hence the total
energy from 1 mole of tripalmitin (3×9981)+1655 = 31598 kJ. Consequently,
the oxidation efficiency is then 21216/31598 = 0.75. The remaining 25% is
64
heat loss.
In ruminant animals considerable amounts of propionic acid, butyric
acid and acetic acid are produced by the process of microbial breakdown of
carbohydrates in the rumen. The intermediary products of carbohydrate
breakdown in the rumen are oligosaccharides and simple sugars. These are
unstable for rumen microorganisms and, in majority, are broken down to
short chain fatty acids (SCFA) also called volatile fatty acids (VFA). First, the
monosaccharides are converted to pyruvate and subsequently to SCFA. The
conversion of sugar to pyruvate and the formation of SCFA generate ATP
which mainly serves as the energy source for microbial protein synthesis.
The conversion of 1 mole of hexose provides 4 moles of ATP, but for the
process is less efficient concerning penthoses contributing only1.67 moles
of ATP. SCFA produced in the rumen pass to the tissues where they partly
serve for fat synthesis and/or glucose and partly undergo oxidation. The
oxidation efficiency of SCFA is about 60% (Table V-4).
5.2.3. Oxidation of proteins
Feed proteins are hydrolysed to amino acids in the intestine, where they are
absorbed and used to build up proteins or are oxidized. Amino acids which
are not used for synthesis are deaminated resulting in ammonia liberation
and the conversion of the amino acids to the corresponding keto acids.
These can be used for glucose synthesis or lipogenesis and oxidized for
energy supply. The final products of amino acid degradation are acetyl-CoA
and, depending on the nature of amino acids, glucose (from glucogenic
amino acids) or ketone bodies (from ketogenic amino acids). The major
glucogenic amino acids are alanine, glutamate and valine, while leucine is
ketogenic. Other amino acids may be partly glucogenic and partly ketogenic.
Most of the ammonia arising from the degradation of amino acids is
excreted as urea. Urea formation requires energy; a total of 4 moles of ATP
is needed per 1 mole of urea formed. Further, about 1 mole of ATP is used
during the excretion of urea. Therefore, in assessing the efficiency of amino
acids oxidation, the energy required for urea synthesis must be considered
in the calculations. The high energy requirement for urea synthesis and
excretion may be one of the main reasons for the great heat output observed
after protein ingestion in the excess of the above mentioned heat require-
ment. Taking aspartate as a model; 16 moles of ATP are formed in the cell
from each mole of aspartate. The energy stored per mole of ATP is 52 kJ,
corresponding to 16 x 52 = 832 kJ/mole aspartate oxidized in the body. The
energy yield of aspartate in the bomb calorimeter is 1568 kJ/mole and con-
sequently the efficiency of the oxidation is 832/1568= 0.53.
Data in Table V-4 summarized the estimates of heat energy equivalents
of high energy phospate bonds formed during the oxidation of several major
energy sources.
65
TableV-4: The enthalpies of combustion (ΔHc), yield of ATP on the complete oxidation, the
energy required for formation of 1 mole ATP (ΔHc/ATP) and the efficiency of oxidation
5.3. Energy-requiring processes (anabolism)

Most of the energy transferred to ATP via the oxidative processes is later lib-
erated and used in the anabolic processes. Quantitatively the anabolism
mainly concerns the formation of carbohydrate (glycogen in the liver and the
muscles; lactose in milk), fats (tissue, milk, foetus, eggs) and protein (tissue,
enzymes, hormones, milk, foetus, eggs, and hair). Genetically the urge of
protein formation is quantitatively much more dominating than that of fat
formation. In young growing animals the body gain may comprise 25% pro-
tein, 70% water and 5% fat, which is mostly structural fat. As the animals
approach maturity the preference toward protein retention decreases while
fat accretion increases; in full grown animals body weight change may con-
sist of less than 5% protein and more than 80% fat. By fat synthesis often
too great an energy supply can be stored as a reserve for later mobilisation
in case of insufficient energy supply from feed. Apart from the long-term
regulation of energy supply by means of reserve fat accretion and mobiliza-
tion, there is a short-term regulation of energy supply. Directly after feed
intake the liver stores a large part of carbohydrate as glycogen, and later on
these stores are broken down to glucose.
Glucose is a substrate for the production of glycogen stores and lac-
tose. In ruminants storing and use of the glycogen are less developed than
66
in monogastric animals. Apart from absorbed glucose, glucose is synthe-

sized from gluconeogenic substances. Lactic acid, amino acids, glycerol, and
- in ruminants - propionic acid are the main gluconeogenic substances. In
lactating cows, 55% of the gluconeogenesis is derived from propionic acid,
20% from lactic acid, 15% from amino acids, and 10% from glycerol.
Glycogen is a complex polysaccharide made up of glucose and has the abil-
ity to add on further glucose units. It is a readily mobilizable storage form
of glucose. The formation of glucogen is a fairly cheap process. Totally,
slightly more than 1 mole of ATP per 1 mole of glucose 1-phosphate is used
for the storage in glycogen. Hence, 1 mole ATP is used to phosphorylate glu-
cose to 1 mole glucose 1-phosphate and one high-energy phosphate bond is
spent in incorporating glucose 6-phosphate into glycogen.
Fatty acids are synthesized in the body and those absorbed from the
intestine are involved in the esterification of fatty acids to triglycerides and
certain phosphatides in depot fat. There are two systems of fatty acid syn-
thesis. The first is a cytoplasmic system resulting in the production of
palmitic acid from acetyl-CoA. The second is a mitochondrial system result-
ing in the elongation of the existing fatty acids by two-carbon addition by
means of acetyl-CoA. Most unsaturated fatty acids are formed by the dehy-
dration of the corresponding saturated fatty acids, except the essential fatty
acids, which cannot be produced by the animal, but must be supplied in the
feed. Three moles of fatty acid and 1 mole of glycerol are used for the com-
plete synthesis of a triglyceride. The glycerol part is formed from glucose by
the reduction of dihydroxyacetone phosphate to glycerol.
Proteins are synthesized from amino acids which become available
either as the end products of digestion or as a result of synthetic processes
within the body. In the first step of the protein synthesis the amino acids
are linked to form peptides in a genetically determined order. Then the pep-
tides are polymerised and folded to protein by means of different linkages.
During all stages of protein synthesis energy is provided by hydrolysis of
ATP and guanosine triphosphate (GTP). The total mole number of ATP and
GTP needed to synthesize one peptide bond is 4. The efficiency of 87% (Table
V-5) is about 30% higher than the measured energy required for protein
retention. The main cause of the large difference between the energy
requirement for synthesis and energy requirement for retention is the pro-
tein turnover (synthesis/degradation). Isotope studies have revealed that
the protein synthesis/retention ratio is not constant. Depending on the age
of the animals, 2-5 times more protein is synthesized than is being stored.
Assuming that the average synthesized/retained protein ratio is 3/1, the
energy efficiency for protein accretion will be only 69% as opposed 87%.
Data in the Table V-5 summarize the estimates of theoretical biochemical
efficiencies for protein deposition, fat and carbohydrate are calculated as
the combustion enthalpy of the product divided by the combustion enthalpy
of the substrate + the energy from ATP synthesis.
67
Table V-5: Nutrient energy employed by theoretical stoichiometric efficiencies in synthesis
The outline of energy providing and energy consuming processes is

demonstrated in FIGURE V-2. The energy in the body is provided by the oxi-
dation processes of carbohydrates, fats and proteins. The oxidation process-
es in the body break down the nutrients to carbon dioxide and water, while
some part of the chemical energy is transferred to the energy-rich phos-
phorus compounds. The remaining part of the energy is used to maintain
constant body temperature and is given off as heat. The greater part of the
energy derived from the energy-rich compounds is liberated for the synthe-
sis of body tissues, while the rest is used for the physical processes such as
activity, feed intake and transport, nutrient absorption from the intestinal
tract, blood circulation, fluid balance maintenance, ion transport etc. and is
given off as heat.
68
FIGURE V-2: Energy-producing and energy-requiring processes in the body
5.4. Methods for the quantitative determination of heat production
The metabolic processes responsible for the energy supply in the body can
be determined by measuring the heat production of the animal. The energy
quantity in the oxidative processes of carbohydrates, fats, proteins and
short chain fatty acids can be measured as heat. In the previous subchap-
ter we calculated that the animal utilizes about 70% of carbohydrate and fat
energy, and 40% of protein energy for its energy-requiring processes, while
the rest is lost as heat. Part of the energy is transferred to the energy-rich
phosphorus compounds and the later hydrolysed energy will also, sooner or
later, be converted to heat. This means that all the metabolic processes can
be quantified on the basis of heat production in the animal.
Heat production can be measured by means of various calorimetric
methods, either directly in a calorimeter or indirectly by measuring the gas
exchange from the animal. The most common indirect method is based on
69
gas exchange (O2-consumption and CO2-CH4-production) measurements

combined with nitrogen excretion measurements in urine (RQ-method) or
combined with carbon-nitrogen balances (C-N-method) measurements. The
results of respiratory exchange and heat production in different species
obtained from gas exchange measurements are shown in Table V-6.
Table V-6: Examples of respiratory exchange and heat production in different species
Methods for the estimation of heat production and retention of ener-

gy described above require the use of a respiration chamber. But attempts
have been made to measure energy retention in a simpler way, based on the
fact that the weight gain of an animal is the result of protein, fat and min-
erals retention. The day-to-day protein and ash retention can be determined
by balance trials. If the weight of water in the living animal can be meas-
ured, the weight of stored fat can be estimated by subtracting the lean body
mass. In practice, total body water can be estimated by the so-called ‘dilu-
tion’ techniques, in which a known quantity of a tracer substance is inject-
ed into the animal, allowed to equilibrate with the body water, and then its
equilibrium concentration determined.
Direct determination of muscle or body protein using body potassium measurement, crea-
tinine excretion, neutron activation analyses and 51Cr-labelled red blood cells has also
been evaluated in body composition studies. The most extensive research has been attrib-
uted to potassium either by using whole body counting (40K) or isotope dilution (42K). Both
techniques are based on the presence of potassium as the major intracellular cation.
Computer tomography and nuclear magnetic resonance (NMR) have been used to provide
cross-sectional images of animals. Although, until now both methods are expensive they
have several application in studies for body composition in live animals. In growing ani-
mals, to obtain better insight in the composition of the live weight gain, the comparative
70
slaughter method is sometimes used (For more details see Chapter III).
The doubly-labelled water (DLW) method uses the principles of indirect calorimetry to meas-
ure total heat production (energy expenditure) from the turnover rates of two stable iso-
topes: deuterium and oxygen 18. Labelling total body water also provides estimates of body
composition and measurements of water outflow rates. It is the first genuinely non-inva-
sive method for measuring energy expenditure in free-living species, providing estimates of
habitual expenditure over a time period of 10-20 days. In principle the original experimen-
tal technique for humans was very simple: a subject merely drank a dose of DLW, and 2
urine samples were collected about 2 weeks apart. However, the DLW method has suffered
from excessive optimism. The fundamental observation making the method possible is that
the oxygen of expired CO2 is in isotopic equilibrium with the oxygen of body water. A
human or animal loaded with 18O-enriched water displays a labelling of expired CO2. This
reaction is catalysed by the enzyme carbonic anhydrase. Hence, in a subject loaded with
isotopically labelled water, the disappearance of 2H from body reflects water output, where-
as the disappearance of 18O represents water and CO2 outputs. The technique involves
enriching the body water of a subject with an isotope of hydrogen and an isotope of oxy-
gen, and then determining the washout kinetics of both isotopes as their concentrations
decline exponentially toward natural abundance levels.
5.5. Partition of energy in the body
The main phenomenon in animal life processes is the redistribution of ener-

gy. The process starts with the ingestion of feed containing organic and
inorganic compounds. Subsequently a varying proportion of the organic
compounds in the ingested feed are hydrolyzed and the products of hydrol-
ysis are subject to either absorption or further microbial degradation. Not
all the feed can be digested and absorbed; the residue is excreted in the fae-
ces. Urea, uric acid, allantoin, purines and other detoxification compounds
have to be excreted with urine. Furthermore caused by the fermentation
processes in the forestomachs and large intestine energy in gaseous form
(CH4, H2) is lost. Then all catabolic and anabolic processes are associated
with energy losses as heat. Therefore when describing the energy metabo-
lism in animals different energy forms are distinguished:
Gross energy (GE) is the amount of energy manifested as heat when

the feed is completely oxidised. GE content of feed is measured in an appa-
ratus known as a bomb calorimeter which consists of a combustion cham-
ber (the calorimetric bomb) resting in an insulated tank of water. A dried,
weighed feed sample is placed in a platinum cup in the bomb. The bomb is
closed hermetically, and 25-30 atmospheres of oxygen are admitted under
pressure, after which the nickel wire is ignited electrically, and the feed
sample burns. Owing to the high oxygen pressure, the combustion process
is explosive and complete. The quantity of heat produced, calculated from
the rise of water temperature, corresponds to the gross energy of the feed.
The content of energy in faeces, urine and in animal products like milk or
eggs is determined in the bomb calorimeter as well. Gross energy of differ-
ent materials varies, but typical values are for proteins, carbohydrates and
fats, 23.86, 17.56 and 39.76 kJ/g, respectively. The differences between
71
these nutrients primarily reflect the C: H ratio and the O and N contents.
For example glucose, C6H12O6, has 1 atom of oxygen per atom of carbon
whereas a fat molecule, for example glycerol trioleate, C57H104O6, has 6
atoms of oxygen per 57 atoms of carbon. Thus fat requires more oxygen for
oxidation and gives off more heat in the process.
Digestible energy (DE) includes not only energy absorbed by the body, but
energy of fermentation and energy of gaseous products (such as methane).
DE is the proportion of the gross energy of a feed which is apparently digest-
ed. DE = GE - FE; gross energy minus energy in faeces (FE) measured in a
bomb calorimeter. Faecal energy includes undigested feed and metabolic
products (intestinal cells, bacteria, and enzymes). DE is determined by
digestibility trials performed according to the same principles as for the
determination of the digestibility quotient of the nutrients.
Metabolizable energy (ME) is the amount of energy available to the animal

for body functions (maintenance, growth, production of milk and eggs, etc.).
Metabolizable energy represents the energy actually capable of transforma-
tion to other forms of energy in the body, regardless of type of transforma-
tion, and whether the transformations are of any use to the animal. ME
includes not only energy available for maintenance and production, but also
heat produced during nutrient metabolism and fermentation. ME is usual-
ly 90-98% of DE for most mammals, however, ruminants have ME coeffi-
cients of 80-90% of DE, and when forages contain high levels of secondary
plant compounds, the value may drop to 73%, such as for marsupials eat-
ing eucalyptus. ME = DE - UE - CH4E; digestible energy less urinary ener-
gy (UE) measured in a bomb calorimeter, and less methane energy (CH4E),
measured by respiration trials. Urinary energy losses originate from waste
products of imperfect nutrient metabolism and endogenous catabolism.
These losses can be substantial for animals consuming plants with a high
level of secondary plant compounds because many of these compounds
cannot be used for energy and must be excreted in the urine. Urinary ener-
gy also includes waste nitrogen (such as urea or uric acid), which increas-
es when energy intake is lower than required for maintenance and the ani-
mal begins to break down muscle (protein) for energy. ME can be measured
according to the same principle as digestible energy, but digestibility trials
have to be supplemented with quantitative collection of urine.
Determination of ME in ruminants involves measurement of the methane
produced by the decomposition of carbohydrates in the forestomachs. It
may be assumed that a full-grown cow on a reasonably large feed ration
produces 500 litres of CH4/day, which is equivalent to 500 l x 39.6 kJ =
19.8 MJ, or 10% of the gross energy. In monogastric animals, however, the
methane production is much less; in pigs, for examples, the methane ener-
gy is only 1-2% of the gross energy.
72
Heat energy (HE), also called heat production or energy expenditure,

is the energy produced by the total metabolism in the body. It is a sum of
energy derived from oxidation of carbohydrate, fat and protein, and fermen-
tation processes. Total heat energy includes heat produced by maintenance
processes (HEm) and the so-called heat increment (HI). Heat increment (HI),
also called dietary induced thermogenesis is the heat produced by the
metabolism of feed and the heat produced by the conversion of the metab-
olized nutrients into animal products, e.g. protein and fat retained in the
body, eggs, milk etc. Heat increment the energy converted to heat due to
inefficiencies in metabolic reactions. This energy is either lost as heat, or
can be counted as maintenance energy if the animal must expend energy to
maintain body temperature (i.e. a homeotherm in a cold environment).
The magnitude of the heat increment depends on the amount of feed
ingested and the composition of the diet. HI increases with increasing ener-
gy supply (amount of feed), because the transport and digestion of feed in
the alimentary tract requires energy. For example, HI constitutes about 20%
of the supply of 30 MJ metabolisable energy in cattle, whereas when the ME
supply is 90 MJ, the HI is about 30%. The heat increment also varies from
feedstuff to feedstuff, depending on structural differences, mainly owing to
varying fibre contents. For concentrate feeds HI ranges from 15-25% of ME,
whereas the range is 35-60% for roughages.
All body functions result in a certain heat production, corresponding
to the amount of energy used in the metabolic processes. This heat pro-
duction forms part of the heat increment and depends on the extent and
type of the anabolic processes. In case of fat retention, for example, the ME
is utilized up to 75%, i.e. 75% of ME available for fat retention is transferred
to the energy retention in fat, while the remaining part is included in the
total HI and is lost as part of the heat production. Similarly, the utilization
of energy in protein retention is about 50%, that is, 50% is lost as heat.
Retained energy (RE) is the amount of energy retained in the body

and/or in milk and eggs (so called energy balance). RE=ME-HE; metaboliz-
able energy less heat production. RE in the body or in eggs is mainly ener-
gy retained in protein and fat, while in milk a substantial amount of energy
is also retained in carbohydrate.
Net energy (NE) is the energy of a feed which is available to the ani-
mal for different life processes; NE=GE-(FE+UE+CH4E+HI). The deduction of
the heat increment of a feed from its metabolizable energy gives the net
value of a feed. The net energy represents the quantity of energy which is
equivalent to the body function caused by the feed concerned; one kJ net
energy of a feed is equivalent to one kJ net energy of a body function,
regardless of whether the net energy of the body function is found in a
retained substance or is released as heat caused by the maintenance
73
processes. The net energy value of the same feedstuff may differ according
to the species of animal by which it is consumed and the purpose for which
it is used. The total NE can be partitioned between net energy of retained
substances, e.g. the sum of energy stored in protein, fat and carbohydrate,
and NE associated with maintenance processes, e.g. the amount of heat
produced by maintenance processes. The partition of gross energy from the
feed into digestible, metabolizable and net energy is summarized in FIGURE
V-3.
FIGURE V- 3: Partition of the gross energy in the body
74
The following diagrams (FIGURES V-4, V-5 and V-6) illustrate the partition
of gross energy from feed in the animal body. The presented values are
examples which are attributed to a substantial variation depending on the
feeding level, feed composition and structure, animal production level, man-
agement, environmental conditions and animal genotype.
FIGURE V-4: Energy metabolism of a dairy cow producing 25 kg milk of 4% fat
FIGURE V-5: Energy metabolism of a growing pig of 50-60 kg of live weight
FIGURE V-6: Energy metabolism of laying hen of 80% egg production stage
75
5.6. Energy and protein utilization and requirements

Nutrients and energy are required for a number of different functions in the
body, both for the vital processes necessary for an animal to survive and for
the productive processes such as reproduction, growth, milk production,
and egg production, growth of wool and fur, and work. In attempting to esti-
mate requirements one must ask three questions:
1. requirements of what kind of animal or animal population ?
2. requirements for what purpose ?
3. how should requirements be specified ?
The first question arises from the great differences among animals regard-
ing their own requirements. Animals of different age, live weight, sex and
production level have different requirements. Furthermore, even when these
differences are excluded, each individual animal still differs in their own
requirements due to genetic constitution which is rather difficult to quanti-
fy. It is often observed in animal experiments that even individuals of the
same age, sex, belonging to the same genetic line and being kept in similar
environment show in-between individual coefficient of variation up to 15-
20% for many nutritional parameters. The second question relates to the
criteria we choose to define requirements. For example, whether it is defined
in terms of animal performance as body gain and feed conversion for gain
or in terms of metabolic responses? The third question requires specifying
terms: whether there are requirements for metabolisable energy or for net
energy, or for digestible protein or for amino acids, or what kind of combi-
nation of energy and protein is preferred?
Although carbohydrate and fats are necessary for a number of phys-
iological functions their main importance is the energy supply. Dietary pro-
teins are mainly used for the synthesis of body proteins but they also con-
tribute to the energy supply. Therefore, in the following description, the
emphasis is put on animals’ requirements for energy and protein for main-
tenance and growth, while the specific use of carbohydrates, fats as well as
vitamins and minerals is described in Chapters XI and XII).
5.6.1. Requirements for maintenance
Knowledge of the energy requirement for maintenance is of considerable sig-

nificance to the evaluation of the energy utilization for different life process-
es like reproduction, growth, lactation, work etc. The energy requirement for
maintenance can be defined as the minimum energy needed by the resting
animal kept in a thermoneutral zone to maintain constant body tempera-
ture and to maintain a dynamic equilibrium for protein and fat turnover. It
corresponds to the amount of energy required by the animal to keep it in
energy equilibrium; it neither stores nor loses body energy, resulting in zero
energy balance. It is the amount of energy corresponding to the heat pro-
duction of the animal when both nitrogen and carbon balance are zero. This
76
definition is acceptable for mature, non-productive animals, but it is diffi-

cult to use for growing or productive animals, in which the energy retention
changes in relation to the stage of growth or production level. The energy
balance will always be above zero in growing or producing animals caused
by retained protein and fat in the body or in the products. In such cases,
the maintenance requirement must be regarded as a theoretical value.
However, this value is necessary in many calculations for the energetic effi-
ciency estimation of energy utilization for growth and other processes.
Energy requirements for maintenance are usually expressed in
terms of either metabolisable energy (MEm) or net energy (NEm) depending
on the feed evaluation system used (see Chapter IX). A number of different
experimental methods are available for the determination of maintenance
requirement from trials with fasting or fed animals. In fasting animals,
relaxed in a thermoneutral environment, heat production decreases in the
course of a few days to a fairly constant level. This lowest and constant level
of heat production is taken as a measure of the energy required to maintain
the vital metabolic processes of the animal. Under the above conditions the
energy is supplied by the breakdown of body tissues. Consequently, the
heat production provides an estimate of the lowest possible turnover in the
animal; this is termed as fasting metabolism or basal metabolism.
The most important conditions that have to be fulfilled for determi-
nation of fasting metabolism are: 1) the animal should be completely
relaxed, since muscular activity increases heat production. 2) The ambient
temperature should be such that it neither increases nor decreases heat
production in the animal (thermoneutral zone). 3) The animal should be in
a good condition at the onset of the experiment. 4) The period of fast should
be sufficiently long for the animal to reach the postabsorptive state, that is,
the digestion and metabolism of the previous meals should be terminated in
order to avoid the complicating effect of the heat increment by food.
Depending on the species a fasting period of 2-4 days would be need-
ed for monogastric animals. However, in ruminants it is futile to estimate
the basal metabolism by fasting experiments. Firstly, because nutrient
digestion and metabolism continue for several days after feeding ceases.
Secondly, the postabsorptive state causes cessation of microbial activity in
the rumen, which physiologically is an abnormal condition. An approximate
value for minimum metabolism in ruminants is sometimes estimated by
measuring the heat production after long enough time has elapsed to
ensure that the metabolism in the gastrointestinal tract is very low, that is
when the methane excretion has reached a low and constant value, and/or
when the respiratory quotient (RQ) has decreased to 0.7, which indicates
that energy is being obtained from body fat reserves.
Since the middle of the XIXth century it has been known that fast-
ing heat production is related to body size. RUBNER developed the concept
that the heat given off by all warm-blooded animals in the late fasting peri-
77
od is directly proportional to their body surface, referred to as the “Surface

law.” In view of the difficulties and uncertainties involved in measuring sur-
face area, formulas were devised for computing it from body weight, recog-
nizing that surface was proportional to some fractional power of weight,
between 0.5-0.9. On the basis of a series of results of estimates of energy
metabolism in animals ranging in size from mice to elephants, BRODY found
that fasting heat production (HEf) is a function of live weight (LW) raised to
the 0.73 power, since log HEf, kcal/day = 0.73 × log LW, kg + 1.845, which
is convertible to a power function: HEf, kcal/d = 70.5×LW, kg0.73. Based on
BRODY’s principles, KLEIBER (1947) proposed a definition of the metabolic live
weight as kg LW raised to the 3/4 power to be used as a unit of reference
when comparing basal-metabolism data of mature animals of different
species, size and live weight. It is now generally accepted to use the expo-
nent 0.75 both for growing and mature animals in order to facilitate com-
parison between different species, breeds and individual animals. Recently
other exponents have been introduced, i.e. the Agricultural Research
Council recommends 0.63 for pigs.
The energy requirement based on the heat production of a fasting ani-
mal is in principle equivalent to net energy for maintenance, HEf = NEm. This
equation may be translated to the requirement for the metabolisable energy
(MEm) if the coefficient of utilization (km) of the ME for the processes of main-
tenance, MEm = HEf/km, is known (km is often indicated as 0.8, correspon-
ding to 80% utilization of ME for maintenance). When the basal metabolism
of mature animals was stated as 70.5 kcal × LW, kg0.73, it was supposed to
be a theoretical value for net energy being about 300 kJ/kg0.75, or for
metabolizable energy 375 kJ/kg0.75 (since 300/0.8 = 375). However, the
values of fasting heat production are average values, and there are sub-
stantial differences between species, ranging from 340 (growing rat) to 600
(growing pig) kJ/kg0.75 (CHWALIBOG et al. 2005), as well as there is a con-
siderable genetic variation within species.
Fasting experiments have also been used to estimate the mainte-
nance requirement in growing animals. For example, in measurements with
growing rats fasting heat production was 340 kJ/kg0.75. In chickens a
mean HEf was 370 kJ/kg0.75, equivalent to a maintenance requirement of
metabolisable energy of 460 kJ/kg0.75. However, in growing pigs in relation
to metabolic weight fasting heat production was 50% higher in pigs weigh-
ing 20-30 kg than in pigs of 50-100 kg, and the following formula from 20
to 100 kg LW has been suggested: HEf, kJ = 3235 + 167 × LW 0.75. On con-
dition that the efficiency of utilization of MEm is 80% (km = 0.8), the main-
tenance requirement for metabolisable energy could be calculated to be
MEm, kJ = 4060 + 210 × LW, kg0.75.
The results of fasting metabolism estimates are considerably lower
(20-30 %) than the values found in feeding trials. The reason is that the ani-
mals’ energy requirement for maintenance decreases during fasting, partly
78
because of reduced muscular activity and partly because of changes in fat,

protein and carbohydrate metabolism. It is, therefore, difficult to translate
values from fasting metabolism in to maintenance requirements of fed ani-
mals. The other method to estimate maintenance requirement is feeding ani-
mals at different feeding levels. In these trials different feeding levels are
used, and if the estimated energy retention (RE) is related to the metabolic
live weight of the animal (RE/kg0.75), it will often show a linear dependence
on the ME supplied (ME/kg0.75), as shown in FIGURE V-7. The linear func-
tion may be described in a regression equation RE/W0.75=-a+b× ME/W0.75,
where the interception of the curve and the x axis, calculated by -a/b, indi-
cates the energy requirement for maintenance, while the coefficient b of the
regression is a measure of utilization of ME for energy retention.
FIGURE V-7: Model for determination of the energy requirement for maintenance
(MEm/W0.75), based on regression of RE/W0.75 on ME/W0.75
The validity of the results from the one-dimensional regression for

the determination of maintenance requirement by feeding trials with differ-
ent feeding levels is subject to certain conditions. The calculations demand
a sufficiently high number of measurements both of the high and the low
energy retention. Since energy retention is the sum of energy retained in
protein (RP) and in fat (RF), the RP/RF ratio should be constant and inde-
pendent of age (live weight) and feeding level. When the measurements
79
relate to growing animals, the constant ratio between protein and fat reten-
tion can only be obtained if the measurements are made at narrow weight
intervals. But it is very difficult, if not impossible, to obtain the same RP/RF
at different feeding levels in growing animals, because they try to maintain
the greatest possible protein retention at the cost of fat storage. Therefore,
multiple regressions are often used for simultaneous determination of
maintenance requirement and of energy utilization for protein and fat stor-
age, as demonstrated above. Results of Danish experiments on mainte-
nance requirements determined by fasting and feeding trials are shown in
Table V-7.
Table V-7: Danish results concerning energy requirements for maintenance

(from balance and respiration experiments)
In a similar way protein requirements can either be estimated from

fasting or from feeding trials. With the first method it is assumed that in
theory endogenous N excretion is equivalent to N requirement for mainte-
nance. During the fasting period, N-excretion in faeces is negligible,
whereas considerable amounts of N are excreted in the urine. During the
first days of fast the urinary N loss decreases until the nitrogen originat-
ing from the previous N supply has been excreted, after that the N excre-
tion will be fairly constant. The urinary nitrogen excreted by an animal
deprived of feed nitrogen at this minimum level is known as the endoge-
nous urinary nitrogen. BRODY and associates also found that the amount
of endogenous urinary nitrogen is proportional to the basal metabolism,
which means that endogenous urinary nitrogen may be expressed as a
power function of the live weight of the animal: Endogenous Nurine, g/d =
0.146 × LW, kg0.72. It appears that the exponent in this formula is practi-
cally the same as for the energy metabolism, and the formula might as
well be: Endogenous Nurine, g/d = 0.15 x× LW, kg0.75. For example, the
maintenance requirement of a cow weighing 500 kg with a DM intake of
10 kg/d, will be 320 g digest. crude protein/day. Assuming a protein
80
digestibility of 75%, we get 320/0.75 = 426 g crude protein. It has to be

kept in mind that the above calculations for cattle are based on several
assumptions which may be critical. As mentioned before the endogenous
N excretion can be esteemed by feeding of N-free diets and measuring uri-
nary and faecal nitrogen excretion. This can be done over a short period in
monogastric animals, however, when a protein-free diet is given to rumi-
nants the micro flora is deprived of NH3, fermentation is ceased and as a
consequence feed intake decreases. When feed intake is decreased below
maintenance energy requirement, the N excretion no longer represents tis-
sue maintenance need.
Estimations based on fasting trials and N-free diets are unsuitable for
growing animals, and the results can probably not be used under practical
feeding conditions. Therefore, feeding trials involving growing animals are
conducted according to the same principles as determinations of energy
requirement for maintenance. Different N-levels are given, and the retained
nitrogen (RN) is related to the digested nitrogen (DN), both expressed per kg
metabolic weight. In order to obtain linear dependency, the calculations are
made at narrow weight intervals. The point of intersection of the curve with
the x-axis indicates the quantity of digestible nitrogen required by the ani-
mals to ensure N equilibrium (RN = 0). Consequently, regression calculation
may be done according to the following model: RN, mg/LW, kg0.75 = -a + b
× DN, mg/kg0.75, where -a/b expresses the animal’s nitrogen requirment for
maintenance (Dam/LW, kg0.75) within a defined weight range. By means of
the described method THORBEK et al. (1984) found that for pigs weighing 20
- 120 kg the daily N requirement for maintenance can be expressed by the
equation: DNm, g/d = 2.69 + 0.160 × LW, kg0.75. The calculation presup-
poses a sufficient intake of lysine and methionine + cystine. In Table V-8 the
protein requirement for maintenance for growing pigs is compared to the
calculated requirement based on fasting trials and N-free feed and the equa-
tion suggested by THORBEK. It will appear from this table that the require-
ment calculated from fasting trials is underestimated for young pigs and
overestimated for older pigs as compared to the calculations based on the
equation.
Table 8: Maintenance requirement for digestible protein (g/d) in growing pigs
81
However, the method of extrapolation of endogenous N excretion from dif-

ferent levels of protein intake, when applied to cattle, may be questionable
to a high net uptake and recycling of urea with low protein diets. The main-
tenance requirement for an amino acid is usually taken to mean the amount
that must be supplied in the diet in order to maintain constant the body
nitrogen content. Such estimates are usually made by giving diets specially
limiting in one amino acid at a range of intakes supporting nitrogen reten-
tion close to zero.
5.6.2. Requirements for growth
It should be kept in mind that there is more to growth than a change in live
weight. The specific development of the animal should also be considered,
weight gain says nothing about that. The pattern of live weight growth
changes with the development of the animal. We distinguish between three
stages of growth: a) development of bones and organs (heart, liver, kidney
etc.); b) development of muscles and c) fat retention. As demonstrated below
in first stage the growth and development of bones dominates over other tis-
sues, in the second stage the growth of muscles and in the third stage the
growth of fat tissue are dominant. Finally, when an animal is mature the
only tissue which may develop is the fatty tissue.
In contrary to the other tissues the fatty tissue is synthesised dur-

ing the whole life. However, during the growth period the visceral, subcuta-
neous and intermuscular fat are synthesised prior to intramuscular fat, the
latterbeing the only fat retained in the adipose tissue of adult animals. The
above grouping is not changed, but the length of the individual periods
varies depending on the feeding intensity. If a varying fattening level is left
out of account, attention should be given to the influence of the nutrient
supply, when the animal - depending on the quantity and quality of a
ration, may attain the adult weight in the course of a shorter or longer time.
A characteristic feature is that the chemical composition of the animal body
changes considerably during the growth period, in the way that the water
percentage decreases, the fat percentage increases, while the protein and
82
ash percentages remain fairly constant. For example a newborn calf con-
tains on average 74% water (in some cases, even 80% water). This means
that water alone weighs about 30 kg in a newborn calf, and only 10 kg con-
sists of dry matter. The water percentage decreases with the growth of the
animal, and an adult cow in medium condition may be assumed to contain
60% water. Fattening decreases the water content further to about 50%,
and in cases of extreme fattening the water content has been found to be as
low as 40%, i.e. the water percentage is halved as compared with that of the
newborn animal. Water is present in all body tissues, but it is far from even-
ly distributed. Meat contains 72-78% water, whereas bone contains only
about 45%.
Since the proteins are present in all cells in the entire body, it is nat-
ural that the protein percentage remains practically unchanged, about 15-
18%, from birth to maturity. This constant protein percentage in the body
does not mean that the retained protein values in the growing animal are
also constant, on the contrary, the capacity to retain protein (to build up
meat) changes in the course of the growth period, so that young growing
animals retain increasing amounts of protein up to a certain age, when the
retention decreases. Carbohydrates only make up about 1% of the animal
body, and the level is fairly constant during the growth period. The growth
processes do not only include synthesis and product retention. A certain
percentage of the synthesized components are degraded again, e.g. about
2/3 or more of the protein pool is broken down. This means that only 1/3
or less of synthesised protein will be retained. Therefore, in young growing
animals the growth rate is determined by the difference between synthesis
and degradation of musculature, fatty tissue, bones etc. The partitioning
(distribution) of substrates in growing animals is summarized in FIGURE V-
8.
FIGURE V-8: Substrate distribution in growing animals
For example, in a newborn pig protein synthesis constitutes about

15% of the muscular mass/day. But since the protein degradation at the
83
same time amounts to about 9%, the actual increase of the muscular mass
is about 6% per day. In other words, the daily protein retention is 6/15 =
40% of the synthesis. But in a pig weighing about 150 kg the protein reten-
tion has been reduced to 15% of the protein synthesis. Consequently, in
adult animals synthesis and degradation of protein will be equal, so reten-
tion will be zero.
Calculation of the energy and protein requirements for growth is
based on the amounts of protein and energy retained daily. These values can
be estimated either by means of balance experiments or from slaughter
experiments, where the animals are killed and analyzed at different ages
and weights, or through metabolic measurements. When the values for
nutrient and energy utilization for growth are known, the requirements can
be estimated.
Since most of the energy evaluation systems used today are based
on metabolizable energy, the following description will primarily deal with
metabolisable energy requirements. As shown in FIGURE V-9 distinction is
made between ME for maintenance (MEm) and ME for growth (MEg). ME for
growth may further be divided into ME for protein retention (MEp) and ME
for fat retention (MEf). In growing animals the retained energy is stored in
protein and fat. If the energy retained in protein is termed RP (retained pro-
tein energy) and the energy retained in fat RF (retained fat energy), and the
total energy retained is RE = RP + RF, the coefficient of utilization of ME for
RP (MEp) and RF (MEf) may be expressed as kp = RP/MEp, and kf = RF/MEf
or by kg = RE/MEg. In order to evaluate the utilization of energy for growth
and other productions we have to know the energy requirement for mainte-
nance, since MEg = ME (total) - MEm. As discussed before results of fasting
trials with growing animals may be inadequate. Therefore, the maintenance
requirement is often estimated by trials using different feeding levels. Such
a method does not only give a value for maintenance, but also a value for
the utilization of energy.
84
FIGURE V-9: The partition of metabolizable energy into ME for maintenance (MEm), growth
(MEg) and energy retention (RP), fat (RF) and total energy retention (RE). Efficacy of ME for
RP (kp), RF (kf) and RE (kg)
The total coefficient of utilization for growth (kg) can be determined

by a one-dimensional regression (FIGURE V-9). The partial efficiencies of
ME utilization for retention of protein (kp) and fat (kf) can be calculated by
means of multiple regressions. A multiple regression model includes energy
requirement for maintenance as well as utilization of energy for protein and
fat retention. Generally, the model may be described by the following equa-
tion:
ME = b1 × LW, kg0.75 + b2 × RP + b3 × RF
where b1 × LW, kg0.75 is the maintenance requirement for ME, 1/b2 = kp
and 1/b3 = kf. It is generally agreed that the efficiency of ME for fat ener-
gy retention is about 75%, whereas the energetic efficiency of protein ener-
gy retention varies considerably, from 40% in cattle to 60-70% in pigs and
chickens. The coefficient of utilization of ME for energy retention in fat of
75% means that for each kJ retained in fat 1.3 kJ ME has to be available
85
for retention (MEf), or each gram stored fat requires 52 kJ MEf (1.3×39.8,
since 1 g fat contains 39.8 kJ). For the energy retention in protein the uti-
lization of 60% corresponds to a requirement of 1.7 MEp per kJ stored in
protein, or 40 kJ MEp per g stored protein (1.7×23.9, since 1 g protein con-
tains 23.9 kJ).
Procedure for estimation of the energy requirement for maintenance
and growth is exemplified below for pigs from 20 to 100 kg LW. The daily
protein retention is calculated from nitrogen balance, while fat retention
from carbon and nitrogen balances. Assuming the coefficient of utilization
of ME for energy retention in protein is kp = 0.68, whereas for energy reten-
tion in fat is kf = 0.75, we can calculate that 1/0.68 = 1.5 kJ MEp is required
for each kJ retained protein, and 1/0.75 = 1.3 MEf is required for each kJ
stored fat, which is equivalent to 36 kJ MEp per gram protein and 52 kJ MEf
per gram fat. Consequently the energy requirements for retention shown in
Table V-9 are calculated by multiplying by 36 and 52 for protein and fat,
respectively. The requirement for metabolizable energy for maintenance is
calculated from the equation: MEm, kJ/d = 3140 + 360 LW, kg0.75 The cal-
culated values for ME for maintenance and growth were converted to the net
energy based feed units for growing pig (FUgp) by dividing by 12.5, because
the mean content per FUgp was 12.5 MJ ME (see Chapter IX). For compar-
ison, the Danish allowance (at a daily weight gain of above 750 g) was also
included in the Table V-9.
Table V-9: Calculation of the energy requirement in growing pigs
The estimation of protein requirements of growing animals requires

knowledge about the animals’ ability to retain protein and the efficiency by
which the feed protein can be utilised. The efficiency by which feed protein
can be utilized for protein retention depends on a variety of factors such as
digestibility, amino acid composition and energy supply. The amount of pro-
tein an animal is able to retain during growth will depend on its physiolog-
ical condition. This means that it is not possible to force protein retention
86
beyond a certain maximum, regardless of the amount of protein in the feed.

If the animal is given more protein than it is able to store, the excess
nitrogenous part will be excreted in the urine and thus wasted.
Since the protein requirement for growth can be defined by the
amount of protein retained during growth, determination of maximum pro-
tein retention is used as a basis for estimation of the protein requirement.
For many years it has been a subject of discussion whether the maximum
protein retention in growing animals is a function of the protein and ener-
gy supply, or a function of the ability of the cells to form protein. The two
points of view can be squared by assuming that during the first part of the
growth period the cells have such a high capacity to produce protein that
maximum retention cannot be attained, so the retention becomes a function
of the energy and protein supply. As the capacity of the cells to produce pro-
tein declines with age, there will be a time when the animal can get suffi-
cient energy and proteins through the feed, and maximum protein retention
can be obtained, which will be a function of the age of the animals (live
weight). Therefore, maximum protein retention in growing animals can, in
principle, be determined as a function of the animals’ age/weight under
such conditions when an increasing supply of proteins, amino acids and
energy is unable to increase the protein retention.
Generally, there are two major approaches to describe the pattern of
protein retention (RP) in growing animals. The linear increment up to a certain
plateau, being kept constant over a period and then declining until maturity,
has been suggested from several experiments. A second approach is based
on a non-linear relationship between RP and live weight of growing animals.
Different allometric equations have been proposed to describe the pattern of
RP and to predict the maximum rate of RP during growth. Quadratic functions
of RP in relation to LW, to metabolic body mass or in relation to metabolisable
energy have been used. Also non-linear relationships are commonly used in
the mathematical models of growth to predict the potential RP.
By means of nitrogen balance experiment it is possible to estimate the quantity of
protein the animal should get during its growth period to obtain maximum protein reten-
tion corresponding to maximum meat production without undue waste of proteins. The
procedure for growing pigs is exemplified below. From nitrogen balance measurements
(CHWALIBOG et al., 1996) with Danish Landrace during the growth period from 2 to 120 kg
LW the following relation between retained protein and live weight was obtained: RP, g/d =
11.55 × LW, kg0.75 - 0.185 LW, kg1.50. The equation has been used to demonstrate the
pattern of the RP curve during the period of growth, showing zero retention at conception,
a summit of 180 g/d at 98 kg LW and zero at 250 kg LW. The individual values of digest-
ed protein (DP) and RP measured in these experiments yield an average efficiency of pro-
tein utilization, RP/DP = 0.60. Then the requirement of digested protein to obtain maxi-
mum RP during the growth period from 2 to 120 kg LW was calculated by using the func-
tion for maximum RP and the efficiency of 60 % (Table V-10).
87
Table V-10: Digestible protein (DP) requirement for maximum protein retention (RP)
The above values can hardly be used as a general requirement for all growing pigs,
because there may be considerable variations between breeds, sex and hereditary develop-
ment regarding the ability of the animals to store protein, besides a number of external fac-
tors, such as composition of the feed, feeding level, feeding technology, housing, climate
etc.
Dietary requirements for protein are actually needs for the amino acids con-
tained in the dietary protein. Amino acids are used by animals to fulfil a
diversity of functions. For example amino acids, as proteins, are primarily
constituents of structural and productive tissues, such as skin, hair, feath-
ers, bone matrix, and ligaments, as well as of the soft tissue, including
organs and muscles. Digested amino acids and peptides also serve a variety
of metabolic function and are precursors of several non-protein substances
in the body. Because body proteins are in a continuous dynamic state of
synthesis and degradation (protein turnover), an adequate intake is
required. If the supply of protein/amino acids is below requirements there
is a withdrawal of protein from less vital body tissue (muscle) to maintain
the functions of more vital tissues (liver, kidney, etc.) resulting in reduction
or cessation of growth. In growing animals, the majority of absorbed protein
is deposited in the carcass. However, in young, fast growing animals meta-
bolic organs like digestive tract, liver and kidneys also have a high priority
for protein deposition. There are 22 main amino acids in body proteins, and
all are physiologically necessary. Some of the amino acids are synthesized
in the body, and the rest are referred to as indispensable or so-called essen-
tial amino acids (Table V-11), which have to be present in the feed in the
necessary quantity. If the non-essential amino acids are not supplied by the
diet, they must be synthesized to a significant extend from non-essential
amino acids by the body. Therefore the presence of adequate amounts of
non-essential amino acids should not be overlooked and stating dietary
requirements for both protein and essential amino acids is an appropriate
way to ensure that all amino acids needed by the organism are provided.
88
Table V-11: Classification of amino acids for growth in pigs and poultry
1 Semi-essential amino acids are usually not essential, but are necessary for optimal growth;
2 Glycine is only required by poultry for the synthesis of uric acid.
The most important factor affecting the efficiency of protein utilization

is the dietary balance of amino acids. According to LIEBIG’s “Law of
Minimums” the undersupply of a single essential amino acid will inhibit the
responses to that supply in an adequate amount. In monogastric animals
the supply is greatly influenced by the diet, but in ruminants the microflo-
ra has a considerable effect on the pattern of amino acids absorbed. In order
to fulfil the requirements for dietary amino acids in monogastric animals,
the concept of the “ideal protein” has been developed as the basis for pro-
tein requirements for pigs and poultry. It was suggested that it should be
possible to establish an optimum balance of essential amino acids which,
when supplied with sufficient nitrogen for the synthesis of non-essential
amino acids, would constitute the ideal protein. Lysine is supposed to be
used as the reference amino acid and the ideal ratios of essential amino
acids to lysine are used as a basis. While such a concept has been devel-
oped in pigs and poultry, it is equally applicable to other species. Table V-
12 shows the optimum balance of essential amino acids in the ideal protein
required for growing pigs.
89
Table V-12: Estimated amino acid requirement for maintenance and growth in pigs in relation to the lysine
requirement, as well as amino acid content in the body and in sow’s milk
Legend: CP: crude protein; MEt= methionine; Cys= cystine
In order to determine ratios of the essential amino acids relative to

lysine it is necessary to establish requirements for different functions, e.g.
maintenance and growth, and utilization of amino acids for these functions.
Therefore it is necessary to know composition and rate of protein accretion.
Putting together the individual requirements of each amino acid will give an
idea of an optimum amino acid composition—ideal protein - necessary to be
supplied with the diet.
When applying the concept of the ideal protein it has to be kept in
mind that not only deficiency of only one of the indispensable amino acids
will limit protein utilization, but also a deficiency or surplus of indispensa-
ble in relation to the dispensable amino acids may also inhibit protein uti-
lization and hence growth. For example, in the conventional feeding of pigs,
where an increasing amount of the protein originates from cereal grain, pro-
tein utilization decreases. One important cause of this effect is that cereals
contain inadequate amounts of indispensable amino acids in relation to the
dispensable amino acids such as glutamine and proline. Experiments have
shown that the indispensable/dispensable amino acid ratio for optimum
protein retention in pigs should be at least 45:55.
In a number of broiler chicken studies of amino acid requirements for
broilers the response curves for growth and protein retention were estab-
lished in relation to different quantities of amino acids and the interrela-
tions between the individual amino acids. The results of the experiments are
shown in Table V-13.
90
Table V-13: Suggested ideal protein requirements for broilers, ratios expressed relative to lysine
1 Ratios based on a total requirement basis (NRC, 1994)

2 Illinois Ideal Chick Protein based on a digestible requirement basis (BAKER and HAN, 1994)
In principle, protein metabolism at the tissue level is not different

between ruminant and monogastric species. Early tracer studies indicate
that the essential amino acids are the same in cattle and sheep as in pigs.
The ruminants, however, differ from other species in a way that the amino
acids absorbed in the small intestine are highly dependent on the complex
metabolism in the forestomachs and less on dietary supply. The symbiotic
relationship between rumen bacteria and the host animal has created many
problems of understanding the amino acid requirements in ruminants. First
of all there were problems of estimating the maintenance protein require-
ment or minimal N excretion and then to estimate the utilization of micro-
bial amino acids. The optimal amino acid composition for growth and lac-
tation is summarized in Table V-14 (see also Chapter X).
Table V-14: Calculated optimum ratio of the five limiting amino acids for growth and lactation in ruminants,
g amino acid/100 g total amino acids (ØRSKOV, 1991)
91
FOR FUTHER READING
ARC (1981): The nutrient requirements of pigs. Commonwealth Agricultural Bureaux.

Farnham Royal, Slough
Baker, D.H. and Han, Y. (1994): Ideal amino acid profile for chicks during the first three
weeks posthatching. Poult. Sci. 73, 1441-1447.
Blaxter, K.L. (1989): Energy metabolism in animals and man. Cambridge University Press.
Chwalibog, A. (1991): Energetics of animal production. Research in Copenhagen, review
and suggestions. Acta Agric. Scand., 41, 147-160.
Chwalibog, A., Jacobsen, K. and Thorbek, G. (1996): The pattern of protein retention in
pigs from 2 to 120 kg live weight. J. Anim. Physiol. a. Anim. Nutr. 49, 181-186.
Chwalibog, A., Jakobsen, K., Tauson, A-H. and Thorbek, G. (2005): Energy metabolism and
nutrient oxidation in young pigs and rats during feeding, starvation and re-feeding.
Comp. Biochem. Physiol., Part A, 140, 299-307.
Kleiber, M. (1975): The fire of life, an introduction to animal energetics. Robert E. Krieger
Publishing Company Huntington, New York
Sawosz, E., Chwalibog, A., Niemiec, T., Kosieradzka, I. and Skomial, J. (2005): Effect of
nutrition on oxidative stress. J. Anim. Feed Sci., 14 (Suppl. 1), 87-97.
Thorbek, G., Chwalibog, A. and Henckel, S. (1984): Nitrogen and energy metabolism in pigs
of Danish Landrace from 20 to 120 kg live weight. Norms for protein and energy
requirement for maintenance and growth. Beretning fra Statens Husdyrbrugsfors.
563, p. 114
Ørskov, E.R. (1991): Progress and new frontiers in knowledge of protein nutrition in rumi
nants. Proceedings 6th Int. Symp. Protein Metabolism and Nutrition. EAAP Publ.
No. 59, Vol. 1.pp 80-87.
92
Chapter VI
REGULATION OF ENERGY METABOLISM

AND FEED INTAKE
6.1. Overview of endocrine control system of

energy balance
6.1.1. Hypothalamo-pituitary level
6.1.2. Thyroidal level
6.1.3. Leptin and ghrelin

6.2.1. Regulation of the feed intake
6.2.2. Prediction possibilities of feed intake
6.2.3. Fulfilment or fill unit (FU) or Fill Unit
(UE) or Satiety Value (SV)
CHAPTER VI: FEED INTAKE REGULATION
o maintain instantaneous and long-term energy equilibrium of organ-
T ism, a very complicated, fine tune regulatory network developed dur-

ing evolution, which includes both endocrine and physiologic regula-
tion of energy metabolism, feed intake and heat production.
6.1. Overview of endocrine control system of energy balance

© Peter Rudas
6.1.1. Hypothalamo-pituitary level

The most classical endocrine regulatory pathway governing energy metabo-
lism is the hypothalamo-hypohyseal system. Since the description of this
classical pathway our knowledge has been changed tremendously. The clas-
sical negative and positive feedback mechanisms involve effects of target tis-
sue products on the hypothalamus and on the hypophysis like sex steroid
effects on GnRH, FSH and LH synthesis and release, thyroxin and tri-
iodothyronine effects on TRH and TSH production, IGF effect on GH release
and inhibition of CRH and ACTH production by glucocorticoids. Examples
for positive feedback mechanisms are estrogens induced LH release in
mammals or progesterone induced LH-surge in the fowl. More and more
data prove, however, that beside these classic feedback regulatory pathways
many peripheral signals may interact with the hormone production of the
pituitary gland. Among these signals are peptides that also occur in the
hypothalamus, but are also produced by other tissues (pancreatic and gas-
tric somatostatin). Others can not be found as a product of the hypothala-
mus and solely come from the periphery: leptin and ghrelin. The former is
produced by the adipose tissue and the latter is produced primarily by the
stomach.
Because of the space limitations the above classic pathways are not
discussed here in more depth, instead we concentrate on the novel aspects
of the unconventional regulatory mechanisms influencing energy metabo-
lism, namely that of the Folliculo Stellate Cell (FSC) activity. FSC as special
components of the pituitary has not received attention for a long time, since
no trophhormone production is bound to these types of cells. The unique
web of the FS cells that are connected by gap junctions makes up a dense
cellular network throughout the pituitary. It has been suggested earlier that
94
they can act as stem cells, since they show mitosis and could transform into
granular or chromophilic cells according to the concept of a “cell renewal
system”. As knowledge about these cells has accumulated more and more
functionality was ascribed to them. Since about 1990 when ALLAERTS and
co-workers revisited the morphological and functional aspects of these cells
the physiological importance of FSC became the target of intensive research
in mammals. It is now clear that the FS cells strongly influence hormone
producing troph cells of the pituitary and that communication of the FS
cells among each other may involve paracrine and electric circuits. The
presence of spontaneous, voltage- and inositol 1,4,5-trisphosphate-induced
intercellular calcium signals that were abolished by gap junction channel
blockers and excitability of these cells speak in favour of this. Accordingly,
the synchronization of intrapituitary electrical and calcium signalling may
be one of the major features of the FSCs. Isoproterenol, prostaglandin E2,
bradykinin and lysine vasopressin are able to stimulate active ion transport
across FS monolayer in the bovine indicating that the ion transporting FS
cells may be important in local regulatory functions. The action mechanism
may involve the increase of blood supply to selected troph cell populations by
producing vascular endothelial growth factor (VEGF), too.
Using radiolabelled iodocyanopindolol was possible to identify and
characterize beta-adrenergic receptors in bovine pituitary folliculo-stellate
cells grown in culture. Later, the occurrence of FS cells were described in
domestic species (goat, bull, sheep and foal). Steroid mediated effects on
prolactin release modulated by FS cells were shown too. In chickens the
developmental aspects of FS cells were clarified using differential immuno-
histological markers for hematopoietic, hormone-producing, and folliculo-
stellate cells. It was found that specific FS cell populations appear only after
hatching.
The shock protein (metallothionein) immunoreactivity was detected
recently of pituitary FSC in the bovine. It was postulated that, besides the
protective action of shock proteins against free radicals, hypophyseal metal-
lothioneins might be involved in the regulation of the release of hypothalamic
peptide hormones.
There are marked increases in colloid-filled follicles in the anterior
pituitary gland of the senescent porcine pituitary. These colloids as shown
by Percoll gradient centrifugation and glycoprotein analysis to be of two
types, albumin fragment and clusterin containing colloids. FSC activity and
aging are thus closely coupled in the pig and this might serve as a pituitary
model of ageing in other species, too. The occurrence and importance of FS
cells in the pituitary of pigs was also described recently. A close link
between the activities of FS cells in the fowl with energy metabolism was
described in moulting. With specific immuno-histochemistry it was demon-
strated that FS cell population increases at the final stage of fasting induced
moulting and they may be involved in control of pituitary functions. The impor-
95
tant paracrine function of the folliculo-stellate cell in the control of energy

metabolism is underlined by the fact that leptin is expressed in FS cell line.
The finding of leptin and of the long isoform of leptin receptor in these cell
lines implicates an autocrine/paracrine loop in the production and regula-
tion of the hormone that is a principal signal of the energy status of the body
(JIN et al. 2000).
Beside the FS cell activity there is another important issue that
changes our view of the classical “hypothalamo-hypophyseal-target
endocrine organ” relationship. This is that at the level of troph hormone
(TSH, FSH, LH, GH etc) producing cells, on which the multitude of signals
of hypothalamic, folliculo-stellate and of peripheral endocrine gland is orig-
inated there are distinct subsets of secretagouge cells. Since in the last
years this diversity of troph hormone producing cell populations is most
apparent in the case of growth hormone (GH) we shall summarize some new
results in this direction. Isoforms of growth hormone probably produced by
different cell types can be lypolytic or antilypolytic. Altered biological activ-
ity of glycosylated and non glycosylated variants have also different biolog-
ical availability since their disappearance rate and their binding character-
istics to hepatic GH-receptors is markedly different. These and many other
data support the view that the heterogenic effects of growth hormone may
be explained of charge, size and structural isoforms secreted by different
subsets of GH producing cells in the pituitary.
In summary the first novelty at the pituitary level according to the
facts seen above is that beside the secretagouge cells governing energy
metabolism (thyrotrophic and somatotroph cells in the first place) are not any
more considered to be independent of their local micro-environment within the
pituitary: a co-ordinating cell population, namely the folliculo-stellate cell
line may play a crucial role in the regulatory circuit. In the last ten
years it has also become clear that the classical “releasing factor (inhibiting
factor) - troph hormone secreting cell” relationship is replaced by the “many
endocrine-parapcrine factors influencing many subtypes of secretagouge
cells” producing isoforms of pituitary troph hormones.
6.1.2. Thyroidal level

One of the major roles of thyroid hormones is the effect on morphogenesis,
especially in lower vertebrates. This aspect will however not deal with here,
since the major role of the thyroid system in mammals and birds is the con-
trol of energy expenditure on heat production (GREER, 1974). The classical
hypothalamo-hypophyseal-thyroid feed back mechanism in regulating thy-
roid economy has first been proved that the thyroxin (T4) reaching the plas-
ma from the thyroid gland can normally be converted to tri-iodothyronine
(T3), which means that T3 can not only be of thyroidal, but also of periph-
eral origin. It is clear today that, however, T4 is the major product of the thy-
roid gland it should be considered as the prohormone of the final, active
96
form of triiodotyhronine (T3) which is mainly produced by the peripheral

(non-thyroidal) cells in the body. Deiodinase systems and other factors like
sulphation etc. are also known. In short the intracellular events of activa-
tion and inactivation entail the followings. In birds, as in mammals, so far
three types of deiodinases have been identified, type I (D1), type II (D2) and
type III (D3).
Sulphation is the other important step in making more or less thy-
roid hormone available to the final action. Conjugation of sulphate to the
hydroxyl group on the outer ring is a regulated process by which the avail-
ability of thyroid hormones to deiodination is regulated. Accordingly sulpha-
tion is another possibility to fine tune the amount of active thyroid hormone
(3,5,3’-triiodothyronine) available for the nuclear receptor system.
Sulphation of T4 can increase the inactivation of this hormone by inner ring
deiodination or decreasing the inactivation by inactivating outer ring deiod-
ination while sulphated form of T3 stimulates its degradation.
As for the heat generation of cells it is of crucial importance that it is
not only the thyroidal thyroid hormone production that determines the
effectiveness of heat generation, but more importantly the bioavailability of
thyroid hormones. This new approach was conceived by many authors, who
became more and more convinced that the peripheral conversion (activation
and inactivation) of thyroid hormones within the target - heat generating -
cells is a key factor in the hormone action. Many effects of thyroid hormones
can be explained by local deiodination and sulphation, but the uptake rate
and elimination of these hormones are also of crucial importance. This has
clearly been shown by a series of experiments investigating the unique han-
dling of thyroid hormones by the brain tissue in chickens. The increased
uptake of thyroid hormones into the brain in hypothyroid chickens in these
experiments (RUDAS et al. 1993, 1994) speaks in favour of a specific, regu-
lated transport mechanism of thyroid hormones that is able to increase its
activity when neuronal hypothyroidism is to be avoided. Indeed it was found
by others that the transport of thyroid hormones is not a simple diffusion
process but it is determined by specific transporters that belong to the
MTC8 transporters family.
The final effect of thyroid hormones is dependent on the availability of
thyroid hormone receptors. Thyroid hormone receptors preferentially bind
3,5,3’-triiodothyronine (T3). They belong to the steroid/retinoic acid/D-hor-
mone super family and two major isoforms of them are known: TRa and TRb.
Even if the level of receptor proteins was shown not to be closely related to
the actual level of the mRNA in some cases, one can conclude from our
studies that the changes in receptor expression is a most likely candidate
for adaptation to energy restriction in chickens. The final, direct action of
the activated thyroid hormones to increase heat production depends on the
increase of the expression of UCPs (uncoupling proteins) and on events at
the mitochondrial molecular level. The practical outcome of the sophisticat-
97
ed activation-inactivation and availability of thyroid hormones described

above on the energy metabolism of animals will be elucidated in one species
on the chicken based experiments by BARTHA et al. (1989 and 1998).
These studies suggest that food/energy deprivation triggers a mech-
anism by which thyroid hormone activation is diminished by the peripher-
al cells. The decreasing production of the active hormone (3,5,3’-triiodothy-
ronine, T3) allows conservation of metabolic fuels. The major trigger of nor-
malization of this adaptation after refeeding is a glucose dependent path-
way.
6.1.3. Leptin and ghrelin

About fifty years ago it was suggested that there should exist a peripheral
signal that informs the brain about the actual level of energy stores in the
body. This supposition was proven true later by finding the signal, leptin
that is produced in the adipose tissue and elsewhere (ZHANG, 1994). The lep-
tin gene is highly conserved in different species and consists of about
15,000 base pairs and contains 3 exons, which are separated by 2 introns.
The structure is similar to the cytokine family of peptides as IL-6 GH. The
primary source of leptin is the adipose tissue but nowadays it is discovered
in more and more diverse tissues included cattle (SAYED at al. 2003). Leptin
may be found in the circulation in the free form or bound to binding pro-
teins, and this feature seems to be species specific.
The leptin receptor has a single membrane-spanning domain and
exists in different isoforms that derive from alternative splicing of mRNA
(SAYED et al., 2003). All isoforms have similar ligand-binding domains but
differ at the C-terminus, the intracellular domain. The Ob-Rb, which con-
tains a long intracellular domain, is the only isoform with both of the pro-
tein motifs necessary for activation of the kinases and signal transducers
and activators of transcription.
The original overall metabolic role of leptin to decrease appetite
and increase energy expenditure was based on the following observations.
a) Leptin (ob/ob mice) and leptin receptor mutants (db/db mice, fa/fa
rats) have lower core body temperatures, are more susceptible to cold stress
soon after birth, and are hyperphagic and obese by the time of weaning.
b) Daily intraperitoneal (ip) injections of recombinant leptin reduce
food intake and body weight and increase energy expenditure in ob/ob mice
and in wild type rodents but no effect on db/db mice.
c) The long isoform of leptin receptor is predominately expressed in
the hypothalamus especially the arcuate, paraventricular and ventromedial
nuclei that control both food intake and sexual behaviour.
d) The high affinity leptin transport in the hypothalamus and choroid plexus
is playing a key role in regulating leptin entry into the CNS and CSF.
Regulation of feed intake, energy balance and neuroendocrine func-
tion by leptin is thought to be mediated by differential expression of a net-
98
work of hypothalamic peptides with orexigenic (anabolic) and anorexiogenic

(catabolic) effects. The orexigenic neuropeptides are neuropeptide Y (NPY),
agouti-related peptide (AgRP), melaninconcentrating hormone (MCH),
galanin and orexins, while anorexigenic peptides include corticotropin-
releasing hormone (CRH), pro-opiomelanocortin (precursor of -melanocyte
stimulating hormone MSH), cocaine and amphetamine-regulated transcript
(CART) and thyrotropin releasing hormone (TRH). This concept is support-
ed by the co-localization of many of these neuropeptides neurons with lep-
tin receptors. Leptin acts centrally to inhibit the effects of NPY by inhibiting
its synthesis in the arcuate nucleus of the hypothalamus. Leptin adminis-
tration to the hyperphagic ob/ob mice, wild type mice and sheep results in
a rapid reduction in NPY mRNA abundance, protein secretion and reduced
food intake before any change in body weight. Both NPY- and galanin-
induced feeding behaviours were completely inhibited by pre-administration
of leptin.
The anorexigenic neuropeptides include:

1. the corticotrophin-releasing hormone (CRH)
2. Melanocortins (MC) that is cleaved from the precursor molecule
POMC that is expressed in the ARC of the hypothalamus and pituitary
gland. The agouti-related protein AgRP is another orexigenic peptide secret-
ed by the ARC and acts as melanocortin receptor antagonist.
3. Cocaine-amphetamine regulated transcript (CART) containing neu-
rons are widespread throughout the brain, including the ARC, LH and PVN.
Recombinant CART injected rats tended to inhibit normal and previous
starvation-induced feeding
4. Thyrotropin-releasing hormone (TRH). Leptin activates directly the
TRH neurons in the paraventricular nucleus or indirectly by increasing the
production of the MC4R ligand (melanocyte stimulating hormone -MSH) to
regulate TRH expression on the level of TRH promoter and thus influencing
thyroid hormone release, too. FIGURE VI-1 vizualizes the most important
effect of the leptin.
99
FIGURE VI-1: Regulatory role of adipocytes in the mammalian organism
The intracellular signalling pathway induced by leptin via its receptor

in different tissues reflects a wide range of effects. JAK (Janusactivated
kinase) family members initiate an intracellular signalling cascade. Some
isoforms of the receptor may contain intracellular tyrosine residues, which
interacts with STAT (signal transducers and activators of transcription).
STAT protein enters the nucleus and interacts with specific DNA elements
in the promoters of target genes to regulate gene expression. Leptin may
also induce the MAPK (mitogen activated protein kinase) system. Leptin
induces transient expression of SOCS-3 (suppressor of cytokine signaling-
3) peptide. SOCS proteins act in a typical negative feedback loop and can
inhibit leptin signalling. IRS (insulin receptor substrates) phosphorylation
and their interaction with PIK phosphatidylinositol-kinase are modulated by
leptin. Many other signalling proteins are also regulated by leptin (PKB, pro-
tein kinase B; PKC protein kinase C; NO nitric oxide; cyclic AMP; actin;
ROS, reactive oxygen species; and prostaglandins.
Ghrelin. Growth, metabolism and energy balance of the body is

mainly regulated by the growth hormone. According to the classical feed-
back pathway (see earlier) it seemed to be under the control of only two
hypothalamic factors GH-RH (growth hormone releasing hormone) and SRIF
(somatotropin release inhibiting hormone or somatostatin). A family of pep-
tides has been discovered that they were able to stimulate growth hormone.
These peptides were called growth hormone secretagogues (GHS). Soon later
secretagogue receptors (GHSR) were also identified in the hypothalamus,
but interestingly enough for a longer time, there were no natural ligands
coupled to these receptors. Later a peptide has been isolated from the stom-
100
ach that was able to stimulate GH-release. In domestic mammals it was the
elegant experiment with hypothalamic-pituitary stalk-sected (HST) pigs that
has proven the presence of a non hypothalamic factor that is able to stim-
ulate growth hormone release.
The peptide was named ghrelin. Ghrelin is produced in the chromogranin
A-immunoreactive endocrine cells within the mucosal layer of the fundus in
most species. Ghrelin, possesses a hydrophobic moiety: an octanoylated lin-
ear chain and the ester bond that links the alkyl chain to the serine (Ser3)
side chain. Without this side chain the peptide itself can not bind to its
receptor. Therefore the octanoylated form is also called “active ghrelin”.
Introducing ghrelin into the already complicated picture of energy metabo-
lism seems to be filling the gap between gastrointestinal regulation and
energy expenditure and between the short term regulation of energy metab-
olism by thyroid hormones and a long term balance keeping mechanism.
Ghrelin not only stimulates GH secretion but also stimulates feed intake by
stimulating the hypothalamo-hypophyseal system.
The neuroendocrine system in regulating pulsatile GH secretions may
be influenced by ghrelin in coordination with a number of other peptides of
local hypothalamic or systemic origin. The arcuate nucleus and ventrome-
dial nucleus of the hypothalamus are containing the highest amounts of
ghrelin (GHS-R) receptor expression. (The arcuate nucleus contains neu-
rons for orexigenic peptides (i.e. NPY, AGRP and POMC). The orexigenic
effects of ghrelin can be blocked by chemical ablation of the nuclei of the
hypothalamus. The central physiological effect of ghrelin is to stimulate feed
uptake while increasing growth hormone secretion. In this way it is the only
endocrine and stomach derived factor at present that stimulates appetite
and induces a positive energy balance leading to body weight gain together
with its ability to stimulate GH secretion. Beside this central role it acts on
a number of tissues directly via GHS-R receptors. It was shown that it stim-
ulates gastric acid secretion and gastrointestinal motility, it interacts with
the sleeping pattern in animals, it modulates cell proliferation and survival,
it influences cardiac function and vascular resistance, it influences the
exocrine and endocrine pancreas, it acts on the immune system and acts on
the lactotroph and corticotroph cells in the pituitary, and also, it acts on the
adipose tissue, too. All these effects are coordinated into one direction: to
stimulate anabolic pathways.
Ghrelin as a gastrointestinal peptide contributes to the regulation of
diverse functions of the gut-brain interrelationship and thus ghrelin itself
and/or agonists and antagonists of it will have substantial clinical impact
in veterinary medicine, too.
101

Introduction
As a result of evolutionary development the behaviour of „feeding on ener-
gy” has been developed. This means that voluntary feed intake is adjusted
to the actual energy needs of the organisms. Thus, dry matter intake is
more likely obtained from a low concentrated feed mixture, and less likely
from a high energy density feed. Domestication has modified these charac-
teristics (selection for higher growth rate in swine and broiler chicken),
but it can be seen in its clear form in laying hen, rabbit and rodents (FIG-
URE VI-2).
FIGURE VI-2: Effect of deietary energy concentration on voluntary feed intake
The fibre concentration has the highest influence on energy density

of feeds, therefore the closer the range of fibre in the feed mixture of a
given species is, the more accurate prediction of voluntary dry matter
intake can be predicted. Fibre concentration shows the largest deviations
in ruminant rations (12-30% for intensive (dry-lot) beef cattle and dry cow,
respectively); in case of monogastric range is narrowed, consequently, pre-
diction possibilities in pig and poultry are better than in case of rumi-
nants. In practice, feed intake may be influenced both in positive and in
negative directions. In case of fast growing meat animals the stimulation
of feed intake may have some advantages (e.g. using of feed flavours), but
in some special cases (e.g. retarding sexual maturity, increasing lean meat
102
percentage) reducing the appetite may be the goal. If regulation is func-

tioning in primary forms, animals can be fed ad libitum (e.g. in growing
rabbits).
6.2.1. Regulation of the feed intake

Concerning the regulation of feed intake qualitative (WHAT TO EAT) and
quantitative (HOW MUCH TO EAT) questions arise. Feed selection is influ-
enced by taste, smell (humans, rats, cats), temperature and humidity (new-
born animals, adult carnivores), imitation (sheep and poultry) and jealousy
(sick dogs). A special case of feed selection is the intake of own faeces or cae-
cum content (physiological coprophagy and caecotrophy) as well as forms of
allotriophagia (feeding on faeces of other species, dirty, debris etc.). They
used to exist technologies for feeding of processed bedding of a species to
another one (e.g. poultry bedding for ruminants).
To determine the amount of ingested feed, there are long-term and
short-term regulations. The aim of long-term regulations is to maintain
healthy live weight, body composition and energy and nutrient depots.
Long-term regulation is of homeorrhetic character and basically stands
under the control of central nervous system but in their turn, peripheral
hormones (leptin and ghrelin) also play important role. The function of
adipocytes and their leptin production should be highlighted, because their
number is congenital (cellularity) and leptin production is genetically deter-
mined (see ob/ob and fa/fa genotypes). One of the key factors of long-term
regulation is that metabolites of fat tissue (leptin) and gonads (sexual
steroids) decrease feed intake (see belling of deer, heat of sow). Circulating
growth hormone (GH), thyroid hormones, ratio of glucagon to insulin are
significant in post calving fat mobilisation and in the appetite of high yield-
ing dairy cows. (The role of growth hormone in poultry is different, because
it would rather promotes fat accumulation.)
The goal of short-term regulation is the maintenance of milieu
interieur, covering momentary needs. Hunger centre in hypothalamus is
continuously active, which is, among others, blocked by gastrointestinal
hormones (ghrelin, cholecystokinin=CCK). Actual feed intake may be modi-
fied by preference, habituation and feed refusal factors (e.g. T–2 toxin, some
drugs). Daily feed intake is determined by physical (mechanical), physico-
chemical (osmotic pressure and pH of ruminal and gastrointestinal content,
blood glucose, amino acids and volatile fatty acid level), environmental tem-
perature and endocrine background. Appetite of patients with fever (pigs,
dogs and cats), such as fishes, amphibians and reptile, in cold environ-
mental environment decreases. In practice, the effect of environmental tem-
perature on feed intake should be taken into consideration mostly in case of
poultry, dairy cows (heat stress) and beef cattle (cold climate).
Natural control of feed choice and intake in animals is generally used
only in case of drinking water and salt. Although poultry is able to select
103
feed necessary for covering its amino acid requirement, but it cannot be
employed in practice. In case of the majority of micronutrients (e.g. phos-
phorus for cattle) the taste has stronger effect on intake than the real needs
of the organism.
6.2.2. Prediction possibilities of feed intake

The main character of feed intake regulation can be concluded from fibre
(ballast) content. For low density feed simple physical control (capacity of
gastrointestinal tract) is predominant. Parallel to the increase of energy den-
sity chemical regulation receive more and more importance. In case of prac-
tical feed mixtures both forms of regulation play an active role. The most
complex is the regulation of cattle feed intake where both pH, osmotic pres-
sure, fullness of rumen and efficacy of fermentation as well as outflow rate
and digesta passage have a dominant role.
For prediction of feed intake in case of CATTLE the following equations
are used:
FIroughage= 36 V + 540
where: FI = roughage intake in gram;
V = rumen volume, litre.
DMIdairy= 0.01 × W0.75 × (0.225 × FCM + 8.796)
DMIbeef= W0.75 × (0.0355 NEm - 0.026 NEm2- 0.0196)
where: DMI= dry matter intake, kg;
FCM= 4% fat corrected milk;
W= live weight, kg,
W0.75= LW, kg0.75
NEm= energy concentration of feed mixture,
MJ/kg DM.
For LAYING HEN the dairy energy intake can be predicted with consid-
eration of environmental temperature, too:
ME = W (586-8.4 T) + 8.4 Ep +21 W;
where ME = energy intake during 24 hours in metaboli-
zable energy, kJ
T = environmental temperature, oC;
E = daily egg production, g;
W = average daily gain, g.
If that is the case, for laying hen it is advisable to use the winter and
the summer feed mixture with different energy and nutrient density,
because heat stress decreases voluntary feed intake. Thus, protein, miner-
al and vitamin levels should be increased.
104
6.2.3. Fulfilment (FU) or Fill Unit (“unité d’encombrement=UE”) or

Satiety Value (SV)
All of the above described prediction approaches aresimplified, because the
modifying effect of fibre fractions has not been taken into consideration.
When feeding cattle on feedstuffs of different fibre composition voluntary
dry matter ingestion is significantly differed: the highest dry matter intake
has observed in case of feeding on grass and the lowest in case of feeding
on straw. The reason of this phenomenon is that animals detested different
fibre composition of the offered roughages. In order to obtain a more accu-
rate prediction INRA (1978) introduced FULFILLMENT UNIT (UNITÉ D’ENCOMBRE-
MENT). This unit expresses dry matter intake of a „standard ruminant” from
a „standard grass” and expected voluntary dry matter intake from other
roughages that are expressed in the proportion of this etalon. Thus, two dif-
ferent feedstuffs of the same fulfilment unit have the same decreasing effect
on appetite. This type of prediction gives more accurate results than those
of based on the percentage of live weight.
During classical experiments cows were fed ad libitum on different
feedstuffs. Daily meadow grass intake was 60 kg (i.e. 16 kg dry matter) and
rumen content weighed 74 kg. Feeding on straw the corresponding values
were 7 kg (6 kg dry matter) and 107 kg. In case of giving total mixed ration,
the feed intake amounts to 74 kg (19 kg dry matter) with the rumen content
of 84 kg. It is clear, that in certain situations, dry matter-based prediction
for ruminant are not appropriate.
Explanation lies in the chewing movements and rumen fullness,
lignin being the keyword. Feed intake has a negative correlation with lignin
content. More lignin proportion involves more chewing activity and it
requires more time. Efficacy of fibre degradation is determined by the rela-
tive measure of the rumen and intestines, which explains the differences
between the ruminant species. Nitrogen and energy supply of cellulolytic
bacteria have also a great impact. If rumen nitrogen balance is positive,
microbial fibre degradation is more efficient and dry matter intake increas-
es.
In basic trials a wether of 60 kg of live weight ingested 1.8 kg fresh
grass daily, while from wheat straw only 0.7 kg. Thus, feed intake is funda-
mentally influenced by its physical structure. For SHEEP the UEM (M=sheep,
mouton) is used, where the unit is the medium grass (15% crude protein
and 25% crude fibre in dry matter) of 1 kg of dry matter. Daily intake of
standard wether from this standard grass amounts to 1.62 kg DM (i.e. 75
g/W0.75). For DAIRY COWS the UEL (L= milk, „lait“) has been proposed: medi-
um frame, medium yielding cow that ingests 17 kg DM (i.e. 140 g/W0.75)
from the standard grass. The UEB (B=cattle of other categories, „bovin“) is
applied for non-lactating cattles which ingest 8.5 kg DM (i.e. 95 g/W0.75)
from the standard grass. The intake capacity of the MILK GOAT is situated
between dairy cow and beef cattle, having 123 g/W0.75 average dry matter
105
intake from the reference grass.

For example, UEM value of the reference grass for the standard
wether is 1 and its means a dry matter intake of 1.62 kg. Dry matter intake
from young grass and from oats straw appear 1.8 and 0.7 kg, respectively.
It means that fulfillement unit (FU) of young, tender grass and oats straw
equal to 1.62/1.80=0.9 and 1.62/0.7=2.3, respectively. In other words, sati-
ating effect of tender grass is smaller by 10% and that of straw is greater by
2.3 times than that of the reference grass. Fulfilment units of the most
important ruminant feedstuffs are listed in Table VI-1.
In Denmark the developed version of FU is in use: the “Feed Intake
Potential” (FIP), expressed as (FU/cow/day) is a function of lactation num-
ber, day in milk and days pregnant. THOMAS (2004) applied a stepwise mul-
tiple regression analysis in developing the Feed into Milk (FIM) system.
Variables of this equation are forage intake potential (FIP in g/W0.75), con-
centrate dry matter intake (CDMI in kg/day), week of lactation (WOL), for-
age starch concentration (FS in g/kg DM) and the crude protein concentra-
tion of the concentrate (CCP in g/kg DM). Although the physiological basis
of these systems is the same, the FU, FIP and FIM systems have different
meaning and are expressed in different units (TAMMINGA, 2005).
Table VI-I: Fulfilment unit (FU) of roughages (INRA, 1978).
*UEM = sheep FU (unité encombrement mouton)

** UEL= dairy cow FU (unité encombrement laitière)
*** UEB = cattle FU (unité encombrement bovin)
Energy density expresses the amount of feed energy (in French sys-
tem: UF, unité fourragère) per fulfilment unit, i.e. UF/UE.
While partly replacing roughages and silages by concentrated feed
(e.g. cereals), the feed intake depression of total mixture decreases.
106
The SUBSTITUTION NUMBER shows the fall in roughage intake after mix-
ing a unit of concentrate into the silage (FIGURE VI-3).
FIGURE VI-3: Interrelationships of roughage and concentrate ratio and total dry matter
intake (after JARRIGE, 1978),
Values of substitution number for concentrated feeds have been

measured between 0.2 and 0.6, which means that mixing of 1 kg concen-
trate decreases roughage/silage dry matter intake by 0.2-0.6 kg. (Thus,
total dry matter intake to an optimum point improves.) UK recommendation
(AFRC) defines this phenomenon inversely: addition of 10% concentrate
increases total dry matter intake by 0.12-0.5 kg.
FOR FURTHER READING
Agricultural and Food Research Council (1991): Technical Committee on Responses to

Nutrition. Report No. 8, Voluntary Intake of Cattle. CAB. Wallingford.
Andrieu, J., Demarquilly, C. and Wegat-Litre, E. (1981): Prévision de la valeur nutritive des
aliments des Ruminants. Éd. INRA publications, Route de St-Cyr, 7800 Versailles
107
Bartha, T., Rudas, P., Fekete, S. and Pethes, G. (1989): Restricted feed intake influences
thyroid hormone production and peripheral deiodination in chickens. Acta Vet. Hung.
37, 241-246.
Daniel, J.A., Whitlock, B.K., Baker, J.A., Steele, B., Morrison, C.D., Keisler, I. and Sartin,
J.L. (2002): Effect of body fat mass and nutritional status on 24-hour leptin profiles
in ewes. J. Anim. Sci. 80, 1083-1089.
Forbes, J. M. (1986): The voluntary food intake of farm animals. Longman. London.
Horvath, E. and Kovacs, K. (2002): Folliculo-stellate cells of the human pituitary: a type of
adult stem cell? Ultrastruct. Pathol. 26, 219-28.
Jarrige, R. (Ed.) (1988): Alimentation des bovins, ovins & caprins. INRA. Paris. 29-56.
Jin, L., Zhang, S., Burguera, B.G., Couce, M.E., Osamura, R.Y., Kulig, E. and Lloyd, R.V.
(2000): Leptin and leptin receptor expression in rat and mouse pituitary cells.
Endocrinology. 141, 333-339.
Kennedy, G.C. (1953): The role of depot fat in the hypothalamic control of food intake in
the rat. Proc. Roy. Soc Lond., 140, 579-592.
Kojima, M., Hosoda, H., Date, Y., Nakazato, M., Matsuo, H. and Kangawa, K. (1999):
Ghrelin is a growth-hormone-releasing peptide from stomach. Nature. 402, 656–660.
Nakazato, M., Murakami, N., Date, Y., Kojima, M., Matsuo, H., Kangawa, K. and
Matsukura, S. (2001): A role for ghrelin in the central regulation of feeding. Nature.
409, 194-198.
NRC (1987): Predicting feed intake of food-producing animals. National Academy Press.
Washington, D. C.
Rudas, P., Bartha, T. and Frenyo, V.L. (1994): Elimination and metabolism of triiodothy
ronine depend on the thyroid status in the brain of young chickens. Acta Vet. Hung.
42, 465-476.
Sayed-Ahmed, A., Kulcsár, M., Rudas, P. and Bartha, T. (2003): Expression and localiz
ation of leptin and leptin receptor in the mammary gland of the dry and lactating
non-pregnant cow. Acta Vet. Hung. 52, 97-111.
Tamminga, S. (2005): Challenges in dairy cattle nutrition in the 21th century. in
Babinszky, L. (Ed.): New challenges in 21th century animal nutrition. Kaposvár, p. 47-65.
Thompson, N.M., Gill, D.A.S., Davies, R., Loveridge, N., Houston, P.A., Robinson, I.C. A. F.
and Wells, T. (2004): Ghrelin and des-octanoyl ghrelin promote adipogenesis direct
ly in vivo by a mechanism independent of the type 1a growth hormone secretagogue
receptor. Endocrinology. 145, 234-242.
Wang, G, Lee, H.M., Englander, E. and Greeley, G.H.Jr. (2002): Ghrelin – not just another
stomach hormone. Regul. Pept. 105, 75-81.
Zhang, Y., Proenca, R., Maffei, M., Barone, M., Leopold, L. and Friedman, J.M. (1994):
Positional cloning of the mouse obese gene and its human homologue. Nature. 372,
425-432.
108
JAV_7_VND_FSGy:MINTAÚJ.qxd 2008.09.03. 14:35 Page 109
Chapter VII
DIGESTIBILITY OF NUTRIENTS
7.1. General considerations regarding digestibility

and its determination
7.1.1. Excretion by faeces
7.1.2. Determination of digestibility
7.1.3. Digestibility by difference
7.1.4. Determination of the true digestibility
7.2. Factors affecting digestibility
7.2.1. Endogenous factors
7.2.1.1. Species of animal
7.2.1.2. Age
7.2.1.3. Movement
7.2.1.4. Habituation
7.2.1.5. Climate
7.2.1.6. Caecotrophy
7.2.1.7. Physiological state
7.2.1.8. Gender
7.2.1.9. Stress
7.2.2. Exogenous factors
7.2.2.1. Environmental temperature
7.2.2.2. Feeding level
7.2.2.3. Composition of feed mixture or daily ration
7.2.2.4. Biologically active substances, feed additives
7.2.2.5. Feed processing
7.2.2.6. Feeding technique
7.3. The concept of ileal digestible amino acid and ideal protein
in swine and poultry nutrition
7.3.1. Digestibility of amino acids in pigs
7.3.2. Determination of ileal digestibility with different
techniques
7.3.4. Application of ileal digestible amino acids in the
formulation of swine diets
7.3.2.1. Digestibility of amino acids in poultry
7.3.2.1. Determining the digestibility of amino acids with
different methods
7.3.2.2. Ideal protein concept in poultry nutrition
7.3.2.3. Diet formulations on the basis of digestible amino
acid concept
109
Chapter VII: DIGESTIBILITY OF NUTRIENTS
ne of the most important characteristics of feedstuffs and feed mix-
O tures is their digestibility, which basicly determines the available

proportion of the ingested major nutrients. The part of the feed
which can pass through the intestinal wass into the blood or lymphatic cir-
culation is considered digestible (owing to splitting, emulsion formation
etc.). Generally, digestibility is measured indirectly as a difference between
the amounts of compounds, that are ingested by feed intake and ejected in
the feaces. It is called apparent (faecal) digestibility. In equation:
DC=(gram nutrient eaten – gram nutrient in faeces)/nutrient eaten × 100,
In which DC is the digestion coefficient, as the percentage of nutrient con-
sumed which did not appear in the faeces.
Part of the absorped compounds will be re-excreted into the intestin-
al lumen (metabolic proportion), besides, faeces may contain detached cells
of the intestinal mucosa as well as tresidue of digestive juices. Calculation
taking into consideration the above mentioned factors gives the so called
true digestibility. Both apparent and true digestibility are given in percent-
age, in form of digestion coefficient (DC).
7.1. General considerations regarding digestibility and its

determination
7.1.1. Excretion by faeces
Faeces basically consists of the indigestible portion of the feed which owing
to physical or chemical (indigestible complex building) charcteristics cannot
be absorped. Furthermore, residue of digestive enzymes, bile salts and pig-
ments (bilirubinâstercobilin), detached epithelial cells and microorganisms
are also present in the faeces.
There is a nitrogen and mineral seccretion from the body into the
intestinal lumen through the secretory glands and cells. The higher is the
metabolic faecal losses in nitrogen the higher is the feed intake and the
lower the digestibility of organic matter. Compounds undergoing microbial
fermentation in the large gut produce fermentation and putrefaction prod-
ucts (indol, scatol, mercaptane etc. see Chapter VIII, Biogenic amines)
which are responsible for the smell of faeces. Water content – determining
consistency – may move between large limits (50-80%).
Amino acids are absorpted from small intestine. Caecal and colonic
110
microflora modify amino acid composition of the digesta from small intes-
tine. This failure can be eliminated by calculating the absorped proportion
as the difference between the ingested feed and ileal content istead of the
difference between the ingested feed and faecal content (apparent ileal
digestibility). When the endogenous part of ileal digesta is also considered,
we receive data of true ileal digestibility (Table VII-1).
Table VII-1: Apparent and true faecal (ADC) and true ileal (TDC) digestion coefficients in pig
% of the two most important protein sources
From the table it is clear that values of true ileal digestibility were
higher by 2-8 percentage unit than those of apparent faecal digestibility (for
more details see section 7.3.)
7.1.2. Determination of digestibility

Digestibility is commonly determined by means of in vivo feeding trials. For
measuring the digestibility of organic matter, protein and fibre in vitro
methods are also applied. In case of ruminants in vivo methods involving
living rumen content are also accepted (TERRY and TILLEY, 1963;
VERESEGYHÁZY and FEKETE, 1987).
During animal digestibility trial the amount and chemical composi-
tion of ingested feed and the refused feed (“orts”) should be measured. The
start and length of faeces collection should be fixed in such a way that sam-
ples reflect the digestibility of the tested material. In monogastric animals
the use of colouring agents (e.g. carmine, fuchine) help in setting the appro-
priate time.
In case of total collection trial after an adjustment period a so called
preliminary period of several days follows in which the same feed or ration
to be investigated is fed in the same weighed amounts daily as in the col-
lection period or actual experiment. This is for the purpose of removing from
the digestive tract all residues of previous feeds and also establishing as
uniform a rate of passage of feed products and egestion of faeces as practi-
111
cable, relative to feed intake. In the collection period (4-14 days) the whole
amount of faeces is gathered. The length of collection period necessary to
obtain reliable results depands upon the species, longer periods being nec-
essary in case of herbivorous, especially ruminants, than for other animals
because of the daily variations in the amount of faeces. In general, the
longer the period of collection, the more accurate the results are.
The advantage of the indicator method is that the total faeces are not
collected and weighed but merely sampled and analysed. This departure
from the former method of determining digestibility has been referred to as
the clue, indicator, tracer, ratio, reference, inert reference substance or
index method. By this method, in addition to the chemical analysis of the
usual proximate nutrients, the content in the feed and in the faeces of a very
indigestible reference substance is determined. The substance may be a
natural constituent of the feed or to be added to it, or both. Substances pre-
viously used for this purpose have been ferric acid, chromic oxide (also
called chromic sesquioxide or chrome green), lignin, silica, crude fibre, fae-
cal nitrogen and chromogen, a naturally occurring plant pigment.
The basic idea of this method is, that the digestibility is calculated
from the relation between the nutrients and the indicator substance in the
feed and in the faeces. The digestion coefficient is computed by using the
change in the ratio of each nutrient with reference to the special indigestible
substance in the feed and in the faeces. An unabsorbable material is used
as an index, that is, a substance which is recovered quantitatively in the
faeces.
Digestibility of a specific nutrients can be calculated using the follow-
ing equation:
Cfaeces—Cfeed
DC= ————————-——- × 100
Cfaeces
where: DC=(apparent) digestion coefficient, %; Cfaeces=concentration

of indicator in faeces, related to the nutrient to be tested and Cfeed=concen-
tration of indicator in feed, related to the nutrient to be tested.
For example. if the concentration of indicator in feed dry matter is 1%
and 1.2% in the faeces dry matter and the ingested feed contains 20% crude
protein in dry matter and the excreted faeces has a crude protein level of 10
in dry matter, then the digestion coefficient of crude protein is 58.3%
(1.2/10-1/20)/(1.2/10)×100=
=(0.12-0.05)/0.12×100=(0.07/0.12)×100=58.3).
Required characteristics of indicator substances
-innocuity to animals,
-it should not modify digestive processes
-total insolubility (indigestibility), or invariable knownabsorption rate,
-homogenous distribution in feed,
112
- analytical tracebility.
Commonly used indicators are: the added chromium-oxid (Cr2O3),
iron (III)-oxid (Fe2O3), barium-sulphate (BaSO4), poly-etilen-glycol (PEG), as
well as silica (SiO2), acid insoluble ash (AIA), lignin and plant pigments com-
ponents of feed.
7.1.3. Digestibility by difference (indirect determination method)

In the tirals decribed above, digestion coeffients were determined while the
feed in question was fed exclusively. The determination of the digestibility of
feedstuffs that cannot be fed alone (e.g. low fibre cereals for rumints, fish
meal for pig etc.) is somewhat more complicated. The trials of digestibility
by difference has three types.
a) Simple trial of digestibility by difference: in an experiment the feed
in question is fed with a base feed and their digestibility is determined by
difference. The difference is calculated and is assumed to be digested from
the unknown feed. The usual method is to conduct two trials. In one trial
the digestibility of the base diet (e.g. roughage) is determined and in the sec-
ond the digestibility of a combination of the test feed (e.g. corn) and the
roughage is determined. In this procedure the digestibility of a roughage as
a basal ration is usually determined first and then the feed under consider-
ation is added to the roughage for the second test. In this ssubsequent trial
the same basal feed plus the new feed are fed together. The total faecal out-
put is measured as usual. Then the assumption is that the nutrients in the
original basal ration show the same percentage digestibility as they did in
the first trial when it was fed alone and the amount of nutrients in the fae-
ces in the second digestion trial is presumed to belong to the basal portion
of the ration. The remaining faecal output is assumed to have come from the
feed being tested. By considering the difference between tha values obtained
for the roughage alone and for the combination, coefficients for the
digestibility (DC) of the other feed are calculated by difference.
(M-B)
A= ——————————- + B
a
where: A= DC of added (to be tested) ) feed, %,
M= DC of feed mixture (base+added), %
B= DC of base feed, %,
a= proportion of added feed in the mixture, %.
The lower is the proportion of the feed to test the more experimental
animal should be involed in the trial to achieve acceptable accuracy.
b) Regression method is used when after having determined
digestibility of base feed, the feed in question is added in different concen-
trations (multiple substitution). Based on a linear regression between the
amount of ingested nutrients from the test feed and the digestibility of feed
mixture the digestion coefficients are calculated. This methods is used when
adding more than 30% to the base feed would be unhealthy. The equations
113
for calculation are as follows: y= a + bx, where
nutrient intake from feed II

y = —————————————————————
digestible nutrient in the whole mixture
nutrient intake from feed I

x = —————————————————————
digestible nutrient in the whole mixture
Digestion coefficient of the nutrient in question in feed II= 1/b and in

feed II -b/a.
c) Digestibility by difference using reference material. In the first trial
digestibility of a mixture of the base diet and a reference material is deter-
mined. Reference material has a known or 100% digestibility, for example
starch. In the second trial the feed to test is added in the account of refer-
ence material. Advantage of this method is a unchangeable dry matter
intake.
7.1.4. Determination of the true digestibility

For a more accurate calculation there is a need for the knowledge of true
digestibility data. To obtain these the metabolic portion of faeces should be
measured. Pepsin-hydrochloric acid soluble faecal protein (N) is assumed to
derives from the body. Exretions in animals deprived of feed or fed on pro-
tein-free diet reflects metabolic faecal losses, and the use of isotopes is also
possible.
7. 2. Factors affecting digestibility

Factors influencing digestibility, may be classified into two groups, those,

related to the living organism (endogenous), and those of the environment
(exogenous).
7.2.1. Endogenous factors
7.2.1.1. Species of animal. Digestive capacity of animals significantly differs
in different animal species. The main reason for this is the structure and
function of the digestive system. The largest species differences can be
found in case of the vegetable fibre. Omnivore and carnivore animals digest
fibre much less than ruminants. Between these value are the caecum and
colon fermenter animals (horse): FIGURE VII-1.
114
FIGURE VII-1: Relation between cellulose digestibility and body weight (after ENGELHARD et
al. 1985)
High protein and low fibre feed are better utilized by monogastrics
than by ruminant because in the rumen some of the nutrients undergo
microbial degradation and this means some nutrient losses (e.g. in form of
methan and ammonia). Table VII-2 gives the digestibility data of a meadow
grass.
Table VII-2: Digestibility of organic matter and fibre in meadow grass, %
Gastrointestinal tract is very similar in the different ruminant

species. Differences in digestion (more effective digestion of goat and deer)
derive from the varying ratios of rumen volume to body size. Within a
species there are only tiny (1-2 percentage units) differences owing to breed,
lines and types.
7.2.1.2. Age. Monogastric suckling animals generally have lower starch and
protein digestibility (except mother milk’s), having a special set of digestive
115
enzymes and only gradually establishing gut microflora. In ruminant there

is even more difference between young, functionally monogastric animal
(newborn calf, lamb) and the older, already ruminant one (3-5-month-old
calf or lamb).
Digestion of adult animals in terms of digestive physiology (2-month-
old piglet, 3-4-month-old calf, half-year-old foal, 8-9-week-old rabbit) prac-
tically does not change as a function of age, except for the very old and gen-
erally weak animals’ decreased digestive capacity.
Precocial (nest leaving) birds (e.g. chicken) hatch with mature diges-
tive system, whereas altrical (nest living) species (pigeon, parrots) at begin-
ning should be fed on crop milk or predigested material.
7.2.1.3. Movement. Physical activity or exercise may delay stomach empty-
ing, fasten gut peristaltic movement, but its overall effect on digestion is
small.
7.2.1.4. Habituation. After adjustment some feed are better digested,
because both the rumen and intestinal microbe population and the host
enzyme production adapt. For example the raw starch digestibility in rat
improved from 43% up to 59% during a month.
7.2.1.5. Climate. There are some publications reporting about a less effec-
tive digestion in winter of sheep, probable owing to the cold environmental
temperature.
7.2.1.6. Caecotrophy. Physiological coprophagy (caecotrophy) may modify
protein digestibility in rabbits fed on low protein-high fibre diet. In this case,
if caecotrophy is inhibited (e.g. by means of a collar), the measure of protein
digestibility may fall by up to 30 %. This can be explained by the fact that
the rabbit regulates caecotrophy intake according to [protein minus fibre]
the level of the daily ration (FEKETE, 1985). The daily caecotrophy excretion
practically is stable.
7.2.1.7. Physiological state. In the second half or last third of gestation
digestion coefficient generally decreases. This is due to an increased activi-
ty of hind gut glands to compensate slower gut peristalsis by a higher water
content of faeces. Digestion of lactating-suckling animals first improves,
then, owing to the elevated dry matter intake, generally decreases.
7.2.1.8. Gender. There are no significant differences between the digestion
coefficients of different sexes. Thus, in practice there is no need to take it
into consideration (This is the body composition, which is highly influenced
by gender.)
7.2.1.9. Stress. Although long-lasting (1-2 weeks) stress situation may
decrease digestion coefficients by 1-2 percent unit, its main effect is rather
the change of sodium to potassium ratio in faeces and caecotrophy (FEKETE
et al. 1988).
116
7.2.2. Exogenous factors

7.2.2.1. Environmental temperature
Environmental temperature does change digestibility of nutrients. Rabbits,
kept at 18oC, had digestion coefficient higher by 2-7 percentage units than
rabbits kept in heat stress conditions (34oC), the relative humidity being the
same 65%. If the environmental temperature is less than 20oC, efficiency
digestion of dairy cows worsen. A fall of 10oC is reflected as a 1.8 percnt
unit decrease in organic matter digestibility (NRC, 1978). Data of nutritive
value in NRC (1984) tables are valid only within thermoneutral zone and
according to the actual environmental temperature correction that has to be
made:
B= B + B(0.001(T-20), where
A = corrected (adjusted) value
B = base value
T = environmental temperature, oC.
So, for instance, on 0oC digestibility will fall by 2%. Cold fastens
digesta passage time decreasing available digestion and absorption time.
7.2.2.2. Feeding level
The higher the feed intake, the lower its digestibility. This effect is especial-
ly pronounced in ruminants, where the daily dry matter intake may vary
over a wide range. Increasing feeding level will primarily affect digestibility
of n-free extrac and fibre (see role of discount factor in computing NEl
value).
7.2.2.3. Composition of feed mixture or daily ration
Increasing protein levels – in a continouesly decreasing fashion- will
improve protein (and in some cases fat, carbohydrates and fibre) digestibil-
ity. This phenomenon is referred to as the lift of digestion depression, which
means that the extra protein (nitrogen) serves to supply rumen and gut
microbes. In pigs, from the chemical composition of feed, deviations in pro-
tein digestibility can be put down to protein in 25.5%, to fibre in 27.1% and
to N-free extract in 39%.
Increasing fat/oil level in feed mixture of ruminants ma lead to a fall
in fibre digestion. From this aspect low melting point oils are more effective,
encapsulating fine feed particles preventing cellulolytic bacteria from
attaching to them.
Vegetable fibre significantly decrease digestibility of each major nutri-
ents, including certain fibre fractions. The extent of decreasing effect basi-
cally depends upon the species of animal. Thus, 1% increase in fibre level
will decrease digestion coefficient of organic matter by 0.85% in ruminants,
by 1.6% in monogastric mammals and by 2.3% unit in poultry.
N-free xtract (N-f.e.). High sugar and starch level through lactic acid
production may upset intestinal eubiosis, balance of ruminal microbes,
with a concomitant decrease in protein, fibre and carbohydrates digestion
coefficients. This digestive depression may be important primarily for rumi-
117
nants.
Water content practically does not affect digestibility. The mildly dried
hay has the same digestion coefficients as the original wet grass. (Water
content will influence dietetic effect of feedstuffs and feed mixtures.)
7.2.2.4. Biologically active substances, feed additives
Antinutritive substances (e.g. trypsine inhibitor, lectines etc.) decrease pro-
tein digestibility. Salt, feed flavours increase voluntary dry matter intake,
but have no influence on digestibility. Surface active materials (e.g. bolus
alba, medicinal charcoal, bentonites etc.) generally decrease digestibility
because they are able to bind and eliminate nutrient molecules from body.
Purgatives primarily decrease fat and to less extent digestibility of
proteins. By means of commercial enzyme preparation digestibility of pro-
teins (proteases), N-free extract (glucanases) and fibre (cellulose) can be
increased. Thyreostatic agents, like ammonium perchlorate by slowing
down digesta passage rate may improve digestibility. Copper sulphate (espe-
cially in pig), antibiotics and antimicrobial substances (carbadox) – prima-
rily by means of influencing gut flora – improve organic matte (in the first
line protein) digestibility.
7.2.2.5. Feed processing
Ruminants generally chew thoroughly their feed, thus chopping of feed-
stuffs hardly has any effect on digestibility. In contrast, grinding of hard or
small grains (corn, sorghum, milo) will improve digestibility. Too fine grind-
ing of green meals (grass, alfalfa meal) may lead to digestive depression,
because small feed particles pass rumen and gastrointestinal tract very
quickly.
Healthy horses chew thoroughly; hence crushing oats does not mean
an advantage for them. In case of sick horses or individuals with tooth
decays crushing may improve digestibility. Crushing or coarsely grinding
may also have a positive effect if horses are fed from the same manger and
they usually chew quickly and superficially competing with one another.
Grinding of feed has no effect on digestibility at rabbits, but may worsen
dietetic effect.
Digestion coefficients of grounded corn and milo organic matter in pig
are by 7 and 29 percent unit higher than those of whole grain. Even grind-
ing of coarsely ground barley meal may improve digestion coefficients when
fed for pigs.
In case of poultry grinding of grains or seeds in general has no effect
on digestibility, because grains and seed become sufficiently soft in the crop
and gizzard for subsequent digestion. Fine grinding of green meals from 4
mm of particle size to 1 mm did improve digestibility.
Well chosen and executed heat treatment (drying, boiling, cooking,
extrusion, toasting, infra red treatment, use of autoclave etc.) by inactivat-
ing heat sensitive antinutritive compounds and pre-treating of starch do
improve digestibility. Overheating, by means of Maillard reaction may
118
decrease the digestibility of protein, N-free extract and fibre.

Soaking generally has no effect on digestibility with the exception of
dog and coypu, for them soaking improve the digestion coefficient of starch
(effect of amylase). Deep cooling does not influence digestibility. Effect of
pelleting depends upon its method of realization (time and temperature).
Ensiling forages – except the first phase when readily fermentable and
therefore potentially digestible carbohydrates are consumed – have no effect
on digestibility. Mixing feedstuff may have an effect on digestibility if
extreme associative effects occur. Alkali or ammonia treating of straws, sac-
charifying wooden materials by calcium hydrosulphite significantly increase
digestibility of fibre, especially for ruminants and horses.
Fibre digestibility can be improved by the application of SFF process
The solid state fermentation (SSF) is a new version of the old Chinese „koji”
process. The Alltech SSF process uses selected Aspergilus and Rhizopus
fungi. The main changes during the SSF process are the convesrsion of
cellulose to glucose, the increase of DE and ME value and an improve of the
availability of phosphorus. Processed fibrous by products (e.g. wheat bran)
get appropriate to be included into the pig and poultry diet. Using Allzyme
SSF® in poultry, improved the N digestibility and P availability; at the
finisher pigs increased performance parameters by increasing energy and P
availability (RUTZ and RIGOLIN, 2008).
7.2.2.6. Feeding technique
Feeding frequency or change of feeding order of the daily ration different
component generally has no effect on digestibility. Also, there is no effect of
chopping and feeding in manger ad libitum when compared with grazing.
7.3. The concept of ileal digestible amino acid and ideal pro
tein in swine and poultry nutrition
© Laszlo Babinszky
7.3.1. Digestibility of amino acids in pigs
With the need of more accurately meeting the amino acid requirement of
pigs the study of ileal digestibility of amino acids has become an important
area of research in the past fifteen years. The digestibility of dietary proteins
and amino acids used to be characterized with the apparent faecal
digestibility coefficient - similarly to other nutrients. However, the findings
of digestion-physiology research works prove that the intestinal flora in the
colon simultaneously synthesizes and catabolizes protein. This is the rea-
son why the faecal digestibility of dietary proteins will in some cases under-
estimate, in others overestimate the real value.
Consequently, ileal digestibility of proteins and amino acids is used in
many countries. This method may seem to have a disadvantage in that it
does not take into account the amount of amino acids absorbed from the
119
large intestine. The results of relevant studies show, however, that this is
only an apparent source of error, as the various nitrogen bonds are
absorbed from the postileal section (colon) almost exclusively in the form of
ammonia, thus they do not participate in protein synthesis and are simply
excreted with the urine. From the point of animal nutrition therefore the
amount of amino acids absorbed until the end of the small intestine (ileum)
is important only.
DIFFERENCES AMONG APPARENT, STANDARDISED AND TRUE ILEAL DIGESTIBILI-
TY
It is generally accepted that the apparent ileal amino acid digestibility coef-
ficients are dependent on the amino acid content in the assay diet.
Apparent ileal amino acid digestibility inrease curvilinearly with increasing
amino acid content in the test diet.
The word „apparent” refers to the fact that the coefficients are not
adjusted for endogenous nitrogen and amino acid losses. The limitation to
the use of apparent ileal amino acid digestibility is that digesta collected at
the end of the small intestine contains large quantities of endogenous pro-
tein. As illustrated in FIGURE VII-2, the endogenous amino acid losses can
be separated into a basal (minimum) and an addicional specific loss.The
basal loss is non specific and related to dry matter intake. Howewer, the
specific loss is related to inherent factors in feedstuffs, e.g. fiber and anti-
nutritive factors etc. (PACK et al. 2002).
FIGURE VII-2: Origin of amino acid losses at the terminal ileum
The amounts of basal endogenous protein or amino acid losses in

ileal digesta can be determined by different methods such as feeding protein
free diet, feeding diets containing protein sources that are assumed to be
100% digestible with complete absorption of amino acids and the regression
technique. If corrections are made for basal endogenous amino acid losses,
the standaardised ileal amino acid digestibility coefficients (SDC) can be cal-
120
culated with the following equation:
AA intake–(AA excreted–basal endogenous AA)

SDC, % = —————————————————————— × 100
AA intake
Standardized ileal protein and amino acid digestibility has the advan-
tage over both apparent and true digestibility in that it represents a funda-
mental property of the individual feedstuffs, namely standardized digestibil-
ity values include any variation of the endogenous fraction related to the
feedstuff itself. The effect of dietary amino acids on their respective appar-
ent, standardized and true ileal digestibility coefficients can be seen is
FIGURE VII-3.
FIGURE VII-3:Changes of apparents, standardized and true ileal amino acid digestibilities
as a function of dietary AA intake
7.3.2. Determination of ileal digestibility with different techniques

There are several methods for measuring the ileal digestibility of amino
acids. The majority of these methods require surgical operation prior to the
start of the trials to obtain digesta (chyme) from the terminal ileum from
which to determine the ileal digestibility.
Ileal digestibility studies without cannulation techniques
a) Post mortem digestibility studies: Before the expansion of cannulation
techniques the only possible means for determining the amount of nutrients
absorbed from the intestinal sections was the analysis of digesta samples
collected from the intestine after the trial animals were slaughtered. The
disadvantage of this method is, that the animals can be used in a single trial
only. As for the reliability of the trial results the biggest shortcoming of the
method is, that the electric shock or other impulse occurring at the moment
of killing will cause a shedding of mucosa in the intestine to an extent that
it may provide incorrect information about the digestibility of proteins and
amino acids. For this reason the method is used extremely rarely by now.
121
b) Ileo-rectal anastomosis (IRA):

The process based on ileo-rectal anastomosis allows total digesta collection
without the insertion of a cannula. With this surgical method the end of the
ileum is connected directly
to the rectum with the
removal of the cecum and
colon (FIGURE VII-4) and
thus the digesta from the
ileum is excreted through
the rectum and can be col-
lected quantitatively. A
doubtless advantage of the
process is the possibility of
collecting digesta directly,
but it has the disadvantage,
that by removing a part of
the intestinal tract the
nutrient and vitamin supply
of the body suffers and also
the electrolyte balance can
be easily upset.
FIGURE VII-4: Stages in the sur-

gical procedures used to estab-
lish an ileorectal anastomosis in
pig. A) caecum removed; b) colon
cut; c) ileum, including ileo-cae-
cal valve connected to the cae-
cum
122
Ileal digestibility studies with different cannulation techniques

a) Simple T-cannula method: the simplest method for representative digesta
collection is enabled by the so-called T-cannula, for the application of which
several research teams have developed various procedures (FIGURE VII-5).
A common characteristic of all procedures is that they necessitate the use
of an indicator (marker), because the T-cannula is not suitable for quanti-
tative digesta collection.
FIGURE VII-5: Simple T-cannulatiopn technique in pig
The collection of the digesta at regular intervals (at least 4 times daily) dur-
ing the collection period is enabled by a polythene bag attached to the can-
nula.
b) Re-entrant cannulation: This procedure allows quantitative collection of

the ileal digesta. The method essentially involves the insertion of an ileum-
ileum or ileum-caecum cannula in the animals subject to the purpose of the
study (FIGURE VII-6). These cannulas can be perceived as an artificial
intestinal section with a valve, leading the ileal digesta outside the body per-
mitting its collection and quantitative weighing; following which the entire
digesta or part of it is returned to the intestinal tract. Disadvantages of the
method are the complexity of the required surgery, and also the risk of can-
nula blockage, which may primarily be attributed to inadequate intestinal
peristalsis at the cannula insertion point and/or to the high fibre content of
the test diets.
123
FIGURE VII-6: Re-entrant cannulation technique in pig
c) PVTC (post valve T-cannula) method: The procedure was developed by

LEEUWEN et al (1988). With this surgery operation a specially shaped T-can-
nula is inserted in the caecum, upon the opening of which the end of the
ileum is sucked into the cannula by the vacuum generated and with the
control of the ileo-caecal valve the entire digesta is voided outside. When the
cannula is shut the digesta passes through the gut without interruption. A
major advantage of the method is, that the necessary operation is relative-
ly simple, the peristalsis of the small intestine is undisturbed and it allows
quantitative collection (FIGURE VII-7).
FIGURE VII-7: PVTC (post valve T-cannula) technique in pig
The digesta is collected in a polythene bag attached to the cannula.
124
d) Mobile bag technique: PETRY and HANDLOS (1978) were among the first to
apply the mobile bag (or nylon bag) technique in the evaluation of swine
feeds. The ileal digestibility of crude protein and amino acids can be meas-
ured with the improved PVTC procedure. In this method the test feed sam-
ples following their in vitro or in vivo incubation are placed in small bags
and introduced into the GIT through the duodenal cannula, and are then
collected with the digesta which is voided through the PVTC cannula. The
bags are cleaned, weighed, and then their homogenized content is analyzed.
Advantages are the rapidity, accuracy and relatively small substance
requirement of the procedure. Another important benefit is, that already at
the initial stages of the e.g. plant selection work - when the amount of test
substance is still very limited - the digestibility of protein and/or AA of a
given line or cross combination can be determined with quite high accura-
cy. A notable disadvantage of the method is, that due to the small amount
of test substance remaining in the bag, nutrients which require relatively
large samples for their quantitative assay (e.g. Weende analysis) can not be
determined with this procedure.
Important characteristics of studies based on the cannulation technique

Table VII-3 summarizes the more important features of study methods using
various cannulation techniques on the basis of work carried out by the
University of Kaposvár, Department of Animal Nutrition.
Table VII-3: Important characteristics of methods based on different cannulation tech-

niques (weaned piglet, growing and finishing pig) (TOSSENBERGER and BABINSZKY, unpub-
lished data)
The data in Table VII-3 show, that the implantation of the simple T-
cannula requires the shortest time for surgery (20-25 minutes / animal),
and due to the implantation of the two cannulas (duodenal and PVTC can-
nula) the surgery preparation of the mobile bag technique lasts the longest
(40-45 minutes/animal). Post-operation recovery is almost of the same
duration, 5-8 days, except for the mobile bag technique. The length of the
adaptation period is 6-7 days, irrespective of the surgery method. There is
125
a difference in the length of the collection period, however, as it changes

between 2 and 4 days, subject to the method applied. In addition there are
also significant differences in the frequency and duration of daily digesta
collections.The animals are housed individually in metabolic cages during
the adaptation and collection periods of these studies as well. The ileal
digestibility of nutrients (e.g. protein or AA) is calculated in accordance with
the contents of Sections 7.1.1. and 7.1.2.
In summary it can be concluded, that all procedures based on can-
nulation techniques provide reliable data about the ileal digestibility of
dietary proteins and amino acids. When selecting the method, however, the
composition of the test diet, the age of the animal, and the skill of the team
conducting the study need to be taken into account.
THE DIFFERENCE BETWEEN FAECAL AND ILEAL DIGESTIBILITY

When evaluating the digestible amino acid content the question arises,
whether ileal digestibility provides a more precise information about the
digestibility of amino acids than the data based on faeces collection. Studies
show, that there is a considerable difference for instance between the faecal
and ileal digestibility of the crude protein, lysine, methionine and cystine
content of full fat soya, which difference should definitely be considered in
the diet formulations (Table VII-4).
Table VII-4: Digestibility of crude protein and amino acid content of full fat soya in growing pigs (BABINSZKY,
2002)
Overall it can be concluded that the digestibility measured along the

full length of the intestinal tract (faecal digestibility) shows a higher value
than ileal digestibility does. The difference between the two sets of data is
primarily attributable to the energy supply of the colon microflora, because
when energy is the limiting factor in the colon, the microorganisms utilize
the indigestible protein content of the digesta as a source of energy, and
consequently the apparent faecal digestibility of protein will be higher. If
there is no energy deficiency in the colon, the microbes produce bacterial
protein from the N compounds of the digesta by way of de novo synthesis.
Thus the protein excretion via faeces will increase and the digestibility coef-
ficient will be lower.
126
Imperfectly conducted feed processing procedures may result in

major changes of the digestibility of crude protein and amino acids. Results
of growing pig trials from the Netherlands show, that while the faecal
digestibility of crude protein, lysine, methionine and cystine in soybean
meal did not reflect the defects of toasting (the digestibility of crude protein
and amino acids was the highest in the over-treated feedstuffs) its effect on
ileal digestibility was obvious (Table VII-5).
Table VII-5: Effect of toasting on the faecal and ileal digestibility of crude protein, lysine,
methionine and cystine in soybean meal (%) (van WEERDEN, 1985)
The data of the Dutch work in reference and of other studies as well prove
that when the objective is the evaluation of the quality of dietary protein, the
measurement methods based on ileal digestibility are considerably more
sensitive than those based on the collection of faeces.
To summarize the information on the ileal digestibility of amino acids the

following conclusions can be drawn:
127
Application of the ideal protein concept in swine nutrition

The necessity of applying the so-called ideal protein concept both in diet for-
mulation and in the substitution of protein sources has become increasing-
ly important in recent years. The development of the “ideal protein” theory
for swine has been a subject of work in the UK already since the early eight-
ies. Those studies, however, were still based on the total amino acid require-
ment of the animals. Still earlier, several authors already expressed the
amino acid requirement of pigs by listing the necessary amount of each
amino acid as a percentage of lysine. LOEN (1966) was among the very first
to determine the total lysine requirement of fattening pigs in relation to the
starch value, and to specify the amount of the other amino acids in per-
centage of lysine.
In the meantime an adequate database was generated for the ileal
digestible amino acid content of various feedstuffs as well, and consequent-
ly the concept of ideal protein could also be modified, i.e. it could be made
more accurate. LANGE (1992) highlighted the fact, that when the ileal
digestibility of amino acids is used in the calculation, the profile of ideal pro-
tein is partially altered. The biggest change is seen in the case of threonine
(Table VII-6). This can be explained by the fact, that the threonine portion of
ileal endogenous protein is relatively high. The high level of endogenous
threonine is the reason why the digestibility of threonine measured at the
end of the small intestine remains significantly below the ileal digestibility
of lysine.
Table VII-6: Essential amino acid profile in ideal protein (lysine: 100 %) based on the total and ileal digestible
amino acid content (WANG and FULLER, 1989)
The amino acid profile of ideal protein differs in maintenance from

that in growth (production). According to the relevant studies the ideal pro-
tein needs to contain a higher portion of methionine and cystine, and also
threonine and tryptophane to meet the amino acid requirements for main-
tenance, compared to the needs of weight gain (Table VII-7). The practical
implication of this is, that the ratio of these amino acids should be increased
in the ideal protein in order to cover the amino acid requirement of mainte-
nance, because with the increase of live weight also the protein and amino
acid requirement of maintenance increases.
128
Table VII-7: Essential amino acid profile in ideal protein (lysine: 100 %) for maintenance
and weight gain (HENRY, 1993)
With the purpose of meeting the amino acid requirements more accu-
rately, BAKER and CHUNG (1992) proposed to specify the amino acid compo-
sition of ideal protein for three different body weight categories (between 5
and 100 kg). The recommendations are summarized in Table VII-8.
Table VII-8: Percentage of essential amino acids in ideal protein (lysine: 100 %) for pigs in
the growing and fattening stages (BAKER and CHUNG, 1992)
In addition to weaned piglets and growing-finishing pigs, the litera-

ture also provides recommendations for the percentage composition of
amino acids in the diet of pregnant and lactating sows (Table VII-9). These
recommendations should be taken into account in the diet formulations.
129
Table VII-9: The ideal pattern of amino acids during pregnancy and lactation (CLOSE,
1995)
7.3.4. Application of ileal digestible amino acids in the formulation

of swine diets
In the field of practical feeding it is also important to know the relationship
between the fattening performance and the method of diet formulation, i.e.
whether the diet was formulated on the basis of faecal or ileal digestibility
of the crude protein. According to Belgian work there is a very strong cor-
relation between the ileal digestibility of crude protein and the weight gain
and feed conversion rate (FCR), while the strength of relationship between
faecal digestibility and the two parameters mentioned is only medium or
even looser (Table VII-10).
Table VII-10: Correlation between the ileal and faecal digestibility of dietary crude protein
and the weight gain and FCR of growing-finishing pigs (DIERICK et al. 1987)
These data allow the conclusion, that the amino acid requirement of
growing and finishing pigs can be satisfied better if the requirements are
specified as ileal digestible amino acids. There has been an increasing num-
ber of publications reporting of such studies in latter years. As for the diet
formulation it should also be noted, that the ileal digestible amino acid con-
tent represents an advantage over the faecal digestibility data only in case
the diet does not consist of corn and extracted soybean meal exclusively,
because when the amino acid requirements of pigs were determined, the
trial animals were fed with a corn and soybean meal based diet (NRC, 1998).
In all other cases (e.g. when feeding multi-component diets, or in the sub-
stitution of protein sources) using the data based on ileal digestibility may
yield practical benefits also in the diet formulations.
130
Another important fact to know in the formulation of swine diets is,

that also the energy supply of the animals is crucial for the adequate uti-
lization of amino acids. A multitude of relevant trial findings prove that the
growth performance and the chemical composition of the carcass (the pro-
tein : fat ratio; i.e. the quality of the meat) is influenced not only by the
genetic potential, but also by the amino acid : energy ratio of the diet.
As lysine is generally the first limiting amino acid for pigs, all efforts should
be made in the diet formulations to achieve the best possible lysine : ener-
gy (digestible energy, DE) ratio in the interest of enhancing protein deposi-
tion. The trial results prove, that during the first phase of fattening (25-60
kg) the lowest level of fat deposition can be expected in the case of an 0.6 g
ileal digestible lysine/MJ DE ratio. In the second phase of fattening (60-105
kg) this ratio decreases to the level of 0.5 g ileal digestible lysine/MJ DE
(BABINSZKY, 2002).
THE THEORETICAL BASIS FOR SUBSTITUTING PROTEIN SOURCES

The need to replace the protein components of pig diets with alternative pro-
tein sources has been considered more than once during the last decade.
The demand for substitution is attributable to numerous causes (economi-
cal, animal health, certain traditional local feeding habits, etc.). The earlier
basis for substituting protein sources was the so-called protein equivalence.
According to the results of the latest research work however, the substitu-
tion of protein sources is now made on the basis of the ileal amino acid con-
tent of dietary proteins.
When replacing protein sources care should be taken to maintain the
appropriate lysine/digestible energy ratio and to include in the diet all other
amino acids important for the pig at levels which meet the ideal protein con-
cept. In this section we present a model- calculation for substituing protein
sources in pig nutrition. As there are different recommendations for the
composition of the ideal protein, we decided to use in the diet formulation
the NRC (1989) requirements which represent an average value.
In case we intend to replace a protein source of animal origin (meat and
bone meal, mixed animal protein meal, etc.) with a vegetable protein source,
we should focus not only on the difference in the nutrient contents of the
protein source (e.g. ileal digestible amino acid), but also on the difference in
their phosphorous content (FIGURES VII-8, 9 and Table VII-11).
131
FIGURE VII-8: Ileal digestible lysine content of some protein sources for pigs (NRC, 1998)
FIGURE VII-9:Ileal digestible methionine plus cystine content of some protein sources for
pigs (NRC, 1998)
According to the data of FIGURES VII-8 and 9 there is a substantial differ-

ence between the ileal digestible lysine, methionine and cystine contents of
the various protein sources. The proper method to follow in the replacement
therefore is to calculate with these amino acid data in the diet formulation.
132
Table VII-11: Total and digestible phosphorous content of some important animaland veg-
etable protein sources (CVB, 1999; DLG, 1999)
As it can be seen from Table VII-11 the total and digestible phospho-
rous content in animal protein sources is substantially higher than in the
vegetable protein sources. This warns us, that when replacing animal pro-
tein sources with vegetables ones it is essential to provide phosphorous
supplementation also. The question arises whether phosphorous supple-
mentation should be based on total phosphorous or on digestible phospho-
rous. The latest research results indicate that the phosphorous require-
ments of pigs can best be satisfied when it is determined as digestible phos-
phorous. In case of a vegetable phosphorous source it should also be con-
sidered, that most of its phosphorous exists in the form of so-called phytate
bonds, and thus only a very limited portion of it is available to pigs.
Consequently, when we wish to replace an animal protein source with a veg-
etable one instead of adding extra phosphorous to the diet we should incor-
porate phytase enzyme in it. According to our own studies and on the basis
of international data the recommended level of phytase enzyme supplemen-
tation for the corn - soya based diet of fattening pigs should be 500 U/kg
diet (BABINSZKY, 2001).
FEED FORMULA ALTERNATIVES FOR SUBSTITUTING PROTEIN SOURCES

Bearing in mind the general rules discussed above we performed some
model calculations for the substitution of various protein sources. In the
preparation of feed formula alternatives run on computer, both for Phase 1
(20-60 kg) and for Phase 2 (60-105 kg) diets, we first determined the ileal
digestible (ID) lysine/DE ratio, which as mentioned in this Section is 0.6 g
ID lysine/MJ DE between 20 and 60 kg, and 0.5 g between 60 and 120 kg,
and next, using the NRC (1998) requirements, we determined the ileal
digestible methionine, cystine, threonine and tryptophane levels relative to
the lysine level.
Phytase enzyme (Natuphos) was incorporated in the concentrate due
to the reasons already discussed. It can be seen in Table VII-12 that during
133
the first phase of fattening (20-60 kg) we substituted the meat meal in the
concentrates partly with extracted soybean meal with 48 or 46 crude pro-
tein content, or with full fat soya, peas, lupine, sunflower seed meal or dou-
ble zero rapeseed, so that the energy, protein and digestible phosphorous
content of the diets should be identical and the ratio of amino acids to lysine
should follow the profile of the ideal protein. Diets for the second phase of
fattening (60-105 kg) were formulated on a similar basis (data are not
shown).
Table VII-12: Percentage composition and calculated nutrient content of diets (20–60 kg)
(BABINSZKY, 2001)
* ileal digestible
134
Fattening trials
The question is, that if protein sources are substituted according to the con-
cept discussed above can we expect the performance and slaughter quality
of our animals not to deteriorate. In order to investigate this issue we joined
forces with the Wageningen University Department of Animal Nutrition to
conduct an individual model fattening trial involving 100 animals, and in
this trial we substituted a part of the extracted soybean in the diet with
peas, sunflower seed and fish meal (Table VII-13).
Table VII-13: Composition (%) and nutrient content of diets (SZABÓ et al. 2000)
*Barley, wheat, molasses, and supplements
The results of the study show, that when the substitution of protein sources
is made in accordance with the concepts discussed above, no adverse
changes in the fattening performance and quality of meat should be expect-
ed (Tables VII-14, 15).
135
Table VII-14: Weight gain and feed conversion rate of the trial groups (SZABO et al. 2000)
Table VII-15: Slaughtering parameters of the trial groups (SZABÓ et al, 2000)
1 intramuscular fat
2 pH value in the musculus longissimus dorsi 45 minutes after slaughtering
3 pH value in the musculus longissimus dorsi 24 hours after slaughtering
Summarizing the data of the literature and the results of the studies
here presented it appears, that the animal protein source in the pig diets
(bone and meat meal, fish meal, etc.) can be successfully substituted with
vegetable protein sources provided this substitution is made on the basis of
the ileal digestible amino acid content of the feed ingredients and in compli-
ance with the ideal protein concept. Consequently we first need to determine
the digestible energy content (DE) of the diet to be fed, and then select the
appropriate ileal digestible lysine/DE ratio. Finally the ratio of lysine and
the other ileal digestible amino acids should be determined in accordance
with the ideal protein concept. It is essential to provide phosphorous sup-
plementation in the diets. In order to meet the phosphorous requirements
more accurately the digestible phosphorous content of the feed ingredients
should be used in the calculations. When a vegetable protein source is used
the recommended level of phytase supplementation is 500 U/kg of diet. If
the substitution is implemented in accordance with the foregoing sugges-
tions, we need not expect any decrease of production or deterioration of the
meat quality.
7.3.2. Digestibility of amino acids in poultry

7.3.2.1. Determining the digestibility of amino acids with different methods
136
Nutritionists already in the sixties were concentrating on the question of

how to determine the digestibility of amino acids in poultry diets and meet
more accurately the amino acid requirements of poultry. No significant
progress has been made in this field in the last 30 years, and only a rela-
tively small number of papers have been published. The development of
cannulation techniques, however, seems to offer a new approach for solving
the problem. In the past decades the various recommendations and require-
ment levels applied in the diet formulations used to be calculated on the
basis of total amino acid content in the field of poultry nutrition as well. On
the basis of research results it has also become apparent, that the amino
acid requirement of poultry could be best satisfied, if - similarly to swine -
the calculations were made with the so-called available amino acid content
of the diets.
The availability of each amino acid is that part of the total amount in
the diet which is actually supplied to the sites of protein synthesis.
However, the current methods for assessing the amino acid availability of
feedstuffs for pigs and poultry have drawbacks:
The recent results show that the use of digestibility values in practical
diet formulation is the closest to a system based on availability.
There are several methods available for determining the digestibility of
amino acids in poultry.
A) Method based on the collection of excreta (dropping digestibility)

One of the most frequently used digestibility studies is the method based on
the collection of excreta. When this technique is applied (FIGURE VII-10),
the excreta voided through the cloaca is collected quantitatively, and then
chemical methods are used to separate the faecal and urinary nitrogen con-
tent of the excreta from each other. An apparent disadvantage of the method
is, that its accuracy is limited, furthermore it does not take into account any
potential bacterial activity in the caecum and colon.
137
FIGURE VII-10: Dropping digestibility (intact bird)
B) Method based on the removal of the caeca (caecectomyzation)

In the application of this method the paired caeca of the birds are removed
eliminating thereby the potentially distorting effect of bacterial activity tak-
ing place in the caecum (FIGURE VII-11). The surgery is performed in anes-
thesia. A disadvantage to be noted is, that after the caeca are removed, bac-
terial activity may still be present in the remaining caecal stump. A similar
problem should be expected in the colon as well. Moreover it also necessi-
tates the assumption that the amino acid content of the urine is negligible.
The application of the method therefore requires numerous assumptions to
be made before the digestibility data can be accepted.
FIGURE VII-11: Dropping digestibility (caecectomised bird)
C) Methods based on cannulation techniques

Determining the digestibility by collecting the faeces (colostomization)
Application of this method requires cannulated birds. During surgery after
the abdominal cavity is opened an incision is made in the colon at about 15
mms from its sphincter, and then a simple T-cannula is inserted in the ter-
minal colon (FIGURE VII-12). The advantage of the method is, that there is
no need to separate the faeces from the urine by chemical methods. The
resulting digestibility coefficients have a good repeatability and the data rep-
resent the apparent digestibility of amino acids well. When the nitrogen and
138
amino acid content of the faeces is adjusted with the endogenous portion,
the true digestibility of amino acids can be calculated (see: Section 7.1.2.).
Birds prepared in this manner are also suitable for conducting N balance
studies.
FIGURE VII-12: Faecal digestibility (cannulated/colostomised) birds
D) Determining the digestibility by collecting the ileal digesta

Some researchers argue, that similarly to swine it is the ileal digestibility of
amino acids which provide the most reliable data for poultry species. There
are two methods known for determining the ileal digestibility:
I. Although no cannulated birds are required for the first method (post
mortem digestibility studies) it is still necessary to discuss it here, as ileal
digesta is collected during these studies. In these studies the birds are
killed, and the digesta collected from the last section of the small intestine
is subjected to the necessary chemical assays. The problems pertaining to
the post mortem studies are the same as those discussed with swine
(Section 7.1.2.).
II. The cannulation technique appears to be the most reliable method for
determining the ileal digestibility of amino acids. During the surgery opera-
tion preparatory to these studies the ileum is separated from the postileal
section of the GIT, and then a simple T-cannula is inserted in its terminal
section (FIGURE VII-13). The cannula allows the quantitative collection of
the digesta and the calculation of the apparent ileal digestibility of the
amino acids. Similarly to the method based on the collection of faeces,
adjustment for endogenous nitrogen (amino acid) can be made in this case
as well, which will result in the true ileal digestibility of the amino acids (see
Section 7.1.2.).
139
FIGURE VII-13:Ileal digestibility ( cannulated bird)
It should be noted, however, that these procedures should only be con-

ducted with adult birds; the various cannulas cannot be implanted into
young birds.
E) Important characteristics of methods based on the different cannulation

techniques in poultry
Experiences gained with the various surgery techniques and the related
studies are summarized in Table 16.
Table VII-16: Important characteristics of methods based on different surgery techniques

(adult poultry) (TOSSENBERGER and BABINSZKY, unpublished data)
Data in the Table VII-16 show, that while cecectomy or the insertion
of the colon-cannula require 15-20 minutes, the implantation of the ileal-
cannula takes 30 minutes. The length of the post-operation recovery period
differs by the surgery technique applied, and varies between one to two
weeks. The length of the adaptation and collection period of the digestibili-
ty study is 4-5 days and 4 days, respectively, regardless of the method.
During the trial (adaptation and collection periods) the birds are placed
individually in metabolic cages. The digestibility of nutrients is calculated in
accordance with the contents of Sections 7.1.1. and 7.1.2.
140
DIGESTIBILITY OF AMINO ACIDS MEASURED WITH DIFFERENT TECHNIQUES

The digestibility coefficients determined with different methods may differ
significantly from each other in many cases. Table VII-17 summarizes the
results of the study conducted by BRAGG et al (1969), who determined the
digestibility of amino acids in sorghum using cannulated birds. Their data
were compared to the results from digestibility studies based on the collec-
tion of excreta (dropping digestibility). The results of the trial series show,
that in the case of some amino acids (lysine, cystine, threonine) there are
major differences between the values measured by the two digestibility
methods. Similar differences can be found in the ileal digestibility and drop-
ping digestibility of amino acids in soybean meal also (Table VII-18) (BIELORI
and IOSIF, 1987).
The data in Tables VII-17 and VII-18 therefore highlight the fact, that
there may be significant differences in the digestibility of amino acids of var-
ious feed ingredients, depending on where, in which intestinal section the
values were measured.
Table VII-17: True digestibility of amino acids in sorghum based on the collection of faeces
and excreta (BRAGG et al. 1969)
Table VII-18: Ileal and dropping digestibility of selected amino acids of soybean meal
(BIELORI and IOSIF, 1987)
At present there are relatively few data available on the ileal digestible
amino acid content of various feed ingredients. The Dutch Central Bureau
for Animal Nutrition (CVB, 1997) has published informative figures which
can be used well in the diet formulations. Table VII-19 contains the total and
ileal digestible amino acid content of some feed ingredients.
141
Table VII-19: Total and ileal digestible amino acid content (g/kg) of selected feed ingred
ents (CVB, 1997)
It appears from the data presented also, that there are substantial dif-
ferences in the digestibility of amino acids of certain feed ingredients. Thus
for instance, the digestibility of lysine in corn is only 61%, while that in
wheat is 83%. Similar differences can be measured in the digestibility of
lysine in meat and bone meal (73%) and in fish meal (91%). These discrep-
ancies warn, that the amino acid requirement of poultry can be meet with
more accuracy if instead of calculating with the total amino acid content of
the feed ingredients, the digestible amino acid content is applied; as the
birds are able to use for protein synthesis only that portion of the amino
acids which was absorbed from the ileal section of the GIT. When discussing
the ileal digestible amino acid content, the question may arise as to whether
the differences between the ileal digestible amino acid content and the
digestibility values based on collection of excreta also exist when complete
concentrates are fed. The data in Table VII-20 show, that the differences
seen in the case of feed ingredients should be expected for compound feed
as well (TOSSENBERGER and BABINSZKY, 1998).
Table VII-20: Total and apparent digestible amino acid content of grower diet for broilers*
(TOSSENBERGER and BABINSZKY, 1998)
*Corn-soy based diet: 13.4 MJ ME/kg feed; 19.8% crude protein
In an other study on birds fed compound feeds no significant differ-

ence was found in the digestibility values determined by colon and ileum
cannulated birds. However, digestibility values measured on the basis of
excreta collection (intact birds and caecectomized birds) were significantly
142
lower for all amino acids tested compared to the digestibility measured at
the end of the ileum or colon. This finding can be attributed to urinary AA
excretion. The use of ileum or colon cannulated birds can both be recom-
mended for the purpose of determining the amino acid digestibility of poul-
try diets (BABINSZKY and TOSSENBERGER, 2005).
7.3.2.2. Ideal protein concept in poultry nutrition

In poultry nutrition the optimal dietary protein allowance is one which
equally satisfies the essential and non-essential amino acid requirements of
the birds so that at the same time it contains no surplus of any of those
amino acids (BABINSZKY and VINCZE, 2002). Such a diet, however, can only be
produced by synthetic or semi-synthetic methods. In practice, an allowance
can be considered ideal, when its protein content is the closest to the opti-
mal composition beside the use of the least expensive raw materials.
When the amino acid composition of the ideal protein allowance is
determined, it should not be forgotten, that the diets must contain not only
all essential amino acids but also the total quantity of non-essential amino
acids pro rata to the energy content of the diet. This is necessary, because
the deficiency of any essential amino acid, or of the total combined quanti-
ty of non-essential amino acids compared to the requirement will diminish
the protein utilization. The lower protein utilization then leads to poorer
growth rates and feed conversion rates. The composition of ideal protein is
given in percentage relative to lysine for poultry nutrition as well.
When the amino acid composition is determined for poultry, however,
most of the problems are encountered in the estimation of sulfur-contain-
ing amino acid requirements. The reason for this is the feathering which
changes with age. It should also be noted, that further studies are needed
to determine the ratio of some essential amino acids to lysine. Several trials
are being conducted at present with this purpose. Table VII-21 provides a
recommendation for the composition of ideal protein by age groups.
143
Table VII-21: Recommended ideal protein composition for broilers by age grou*
(BARNA, 1999)
*Ratio of true digestible amino acids to lysine
7.3.2.3. Diet formulations on the basis of digestible amino acid concept

The application of the digestible amino acid content in the diet formulations
can only be justified professionally when the associated benefits are realized
in production also. For this reason it is especially important to become
acquainted with the results of field trials how the performance of broilers
and the economics of production changed when the diet was formulated on
the basis of digestible amino acid content. ROSTAGNO et al. (1995) in their
broiler studies fed three different compound feeds (HD, LD and LD+AA) both
in the starter and in the finishing phase.
In the first treatment (control) the diet was formulated with a highly
digestible amino acid content (HD). The diet fed in the second treatment
contained a very poorly digestible amino acid portion (LD); while in the third
treatment the diet of the second treatment was supplemented with crys-
talline amino acids on a digestible amino acid basis, so that the digestible
amino acid content of the diet fed in this treatment equaled the digestible
amino acid content of the control diet (Table VII-22).
144
Table 22: Nutrient content of the diets (%) (ROSTAGNO et al. 1995)
* HD: highly digestible amino acid, LD: low digestible amino acid, AA: amino acid supple-
mentation **d=digestible **M+C=mathionine+cystine
The results of the broiler growing trial are summarized in Table VII-23
The trial data show, that when the amino acid content of the diet is poorly
digestible, but it is supplemented on a digestible amino acid basis, the per-
formance of the broilers will be at least the same as that achieved with a
highly digestible amino acid containing diet, without harming the econom-
ic efficiency of the production. The application of industrially manufactured
amino acids, however, should be preceded by economic calculations.
Table VII-23: Summary of results of the broiler growing trial (ROSTAGNO et al. 1995)
In summary it can be concluded, that
145
FOR FURTHER READING
Babinszky, L. (2002): The status of protein management in Hungary and the strategy for
development. Agricultural Class of the Hungarian Academy of Sciences. Agroinform,
Budapest. 207 pp. [in Hungarian with English summary].
Babinszky, L. (1996): The feed- to food to environment chain. Possibilities in nutrition to
improve meat quality and to reduce nitrogen and phosphorus excretion in pigs. In:
Aminal Production, Healthy Nutrition, Environment. 4th Int. Symp. „Animal Science
Days”. 8-10 September 1996. Kaposvár. Supplement of Szaktanácsok. 7-23.
Bragg, D. B., Ivy, C. A. and Stephanson, E.I. (1969): Method for determining amino acid
availability of feeds. Poult. Sci. 48, 2135-2137.
Close, W.H. (1995): Energy and protein supply and reproductive performance of sows:
relevance to the development of practical feeding strategies. In: L. Babinszky (ed):
Energy and protein supply and their effects on the production of monogastric and
ruminant animals. 4th Intern. Symp. on Animal Nutrition Kaposvár, Hungary. Proc.
Pp. 49-60.
Fekete, S. and Bokori, J. (1984): Influence of fibre and protein level of rations on the spon-
taneous caecotrophy of rabbits (in Hungarian with English summary). Magyar Álla-
torvosok Lapja. 39, 295-298.
Fekete, S. and Gippert, T. (1985): Effect of crude fiber on protein utilization by rabbits. J.
Appl. Rabbit Res.. 8, 31-38.
Fekete, S. and Lebas, F. (1983): Effect of a natural flavour (thyme extract) on the
spontaneous feed ingestion, digestion coefficients and fattening parameters (in
Hungarian, with English summary). Magyar Állatorvosok Lapja. 38, 121-125.
Lebas, P. and Collin, N. (1976): Méthodes d’études de la digestibilité des aliments chez le
lapin. I. Durée de périodes de collects. Sci. Tech. Anim. Sci. 1, 71-77.
Leeuwen, van P., Babinszky, L., Verstegen, M.W.A. and Tossenberger J. (2000): A
procedure for ileostomisation of adult roosters to determine apparent ileal digestibility
of protein and amino acids of diets: Comparison of six diets in roosters and growing
pigs. Liv. Prod. Sci. 67, 101-111.
Rutz, F. and Rigolin, P. (2008): The quest for improved fiber utilisation. Feed Intern. April,
pp 16-18.
Rostagno, H.S., Pupa, J.M.R. and Pack, M. (1995): Diet formulation for broilers based on
total versus digestible amino acids. J. Appl. Poult. Res. 4, 293-299.
Schiemann, R. (1981): Methodische Richtlinien zur Durchführung von Verdauungs-versu-
chen für die Futterverschtzung. Arch. Tierernhr. 31, 1-19.
Wang, T.C. and Fuller, M.F. (1989): The optimum dietary amino acid pattern for growing
pigs. 1. Experiments by amino acid deletion. Br. J. Nutr. 62, 77-89.
146
CHAPTER VIII: BIOLOGICALLY ACTIVE PLANT PRODUCTS
Chapter VIII
BIOLOGICALLY ACTIVE SECONDARY

PLANT PRODUCTS
8.1. Antinutritional factors (ANFs)

8.1.1. Original functions of ANFs in the plant
8.1.2. Chemical characteristics of ANFs may be various
8.1.3. Major physiological effect in animals
8.1.3.1. Decrease in protein digestibility and utilization
8.1.3.2. Antigenic effect
8.1.3.3.3. Disturbed mineral metabolism
8.1.3.4. Phyto-oestrogens
8.1.3.5. Natural antivitamins
8.1.3.6. Photosensitization
8.1.3.7. Other effects (miscellaneous
8.2. Plant poisonings and selected phytotoxins
8.2.1. General considerations
8.2.2. Habit
8.2.3. Environment
8.2.4. Animal species
8.2.5. Exposure
8.2.6. Selection and ingestion
8.2.7. GI-environment and absorption
8.2.8. Distribution, detoxification and toxication
8.2.9. Adaptation
147
8.1. Antinutritional factors (ANFs)

Antinutritional factors (ANF) are compounds which prevent the maagnitude

of the animal production one would expect given the chemical composition
of the ingested feed. Typically when aanimals consume an ANF-containing
ration, the feed intake, the daily gain and feed efficiency, as well as milk and
egg production decrease. Affected animals usually do not show clinical
symptoms or signs of poisoning, but may suffer reduced immunocompe-
tence and reproductive failure. Pioneering trials by OSBORNE and MENDEL
(1917) demonstrated that growing rats, fed on raw soybean, became stunt-
ed. ANFs haave been in the limelight during the last few decades because
more and more plant protein has been used to replace more limitedand
expensive supplies of animal protein used in the animal nutrition. Except
for sunflower and safflower seed, the concentration of ANFs has a close pos-
itive correlation with the protein content of the plants. Moreover, during
some feed industrial process, the ANFs are enriched. For example, the
extraction of oil from oilseeds increases the concentration of proteinase
inhibitors, and alfalfa leaf protein extraction results in a higher saponin
level. Feed laws regulate the highest permitted level of the most important
antinutritional factors (see Chapter II).
Classification of ANFs may take place according to their original func-

tion, chemical composition and their influence on the animal organism.
8.1.1. Original functions of ANFs in the plant
a) Vegetable nutrients (e.g. carbohydrates) are stored in plants bonded to

phyto-haemagglutinins or lectins.
b) Another class includes compounds that inhibits the germination of grains
and seeds, aand factors that facilitate the breakdown of stored nutrients.
c) Still more directly protect plants against birds (e.g. tannin), bacteria, and
pests. For example the trypsin inhibitor of legume seeds against the
destroying effect of insect infestation (e.g. weevils have trypsin-type diges-
tion), or the alfalfa saponin against the attack of Trichoderma viridae fungi.
Consequently, genetics should also take into consideration the ANF
content of the potential feed plants. Some ANF occurs in every individual
148
plant, but higher amounts aare found in the leguminous seeds (proteinase
inhibitors), in the vegetative parts of legumes and potatoes (saponin, sola-
nine, phyto-oestrogens) and in all organs of cruciferous plants (thyreostatic
compounds of rape, cabbage etc.)
8.1.2. Chemical characteristics of ANFs may be various
-Proteins and its derivates: proteinase inhibitors, biogenic amines, goitro-

genic peptides, phyto-haemagglutinins and enzymes.
-Alkaloids, like quinolizidine, lupinin and sparteine in lupine seed, vicine
and convicine in bean (see favism).
-Steroids: saponin in alfalfa, solanine and phyto-oestrogens in potato plant.
-Saccharides: pentosan in rye, raffinose and stacchiose in legume seeds.
-Phytate: inositol-hexa-phosphates in cereal grains.
-Glutamic acid-antagonists like the N-oxalyl-diamino-propionic acid (OADP)
and the N-glutamyl-aminopropionitryl (GAPN) in vetch and non-protein
amino acids in cabbage, onion and garlic.
-Silica: rice, rice bran, sedges, rush and cane.
-Unsaturated fatty acid ether like the erucic acid in the rape seed.
-Binaphtol derivates like gossypol in the pigment of cottonseed.-Phenolic
compounds: condensed tannic acid in (broom) sorghum and barley grain, in
rapeseed and horse chestnut.
8.1.3. Major physiological effect in animals
8.1.3.1. Decrease in protein digestibility and utilization

a) PROTEINASE INHIBITORS of leguminous seeds (e.g. different bean and pea
species and soybeans) decrease the protein digestion and utilization in fish,
birds and mammals. Perhaps the most commercially important is the
trypsin inhibitor of soybean. Trypsin inhibitor activity of beans are 30 to
40% and in case of the peas 10 to 15% that of the raw soybean. Most of the
unfavourable effects can be eliminated by wet heat treatment. The activity
of trypsin inhibitor can be measured directly or, indirectly by monitoring
urease activity. The most important proteinase inhibitors of the soybean are
as follows: Kunitz-factor (Fl, F3 fractions), Bowmann-Birk-factor, SBTI
(Soybean Trypsin Inhibitor) A1, B, B2 and the S-inhibitor (isolated by sedi-
mentation). These compounds do differ in molecular mass (14,000 to
20,000) and in heat lability (Bowmann-factor is heat-stable; Kunitz-factor is
heat-labile). This variation is the explanation of some contradictory experi-
mental results (CSÁKY AND FEKETE 2004a). The main point of the effect is the
inhibition of the activity of trypsin, chymotrypsin, elastase and plasmin.
Inhibitor molecules bind by adsorption to trypsin, chymotrypsin and argi-
nine molecules and are excreted with them in the faeces. This is a signal to
increase pancreozymine and cholecystokinin blood concentration, stimulat-
149
ing the pancreas to produce more and more proteinase enzyme. As a con-
sequence, pancreas hypertrophy develops (Table VIII-1). These enzymes
have a high cystine content, which means a loss to the organism and the
need for dietary increases in supplementation of sulphur-containing amino
acids. (Soybean is poor in methionine to begin with, related to its lysine con-
tent.)
Table VIII-1: Pancreatic hypertrophy in rats fed raw and heat-treated soybean meal
(Initial LW: 100 g; experimental feeding: 10 days) (GREEN et al. 1986)
Renewed role of soybean in the animals` protein supply. Since the

outbreaks of BSE, the authorized use of animal protein meals are limited in
the feeding of production animals. The possible replacement sources are the
soybean, other oilseeds, legumes, rape- and sunflower seed. However, the
production of the latter crops are limited by the Blair House Agreement.
Consequently, the use and importation of soybean in the EU continues to
increase. Seed of soybean (Glycine soja) contains 37 to 39% crude protein,
19 to 21% ether extract and 25% N-free extract (carbohydrates). Except the
relatively low S-containing amino acid levels, its amino acid content is very
favourable: 31 g/kg lysine, 45 Lys% Met+Cys, 22 Lys% Trp and 52 Lys% Thr
in extracted solvent soybean meal of 48.5% crude protein. The prerequisite
for inclusion of soybean in the diets of monogastric animals is the inactiva-
tion of ANFs. A series of more or less effective processes is known: heat
treatment (dry, wet, under pressure, extrusion), enzyme treatment etc.
HULLÁR et al. (1998) compared the digestibility of diets, containing raw and
extruded soybean meal. Ground chicken carcass served as control. Cats
digested the untreated soybean-containing feed mixture very poorly, but by
extrusion improved the digestion coefficients up to the level of ground car-
cass (Table VIII-2).
150
Table VIII-2: Effect of extrusion on the digestion of soybean based diet in cats (HULLÁR,
FEKETE and SZűCS, 1998)
Legend: DC=digestion coefficient, DM=dry matter, CP=crude protein, EE=ether extract and
NFE=N-free extract
Nevertheless, there are always complaints relating to unfavourable

effects of using soybean in animal feeds. The causes of negative results,
related to the feeding of soybean, may derive from the failures of heat treat-
ment (ineffective or overheating), lack of knowledge of appropriate applica-
tion (ignoring possible antigenic property i.e. the interaction of soybean pro-
teins with the intestinal flora and the gut mucosa (CSAKY et al. 1999)) and
ineffective quality control (disproportionate sampling or choice of inappro-
priate methods). The heat-stable antigenic proteins are considered as the
most harmful to the animal (see below).
Quality control methods include: the measurement of urease activity
(indicates the ineffectiveness of heat treatment); the potassium hydroxide
protein solubility (KOH-PS) (a good indicator of overheating), and the pro-
tein dispersibility index (PDI) which provides information about both fail-
ures. In conclusion, optimum nutritional use of soybeans requires the neu-
tralization of antinutritional factors, setting of the amino acid balance, sup-
plementing the lacking minerals (Fe) and vitamins (B12), assuring the quan-
tity of omega-3 fatty acids, preventing allergenic effect and assuring myco-
toxin-free state.
b) LECTINS or phyto-haemagglutinins (PHA) are proteins, mostly in
leguminous seeds, with a high affinity to sugars and glycoproteins. Lectins
resist proteolytic digestion in vivo. They can be linked to the sugar mole-
cules (polymannose-type side chains in the lower region of the intestinal villi
and more complex sugars of the upper part of the villi) of the gut mucosa
and may alter absorption of nutrients. For the detection of lectins an in vitro
system is used which is based on their ability to agglutinate red blood cells
by linking to the glycoprotein of the cell wall. This is the origin of their syn-
onym, PHA. In cell culture, they stimulate the mitosis of T-lymphocytes.
According to the data of PUSZTAI (1997) the genetic engineering of plants may
modify the characteristics of their surface antigens, too. This may alter the
linkage of lectin and the gut mucosa, causing diarrhoea.
c) SAPONIN. The most common is the alfalfa saponin, accumulating
during the development of the vegetative parts of the plant to protect it
151
against the fungi. The protective effect is based on its physical structure,
which is able to decrease the surface tension. If contacting red blood cell, it
haemolyses them and this is one of the tools used to detect them in vitro.
Since the cholesterols of the feed adsorb them in the gut lumen, their
absorption is trifling and therefore in vivo they do not cause haemolysis.
Nevertheless, by changing the surface tension on gut lining, saponins
decrease protein absorption and even may cause enteritis and diarrhoea.
d) SOLANINS, these bitter poisonous alkalods, are typically found in the
vegetative part, sprout and tuber peel of potato. They can be eliminated by
soaking or neutralized by heat treatment. In lower concentration saponins
can be detected in tomato and nightshade, too.
e) PHENOLIC COMPOUNDS, such as condensed tannin or proanthocyani-
dins are found in the sorghum and amaranth grain, rapeseed and in the
leaves of sunflower. They are able to cause denaturation of proteins and the
free amino acids, above all lysine and arginine. These processes lead to a
disturbed protein digestion.
f) SILICA of rice bran, sedges, bulrush and cane significantly decreas-
es the digestibility of organic matter above all that of the proteins and dam-
age tooth enamel of animal during ingestion.
8.1.3.2. Antigenic effect

Antigenicity of protein has been found in soybean (GLYCININ, A-, B- and G-CON-
GLYCIN), beans and pea. They are storage proteins and resist conventional
heat treatments and digestion and are able to enter the bloodstream, induc-
ing humoral immune response (CSÁKY and FEKETE, 2004b). The chronic
hypersensitivity is more common than the acute hypersensitivity reaction.
Since compounds, causing antigenicity, are heat-stable, they cannot be
totally eliminated by the actually used heat treatments, chemical or enzy-
matic process would be required (e.g. hot aqueous ethanol extraction).
Consequently, some low antigenicity can be expected to be present in the
commercial extracted soybean meal batches.
8.1.3.3. Disturbed mineral metabolism

-PHYTASE is an important phosphorus containing compound in cereal
grains. This bound phosphorus, however, has a low availability for the
monogastric animals. Rabbit and pig are able to partially use this phos-
phorus too, having some phytase activity in their gut mucosa. The utiliza-
tion of phytic phosphate is the lowest in poultry; this is the reason that
nutrient recommendations for poultry give not only the total but also the
available phosphorus requirement. Inositol-hexaphosphate, as a chelating
agent, binds several minerals and secondary zinc and manganese deficien-
cy may develop. By supplementing feed with phytase enzyme of bacterial
origin, the absorption of the mentioned minerals can be considerably
improved.
152
-OXALATES are predominantly present in some plants (spinach, wild

amaranth, garden sorrel, sugar beet leaves etc.) and decrease the calcium
absorption. Calcium oxalate crystals may cause kidney damage or precipi-
tate as uroliths (calculi).
-GOITROGENIC or THYREOSTATIC PEPTIDES (glycosinolates or sinigrin).
These compounds are frequently present in cruciferous plants (family
Cruciferae, above all Brassica genus. e.g. cabbage, broccoli, kale, water-
cress, sweet alyssum, rape etc.). They have a thioglucose core, linked to dif-
ferent R- groups, which gives a variety of compounds of similar biological
effect. After having removed the glucose part from the glucosinolates by
means of the myrosinase or thioglucosidase enzyme (in the plant or in the
intestinal tract), the remaining aglycones, after further degradation, provide
a series of molecules. The latter compounds (e.g. goitrin, vinyl-thiooxa-
zolidon, thiocyanates, isothiocyanathes, nitriles etc.) inhibit the iodine
uptake of the thyroid gland and may cause even goitre. These compounds
are fairly resistant to the heat treatment and therefore the genetic selection
has been used to produce rape varieties, low in erucic acid and thyreostat-
ic substances, commonly called double zero rape (seed), or canola. The
triple zero varieties do not contain tannic acid either. Raw soybean and
peanut have also some thyreostatic compounds.
-CYANOGENIC GLYCOSIDES (e.g. linamarin, durrin) can be found, among
others, in linseed, cassava and sorghum plant. During digestion the prus-
sic acid may get released, absorbed and binds the iron of citochrom oxydase
enzyme, causing oxygen deprivation of tissues and suffocation. (Gossypol
may be classified into this group, too, having iron-binding capacity.)
8.1.3.4. Phyto-oestrogens
The most important representatives of this group are of isoflavonoid-struc-
tured, like genistein, daidzein, glycitein, biocanin A, formononetin and
cumoestrol. Plant oestrogens are most frequently found in leguminous
plants, like soybean, clovers, alfalfa but they could be detected in green
grass, too VETTER, 1991). Their biological influence explains the stimulating
effect of the spring pasture on the sexual activity of turned out grazing ani-
mals (KÉGL, 1991). If they are ingested in high quantity (e.g. eating subter-
ranean clover, Trifolium subterranum), they will cause reproductive failures,
oestrogen syndrome (vulva oedema, cystic ovaries and abortion). There are
initiatives to apply phyto-oestrogens against postmenopausal osteoporosis.
8.1.3.5. Natural antivitamins

Antivitamin may be any substance that prevent a vitamin (by destruction,
binding or competitive antagonism) from normal metabolic functioning.
Antivitamin A is the lipoxygenase enzyme of the raw soybean, destroying
carotinoids and vitamin A. Feeding of young, growing chicks on raw soy-
bean, rickets (rachitis) develops because of the antivitamin D uptake.
153
Gossypol (antivitamin E), which can be found in bean and cottonseed, con-
siderably decreases the tocopherol level of dairy cows), causing infertility
(FIGURE VIII-1). In poultry, by binding the iron in the gut lumen, may cause
anaemia and also reduces egg weight. In preruminant calves, lambs and
monogastric animals, high dosages of gossypol are toxic.
Dicumarol content in stuffy sweet clover (Melilotus officinalis) and some rat
poisons (e.g. Warfarin) interferes with vitamin K in blood clotting. As a
result, blood forms soft swellings beneath skin on different part of the body.
Acetylpyridin and indolacetic acid in corn and millet grain has antiniacin
effect. Overwhelming feeding on corn makes mammals especially prone to
niacin deficiency, because corn grain is poor in the niacin precursor trypto-
phane, too. Thiamine antimetabolite can be found in bracken fern, rock
fern, horsetail or scouring rush (Equisetum spp.), while many fish species
(Table XXIX-3) and the body of one-day chicken contain thiaminase
enzyme. Linatinen, a dipeptide of linseed has antivitamin B6 activity
(antipyridoxin), contributing to growth depression in young chicks.
Antivitamin B12 of raw soybean may cause anaemia, especially in cat.
Although it is not of plant origin, to make this passage fairly comprehensive,
the avidin of raw egg white worth mentioning as an antibiotin. It acts by
binding biotin in the gut lumen and prevent it from absorption.
8.1.3.6. Photosensitization
a) PRIMARY PHOTOSENSITIZATION. Ingestion of buckwheat (Fagopyrum escula-
tum) may cause fagopyrisms, that of St. John`s wort (Hypericum spp.):
hypericism. Chemical compounds, responsible for the photosensitization
are genuinely present in plant; after digestion and absorption, part of them
accumulates in the skin, where they are activated by the direct sunshine.
BAJMÓCZY and GLÁVITS (1988) reported about photosensitization in lambs fol-
lowing green millet grazing.
b) SECONDARY PHOTOSENSITIZATION based on a liver damage, when the
decreased functioning capacity is not sufficient for eliminating the chloro-
phyll metabolite, the phylloerythrine. The latter, getting in the skin and acti-
vated by the UV radiation of the sun, causes more or less severe dermatitis,
superficial necrosis of white or light skinned parts of animals. It occurs
more frequently following the ingestion of alsike or Swedish clover (Trifolium
hybridum: trifoliosis) and less often with alfalfa and yellow vine or catheat
(Tribulus terrestris: tribulosis). These effects should be taken into consider-
ation during differential diagnosis of cases of sheep with swellings on the
head (“big head”), suspected to be caused by Clostridium chauveyi bacteria
and in cases of cattle the salivation, blisters and lesions on muzzle and in
buccal cavity, otherwise thought to be caused by mouth-and-foot disease
(MFD.
154
8.1.3.7. Other effects (miscellaneous)

The LIPOXIGENASE enzyme produces the oxidation and getting oil rancid with-
in the seed of beans (“beany taste”). The enzyme POLYPHENYL-OXYDASE devel-
ops during preservation a brown colour on the cut surface of fruits, like
apple or banana. Most of the leguminous seeds (bean, lentil and pea) have
non-starch carbohydrates (NSC), included oligosaccharide content. Some of
the latter compounds (RAFFINOSE and STACHIOSE) may provoke flatulence,
being undigested in the small intestine, thereby fermented in the caecum
and colon, producing excessive intestinal gas. TOXIC AMINO ACIDS of vetch
(Lathyrus spp.) may provoke lathyrism, which can manifest either in bone
alterations and decreased connective tissue tensile strength or in troubles
of the nervous system (neurolathyrism). In the Brassica plants, onion and
garlic can be found the non-protein amino acid S-METHYLCYSTEIN SULPHOXIDE.
If this compound undergoes rumen fermentation, the released haemolytic
dimethyl-disulfide may cause anaemia in ruminants. Leaves of the tropical
Leucaena leucocephala CONTAIN the tyrosine-analogue MIMOSINE, with possi-
ble nervous effect and causing loss of hair. As an arginine analogue, the
CANAVANINE is produced in the alfalfa sprout, and if ingested in higher
amounts, may cause amino acid imbalance aand the symptoms of lupus.
ERUCIC ACID, present in the seeds of the wild rape varieties causes taste fail-
ures (unpleasant, scratchy taste feeling); moreover, it is suspected to be car-
cinogenic. QUINOLIZIDINE DERIVATIVES (LUPININE, ANGUSTIFOLINE, SPARTEINE and
LUPANINE) in lupine seed are responsible for the bitter taste and at the same
time, they initiate liver damage (hepatotoxicity), birth defects and neurolog-
ical symptoms. SINAPINE is an ester of sinapic acid and cholin, present in the
rapeseed. Owing to bacterial fermentation, in the caeca of fowls sinapin will
be transformed into trimethylamine. Latter gives unfavourable smell and
taste (fishy odour and flavour) to the brown-shelled eggs in laying hen.
BIOGENIC AMINES are produced through the decarboxylation of amino
acids both in plants and in animal body and in their products. According to
the number of amines in the molecules they are mono- or polyamines. One
group of monoamines (tyramine, octopamine, α-phenyl-ethyl-amine) stimu-
lates gut motility, peristalsis and secretion. Polyamines have three sub-
classes, the hydrocarbon–structures (putrescine, cadaverine, tyramine, sper-
mine, spermidine etc.), indol derivatives (serotonin, tryptamine, cate-
cholamines) or imidazol derivatives like histamine. Amines are either prod-
ucts of the cell metabolism or they get from outside (food, feed, and gut
microbes) into the organism.
Both structure and physiological effect of biogenic amines show
colourful variety. The spermine and spermidine are indispensable to cell
growth and differentiation. Polyamines have a significant role in the new-
born in helping and controlling the morphological and functional develop-
ment and maturation of intestinal tract. For example, they participate in the
control of enzyme production and regeneration of the mucous membrane.
155
Exogenous biogenic amines are able to substitute the intracellular produc-

tion, for example low concentrations of ingested putrescine and spermidine
have cell proliferating effect.
Harmful and toxic effects appear only at high feed feed biogenic amine
concentrations. The chances of such a process during grass ensiling and
especially haylage preparation are high. Most of the biogenic amines are
formed by amino acid decarboxylating activity of putrefactive bacteria and
also some species of lactic acid strains. Free amino acids released from
plant proteins become available and their decarboxylation takes place by
metabolic activity of bacteria present (BOKORI et al. 1986; STEIDLOVA and
KALAC, 2004). The most common amines like histamine, tyramine, cadaver-
ine and putrescine are produced from histidine, tyrosine, lysine and argi-
nine or ornitine. Polyamines (e.g. spermidine and spermine), derive from
putrescine. To a certain limit, the rumen microflora is able to adapt and
degrade these compounds. At the same time, polyamines are produced in
the large gut, too. Several members of the intestinal flora are capable of
decarboxylating amino acids, but in normal situation the enzymes of the gut
wall oxydase biogenic amines and thereby a barrier is established against
the absorption of large quantities from the digesta. This is the reason, why
bacterial biogenic amines emerging in the intestinal lumen generally do not
cause clinical problems, unless the amount of ingested or locally produced
biogenic amines is too much or the oxydase activity of intestinal wall is
insufficient.
Biogenic amine production during processing and preservation/stor-
age is common is human protein-rich foods (fish, mammals and bird meat)
and even in wines or rotten fruits. Using proteomic approach, PESSIONE et al.
(2005) demonstrated that amine-producing enzyme production (histidine
decarboxylase and ornitine decarboxylase) of lactobacilli is closely depend-
ent on the presence of the concentration of free amino acids present.
FUSI et al. (2004) administered a series of biogenic amines to weaned
Saanen kids. The direct oral administration had more pronounced effect,
than supplementation of feed mixture. Affected kids lost weight, their dry
matter intake decreased and the meat quality (muscle drip loss and colour)
has also been affected. Visceral organs showed histological lesions.
SCHNEIDER et al. (1989) mapped the distribution and concentration of hista-
mine, tyramine, putrescine and cadaverine in the digesta of growing-fatten-
ing pig. Although small amounts of biogenic amines are already present in
the stomach, the colon presented the main section/site of occurrence. By
applying bactericide supplement the concentration of biogenic amines could
slightly be lowered.
BERMUDEZ and FIRMAN (1998) tested the effect of peroral application of
some biogenic amines at a level, which can be considered as typical in ani-
mal by-product meal. Their conclusion is that in these concentrations nei-
ther cadaverine, nor histamine, phenylethylamine, putrescine nor their
156
combination had damaging effect on broiler chicken performance.

Nevertheless, under circumstances of high biogenic amines ingestion or
damaged gut lining, local lesions, vasodilatation, diarrhoea and nettle-rash
(skin eruption) occur both in animals and human (histamine poisoning or
“scombroid fish poisoning”).
FIGURE VIII-1: Correlation of gossipol intake and serum vitamin E concentration in a dairy
herd
157
8.2. Plant poisonings and selected phytotoxins

© Dan L- Brown
Introduction: the case of the falling cows. Some years ago, while running
in the Northern California countryside, I came across a small feedlot. A few
cattle were dead there, others were down and breathing hard. As I
approached the pen, two more cows fell down. The feed bunks were full of
direct cut Sudan grass (Sorghum sudanese), and I was afraid it might be
cyanide poisoning. I found no workers in the area at all, so after trying to
unload the bunkers by hand and watching more cows fall, I broke into the
office (no mobile phones in 1984), called the foreman at home, the vet clin-
ic, and found some needles and tubes and took some blood samples,
because more cows were falling. But instead of the bright red blood I expect-
ed, it looked like chocolate syrup. What had happened here? More impor-
tantly why did it happen? We will return to this case later…
8.2.1. General considerations

In most pastures, rangelands and forage production meadows, there are a
wide variety of plants, many of which are identified in textbooks and refer-
ences as poison plants. Most ornamental plants in the home are toxic as
well. Yet plant poisonings are not daily occurrences. Why? This is a ques-
tion that causes considerable confusion for many animal producers and vet-
erinarians. On the other hand it is often equally hard to figure out why live-
stock can safely graze and browse the same pasture for years and then sud-
denly suffer losses to poison plants or why healthy grazing animals can be
moved from one botanically identical pasture to another and die eating what
looks like the same forage.
The answers to many such puzzles lie in the many other important
factors to consider other than the mere dose or identity of a plant in a pas-
ture. These factors include habit, environment, exposure, animal species
affected, selection, ingestion, GI-environment, absorption, distribution,
detoxification, adaptation and elimination. The purpose of this chapter is to
help animal owners and their veterinarians use knowledge of these other
factors to explain, predict and prevent plant poisonings.
Many of the toxins discussed in the previous part (Chapter (8.1.), and
quite a few others, are found in pasture and forage plants on most farms
and ranches around the world. Which toxic plants are most important dif-
fer by region. Many of the examples used here are from the Western and
Northeastern US and East Africa, but the principles apply to Europe, Asia,
and South America. And since many troublesome plants are invading
158
exotics lacking their natural enemies, you may be surprised to find a local
plant killing stock a half world away.
Toxic plant substances are found in favored cultivated forage plants
as well as weeds. In some locations, recent planting of pasture or hayfields
combined with meticulous weeding or, more commonly, herbicide applica-
tion can create a monoculture or at least a simple mixture of high quality
forage, but that is not common worldwide. Where such simplified ecosys-
tems are common, they are vulnerable to invasion by large amounts of a
single weed and, if such a weed is toxic can wreck havoc on the animals.
For example, the first cutting of otherwise monoculture alfalfa in semi-arid
California is frequently invaded by groundsel (Senecio). Orchards cleared of
vegetation host fiddleneck (Amsinckia intermedia) infestations. Both of these
contain pyrrolizidine alkaloids that can and do cause severe liver damage
when the hay is infested and the orchards grazed, respectively.
8.2.2. Habit
The shape a poisonous plant takes has a profound influence on whether or
not it is dangerous to animals. Leaves on mature trees may be too high for
animals to get a very great dose of toxin. This is common in the case of cas-
cara (Rhamnus cathartica), maples (Acer spp.) and members of the stone
fruit genus (Prunus spp.) animals can survive the few leaves they can reach
from time to time and create a browse line without intoxication. In these cir-
cumstances, the owners may believe the plant isn’t a problem.
Unfortunately, there have been cases where goats were moved from lots
where Rhamnus cathartica has been browsed for years to fields with rich
low-hanging growth of the same cascara trees. Many of these animals died
taking in doses that cause fatal cardiac symptoms. Maple trees may shade
horses for years without problems, but if a red (Acer rubrum), sugar (A.
sachrum) or silver (A.sacharinum) maple falls in a high wind, then there is
suddenly enough vegetation to deliver the gallic acid and associated oxi-
dants and hemolytic agents to that cause the methemoglobinemia and ane-
mia known as red maple disease. When Prunus spp. (wild cherry etc.)
branches are brought down by enthusiastic climbing browsers, or broken
by the wind, all classes of livestock are not only threatened by the sudden
presentation of toxic fodder once out of reach, the attendant wilting causes
the cyanogenic glycosidase amygdalin to mix with glucoside enzyme and the
release of toxic cyanide is enhanced tremendously.
If prostrate plants such as clovers or immature knapweed rosettes
(Centaurea spp.) are growing under a generous over-story of taller more
erect vegetation they may be missed by large, less selective grazers. As the
rosettes get taller and the knapweed bolts, more toxic material may be con-
sumed by horses and brain damage ensues. [Perversely, some horses
become addicted to these dangerous irritating plants and seek out the flat-
ter leaves in subsequent seasons, if they survive that long.] In dry years,
159
clovers (Trifolium spp.) which have been infected by Rhizoctonia leguminico-

la fungus are exposed to grazing when the overstory is removed by grazing.
The slaframine delivered by these plants causes “slobbers”, excessive sali-
vation in equines, ruminants, and recently in alpacas. This not usually life
threatening, but riders get drenched trying to bridle their mounts. Known
as red clover disease, the offending fungus can infect other Trifolium species
as well.
8.2.3. Environment
Interactions between poison plants and the environment are legion. In addi-
tion to wilting after stem damage, cherry trees also release cyanide if
enzyme and cyanogens mix after cell organelle disruption caused by freez-
ing and thawing of leaves that have not undergone autumnal senescence.
Unfortunately this is also true of other cyanogenic plants such as sorghum
species including the grain crops, Sudan grass and Johnson grass (Sorhum
halepense). One of the serious effects of endophyte-infected fescues on cat-
tle is disruption with thermoregulation, a much greater problem for animals
in the hot and humid southern summers than in cooler seasons and loca-
tions.
8.2.4. Animal Species

Poisonous plants are not poisonous to all animals. Most if not all are
attacked by at least one insect species (although exotics invading a new
continent may be free of such pests until a biological control expert reunites
them with natural enemies from their country of origin). There is diversity
among mammalian species in their response to individual toxic plant
species, as well. Ingestion of leaves from maple trees, star thistle (Centaurea
solstitialis), and Russian knapweed (Acroptilon repens) has killed horses,
but no other farm animal. The pyrrolizidine alkaloids found in fiddleneck,
groundsel (Senecio vulgaris) and comfrey pose a problem for cattle and
humans that ingest them, less so for sheep, a bit less for goats and rabbits
don’t seem to be poisoned at all. Yew (Taxus spp.), rhododendrons and azal-
eas, and Kalmia species (lambkill, mountain laurel) kill our livestock, pets,
Asian axis deer, and European fallow deer at small doses if they eat them.
But the North American whitetailed deer eat these with impunity, much to
the chagrin of landscapers. Why?
In some cases, the bases for these differences are known. Horses are
probably more susceptible to oxidant toxins such as the maple leaf complex
due to negligible NADPH production in red blood cells (glucose-6-dehydro-
genase deficiency, like humans with favism) and we have found that the
microbes harbored by whitetails do destroy toxins more than cattle rumen
microbes (although this is probably not the entire reason). Cattle can suffer
emphysema when rumen microbes convert tryptophan released by rapidly
fermenting lush forage into 3-methyilndole. This does not seem to be a
160
problem for horses, which lack rumens. The basis for many differences is
unknown and represents fertile areas for investigation of comparative
metabolism among domestic animals.
8.2.5. Exposure
The practical application of the old cliché about the dose making the poison
is that one can observe the defoliation of patently toxic plants without
observing any effect. Several species of dogbane (Apocynum spp.) are found
in rough pastures throughout the Northeastern United States and poison-
ings can occur, but it is not uncommon to see flocks of sheep strip leaves
from these rich sources of cardiac glycosides without harm, only hungry
animals in rich solid stands will be affected.
8.2.6. Selection and ingestion

The first line of defense against plant poisoning is accomplished by the con-
scious behavior of the animals themselves. Fortunately, most toxic plants
are irritating or taste bad. Or if they do not taste bad at least many poison-
ous plants taste, feel and/or look distinctive to animals. That way, if they
survive the experience of eating some of a poisonous plant once, they can
associate that ill feeling with the sight taste or mouth feel of the plant and
avoid ingestion. PROVENSZA (1986) has demonstrated that this ability can be
exploited to train livestock to avoid perfectly edible plants by training the
animals not to eat them with timed doses of LiCl sufficient to make them
feel sick when they are eating the plant one wishes to protect.
In this way, animals will usually survive to coexist with a range of
toxic plants, even present in dangerous doses, in their grazing areas. This
may not be able to protect the animals, if they are moved to new range with
unfamiliar toxic plants (thus lantana=Lantana camara) killed thousands of
migrating cattle in Ethiopia a few years ago). The self-training strategy also
will not work if the toxic plant makes up much or most of the forage base
as larkspur (Delphinium spp.) does in the Intermountain West of the US or
tall fescue does in the American South. In addition, due to neurological
effects, animals can become addicted to some plants such as wild lettuce or
yellow star thistle (Centaurea solstitialis), the latter all the more amazing
since it is extremely irritating chemically and physically.
8.2.7. GI-environment and absorption

The pH, oxidation state (aerobic, anaerobic, reducing, oxidizing), and gut
structure all affect the transformation and absorption of ingested toxic
plants. In the rumen, microbial activity can detoxify a wide variety of toxins
by catabolizing them before they can reach the acid abomasum. This is one
of the hypothesized roles of the primordial pregastric fermentations that
predated but led to the evolution of the rumen as a digestion site for fiber
and reactor for amino acid conversions and synthesis.
161
To pass the protein/lipid membranes of mucosal tissue without spe-

cial carriers usually requires a small molecular weight and/or a neutral
charge. Depending on the [pKa] of functional groups, a molecule may be
absorbed or not at a given pH. So a compound that is anion at neutral pH
would need an acid environment to be absorbed and cationic organic com-
pounds need basic conditions to lose its charge. This controls somewhat the
site of absorption as a compound makes its way through the rumen (if pres-
ent) to the acid and pepsin of the stomach, back to more neutral pH in the
duodenum, then to the fermentation sites in the lower gut. Mineral cations
(lead, cadmium, etc.) are not reduced to the uncharged ground state by
physiological conditions of the GI tract, so must be carried into the body
through regulated sites that normally facilitate mineral nutrient uptake. In
addition, heavy metals are subject to binding to metallothionine, a protec-
tive gene product that reduces absorption, induced in gut cells (among oth-
ers) by the presence off the heavy metals themselves. Some metals such as
mercury are rendered toxic by covalent combination with organic com-
pounds in the environment. For example, methyl mercury can pass more
easily into the body and dimethyl mercury can pass through latex gloves
and into the skin to cause poisoning without ingestion at all. Some toxins
such as the ionophores (e.g. monensin and lasalocid) act by interaction with
cell surfaces, enhancing the permeability of membranes to ions such as
potassium and sodium enough to disrupt the activity of excitable tissue,
notably cardiac muscle.
The lectins actually bind to the mucosal cell itself and in the case of
ricin (from the castor bean plant (Ricinis communis), have one subunit that
binds the membrane and another that enters the cell and stops protein syn-
thesis. It is thought that one ricin molecule can theoretically kill one gut
cell, which makes it an incredibly efficient and potent poison. Once the gut
cells are destroyed the mucosa, then the serosa are disrupted, and this
toxin can enter the blood stream and do direct damage to other vital organs
as well. Other compounds that cannot lose a charge on enough functional
groups at once, such as zwitterionic amino acids and similar plant com-
pounds also have specific carrier systems that can be borrowed by similar
toxic compounds.
Most absorbed toxins are carried along with absorbed nutrients into
the blood stream to the mesenteric vein and on into the portal vein to the
major detoxification organ, the liver. The most neutral and lipid soluble tox-
ins can enter the lymph and thereby avoid the enterohepatic circulation and
enter the general circulation before the liver has had a chance to act upon
it. The liver would then only see such toxins as were delivered in the arteri-
al blood supply that reaches the liver.
8.2.8. Distribution, detoxification and toxication

One primary general strategy to reduce the effects of toxins is to get rid of
them as fast as possible by making them water soluble again so they can be
162
more readily excreted by the kidneys. This can be accomplished by attach-

ing such compounds as sulfate, glutathione or glucouronide. Before that
can be done, though, many toxicants need to be prepared for these Phase II
conjugation reactions by undergoing Phase I reactions that create a reactive
site on the molecule for the attachment of the conjugate. Sometimes these
Phase I reactions themselves can render the toxin water soluble for easy
transfer to the kidneys and excretion. Unfortunately, the same mixed func-
tion oxidases (including the many liver P450 enzymes) that activate toxins
for further processing can also activate some toxins to become even more
toxic! Thus the pyrrolizidine alkaloids in groundsel, comfrey or fiddleneck
(Senecio vulgaris, Symphytum officinale, Amsinckia intermedia) are turned
into raging pyrroles, and overdoses of acetaminophen are activated, but
soon exhaust the conjugation mechanisms and the active molecules are free
to attack the surrounding liver cells. The estrogenic effects of zearalenone
produced by Fusarium molds infecting corn and barley are increased by a
factor of 10 when converted to the corresponding alcohol by Phase I
enzymes. So, for some substrates, what should have been the first step to
detoxification can be a dangerous toxication reaction. For some common
toxins that act so quickly that they need immediate attention, there are con-
stitutive enzymes that direct specific detoxification. For example, the rho-
danese reaction converts cyanide and thiosulfate to sulfite and thiocyanate.
Many toxic reactions to poisons are mediated by the release of free
radicals, so a number of systems have evolved not to counter toxins direct-
ly, but to prevent and clean up this manifestation of poisoning. Antioxidants
are prominent in this effort, so vitamin E, vitamin C (for animals that don’t
make enough of their own) and various carotenoids (e.g. beta carotene,
lutein, lycopine, quercetin, etc.) and polyphenolics in the diet are helpful. In
addition such enzymes as superoxide dismutase (contains zinc and copper
and others manganese) and glutathione peroxidase (contains selenium) are
important and interact with sulfur-rich cofactors (such as glutathione). It
can be readily seen that deficiencies in these elements and vitamins will
compromise an animal’s ability to protect itself from toxins in the environ-
ment and well-fed animals are often more robust in a toxin-rich environ-
ment.
8.2.9. Adaptation
In some cases, animals and their symbiotic micro-organisms can become
adapted to toxic plants and toxic loads of nutrients. This adaptation may be
evolutionary, taking place over millennia and hundreds of generations or it
can take place in matter of days (microbes) or hours (induced tissue enzyme
systems). Certainly the tolerance of koala bears for the essential oils in
eucalyptus and the proline-rich salivas of animals (e.g. deer or the humans)
that browse tannin-rich plants represent natural selection of the animals.
But a single species of micro-organism that catabolizes the toxic amino acid
mimosine from Leucena Leucephala, can be added to the rumens of animals
163
fed leucana to avoid hair loss by animals feeding on this nutrient-rich plant.
Slow adaptation to whiteball acacia (Acacia angustissima) can prevent oth-
erwise fatal intoxication in sheep. Is a single organism favored or does a
new, more complex set of organisms need to be prevalent? That system is
not completely characterized, but it contains at least two chemically distinct
factors that both must be present for poisoning to occur, either of which
could be disrupted by the organisms. Three possible candidates for the two
factors: 1) acetyldiaminobutanoic acid 2) an enzyme that can release the
lathyrogen diaminobutanoic acid from compound 1 or 3) condensed tannins
that prevent the microbial destruction of that lathyrogen.
THE CORRECT SOLUTION
What about those falling cows? As the reader can now see, so many fac-
tors have to fall in place for a plant poisoning to occur that it seems amaz-
ing that such a thing could happen at all, but it did: The Sudan grass’s
habit was no protection because it was cut down and carried to the cattle.
In fact, since the senior men in charge did not feel young women watching
cattle on weekends were suited to operating the machinery needed to feed
this material and the men were not around on weekends, they fed the cat-
tle three days worth of Sudan grass on Friday to last until Monday creating
an excessive exposure, due to human error. In this case, that human error
was caused by a cultural force (poor management style and sexism) as pre-
vious male weekend help was always trained in operating the feeding
machinery and fed daily. Since they had eaten the same feed for months
without illness or discomfort, the cattle failed to select against this toxic
dose of feed, took advantage of the extra allotment and ate more than one
days ration. Our frantic removal of feed from the bunk reduced the expo-
sure, but too late for most of the cattle. The weather had been warm and
overcast in the field where the Sudan grass was grown on soil still rich in N
applied for previous and current crops. Thus the environment ensured
that the grass would be rich in nitrates. The rumen microbes and the
reduced atmosphere of the GI environment readily converted the nitrate to
nitrite (toxification) and the favorable pH permitted the rapid absorption of
enough nitrite to oxidize the hemoglobin to methemoglobin - too rapid for
natural detoxification and excretion to take place. Of course all of this
was far too fast for any kind of adaptation to take place. When the veteri-
narian arrived, methylene blue was administered intravenously and this dye
prevented the binding of nitrite to the hemoglobin, yet permitted oxygen to
bind and thus, some of the cattle were saved. And other person, included
women are allowed to operate all of the machinery there now, so the cattle
are fed daily. Disruption of any one of the factors involved might have pre-
vented, postponed or reduced the impact of this disaster.
164
FOR FURTHER READING
Cheeke, P.R. (1998): Natural Toxicants in Feeds, Forages, and Poisonous Plant. (2nd ed.)
Interstate Publ. Inc. Danvilles, IL
Csáky, I. and Fekete, S. (2004): Soybean: Feed quality and safety. Part 1: Biologically active
components: A review. Acta Vet. Hung. 52, 299-313.
Csáky, I. and Fekete, S. (2004b): Soybean: Feed quality and safety. Part 2: Pathology of soy
bean feeding in animals: A review. Acta Vet. Hung. 52, 315-326.
Csáky, I., Majoros, G., Zöldág, L. and Fekete, S. (2001): Interaction between anti-soya anti
bodies and Staphylococcus aureus bacteria. Laser particle sizer and nephelometric
studies. – in Hungarian, with English summary. Magyar Allatorvosok Lapja,
123:375-379.
D`Mello, J.P.F. (2000): Anti-nutritional factors and mycotoxins. In D`Mello, J.P.F.(ed): Farm
Animal Metabolism and Nutrition. CABI Publishing. Wallingford. Oxon. P. 383-403.
Esmail, S.H.M. (2003): Nutritional limits to poultry production from some conventional and
unconventional feedstuffs. Poult. Intern. 42(9):43-45.
Gill, C. (2003): Global soybean meal quality. Feed Intern. 24(7):23-24.
Gontzea, I., Ferrando, R. and Sutzesco, P. (1968): Natural antinutritional substances in
feeds - in French. Vigot Freres. Paris
http://www.ncbi.nlm.nih.gov/Structure: VAST. Structure neighbors.
Huisman, J. and Tolman, G.H. (1983): Anti-nutritional factors in animal feedstuffs. In
Haresign, W. (Ed): Recent Adv. Anim. Nutr. London, p 21-37.
Hullar, I., Fekete, S. and Szocs, Z. (1998): Effect of extrusion on the quality of soybean-
based catfood. J. Anim. Physiol. a. Anim. Nutr. 80, 201-206.
165
CHAPTER IX: ENERGETIC FEED EVALUATION
Chapter IX
ENERGETIC FEED EVALUATION
9.1. Measurement of animal needs and nutritive value
9.2. Development of energy evaluation systems

9.2.1. Early history of feed evaluation
9.2.2. Systems based on scientific evidence used in the past
9.2.2.1. Starch equivalent
9.2.2.2. Scandinavian feed unit
9.2.2.3. Oats unit
9.2.2.4. Fraps’ productive energy
9.2.3. Modern feed evaluation systems
9.2.3.1. Digestible energy (DE)
9.2.3.1.1. Pig nutrition
9.2.3.1.2. Rabbit energy evaluation system
9.2.3.1.3. Horse energy evaluation systems
9.2.3.2. Metabolizable energy (ME)
9.2.3.2.1. In MJ ME for all animal species in Sweden
and in the former USSR.
9.2.3.2.2. Beef cattle and small ruminants in Austria,
Germany and Poland.
9.2.3.2.3. A special ME in Germany and Austria: pig
9.2.3.2.4. USA (1998) pig: Mcal ME. :
9.2.2.3.5. Fish and reptile
9.2.2.3.6. Carnivores (dog, cat, ferret, fox and mink)
9.3. Energy evaluation of poultry feeds
9.4. Net energy

9.4.1. Rostock energy evaluation system
9.4.2. The UK ME–NE system
9.4.3. Californian net energy sytem for beef cattle
9.4.5. Beltsville dairy cow system
9.4.5. European cattle systems
9.4.6. Energetic evaluation in sheep and goats
167
he main goal of the energetic evaluation of feed is to make a reliable
T prediction about the estimated animal performance. The knowledge of

energetic value of feed and the animals’ need are prerequisits for
achieving the above formulated aim. The energetic value of feeds should be
given in the same system as the nutrient requirements have been deter-
mined. E.g. using the German DLG data of pigs needs and calculationg by
the US NRC ME value of feeds is not possible. In an ideal situation, the
detailed chemical compostion of feed is known, but in practice, data of the
simplified “approximate” (weende”) analysis are sufficient for a realistic feed
evaluation.
9.1. Measurement of animal needs and nutritive value

Nutrient requirements and nutritive value are the central notion of feeding
and they are reflections of each others. The heve qualitative and quantita-
tive charicters at the same time. Animals’ need are published in recom-
mendations (normes) and the nutritive value of different feedstuffs can be
found in composition tables. By using these recommendation the following
limitations should be kept in mind: 1. average conditions are given, 2. eco-
nomic aspects are not taken into consideration, 3. generally, the physical
characteristics of feed (taste, smell, structure etc.) are neglected and 4. they
are valid in thermoneutral zone.
According to MØLLGAARD (1988) the scientific definitions are as follows:
a) The nutritive value show the maximum possible animal perform-
ance from a unit of the given feed.
b) The published nutrient requirements predict the smallest amount
of feed, which is indispensable for a animal performance. (In this respect
„animal performance“ means maintenance, work and production.)
For expressing nutrient requirements and nutritive value of feed, two
basic approaches exsist. The more theorethical one is based on a very
detailed chemical analisys and the entire list of components is considered
as the nutritive value. The more practical approach gives the total effect in
a cumulative feature, namely in the energy value expressed in calorie or
joule. Thus, the nutritive value of a feed can be evaluated from that point:
which compounds are supplied for the organism, or what is ist energy value.
Supposed that animal needs for specific nutrients (essential amino acids,
168
minerals, vitamins) is covered, the nutrient requirements can be contracted

in the form of energy. This simplification makes possible to use the energy
supply as a predictor of animal performance.
The gross energy (combustion heat) depend only on the chemical
structure alone and only a small proportion can be used by the organism.
By excreting faeces, urine, eructation and flatulance gases an important
amount of energy leaves the animals and the heat production of assimila-
tive chemical processes also imply energy loss. The “ladder” of feed energy
utilisation can be represented in FIGURE IX-1)
FIGURE IX-1. Steps of feed energy utilisation (“ladder” of energy losses)
The several energy evaluation systems basically differ in the choice of

level of that “utilisation ladder”, and also the way of technical measure-
ments and calculation. The principle of digestible and metabolisable energy
169
content is that after weighing the feed intake the energy content of faeces
(digestible energy) and the urine and eructation energy should be sub-
stracted (metabolisable energy). By knowing the heat production or/and the
energetic valu of animal product, the net energy can be calculated (for more
details see Chapter I and V).
9.2 Development of energy evaluation system
9.2.1. Early history of feed evaluation

a) Hay equivalent (THAER, 1804)
b) The total digestibel nutrients (TDN)
The total digestible nutrient can be calculated by the following equa-
tion: TDN,g/kg=dCP+2.25×dEE+dCF+dNFE. The GN (Gesamtnährstoff-
gehalt is very similar but the coefficient of digestible ether extract (fat) is
2.3). In US the TDN can be given also in % (g/100 g).
9.2.2. Old systems based on scientific evidence used in the past

9.2.2.1. Starch equivalent
KELLNER (1905) developed this evaluation system, based on net ener
gy contant. The model of determination used to be the fat production of
adult oxen.
9.2.2.2. Scandinavian feed unit
In the pcartice it can be considered as equal to 0.7 Starch Equivalent.
9.2.2.3. Oats unit
was in use in the former USSR, it equals to 0.6 Starch Equivalent.
9.2.2.4. Fraps’ productive energy
It is also based on net energy and the method of determination was
the burning of the whole poultry body.
9.2.3. Modern feed evaluation systems

CONCEPTUAL CONSIDERATION IN DEVELOPING AN “IDEAL” ENERGY UNIT.
a) Nutrient requirements of animals and nutritive value of feedstuffs
should be expressed in the same way.
b) Should be additive, independent from the raw material
c) Should be exact (of good preditive valeu of animal performance)
d) Should be based on easy-to-determine chemical componants.
e) Should be relatively simple allowing practical use.
As described above, the total energy content (combustion of heat) of a
feed gradually “suffers” losses. Theoretically, each level (DE, ME or NE) may
be the basis of the way of expression of animal needs and feedstuffs value.
It is evident that gross energy cannot be an appropriate energy unit,
because in many cases feedstuffs of high combustion of heat have a vary
low digestibility (e. g. straws). In the majority of countries the digestible and
metabolisable energy has been chosen for pig, horse and poultry, which is
170
not so accurate than net energy but it is technically easier to measure.

Taking into consideration of the digestive characteristics of ruminants, the
metabolisable, or even the different types of net energy systems have been
developed for them. Following the “utilization ladder”, the energetic evalua-
tion system of the different contries are as follows:
9.2.3.1. Digestible energy (DE) expressed in megajoule. In pig nutrition:

Hungary and the United Kingdom; for horses and rabbits in Germany,
Hungary, United Kingdom and United States (in Mcal).
9.2.3.1.1. PIG NUTRITION

Digestible energy content of feedstuffs or feed mixtures can be calcu-
lated by using of the equation elaborated in Rostock (SCHIEMANN et al. 1969):
DE, MJ/kg feed = 0.0242 dCP + 0.0394 dEE + 0.0184 dCF + 0.0170
dNFE, where dCP= digestible crude protein, dEE=digestible ether extract,
dCF= digestible crude fibre and dNFE= digestible N-free extract, each items
in g/kg.
Example: kg corn grain contains 86 g crude protein, 46 g ether extract, 20
g crude fibre and 703 g N-free extract. The approprate digestion coefficients
(in the same order) are: 81, 65, 57 és 94%. Putting these values in the above
described equation:
DE, MJ/kg = 0.0242×0.81×86+0.0394×0.65×46+0.0184×0.57×20+0.0170×
0.94×703 = 14.32 MJ/kg, accordingly l kg of maize grain represents 14.32
MJ digestible energy for pigs.
The new (2002) Danish energetic evaluation system is based on the,
in ileum enzymacically digestible, dry matter and the protein needs are
given in true ileal digestible protein. It has a separate unit for growing-fat-
tening pig (FUgp) and for breeding sows (FUgs). Some examples: FUgp for
barley, wheat and soybean are 1.05, 1.16 és 0.88, respectively.
9.2.3.1.2. RABBIT ENERGY EVALUATION SYSTEM

In the Hungarian Feed Codex (2003) and in the US NRC (1977) the
digestible energy is used, in Europe it is expressed in MJ and in the US in
kcal. The metabolizable energy is also apply for scientific purposes and used
by the leading specialist in France, Spain and Italy.
For the energy content calculation of feedstuffs and feed mixture, the
following equation is proposed: DE, MJ/kg= 0.022 dCP + 0.04 dEE + 0.018
dCF + 0.018 dNFE (Explanation of abbreviations see above, at the pig.)
The metabolizable energy value of a balanced feed mixture can be cal-
culated after (FEKETE and PAPP, 1980): ME = 0.945 × DE.
9.2.3.1.3. HORSE ENERGY EVALUATION SYSTEMS

In the majority of the country the energy system of horses are based
on digestible energy (Germany, Hungary, United Kingdon: in MJ and USA:
171
in Mcal). A net energy based energetic evaluation system has been devel-
oped for horses in France (INRA, 1984); in Roumania it was adapted with-
out significant changes. It is expressed as a barley unit, UFC (Unité
Fourragère Cheval). The maintenance requirement of adult horses is 0.038
UFC×W0.75, i.e. for a horse of 500, 600 and 700 kg of LW 4.0, 4.6 and 5.2
UFC. A 2-hour long work of medium intensity for a horse of 500 kg of LW is
3.2-3.4 UFC. Energy value of some typical horse feedstuffs are as follows:
straws 0.22-0.32, meadow hay 0.38-0.57, alfalfa hay 0.43-0.54, oats 0.88,
barley of reference 1.00 and maize (corn) 1.14 UFC/kg air dry matter.
In Russia the metabolizable energy is used for horse energy evalua-
tion, the form of expression is MJ. The appropriate equation for calculating
the ME content in horse feed is the following (using the above described
abbreviations):
ME, MJ/kg= 19.64 dCP+35.43 dEE+15.95 dCF +15.95 dNFE
9.2.3.2. Metabolizable energy (ME)

9.2.3.2.1. In megajoule expressed metabolizable energy is used for all
animal species in Sweden and in the former USSR.
9.2.3.2.2. In the nutrition of beef cattle and small ruminants in
Austria, Germany and Poland.
9.2..3.2.3. A special form of ME has been developed in Germany for
the pig nutrition and it is also applied in Austria and Poland. This ME is cor-
rected according to sugar (above 80 g/kg DM) and to the BFS i.e. bakteriell
fermentierbare Substanz, mainly hemicellulose and pectin (above 100 g/kg
DM) of the give feedstuff. For the calculation of ME content the following
equation is appropriate:
ME, MJ/kg DM= 0.021dCP+0.0374dEE+0.00144dCF+0.017dNFE-
0.0014SUGAR-0.0068 (BFS-100)
9.2.3.2.4. The USA (1998) pig recommendation gives data in ME,
expressed in Mcal. There is a possibility to correct according to the protein
level:
ME= DE × (96-[0.202 × CP%]) (NRC, 1981), or
ME, kcal/kg= DE×(1.012-[0.0019×CP%]), (NRC, 1998).
9.2.2.3.5. For fish and reptile the application of ME data is also pro-
posed.
9.2.2.3.6. For carnivores (dog, cat, ferret, fox and mink) the use of ME
system is practised (for more details see Chapter XXI).
172
9.3. ENERGY EVALUATION OF POULTRY FEEDS
© Fatma Karakas Ogüz
In the majority of contries the metabolizable energy is in use, either

expressed in kcal or in MJ. More precisely, the application of the AMEn is
common, where “A” stands for apparent, and “n”” means that values are
corrected to zero N-retention. One of the reasons that the ME has been used
for poultry is technical: birds exctetes faeces and urine together and to
measure the metabolisable portion of a feed is easier than the determina-
tion of the digestion coefficiets. To measure of the digestion coefficiet in
birds the use of an artificial anus (for the mechanical separation of faces
and urine) or the chemical determination of the urine-N in the mixed exc-
reta are applied. When measuring the metabolisable energy portion of a feed
the burning value of mixed excreta is measured by using a bomb-calorime-
try. The term “apparent” expresses that the excreta contains not only the
undigested part of the feed, but also enterocytes residue of the digestive
juice which do not derive from the actually measured feedstuff.
To make the comparison of the ME values possible, they are
expressed in the form of zero nitrogen retention. This means that the pro-
tein accretion is considered as oxidized to uric acid supplying energy for the
organism (FIGURE IX-2a and a FIGURE IX-2b). Beigng the combustion heat
of the uric acid-N 34.4 kJ, during the correction each gram retained N
means minus 34.4 kJ discount from the experimentally determined appar-
ent ME valus. (In case of protein mobilisation, e. g. laying hens, the same
value is added to the apparent ME.) It is worth emphasising that from the
biological (growth) and economical pont of view the protein retention is
adventageous, but it is a “lost” one for the energy metabolism.
Knowing the chemical composition of bird feedstuffs and feed mix-
tures the AMEn can be predicted (calculated) by means of the HÄRTEL (1977)
equation:
AMEn, MJ/kg = 17.21×CP+34.82×EE+(18.5 -31.2×CF)×NFE- 3.064,
where CP= crude protein (g/kg), EE= ether extract (g/kg), CF= crude
fibre (g/kg) and NFE= N-free extract (g/kg).
The equation does not give a good prediction in case of sugars, milk pow-
der, sugar beet pulp, potatoe starch, oils and tallow.
Protein needs are generally given in crude protein (N× 6.25). Values in
requirements tables suppose an optimum temperature. In other environ-
mental temperatures both voluntary feed intake and nutrient requirements
will change, consequently, data should be adapted. In this respect the most
important physiological feature of birds is that feed intake is strongly con-
trolled by the energy density of feed mixture, and therefore, all the other
nutrients should be adjusted to the energy level.
173
FIGURE IX-2a: Scheme of the positive N-balance
FIGURE IX-2b: Scheme of the negative N-balance
Legend: Illustration of zero N-retetion corrected metabolizable energy in case of pos

itive N-balance (protein accretion, 2a). ME= DE-urine energy; ME= 100-5= 95 unit; MEn=
ME-20= 75 unit.), and in case of negative N-balance (protein mobilisation, 2b)
174
The European and North American recommendations are based on

the descibed apparent, zero N-retention corrected metabolisable energy
(AMEn) expressed in kcal or in MJ. The French INRA (1984) use N-correc-
tion for 40% (one-day-chicken) and 30% (growing chicks and laying hens)
N-retention.
SIBBALD (1976) developed a system based on “true” metabolisable
energy (TME), which takes of the endogen and metabolic nutrient losses into
consideration. On an average, 1 unit TME equals to 1.097 AME.
In Russia the “energetic feed unit” (ЭKE) is in use, which equals to 10
MJ ME. Calculation of ME for poultry is based on the knowledge of
digestible composition of feeds:
MEpoultry=17.84×dCP+39.78×dEE+17.71×dCF+17.71×dNFE, where dCP=
digestible crude protein, dEE=digestible ether extract, dCF= digestible crude
fibre and dNFE= digestible N-free extract, each items in g/kg.
9.4. Net energy
9.4.1. Rostock energy evaluation system

From 1971 it used to be official for all animal species in the former East
Germany. Similarly to the Starch Equivalent, it was based on fat-producing
net energy. Using multiple regression equations they were able to improve
the accuracy of prediction. In the 90th developed the system to an ATP
based one (CHUDY).
9.4.2. The UK ME–NE system

The new system have been introduced on BLAXTER’s proposal in 1975. The
essential character of this is that instead of the net energy value the
metabolisable energy content of feedstuffs has been tabulated, but using
correction factors, partial net energy (maintenance, milk production) can be
calculated.
9.4.3. Californian net energy sytem for beef cattle

The method developed by LOFGREEN and GARRETT (1968) uses two types of
net energy (“partial net energy”): net energy for maintenance (NEm) and net
energy for gain (NEg). The fundamental data is measured by using indirect
calorimetry (heat production, extrapolated on zero feed intake) for mainte-
nance while body gain of 844 cattle is measured by comparative slaughter
techniques (see Chapter III and V). The actual NRC (2000) beef cattle rec-
ommendation is based on the above pronciples.
9.4.5. Beltsville dairy cow system

Based on MOE and TYRREL (1972) foundings research gives the basics of the
175
today’s NRC-recommendations (2001) for dairy cows. Feeding 543 dairy

cows on 32 different feed mixtures they measured the net energy content in
them. Beside maintenance and milk production the physiological energy
mobilisation and retention have been determined.
9.4.5. European cattle systems

Although the metabolisable energy is really available for the cells of the ani-
mal organism, ist utilisation for maintenance of production mainly depends
upon two factors: the partial transformation factor („k“-value,
k=NE/ME×100)), which is the best for heat production (100%), very good for
maintenance (70-80%), good for milk production (60-70%), mediem for body
gain (30-50%) and bad for building pregnancy tissues and foetus (10-15%).
Among the above described ranges of „k“-value, there is the degree of
rations metabolizability, the „q“-value (q=ME/BE). By increasing the con-
centrate proportion in a feed mixture, the q-value will increase, too, improv-
ing the efficacy of ME-utilisation for net energy, while decreasing heat pro-
duction (FIGURE IX-3).
FIGURE IX-3: Change of the “k”-value in the function of “q”-value
It is worth mentioning the effect of the feeding level (amount of ingest-

ed feed). The increasing feed intake decreases efficacy. To be able to take
into consideration all of the described factors, in the cattle nutrition it
seemed to be advisable to use special net energy units. In Vichy (1976) a
special committee of the European Association of Animal Production made
a decision that based on researches of VAN ES and HARKINS (1975) according
to which new cattle energetic evaluation systems should be developed.
176
Consequently, the today’s systems are based on the same scientific princi-
ples, but the form of expression differs according to each individual coun-
try.
For DAIRY COWS the common ground basis was to consider different
utilisation efficacy factor of metabolisable energy according to physiological
purposes: the partial transformation factor for maintenance (km) is higher
than for milk production (kl), and for growth (muscle and fat accretion it is
even lower. The effect of feeding level and the metabolizability (ME/BE) have
also been taken into consideration. The final mode of expression differ
according to the local pecularities. In Austria, Germany and Switzerland the
unit is the NEL (Netto Energie Lactation) in MJ. In Belgium and in the
Netherland the VEM (Voedereenheidmelk) has been introduced, which is
the net energy laction value, transformed in barley unit of 1650 kcal/kg.
The French INRA system use a reference barley of 1.730 Mcal net energy
laction and the final unit is the UFL (Unité Fourragère Lait). 1 UFL = 1000
VEM = 6.9 NEL. In Hungary the net energy laction is used, expressed in MJ.
The values of Tables correspond to a 3fold feeding unit (approximately 24
kg milk production). For the maintenance of dairy cows 0.3347×W0.75 MJ
NEl and 4.035 g is calculated. During the last two months of gestation the
total need of maintenance and pregnancy is 0.435×W0.75 MJ NE1. For the
production of 1 kg milk with 4% fat 3.10 MJ NE1 and 87 g crude protein are
given. Data are valid for an environmental temperature of 20oC. For high-
yielding cows the application of metabolizable protein (MP) system is pro-
posed.
In case of beef cattle the practice shows more variability. Austria,
Belgium and Germany introduced a metabolisable energy system; in
Switzerland the unit is the Netto Energie Wachstum (NEW) in MJ. In the
Netherland and in France the final form of net energy gain is expressed in
barley unit, called VEVI (Voedereenheid Vetmesten Intensief) and UFV
(Unité Fourragère Viande). 1 UFV = 1060 VEVI = 7.3 NEW. In case of low
growth rate (e. g. growing breeding heifer, wintering beef cattle) the dairy
cow units are used.
In Hungary the maintenance energy requirements of beef cattle is
given in net energy maintenance (NEm, in MJ), those of body weight gain in
net energy gain (NEg, MJ). Extra needs for pregnancy and milk production
are calculated by means of NEm. In every cases (calves, breeding heifers)
both NEm and NEg are in use. Maintenance nedds of beef cattle can be cal-
culated according to the NEm, MJ= 0.3222×W0.75 equation. NEg needs
depend upon the frame size, gender and average daily gain.
9.4.6. Energetic evaluation in sheep and goat

In Austria and Germany the ME system is used for small ruminants. (GFE,
2003) In France, like for other ruminants, they use the UFL (unité four-
ragère lait), UFV (unité fourragère viande), but a proper fulfilment unit
177
(UEM=unité encombrement mouton) has been developed for sheep (INRA,

1988). In the UK the ME (AFRC, 1993) and in the USA (NRC, 2006) the ME,
NEm and NEg have been introduced.
In Hungary the requirement of sheep is calculated similarly to the
beef cattle, i.e. using NEm for the breeding mothers and both NEm and NEg
for growing animals. The numerical value of maintenance requirement for
sheep is lower than that is for cattle: 0.245×W0.75 MJ NEm, which can be
explained by lower heat losses.
FOR FURTHER FEADING
ARC (1980): The nutrient requirements of ruminant livestock. Commonwealth Agricultural

Bureaux. Farnhan Royal. Slough.
ARC (198): The nutrient requirements of pigs. Commonwealth Agricultural Bureaux.
Farnhan Royal. Slough.
Jarrige, R. (1988): Alimentation des bovins, ovins & caprins. INRA, Paris.
Jarrige, R. and Martin-Rosset, W. (1984): Le cheval. Reproduction, Selection, Alimentation,
Exploitation. INRA, Paris.
Kamphues, J., Coenen, M., Kienzle, E., Pallauf, J. Simon, O. and Zentek, J. (2004):
Supplemente zu Vorlesungen und Übungen in der Tierernährung. 10., überarb.
Auflage. Alfeld-Hannover. Verlag M. & H. Scharper
NRC (1981): Nutritional energics of domestic animals. National Academy of Press.
Washington, D. C.
178
JAV_10-1_VND_FSGy:MINTAÚJ.qxd 2008.09.03. 14:42 Page 179
CHAPTER X: PROTEIN EVAULATION- MONOBASTRICS
Chapter X
PROTEIN EVALUATION METHODS AND SYSTEMS

FOR MONOGASTRIC AND RUMINANT ANIMALS
© Carlos CASTRILLO Gonzales, Eva Cenkvari

and Sandor Gy. Fekete
10.1 Monogastrics animals

10.1.1. Amino acid and protein requirement of monogastrics
10.1.2. Procedures for determination of protein quality
10.1.2.1. Chemical analysis of nitrogen and protein
10.1.2.2. Amino acid composition and indices
10.1.2.3. In vitro enzymatic methods
10.1.2.4. Microbiological methods
10.1.2.5. Macrobiological methods
10.2.3. New tendencies in protein evaluation of monogastrics
10.2. Protein requirements of ruminants and protein
evaluation of feedstuffs
10.2.1. Basic aspects of rumen protein metabolism
10.2.1.1. Origin of the amino acids entering the duodenum
10.2.1.2. Factors affecting protein degradability
10.2.1.3. Factors affecting microbial protein synthesis
10.2.1.4. Measurement of feed protein degraded in the
rumen
10.2.2. Consideration of protein evaluation for ruminants
10.2.3. Shortcomings of old crude protein and apparently
digestible crude protein systems
10.3. Overview of protein evaluation systems in different countries
10.3.1. Main steps in the calculation of the metabolizable protein
supply
10.3.2. The UK metabolizable protein system.
10.3.3. The French PDI system
10.3.4. The Hungarian System
10.3.5. Shortcomings of current European systems
10.3.6. Evaluation system of “absorbed protein”
(USA, NRC, 1985).
10.3.7. New aspects introduced in the Cornell (CNCPS)
proposal
179
CHAPTER X: PROTEIN EVAULATION - MONOBASTRICS
I f the energy requirement of the organism is covered, the limiting fac

tors of life manifestations is the protein, included the amino acid sup
ply. The reason for that is that while energy can be got within the phys-
iological ranges from carbohydrates, fats and even from protein, some struc-
tural units of the protein should be received preformed in the feed. This is
why the protein quality is one of the central topic of human and animal
nutrition.
10.1 Monogastrics animals
10.1.1. Amino acid and protein requirement of monogastrics

In the 1980’s, the protein requirement of monogastric mammalians was
expressed in crude protein. Below, it is necessary to make clear some rele-
vant terms and definitons connected to protein evaluation. The method of
Kjeldahl determines the nitrogen content of feedstuffs, which is to be mul-
tiplied by 6.25 to gain the value of crude protein. The proportion of crude
protein, which remains in the organism „apparently” is based on the classic
metabolic trials. It is called digestible crude protein and it is regarded as
digestible true protein which can be precipitated with copper sulphate.
In monogastrics, the protein requirement values can be determined in
two basic ways. One is the factorial calculation, when all the possible and
necessary protein outputs (protein losses) are added. Then the protein
quantity is given, which is necessary to cover the protein losses from trans-
formation (digestion, building the tissues). Another opportunity is to deter-
mine the protein requirement of growth to give diets or different energy and
protein levels, and body gain, feed conversion ratio, body composition and
180
carcass quality are measured.

Monogastric animals have quantitative and qualitative demands
towards protein supply. Protein quality is determined basically by digestibil-
ity, biological value and antinutritive substances possibly occuring.
Digestibility means that feed protein degrades to amino acids and becomes
available for animals via absorption in the small intestine. Biological value
(BV) shows the proportion of digestible protein, which can be built into the
body tissues. The latest has the precondition that the amino acid pattern
should be adjusted to the demands of amino acid synthesis, and it has to
be biologically available (e.g. availability of lysine is decreased by Maillard
reaction).
Roles of amino acids can be well illustrated by the so-called minimum
law of Liebig, which has been developed for the physiology of plants. It
declares that the growth of plants depends always on the essential nutrient,
which is available at the lowest level. Most of the time lysine is the first lim-
iting amino acid is mostly. When its quantity is increased, the amino acid
limits the availability of the others, which is of the second lowest quantity.
These are usually sulphur-containing amino acids like methionine and cys-
tine. The third and fourth limiting amino acids are mostly tryptophan and
threonine, especially in swine and poultry feeding based on maize and soy-
bean.
In practice, it is not possible to consider quantitatively all the essen-
tial amino acids. Therefore, it is introduced into feeding practice that the
quantity of the most important essential amino acid, like lysine, is declared
and its values of requirement are precisely given (in grams per kilogram feed
or grams per MJ DE or in percent of protein). Afterwards, the ratios of amino
acids are determined compared to lysine in the so-called ideal protein (FIG-
URE X-1). Requirement values of these amino acids are given regarding the
proportion of lysine as 100 percent.
181
FIGURE X-1: Composition of ideal protein (after ARC, 1980)
The term ‘IDEAL PROTEIN’ defines a feed protein, which has an adequate
amino acid composition for the amino acid requirements of tissue synthe-
sis. It considers two basic prerequisites, like the digestibility of proteins and
amino acids and availability of the latter ones are excellent. In practice, the
precise quantity of lysine is given, and the quantities of other amino acids
are determined as the proportion of lysine. Table X-1 shows the quantitative
requirements for lysine and the proper ratios of essential amino acids.
Table X-1: Lysine requirement of fattening swines and the proper ratios of essential amino
acids in their feed mixtures
*Maximum 1/3 cysteine and tyrosine can be includeded, respectively.
It is noteworthy that the basis of comparison, the absolute lysine
182
requirement is higher according to the recommendation of ARC (1981).

There is also an approach in which the ratios of essential amino acids are
given as the multiplication of tryptophane demanded in relatively the low-
est quantity. Table X-2 represents four of the most important amino acids
in the significant protein sources of plant origin.
Table X-2: Four of the most important amino acids in the significant protein sources of
plant origin.
*Lysine is considered as 100 percent **Canola rape
It is obvious that soybean contains relatively low levels of sulphur-

containing amino acids, but sunflower meal (extracxted solvent) includes
methionine+cystine, tryptophan and threonine in abundance. It gives us the
opportunity to make the diets complete, e.g. by the partial replacement of
soybean. Rapeseed and cottonseed can serve as a supplemental source of
threonine while alfalfa meal is a good source of tryptophan and threonine.
It is important to note that even the best of proteins of animal origin like the
herring meal does not include adequate S-containining amino acids, there-
fore it can be well supplemented with sunflower meal. According to the com-
position, the ideal proteins gave different results in the practical feeding. A
possible reason for this might be that the digestibility and utilization of pro-
teins, besides their amino acid compositions can also be modified by other
nutritional factors (availability, crude fibre content, antinutritive sub-
stances).
10.1.2. Procedures for determination of protein quality
10.1.2.1. Chemical analysis of nitrogen and protein

Method for nitrogen determination of Kjeldahl. Since all the inorganic and
183
organic nitrogens are converted to ammonium-sulphate during sulphuric

acid destruction, we get a value including also non-metabolizable nitrogen.
We get only an approximate value for protein content, when nitrogen con-
tent is multiplied by a factor of 6.25, since the nitrogen content of proteins
differs from the content of 16%. Therefore, in case of the most significant
feedstuffs, the real factors are applied, like e.g. 5.8 in case of wheat and
6.38 for milk protein. The method described above can be replaced by pho-
tometric measurement (by reagent of Nessler, by biuret etc.)
10.1.2.2. Amino acid composition and indices

CHEMICAL SCORE (CS). Individual amino acids are expressed as a percentage
of certain essential amino acid of a reference protein (e.g. egg, milk, WHO-
protein). The essential amino acid included at the lowest level is called lim-
iting and its value is the chemical score. For example, the lysine level of egg
protein is 7.2% and the one of the wheat protein is 2.7%, so the chemical
score of wheat protein is 2.7/7.2=0.37 or 37%. Table X-3 shows the amino
acid levels recommended as references by FAO-WHO (1985).
Table X-3: Ideal amino acid mixtures recommended by FAO-WHO (1985) for human use*
*FAO = Food and Agricultue Organization; WHO = World Health Organization;

UNU = United Nation University
EAA INDEX (EAAI): essential amino acid index. The value of proteins is char-
acterized by the geometric mean of essential amino acids, while it is com-
pared with the adequate amino acid composition of the whole egg (e.g. EAAI
of barley is 67). Table X-4 shows EAAI values of the most significant protein
feeds and CS values of the limiting acids at the first and at the second and
at the third level. Critics of the indices: they do not reflect the changes in
digestibility and the availability of amino acids. (Available amino acid: pro-
184
portion of digestible protein, which can be utilized for protein synthesis. It

can be detected –among others- by die-binding capacity in the case of basic
amino acids like in the case of lysine.
Table X-4: EAAI and CS values of the most significant protein feeds
*Compared to milk protein
10.1.2.3. In vitro enzymatic methods

Based on the data gained by in vitro enzymatic methods indices can be
developed, which show high correlation with the results of biological meth-
ods. Pepsin Digest Residues (PDR) index shows significant relationship with
the values of NPU (see in 10.1.5.2. below), because pepsine digestion has a
significant role in protein utilization. Pepsine-Pancreatin Digest index (PPD)
is a similar parameter, where digestion performed by pepsine and supple-
mented also by pancreatin in order to simulate the effect of pancreatic juice.
10.1.2.4. Microbiological methods

Basically, they are used for the quantitative determination of amino acids in
protein hydrolisates or for the determination of „Relative Nutritive Value”.
The latter expresses the extent at which protein can promote the prolifera-
tion of microbes.The two main types are as follows: a) protein hydrolisate
can be measured by non-proteolytic bacteria (Leuconostoc mesenteroides,
Streptococcus faecalis) and b) the intact test protein can be measured by
proteolytic bacterium (Streptococcus zymogenes) or protozoon (Tetrahymena
pyriformis), respectively.
10.1.2.5. Macrobiological methods

10.1.2.5.1. Based on the changes in body weight:
a) Tenebrio molitor growth test measures the body weight gain of the larvae
185
of meal beetle. The disadvantage of the method is high sensitivity against

antinutritive factors. b) Protein Efficiency Ratio, PER: body weight gain
caused by one unit of protein intake. Its disadvantage is that the protein
evaluation is based exclusively on the growth and it does not indicate the
quality of protein for maintenance. Determination is to be performed in
standard conditions: 4 weeks feeding ad libitum for male rats 10 percent
protein in the diet. The latter is needed because the same protein has dif-
ferent PER value, tested in various protein concentrations (N-intake).
c) Net Protein Ratio (NPR): it compares the final body weights of test groups
consuming or not consuming protein, and it compares the differences of
body weights to the protein intake. While PER evaluates proteins based
solely on growth, NPR qualifies them taking growth and maintenance
together into account. d) Slope ratio by Hegsted: it is equal to the measure-
ment of NPR but applying some protein levels and mathematical process-
ings, which eliminate the influences of different feed (and protein) intake. It
is referred to as Nitrogen Growth Index (NGI). Its calculation is based on the
determination of the angle of slope decribing the correlation of N-consump-
tion and weight gain.
10.1.2.5.2. Based on the changes in N-content of the organism

N-retention can be determined by the measurement of the consumed and of
the excreted nitrogen, or by that of the increase in N-content of the body
(body analysis). When the amount of nitrogen built in is compared to the
quantity of absorped nitrogen, the index of Biological Value is gained.
Furthermore, when a comparison is made to the total quantity of consumed
N, Net Protein Utilization (NPU) is calculated. Knowing the digestion coeffi-
cient (DC) of protein, NPU can be calculated from BV (and in revers: NPU =
DC×BV.). Table X-5 shows the Biological Values (%) of some significant feed-
stuffs. Note that the Biological Values of a mixtures might exceed the ones
of each component, when their amino acids complement each other.
186
Table X-5: Biological Values (%) of some important feedstuffs
10.1.2.5.3. Based on the metabolism of organs and tissues

It is well known that the sensitivity of organs (organisms) is different
towards protein supply. Some methods are based on the effects of different
feed proteins on regeneration in the organs, which are free of their protein
stores (depleted). Serum protein regeneration: plasmapheresis is induced
(significant quantity of blood is taken, plasma is replaced by physiological
solution of salt, which is then introduced into the bloodstream. PAAR
(Plasma Acid Ratio): concentrations of amino acids are determined in plas-
ma during starvation and then perodically after feeding of a protein source.
Difference between them is written in the numerator and the requirement,
specific for species is in the denominator, respectively. PAAR is calculated
for each essential amino acid. The smaller is the PAAR value of an amino
acid, the more limiting is the protein source fed as test protein. Liver
Protein Utilization (LPU) index is gained by the development of a method
based on liver protein regeneration. Final differences of nitrogen contents
based on the livers of animal groups having protein diets or of non-protein
diets are referred to nitrogen intake, as in the case of the NPU method.
Reverse Urea Value (RUV): the better is BV of a protein source, the less
urea is unutilized, and the lower will be the urea level of blood. Namely, urea
level of blood is negatively correlated with BV.
The sensitivity of organs towards protein supply is as follows: BLOOD >
187
LIVER > MUSCLE > KIDNEY > BRAIN. Muscle protein turnover can be mon-
itored by the urinary excretion of creatinine and 3-methyl-hystidine.
10.2.3. New tendencies in protein evaluation of monogastrics

Calculation with available and absorbable amino acids: technics of deter-
mination in poultry is based on caecotomia and in swine on ileo-rectal
shunt, respectively (see Chapter VII). It is visible from the data of Table X-6,
that the two evaluations are significantly different.
Table X-6: Apparent faecal (AD) and ileal digestibility (true digestibility, (TD) of lysine, %
188
CHAPTER X: PROTEIN EVAULATION - RUMINANTS
10.2. Protein requirements of ruminants and protein evalua-

tion of feedstuffs
Introduction
Purpose of using protein evaluation systems for ruminants

Protein systems try to define, as precisely as possible, both the supply of
amino acids from the gut to the tissues and amino acid requirements to
meet maintenance and production needs. In both monogastrics and rumi-
nants the protein value of the diet is related to the quantity and type of
amino acids reaching the duodenum, the proportion that is absorbed and
how efficiently they can be used to cover amino acid requirements. Both
monogastrics and ruminants need essential amino acids that they cannot
synthesise and also non-essential amino acids that provide ammonia for
transamination reactions. Thus, requirements of absorbed amino acids are
essentially the same in monogastrics as in ruminants. The great difference
between the two is the origin of amino acids reaching the duodenum.
Whereas in monogastrics they are essentially the same as those supplied
with the diet, in ruminants only a variable fraction of dietary amino acids
escapes ruminal degradation and reaches the duodenum, and a great part
of the amino acids arriving in the duodenum come from microbial protein
synthesised de novo in the rumen.
The quantity of microbial protein reaching the duodenum depends
essentially on the availability of energy for microbial maintenance and
growth. Consequently, the protein value of feeds and diets in ruminants is
relatively independent of their amino acid profile, depending more on the
degradability of dietary protein and on the intake of fermentable carbohy-
drate, as well as on other factors than may affect both protein degradabili-
ty and microbial protein yield, such as rumen content outflow rate and frac-
tional outflow rate of particles leaving the rumen. The new protein systems
for ruminants try to take into account the contribution of both microbial
protein and undegraded food protein to the total amino acids reaching the
duodenum, and are based on the recognition of the essential principles of
ruminal and post-ruminal protein digestion and the metabolism of absorbed
amino acids.
189
10.2.1. Basic aspects of rumen protein metabolism
10.2.1.1. Origin of the amino acids entering the duodenum

In ruminants most of their energy requirements are satisfied by the volatile
fatty acids that result from carbohydrate fermentation by rumen microor-
ganisms, mainly bacteria and protozoa. The fermentation of carbohydrates
and, to a lesser extent, of proteins, provides microorganisms with energy, in
the form of ATP, for their maintenance and growth, thus the extent of micro-
bial synthesis depends on energy availability, mainly from carbohydrates.
As microorganisms mostly consist of protein, their growth involves the con-
tinuous synthesis of microbial protein that flows to the duodenum, supply-
ing the largest part of the amino acids absorbed by the host organism in
most feeding conditions. The nitrogen for microbial protein synthesis is
obtained from peptides, amino acids and mainly ammonia, coming from the
microbial degradation of feed nitrogen compounds (true protein and non
protein nitrogen, NPN) and from the urea recycled to the rumen. The sup-
ply of enough degradable protein and NPN to satisfy the nitrogen require-
ments of microorganisms is a major target in diet formulation. These
requirements are directly related to energy availability for microbial growth.
The value of dietary urea and other NPN sources for ruminants
depends on its degradation to ammonia in the rumen by bacteria, which is
carried out quickly, and on the subsequent use of this ammonia for micro-
bial protein synthesis. The ammonia that is not used for microbial protein
synthesis is absorbed in the rumen or further on in the intestines and
excreted as urea in urine or recycled to the rumen via saliva or diffusion
through the rumen wall. Degradation of true dietary protein in the rumen
varies with different protein sources and different processing treatments.
The part that misses degradation supplements microbial protein in provid-
ing amino acids for digestion and absorption in the small intestine. FIGURE
X-2 schematically shows the degradation processes of the dietary nitrogen
compounds and the synthesis of microbial protein that take place in the
rumen.
190
FIGURE X-2: Schematic diagram of the fate of dietary protein in ruminants
The processes of degradation and synthesis that take place in the

rumen determine the quantity and profile of the amino acids absorbed by
the host. In most conventional diets the microbial protein fraction repre-
sents between 40% and 70% of total non ammonia nitrogen arriving in the
duodenum, and the undegraded dietary protein between 20% and 50%. A
small, more constant part (10%-15%) is formed by endogenous nitrogen.
The contribution of the microbial protein to the host protein requirements
depends on the physiological state and level of production. The microbial
191
protein may be enough to cover protein requirements for maintenance and

even the host’s protein requirements at the end of the growth stage and the
beginning of pregnancy, assuming an energy supply with a sufficient diet to
cover energy requirements. Consequently, during these stages it would be
enough to satisfy degradable protein requirements to obtain optimal micro-
bial growth. During stages of higher demands, such as early growth, the end
of pregnancy and early and mid lactation, microbial protein also contributes
to more than 60% of host requirements, but it is necessary to supplement
microbial protein with a considerable quantity of rumen undegradable
dietary protein. From what has been said up to now, we can deduce the
importance of optimizing microbial synthesis in the rumen, and of having a
good knowledge of the mechanisms and factors that affect dietary protein
degradation in the rumen.
10.2.1.2. Factors affecting protein degradability

The degradation of dietary protein in the rumen is carried out by the micro-
bial action of enzymes, chiefly by bacteria but also by protozoa, and proba-
bly fungi. The NPN COMPOUNDS (mainly ammonia, urea, amides and amino
acids) are quickly degraded to ammonia, while the degradation of proteins
takes place more slowly. The process in the case of bacteria requires close
contact between the protein and the bacteria cell, since most microbial pro-
teases are bound to the external surface of the cell wall. This contact occurs
by means of the adsorption of the soluble proteins to the bacterial surface
or by adhesion of bacteria to the insoluble proteins. In the case of the pro-
tozoa, feed particles and also bacteria are engulfed and subjected to the
action of their digestive enzymes. PROTEIN degradation is a multi step
process involving solubilization, extracellular proteolysis, transport of
amino acids and peptides inside the bacterial cell and amino acid fermen-
tation. In many cases protein solubilization is the rate-limiting step but
depending of protein characteristics any of other steps may also limit pro-
tein degradation. Thus casein is hydrolysed very quickly and short peptides
may accumulate temporarily whereas with albumin, a protein that is
hydrolysed slowly, little peptide is detected in the extra cellular nitrogen
pool. The RATE OF DEGRADATION and the potentially degradable protein of foods
in rumen depend on the enzymatic activity of ruminal content and on the
characteristics and accessibility of the protein. Nevertheless, the amount of
protein actually degraded also depends on the retention time of food parti-
192
cles in the rumen.

10.2.1.2.1. Enzymatic activity of ruminal content
Although some ruminal bacteria show a higher proteolytic activity than the
rest, as is the case of Prevotella ruminicola and Peptoestreptococcus, prote-
olytic activity in the rumen seems to be more related to total microbial con-
centration than to the bacterial profile, since nearly 50% of ruminal bacte-
ria have proteolytic activity. Consequently, enzymatic activity depends fun-
damentally on those factors that determine bacterial growth, mainly energy
and nitrogen availability. Low concentrations of ammonia can limit prote-
olytic activity in the rumen due to a concomitant reduction of microbial
growth and, the opposite situation, i.e. high concentrations of ammonia can
cause a decrease in proteolytic activity, perhaps through retro-inhibition
mechanisms.
Moreover, the decrease of rumen pH, along with the supply of high
concentrate diets, may negatively affect the solubility of proteins and thus
their degradability due to the fact that the isoelectric point, or pH at which
the proteins show higher stability, is generally acidic. It is also necessary to
keep in mind that at a pH below 5.5 the growth of protozoa (and therefore
the survival of their symbiotic bacteria) is inhibited.
10.2.1.2.2. Protein characteristics
Regardless of proteolytic activity and the rumen ambience, the rate of degra-
dation of feed proteins depends on their physical-chemical characteristics,
and essentially on their solubility and structure.
SOLUBILITY. A higher solubility allows a higher accessibility of proteins to
bacterial proteases and, in general, the most soluble proteins are the most
susceptible to degradation. It is for this reason that protein solubility in dif-
ferent tampons has been widely used as an indication of their degradabili-
ty in the rumen. From the beginning of the twentieth century proteins have
been classified according to their degree of solubility in different solvents, in
albumins (soluble in water), globulins (soluble in saline solution), glutelins
(soluble in diluted alkaline solutions) and prolamins (soluble in alcoholic
solutions). The proportion of each one of these fractions in feed protein
explains, to a great extent, their potential degradability in the rumen.
STRUCTURE. Solubility is not the only factor that determines the poten-
tial of protein degradability in the rumen, since, for example, casein is bare-
ly soluble and yet it is highly degradable in rumen, whereas lactalbumin
and ovoalbumin, although very soluble, have a higher resistance to ruminal
193
degradation. In addition, the soluble and insoluble fractions of soy bean

meal have shown similar in vitro degradation rates against proteases. The
resistance to ruminal degradation of ovoalbumin seems to be related to its
peculiar primary structure, characterized by an acetyl amino acid at the
ends. Nevertheless, it is the tertiary and quaternary structure, and in par-
ticular the existence of cross-links in the protein molecule that determines
in great part their susceptibility to degradation. The importance of the pres-
ence of disulfide bonds on protein susceptibility to microbial degradation
has been clearly put in evidence by WALLACE (1983), who showed that the
rate of degradation of lactalbumin and albumin of bovine serum, which are
rich in disulphide bonds, was lower than that of casein, which is less solu-
ble but has a very low disulphide cross-links content (Table X-7). When the
disulphide bonds were broken down by means of performic acid oxidation,
the rate of degradation of lactalbumin and albumin increased, becoming
similar or even higher to that of casein. The presence of disulphide bonds is
also one of the reasons that explains the lower degradability of corn protein
(zein) compared to that of wheat (gliadin).
HEAT TREATMENT of proteins produces changes in the molecular struc-

ture of the protein, related to its partial denaturation and Maillard reac-
tions, and the protein becomes more resistant to ruminal degradation. If the
treatment is not too intensive, the changes in protein structure may be
reversible at the low pH found in abomasum, becoming available to enzy-
194
matic digestion in the intestine. Thus, the protein of fish and meat is origi-
nally highly degradable, whereas fish and meat meals are characterized by
their low degradability. The same occurs with soybean protein and other
oilseeds that are more quickly degraded in the rumen when offered as raw
seeds than when offered as cakes after oil extraction, a process that always
involves some heat treatment. Other chemical treatments, such as the appli-
cation of formaldehyde or tannins also reduce rumen degradability of pro-
tein supplements, and the presence of natural tannins in the plants affects
their protein degradability.
10.2.1.2.3. Protein accessibility
Although rumen protozoa have a high proteolytic activity, mainly acting on
engulfed feed particles and bacteria, these are responsible for most rumen
proteolytic activity. Since bacterial photolytic enzymes are closely associat-
ed to microbial cell walls, the degradation of feed protein requires close con-
tact between the substrate and bacteria, and the protection of plant pro-
teins by fibre or starch may hinder their accessibility to bacteria proteolyt-
ic enzymes, making them more resistant to rumen degradation. Many
experiments have found a decrease in the degradation rate of vegetable pro-
tein when the proportion of concentrate in diet increases, which is an effect
that has been mainly associated with ruminal pH decrease, due to the neg-
ative effect of low pH on cellulolytic bacteria growth, and thus on the rate of
plant cell wall degradation.
10.2.1.2.4. Retention time of the feed protein in the rumen
The rate of degradation of the dietary protein in the rumen not only depends
on its potential degradability and enzymatic activity, but also on the time
the protein is exposed to rumen micro organisms, that is, the rumen out-
flow rate of food particles. The effect of retention time of food particles in
rumen on the amount of protein actually degraded depends on its degrada-
tion kinetics. The variation in the retention time affects more those feeds in
which protein shows a high non-soluble but potentially degradable fraction
and a low rate of degradation, as it is the case of soybean meal. By contrast
the effect of rumen retention time is lower in those protein supplements
showing a high rate of degradation, as for example sunflower meal, or a low
fraction of non-soluble potentially degradable protein, as is the case of fish
meal (FIGURE X-3).
195
FIGURE X-3. Kinetics of rumen feed protein degradation with time X – Fish meal, + -
Soybean meal, * - Sunflower meal (x= time, hours; y=dg%);
Feed retention time in the rumen is mainly affected by the level of

feeding, food particle size and density, and the proportion of forage and con-
centrate in diet. The rumen outflow rate of feed particles increases with the
level of feeding. The diminution of forage particle size also trends to increase
rumen outflow rate since it facilitates passage through the reticulum-rumen
hole. By contrast, in mixed diets the reduction of forage particle size trends
to increase the rumen retention time of concentrates, including protein sup-
plements, because the rumination activity and saliva production decrease.
The increase of the concentrate proportion in mixed diets also trends to
increase rumen retention time for the same reasons. Many authors have
also shown a decrease in rumen retention time in ewes at the end of preg-
nancy, due probably to the reduction of the rumen capacity because of the
large volume occupied by the conceptus in the abdominal cavity, although
a hormonal effect may not be discarded.
10.2.1.3. Factors affecting microbial protein synthesis

Microbial growth, and consequently microbial nitrogen entering the duode-
num, is directly related to energy availability in form of ATP and nitrogen
availability as ammonia, amino acids and peptides. For maximum growth
microorganisms also require several minerals, such as sulphur which is
essential for the synthesis of sulphur amino acids, cobalt for the microbial
synthesis of vitamin B12 or phosphorus necessary for the synthesis of
196
nucleic acids. A supplementary supply of niacin in diet has also been shown
to have beneficial effect on microbial growth.
Under conditions of anaerobiosis, the capacity of rumen microorgan-
isms to obtain energy in the form of ATP from feed fermentation is very
small. In these conditions only 10-20% of the fermented organic matter can
be incorporated in the microbial mass. The rest is eliminated as volatile
fatty acids that will constitute the main energy source for the host, and in
form of methane. Most of the energy available to the microorganisms comes
from the fermentation of carbohydrates. Little energy is available from
degradation of proteins, and that obtained from lipid hydrolysis comes only
from the fermentation of the carbohydrate fraction, mainly glycerol and
galactose, because fatty acids are not fermented in the rumen.
From model calculations a microbial nitrogen yield of about 40 g for
each kg of fermented organic matter may be expected in ideal conditions.
The values obtained experimentally in vivo are, in general, lower (mean
value of 32 g microbial nitrogen per kg of organic matter apparently digest-
ed in the rumen), although it varies considerably (Table X-8).
197
The great variability of the values obtained experimentally in vivo may

be due in part to the methodology used to determine the microbial yield,
since it requires the use of digesta flow and microbial markers that intro-
duce a considerable source of error in the estimates. However, the variation
in microbial yield also has a physiological origin.
The main factors affecting microbial yield are nitrogen availability, the
rate of fermentation of organic matter, the synchronisation of energy and
nitrogen supply, and rumen content outflow rate. For an optimal microbial
growth microorganisms need to have enough available nitrogen to synthe-
sise their own protein. Microorganisms use ammonia, amino acids and pep-
tides coming from the degradation and fermentation of food true protein and
NPN and recycled urea, as nitrogen sources. It is generally recognized that
microorganisms are able to capture most of the slowly degradable nitrogen
coming from true protein, whereas nitrogen coming from the NPN and sol-
uble fraction of feed protein is captured with a lower efficiency. For maxi-
mum efficiency of microbial synthesis, the concentration of ammonia in
rumen must be higher than 5-10 mg/100 ml. To reach this concentration
the supply of crude protein in diet must be of the order of 10-14% of the dry
matter, depending on the energy concentration and protein degradability. In
addition, although the most important source of N, and indeed the only
source in the case of the structural carbohydrates fermenting microorgan-
isms, is ammonia, it is necessary for some of the nitrogen to be in the form
of amino acids and peptides to reach the maximum efficiency of microbial
yield.
In general, the efficiency of microbial synthesis decreases when the
rumen content outflow rate decreases, because it means an increase in the
time that the bacteria remain in the rumen, increasing the costs of micro-
bial maintenance in detriment of growth. As previously commented, rumen
content outflow rate is mainly affected by level of feeding, thus microbial
yield increases with level of feeding. The efficiency of microbial synthesis
can also vary according to the source of carbohydrate and its rate of fer-
mentation. When the rate of fermentation is low which happens with diets
rich in structural carbohydrates, the relative cost of microbial maintenance
increases and the efficiency of microbial yield decreases. In fact, it has been
observed experimentally that the efficiency of microbial synthesis is in gen-
eral higher in mixed diets compared to all roughage diets. Efficiency tends
to increase until the proportion of concentrate reaches 50-60% of total diet,
198
but decreases at higher proportions of concentrate. This has been related to

microbial energy spilling when the pH decreases below 6.0, as happens in
diets that are low in fibre, encouraging the production of lactic acid and aci-
dosis. Moreover, feeds rich in volatile fatty acids such as silages, or in fat
such as oleaginous seeds, show a lower efficiency of microbial protein syn-
thesis when expressed by unit digestible organic matter, because neither
the volatile fatty acids nor the long fatty acids from fats are used as source
of energy by the microorganisms.
To reach optimal microbial growth it is necessary not only the supply
of a certain amount of degradable nitrogen per unit of fermentable organic
matter or available energy, but also energy and nitrogen availability should
be synchronized to avoid imbalances throughout the day. An imbalance in
the sense of a higher rate of fermentation of carbohydrates in relation to
that of protein will lead to a decrease in the efficiency of microbial synthe-
sis, below the maximum potential allowed by energy availability. If the
imbalance is due to a smaller rate of fermentation of carbohydrates than
that of the nitrogen source, which may easily occur when most of the nitro-
gen is supplied as NPN, there is wastage of ammonia, which is absorbed and
largely excreted as urea, although some of it is recycled to the rumen. The
recycling of urea may be as great as 60% of nitrogen intake for diets that
are very low in protein which promotes low concentrations of ammonia in
the rumen, and as little as 10% in high protein diets. It is important to bear
in mind that the synthesis of urea in the liver from absorbed ammonia
always involves energy expenditure and that excessive absorption of ammo-
nia can cause toxicity problems and alkalosis.
10.2.1.4. Measurement of feed protein degraded in the rumen

The in vivo measurement of feed protein degradability (dg) involves measur-
ing dietary nitrogen intake (NI), total non ammonia nitrogen reaching the
duodenum (NAN) and the part of this which is of microbial (MN) and endoge-
nous (EN) origin. The rumen undegraded dietary nitrogen is calculated by
the difference between the total NAN flow and the flow of MN and EN, and
protein degradability is calculated as:
CP dg = 1 – [(NAN – (MN + EN))/NI]
This method requires the use of animals cannulated in rumen and in
duodenum, to determine the duodenal flow of digesta and then the duode-
nal flow of NAN, and the proportion NAN of microbial origin. The proportion
199
of endogenous nitrogen in total duodenum nitrogen is not usually measured

but rather is considered as constant (usually 150 g/kg) although it may to
vary between 50 and 200 g/kg. The duodenal digesta flow is very variable
among animals and when only a few experimental animals are used (which
is most often the case in experiments, given the onerousness of the method),
the mean values obtained can vary considerably. In addition, when simple
T-shape, instead of re-entrant, duodenal cannula are used (which is the
most frequent case), the measure of the total digesta and NAN duodenal flow
requires the use of liquid and particle markers because the collected sam-
ple is not necessarily representative of the digesta flowing into the duode-
num, introducing a considerable source of error in determinations. Also, the
determination of the proportion of microbial nitrogen in the duodenal NAN
requires the use of microbial markers (natural compounds specific from
bacteria or protozoa, or external compounds added to diet or infused into
the rumen that are fixed by the microorganisms and reach a constant pro-
portion of their total nitrogen content after a time enough to reach the
steady state). None of these markers are ideal and there is also great
methodological difficulty in isolating a microbial fraction for reference,
which is necessary to determine the concentration of the marker in the
microbial population. Everything introduces another considerable source of
error in the estimates. Moreover, the error associated with the estimation of
the proportion of microbial nitrogen in NAN accumulates in the estimation
of the proportion of rumen undegraded dietary protein, which, in general,
represents a lower fraction than microbial nitrogen and is calculated by dif-
ference.
Consequently, since the in vivo method is onerous and subject to
numerous sources of error, other in situ, in vitro and chemical methods of
estimation of crude protein ruminal degradation have been developed.
However, in spite of the problems commented, the standard method against
which other methods have to be assessed continues to be the in vivo
method.
10.2.1.4. Determination of protein degradability in situ
It consists of the incubation of the food in synthetic fibre bags, which are
suspended in the rumen of rumen-cannulated animals, measuring the pro-
portion of nitrogen disappearance from the bags after different incubation
times. When the proportion of nitrogen that has disappeared (dg) is
regressed on time (t), the relationship can be described by the equation dg
200
= a + b (1 – e-ct) proposed by ØRKOV and MCDONALD (1979), where “a” repre-

sents the proportion of soluble protein, “b” represents the proportion of
insoluble but potentially degradable protein and “c” the fractional rate of
protein degradation (h-1). The extent of dietary protein actually degraded in
the rumen or effective protein degradability (dge) can be calculated consid-
ering the kinetics of protein degradation of each food and the rumen outflow
rate of particles (k, h-1), as: dge = a + b(c/(c+k)).
The rumen outflow rate of particles can be experimentally determined
by using an indigestible marker intimately bound to food particles. After a
pulse dose of marked food, the concentration of the marker in the rumen
content [C] follows first order kinetics, and the evolution of marker concen-
tration on time (t) can be fitted to the equation [C] = A e-kt. It is also possi-
ble to estimate “k” from the evolution of the marker concentration in faeces
over time, which has the advantage that it is not necessary the use rumen-
cannulated animals. Table X-9 shows the variation in the effective crude
protein degradabilbity of different protein sources at different fractional
rumen outflow rates (0.02, 0.05 and 0.08 h-1).
201
Although the in situ method has been adopted by most of the current
systems to evaluate the effective ruminal protein degradability of feedstuffs,
it has some disadvantages, amongst which are: the loss of small undegrad-
ed particles through the bag pores and the contamination of undegraded
residues with microbial protein, which leads to the overestimation and
underestimation, respectively, of protein degradability.
10.2.1.4.2. Laboratory methods
In vitro methods. These methods consist of determining the nitrogen solu-
bility of food in buffer solutions, in solutions containing commercial prote-
olytic enzymes or in rumen liquor obtained from donor animals. They are
easier to apply than the in situ method but in general they are less reliable
than said method.
Chemical methods. There is experimental evidence of the positive
relationship between the degradability of crude protein and the total crude
protein content of forages, given that when forage matures its crude pro-
portion decreases whereas the proportion of nitrogen bound to cell walls
increases. The rate of degradation and the potential degradable nitrogen has
also been related to the proportion of nitrogen that is soluble in borate
buffer or bound to different fractions of the cell wall (see below the CNCPS
approach).
Near-infrared reflectance spectroscopy (NIRS): NIRS has been widely
used in the prediction of the chemical composition of foods and even in the
prediction of its nutritive value. The method is based on the relationship
that exists between the chemical structure and molecular bonds of food and
their near-infrared reflectance. Although it has not been extensively used in
the prediction of rumen nitrogen degradability, the method is very easy to
apply and current data show that the in situ nitrogen degradability of foods
can be predicted with considerable precision and accuracy.
10.2.1.5. Amino acid content and digestibility of microbial and rumen unde-
graded dietary protein
Of the total microbial nitrogen entering the duodenum, only 70-80% is in
the form of true protein, the rest is essentially constituted by nucleic acids.
Although protozoa may represent as much as 30-40% of the microbial bio-
mass in the rumen of animals that are fed mixed diets, most of the micro-
bial nitrogen arriving in the duodenum (85-95%) is of bacterial origin.
Consequently, although the digestibility of the protozoa protein is consid-
202
ered to be higher than that of the bacteria, a constant digestibility of micro-

bial protein is generally assume, regardless of its origin. Most estimates of
the true digestibility of microbial protein are around 85%.
The essential amino acid content is claimed to be relatively constant
and well balanced in relation to milk and tissue protein (Table X-10), and
consequently of a high biological value. However, the proportion of some of
the essential amino acids may vary considerably, methionine and histidine
in particular show coefficients of variation that are higher than 20%.
Considering the amino acid composition of microbial protein and that of
milk or meat, histidine, fenilalanine, lysine and methionine could be limi-
tants.
The essential amino acid composition of rumen undegraded dietary

protein may differ significantly to that of the original diet, since the propor-
tion of essential amino acids in the soluble and insoluble protein fractions
is not the same). The digestibility of the undegraded dietary protein may
203
also vary considerably between foods (Table X-11) and consequently from
diet to diet and has been shown to be inversely related to the proportion of
total nitrogen found in the acid detergent fraction (ADIN), since this N is
considered to be indigestible. Nevertheless, in foods that have been sub-
jected to the action of heat and have undergone Maillard reactions, the
ADIN fraction can increase considerably, without being necessarily indi-
gestible.
The biological value of absorbed protein in the small intestine

depends on its essential amino acid balance and its likeness to the protein
to be synthesised. This balance depends in turn upon the proportion and
amino acid composition of microbial and undegraded dietary protein.
Nevertheless, due to the difficulty of predicting the biological value, and con-
sidering that most of the protein that reaches the duodenum is generally of
microbial origin, all current protein evaluation systems assume a global bio-
logical value for the total protein that reaches the duodenum, although the
majority consider different values according to the function for which the
amino acids are required.
204
10.2.2. Consideration of protein evaluation for ruminants
The significance of protein supply in dairy cow is especially high,

because one third of milk solid content is made up of protein. Therefore, a
cow producing 25 liters of milk daily, excretes ca. 800 grams of protein in
24 hours. It is equal to an achievement of ca. 6 kg of weight gain. FIGURE
X-4. provides a good overview of the turnover of ruminants.
FIGURE X-4: Schematic picture of ruminant nitrogen turnover
Total crude protein requirement can be determined by the so-called

factorial method. It is based on protein requirement for maintenance (fae-
ces, urine, function of perspiratory glands, supplying of N-loss by scaling of
skin), weight gain, foetus development and milk production. The calculated
value (so-called net protein requirement) is expressed as crude protein con-
sidering the possible ratios of absorption and utilization, respectively. Those
two constants are 75 percent and 70 percent for milk production, respec-
tively. If the diet has an above average digestibility, the crude protein
requirement decreases. The reason for that is the reduced loss by faeces.
Efficient ruminal digestion requires a minimum of 11 percent crude protein
expressed in the dry matter of the total diet. In this case, digestibility of
mixed ration decreases and therefore voluntary feed intake is lower. Urea is
produced from the protein fed above the daily requirement, which is excret-
ed via urine and involves N-loss.
The utilization of heat-treated proteins is less efficient. The degree of
205
heat damage can be concluded from the increased N-content of ADF (acid
detergent fiber). The values of feed proteins are also influenced by their sol-
ubility. For optimal utilization, the combination of the feed proteins with
high and low solubility can be recommended. The ‘critical level’ is the crude
protein concentration, above which the ammonia released exceeds the max-
imum capacitity of rumen microbes to utilise it. The limit is 7 percent dry
matter for a ruminant on a diet of mostly fibrous roughages and 13 percent
crude protein in case of rations containing concentrated feeds. (On average,
the limit is considered to be 10 to 12 percent of the dry matter.)
The crude protein fed should meet the N-requirement of rumen
microbes and requirements for amino acids of tissues and of the host ani-
mal as well. Depending on the quality of feed protein, half or even higher
proportion is supplied by microbial protein. A minimum concentration of 3
to 5 mmols per liter NH3 is required in rumen fluid. Below that level, the
rate of microbial growth decreases with a consequent decline in the organ-
ic matter degradation, which in turn, decreases voluntary feed intake. A
part of crude protein requirement can be met by NPN-compounds. The rate
of microbial protein synthesis (MP) from ammonia is proportional with the
energy available in the rumen (8 to 12 g MP/MJ ME). Level of the optimal
urea supply can be determined based on the energy and on the protein con-
tent of feeds and on the ruminal degradability of protein. Protein degrada-
tion itself depends on several factors, like the proportion of roughages in the
mixed ration, the quantity of daily diet and the pre-treatment of feeds (with
special consideration of the effects of heat and chemicals).
Those proteins, which passes by the rumen without degradation and
arrives into the abomasum, is called „bypass” or „escape” or undegradable
protein (UDP) (Table X-12). This proportion of protein is either digested and
absorbed postruminally or excreted via feaces. Ruminal protein degradation
depends on four factors; bacterial population, species of host animal, „incu-
bation” time and the quantity of protein fed to the animal. Therefore,
degradability of the same protein can never be considered constant.
206
Feedstuffs can be classified in three cathegories according to their bypass

character:
- feeds of small bypass values (<40%), e.g. soybean meal, groundnut meal,
- feeds of medium bypass values (40-60%), like cottonseed meal, lucerne
meal, maize meal, by-products of breweries,
207
- feeds of high bypass values (>60%), e.g. meat meal, blood meal, feather
meal and fish meal. This classification can be modified by the (pre)treatment
of feed, animal, feeding, composition and activity of microbial population.
Complexity of the topic is even enhanced by the fact that the amino acid
composition of bypass protein does not reflect the „initial” combination of
amino acids (VERESEGYHAZY and FEKETE, 1989).
The ruminal degradation of feed proteins is generally determined by
an indirect method. Namely, the difference between total feed and microbial
protein is considered as „undegraded protein in the rumen, or rumen unde-
graded protein” (accepted abreviation is UDP or RUP). When it is expressed
as a percentage of the consumed feed protein, the „degradability coefficient”
is calculated. Ther are some opportunities for the measurement of microbial
protein, although the comparison of values received by different methods
might be a source of errors. Accordingly, microbial protein can be estimat-
ed on the basis of measurements with isotope sulphur (35S), with diamino-
pimelic acid (DAPA) as bacterial component and with RNA and D-alanine
(respectively). Since, there are only a few data available by in vivo trials,
efforts being made to measure ruminal degradability of proteins in vitro (in
tube) and in sacco (in small bags suspended in the rumen) methods. When
the bypass character of a feed protein increases in the daily ration, the
demand for NPN-compounds is enhanced. Protein passing by the rumen
does not necessarily meet the requirements of animals, because it might be
of low digestibility in the small intestine, or it does not have the required
amino acid composition or the infiltration of amino acids is not optimal
because of the lack of other feeding factors. However, above a certain pro-
duction level feed protein is not to be degraded in the rumen completely!
Based on the recommendations of ARC (1984), here is an example for the
protein requirement of a cow of 500 kg live weight, producing milk of 4%
milk fat (Table X-13) and different levels of milk production.
208
The most up-to-date standards estimate the protein requirements of

beef cattle by factorial method, too. NRC (1985) regards 4.48 MJ NEm/kg
dry matter as minimum energy concentration in its calculations.
Requirements of weight gain was estimated on the basis of protein infiltra-
tion in an „empty” body (with no ruminal and intestinal content), on the
actual and adult live weights, and on the average daily weight gain. In the
last third of pregnancy 55 g crude protein was calculated for foetus devel-
opment per cow, and 3 to 10 liters of milk with 3.35% milk protein was con-
sidered as typical milk production in the case of beef mother cows. Apparent
digestibility of protein passed into small intestine, is of ca.66%. True
digestibility (corrected by metabolic faecal loss) is applied in the calcula-
tions. Heat treatment decreases the digestibility of fibrous roughages sig-
nificantly. In this case, true digestibility should be determined by pepsine
or by the ADF-technics as mentioned above. Biological value of microbial
protein, among others because of its high nucleic acid content, is between
66% and 81%. Efficiency infiltraion of digestible protein is finally considered
as a combined one (maintenance+production) of 66%. Tables prepared
including the required daily weight gain were based on species, on body
weight and on the demanded daily weight gain.
Composed daily ration should be controlled by the following equa-
tions: UFP = (1.044×TDN-dB)/2.8, where UFP stands for urea fermentation
potential, TDN is total digestible nutrients, and dB refers to the ruminal
degradability of protein. Based upon that the above equation and knowing
the requirements of protein and energy and the degradability of protein in
209
the rumen, the proportion of total crude protein requirement can be calcu-
lated which can be provided in form of NPN-sources. A new feature of this
concept is that it considers the essential role of energy supply in the bacte-
rial protein synthesis.
UFP refers to UREA FERMENTATION POTENTIAL which is supported by the
extra energy produced in the rumen. Negative value of UFP indicates that
rumen receives by protein, which is highly degradable in the rumen; in this
case, NPN-sources are not utilized. If UFP is a positive value, rumen
microbes are able to use urea. At the same time, N-supply of rumen bacte-
ria is also to be considered: a minimum of 90 g crude protein per kg of dry
matter is required. Zero value of UFP shows that the energy content of the
daily ration is just enough to process its NPN-compounds. RUMINAL PROTEIN
BALANCE is basically the inverse of urea fermentation potential. Positive pro-
tein balance means that there is more degradable protein than energy is
available for bacterial protein synthesis. Negative protein balance reflects
nitrogen deficiency. Therefore, supplementtion with an NPN-source is
allowed only in case of negative protein balance or positive urea fermenta-
tion capacity.
10.2.3. Shortcomings of old crude protein and apparently digestible

crude protein systems
Until the nineteen seventies, the protein value of feeds and the protein
requirements of ruminants were expressed in terms of crude protein (CP) or
digestible crude protein (DCP). The reason for establishing the protein value
in terms of CP was that it was recognised that rumen microorganisms
transformed most of the dietary protein in microbial protein, regardless of
the characteristics of the dietary protein, and that the digestibility and bio-
logical value of the microbial protein were relatively constant. DCP was also
used as a measure of the protein value of foods for ruminants in Europe
during the last century, since it was the main form of protein evaluation
used in monogastrics, and it worked reasonably well when well-balanced
conventional diets were used.
However, during the nineteen sixties and seventies, numerous works
revealed that the responses obtained from varying the DCP intake of rumi-
nants was influenced by the source of protein and by other factors, mainly
the energy supply. Moreover, it was seen that the apparent digestibility of
forage protein was closely related to its total protein regardless of its char-
210
acteristics, due to the important contribution of non- alimentary nitrogen,

mainly of microbial origin, to total nitrogen excreted in faeces.
This empirical knowledge and the awareness of the central role of the
processes of degradation and synthesis taking place in the rumen in the
nitrogen economy of the host animal, led to an understanding that the
quantity of truly available protein and amino acids for the host, at duode-
nal level, depended basically on two factors: a) The quantity of available
energy for rumen microbial growth and b) The quantity of feed protein
reaching the duodenum without having been degraded in the rumen. The
new protein systems for ruminants try to take into account the many fac-
tors affecting rumen microbial yield and feed protein degradability in the
rumen, as well as the digestibility of microbial and dietary protein arriving
in duodenum and the biological value of the absorbed amino acids for dif-
ferent functions.
211
PROTEIN EVAULATION - RUMINANTS
212
Chapter
10.3. Overview of protein evaluation systems in different

countries
Introduction. The new systems based on the true protein absorbed

from the small intestine
The concept that the protein value of a feed or a diet depends on the amount
of protein that escapes rumen degradation and the amount of microbial pro-
tein de novo synthesised in the rumen, was proposed at the beginning of the
seventies by different groups in USA, United Kingdom and East Germany.
Since then many systems have being proposed in Europe, USA and
Australia that take into account the different origins of amino acids arriving
in the duodenum. Prediction of the amount of amino acids made available
from these two sources is approached in a variety of ways, usually involv-
ing a number of simplifying assumptions. All new systems also coincide in
the need to consider separately the protein needs of rumen microbes and
those of host animals, the former being covered by dietary degradable pro-
tein and the latter by the amino acids absorbed from the small intestine.
The first formal proposals in Europe came almost simultaneously
from France (INRA, 1978) and United Kingdom (ARC, 1980, 1984), and later
from Scandinavian countries (NKJ-NJF, 1985), Switzerland (BICKEL and
LANDIS, 1987) and West Germany (ROHR, 1987). Some of these systems were
presented and discussed in a joint CEC/EAAP workshop on “Feed evalua-
tion and protein requirement systems for ruminants” held in Brussels in
June, 1986 (CEC, 1987). DE BOER and BICKEL (1988) analyse the situation
of the new systems proposed in Europe at that moment. In 1991, a similar
protein evaluation system (the DVE/OEB-system) was also proposed in the
Netherlands (CVB, 1991), and others were put forward in the USA (NRC,
1985) and Australia (CORBETT et al., 1987). Some of these systems have
already been updated (VÉRITÉ et al, 1987 and INRA, 1988 in France; CSIRO,
1990 in Australia; AFRC, 1992 in UK; TAMMINGA et al., 1994 in the
Netherlands and in Hungary by SCHMIDT et al. 1999).
The INRA system express protein requirements in terms of PDI
(Protéines vraies réellement digestibles dans l’intestine grêle), synonymous
of 6.25 times truly digestible amino N (TDAN). The API (absorbable protein
in the intestine) system introduced in Switzerland derives from the French
system, and the Nordic (NKJ) protein evaluation system also has many sim-
ilarities with the PDI system, using the units AAT (amino acids truly
213
absorbed in the small intestine) and PBV (protein balance in the rumen or
degradable protein supply relative to the need of the microorganisms) to
express the protein value of feed. The Dutch system was also developed on
the basis of the French PDI-system, although elements of other systems
were incorporated; the protein value of feeds and the requirements of dairy
cows are both expressed as the amount of true protein truly digested in and
absorbed from the small intestine (DVE) and, like the Nordic system, each
feed also has a OEB value that shows the balance between the microbial
protein synthesis potentially possible from rumen available protein and that
potentially possible from rumen available energy.
The British systems proposed in 1980 expressed protein require-
ments as apparently absorbed true protein and the protein value in terms
of rumen-degradable crude protein (RDP) and undegraded crude protein
(UDP). It was revised in 1984 changing to truly absorbed protein, although
the feed value continued to be expressed as UDP and RDP. In the last ver-
sion (AFRC, 1992), the requirements are established as “metabolizable pro-
tein” (6.25 x TDAN) and the protein value of feeds is defined by its ERDP
(effective rumen degradable protein) and the DUP (digestible undegradable
protein) content. The German system expresses protein requirements as
“crude protein at the duodenum” and in the USA system (NRC, 1985) the
protein requirements are expressed as “ABSORBED TRUE PROTEIN”. The
Australian system resembles the 1980 ARC system in that it describes the
protein requirements of the animal as “APPARENTLY DIGESTIBLE TRUE PROTEIN
LEAVING THE STOMACH”.
All these new systems, although they differ in their terminology and
in the factors used in the estimation of feed value and animal requirements,
are based on a similar framework and consider similar steps in the estima-
tion of protein available to cover the different animal requirement.
214
10.3.1. Main steps in the calculation of the metabolizable protein sup-

ply (FIGURE X-5).
FIGURE X-5: Main steps in the calculation of the metabolizable protein supply
a) Extent of degradation of dietary protein in the rumen (RDP)
b) Undegraded dietary crude protein reaching the duodenum (UDP)
c) Supply of fermentable substrate for microbial growth
d) Microbial protein yield
e) Efficiency of nitrogen capture by the micro organisms
215
CHAPTER X: PROTEIN EVALUATION - RUMINANTS
f) Proportion of microbial nitrogen present as amino acid nitrogen

(MTP/MCP)
g) Proportion of undegraded dietary protein present as amino acid

nitrogen. All revised systems, except the German and Nordic sytems, con-
sider that all dietary rumen undegraded nitrogen reaching the duodenum is
in form of amino acids. The German system assumes a proportion of amino
acid nitrogen content in undegraded protein of 0.70, whereas the Nordic
system considers a factor of 0.85 and 0.65 for amino acid nitrogen content
in concentrates and roughages respectively.
h) Digestibility of microbial proteins
i) Digestibility of microbial and undegraded dietary amino acids
j) True digestibility of the potentially digestible undegradable nitrogen
k) Protein requirements and allowances.
Table X-14 summarizes the considered factors in the different systems.
216
217
The Cornell University; the Cornell Net Carbohydrate and Protein System
(CNCPS) makes some deviations from the following general scheme, there-
fore it will be discussed in the last part of this subchapter.
10.3.2. The UK metabolizable protein system.
The last version of the UK metabolizable protein system is fully described in

Report no. 9 of the Agricultural and Food Research Council’s Technical
Committee on Responses to Nutrients (AFRC, 1992). It is based on early
proposals (ARC, 1980, 1984) although it incorporates several modifications
in the light of current knowledge. FIGURE X-6 shows a flow chart of the
AFRC model, which is described in terms of N although the advisory model
uses CP (N×6.25) as unit
- Feed protein value
Each feed is defined by two values depending on its CP content, on the effec-
tive rumen degradability of the protein, and on the true digestibility of the
undegraded protein:
1. Effective Rumen Degradable Protein (ERDP, g/kg DM), representing

the protein content available to rumen microorganisms.
2. Digestible Undegraded Protein (DUP, g/kg DM), that represents the

content of truly digestible undegraded protein.
218
FIGURE X-6: Diagram of feed protein evaluation in the Metabolizable Protein (AFRC) sys-
tem
219
1. The ERDP of feeds is calculated from the equation, ERDP = [0.8 (QDP) +
SDP], where QDP represents the quickly degradable protein, calculated as
QDP (g/kg DM) = CP (g/kg DM) × “a”, and SDP represents the slowly degrad-
able protein calculated as SDP (g/kg DM) = CP (g/kg DM) × bc/(c+k).
Constants a, b and c are obtained for each feed by the in situ method after
fitting the proportion of degraded protein degradability values to the model
(dg= a + b(1-exp ct) .
Rumen fractional outflow rate (k), takes the values:
- 0.02 h-1 for cattle and sheep at low planes of nutrition
- 0.05 h-1 for calves, beef cattle, sheep and dairy cows up to 2 × mainte-
nance, 15 kg milk/day.
- 0.08 h-1 for dairy cows yielding >15 kg milk/day
It may by also calculate from the equation: k = - 0.0024 + 0.179 ((1-e (-

0.278L)), were L represents the plane of nutrition as a multiple of mainte-
nance.
The Committee pointed out that possible adjustments to these values

may be made as more information becomes available about the influence of
diet (e.g. forage/concentrate ratio), but to date there is insufficient evidence
to quantify them. Thus, the ERDP may by calculated from the CP content of
feeds, the dynamics of protein degradation in the rumen using nylon bags
and the estimated rumen outflow rate of particles , from the equation:
ERDP (g/kg DM) = CP (g/kg DM) × [0.8 a + bc/(c+k)]
It is assumed that the SDP is used by microorganisms with an efficiency of

1.0 whereas the QDP is used with an efficiency of 0.8.
2. The UDP is calculated from the equation: DUP (g/kg DM) = 0.9 UDP
(g/kgDM) – 6.25 × ADIN (g/kg DM), where UDP = 1 – (QDP + SDP) and ADIN
represents the acid detergent insoluble N.
Thus the UDP content of feeds may by calculated from the CP and ADIN
content of feeds, the dynamics of protein degradation in the rumen using
nylon bags, and the estimated rumen outflow rate of particles, from the
equation:
DUP (g/kg DM) = 0.9 × [(CP, g/kg DM x (1-a-bc/(c+k)) – 6.25 × ADIN,

g/kg DM]
220
It is assumed that the true digestibility of feed undegraded protein is 0.9,

except for the fraction bound to the ADI fraction, which is considered indi-
gestible.
Tabulated values of the CP, ERDP and DUP content of feedstuffs calculated
for 0.02, 0.05 and 0.08 h-1 rumen outflow rates, are given in AFRC (1992,
1993). AFRC (1993) also gives a, b and c values of degradation kinetics.
- Protein requirements
Two types of protein requirements need to be considered:
1) Requirements in ERDP in order to obtain optimal ruminal fermentation

and microbial growth.
2) Requirements in metabolizable protein (MPR) in order to cover the ani-

mal’s requirements.
1) It is assumed that the ERDP is used by the microorganism with an effi-

ciency of 1.0 and, as a consequence, the supply of ERDP must be equal to
the potential of microbial protein synthesis, which depends on the avail-
ability of FME and varies with the plane of nutrition:
ERDP = FME × “y”
The FME content of diet is calculated as the addition of the FME content of
each ingredient, calculated as: FME (metabolic fermentable energy) (MJ/kg
DM) = ME – MEfat – MEferm, where MEfat = 35 kJ/g dietary fat, and
MEferm = 0.1 ME in ensiled feeds or 0.05 in brewery and distillery by prod-
ucts. Tabulated values of calculated FME content of feedstuffs are given in
AFRC (1992, 1993).
The “y” value represents the efficiency of microbial protein synthesis, which
is considered to increase with the rumen outflow rate. The following values
of “y” are suggested for different levels of animal performance:
9 g MCP/MJ of FME for all animals at maintenance level of feeding
10 g MCP/MJ of FME for growing sheep and cattle (2×M)
11 g MCP/MJ of FME for late pregnancy and lactating ewes and lactating
dairy cows (>3 ×M)
The following equation may be also used: MCP/FME = 7 + 6(1-e(-0.35L)),

where L represents the plane of nutrition as a multiple of maintenance.
221
2) The total MPR are calculated as the addition of MPR for the different func-
tions coexisting in the animal (maintenance, tissue protein accretion, pro-
tein accretion in gravid uterus, wool protein accretion or milk protein yield).
The MPR for each function (i) is calculated from the net protein require-
ments (NPi) and assumed efficiencies of utilisation of absorbed amino acid
(kni):
MPR = ΣNPi/kni
Net protein requirements for maintenance include basal endogenous nitro-

gen and dermal losses, and those for other functions depend on tissue
nitrogen retention in wool, live weight gain or conceptus, or on nitrogen
excreted as milk.
The efficiency of utilisation of absorbed amino acids is estimated as kn =

kaai × RV, where:
Kaai = efficiency with which an “ideal” AA mixture is utilised for a purpose.

It is a characteristic of the animal.
RV = relative value. The extent to which the absorbed AA mixture differs

from the “ideal”
Only values for kaai can be predetermined, because values for RV depend
on particular feeding circumstances, especially on the essential AA balance
in the DUP.
Table X-15 shows the ERDP and DUP calculation of some important feed-
stuffs (AFRC, 1992). MPR for certain broad classes of ruminants and phys-
iological states are given tabulated by the AFRC (1992) and AFRC (1993).
222
- Metabolizable protein supply (MP)
The supply of MP with diet to cover the MPR represents the total true
digestible protein of microbial and dietary origin:
MP = Digestible microbial true protein (DMTP) + DUP
DMTP = MCP × 0.75 × 0.85 = MCP × 0.6375, assuming that only a 0.75 of
total microbial protein is true protein and that the true digestibility of the
true microbial protein is 0.85. The MP supply may thus be calculated using
223
the equation:
MP (g/d) = (FME × y) × 0.6375+ DUP
FME, g/d = Σ wi (FME)i, and DUP, g/day = Σ wi (DUP)i, where wi is the

weight of dry matter of feed i, and (FME)i and (DUP)i are the FME and DUP
contents of feed i.
If the requirements of DUP, calculated as the difference between the total

MPR and the DMTP, is zero or even negative, it means that the host MP
requirements are covered by the microbial protein synthesised in the
rumen. In this case only the ERDP requirements need to be covered and
part of them may be supplied with NPN sources such as urea. One gram of
urea supplies 2.3 g of ERDP, considering that 1 g urea = 0.46 g N and that
the efficiency of utilisation of NPN by the micro organism is 0.8 (2.3 =
=0.46×0.8×6.25).
Diet formulation using the proposed UK metabolizable protein system
The following steps are followed for diet formulation:
1. Calculation of MPR requirements to be covered as DMTP + DUP
2. Calculation of minimum ERDP to be supplied to cover MCP requirements
3. Calculation of minimum DUP to be supplied in order to cover the MPR,
not covered by de DMTP
The constraints to be satisfied in order of importance will be:
1. ERDP/ FME > y (9, 10 or 11 g/MJ FME)
2. MP supply > MP requirements, where MP supply = DMTP + DUP
10.3.3. The French PDI system
In the INRA (1988) protein system, the protein value of feeds and animal
requirements are expressed in terms of true protein truly digestible in the
small intestine (PDI). It is based on an earlier version (INRA, 1978), with
some modification mainly in relation to the method of estimation of protein
degradability, which is now assessed by the in situ method, and the pre-
diction of microbial protein synthesis based on the amount of fermentable
organic matter instead of digestible organic matter. A flow chart of the INRA
model is shown in FIGURE X-7
224
FIGURE X-7: Diagram of feed protein evaluation in the PDI system
- Feed protein value. The PDI content of a feedstuff or a diet is the sum of
two fractions: a) the dietary protein undegraded in the rumen, but truly
digestible in the small intestine (PDIA), equivalent to the DUP in the UK sys-
tem, and b) the microbial true protein which is truly digestible in the small
intestine (PDIM), equivalent to the digestible microbial protein in the UK
system.
Each feed is characterised by two PDIM values:
PDIME = amount of microbial protein that could be synthesised from the

energy available in the rumen, when degraded N and other nutrients are not
limiting, and
225
PDIMN = amount of microbial protein that could be synthesised in the

rumen from the degraded dietary N, when energy and other nutrients are
not limiting
As a consequence, two potential protein values are assigned to each feed:
PDIE = PDIA + PDIME, when PDIMN>PDIME
PDIN = PDIA + PDIMN, when PDIMN<PDIME
The lower of these two values is the real value of feed when given alone and
the higher value is the potential value when the feed is complemented with
others to reach an optimal N and energy balance for microbial growth. The
PDIE and PDIN values of feeds are tabulated.
The PDI content of a diet is calculated by adding the PDIE and PDIN
values of each ingredient separately. The lower of the two sums corresponds
to the actual PDI value of the diet. Diets are formulated trying that PDIN
supply = PDIE supply = PDI requirements, taking into account that some of
the PDIN requirements can be covered with NPN sources and some PDIN
deficit may be assumed to be covered by recycled nitrogen.
Calculation of PDI values of feedstuffs
Data for the basic equations for calculating of the PDIE and PDIN values of
feedstuffs were obtained from four individual feed characteristics: the CP
content, the effective degradability of CP in the rumen (deg), the fermentable
organic matter content (FOM) and the true digestibility of undegraded pro-
tein in the small intestine (dsi), together with several other standard coeffi-
cients used to describe the digestive process.
Origin of the coefficients for feed evaluation
- The effective degradability of CP in the rumen (deg): The effective ruminal

feed protein degradability of feeds is obtained from in situ incubation trials
and assuming a constant rumen outflow rate of 0.06 h-1 regardless of
species, diets and levels of intake.
- Rumen undegraded protein reaching the duodenum: The dietary protein

escaping rumen degradation is calculated as 1.11 x (1-deg) to take into
account the underestimation of undegraded protein when determined by
the in situ method due to the escape of some undegraded protein through
nylon bags pores.
226
- Microbial protein synthesis: Although it is recognized that microbial pro-

tein synthesis may be affected by factors other than energy and nitrogen
supply, the PDI system calculates microbial protein synthesis as a function
of the FOM supply, assuming a constant protein yield of 145 g CP/kg FOM).
The FOM supply was preferred to DOM as a predictor of microbial synthe-
sis (this parameter was used in a previous PDI proposal, INRA, 1978), to
take into account that energy from ether extract (EE), undegradable dietary
protein (UDP) and fermentation products in silage (FP) is not available for
rumen microorganisms. FOM is then calculated by subtracting these com-
ponents from the DOM (FOM = DOM – EE – FP – UDP).
The value of 145 g CP/kg FOM was obtained by partitioning the whole
duodenal flow of NAN amongst its three components: microbial nitrogen
(proportional to the FOM intake), UDN and endogenous nitrogen (propor-
tional to the intake of non digestible organic matter –NDOM-). From a data-
base of 405 diets the following equation was obtained:
Duodenal NAN = 23.2±6.3×FOM+1.11±0.07×bag UDN+5.3±1.8×NDOM
The coefficient of regression associated with FOM (23.2 g N, equivalent to

145 g CP/kg FOM), would represent the mean microbial N flow to duode-
num, although the error associated to the regression coefficient was rather
high (±6.3, CV = 27%).
- Potential microbial protein synthesis from degraded dietary N:
The feed contribution to the supply of N for microbial protein synthe-

sis is considered to arise only from the degradable N fraction of feed. The
efficiency of conversion of degraded N to microbial N is assumed to be 0.9,
although for added NPN sources the efficiency is assumed to be 20% lower.
- Amino acid content and digestibility of duodenal protein:
Microbial protein: The amino acid content of microbial protein is considered

to be 0.80, a value within the range of those obtained from microbes col-
lected from the rumen or in vitro cultures (0.75-0.85).
The true digestibility of microbial amino acids is considered to be con-

stant and equal to 0.80, without considering possible differences between
protozoa and bacteria. The adopted value is higher than that proposed in
the 1978 version (0.7), but within the range of those found in the bibliogra-
phy (0.73-0.98) and near to the value obtained when the NAN absorbed in
227
the small intestine from a 367 database was related to the microbial nitro-
gen, the UDN and the endogenous N entering the duodenum (coefficient of
regression associated with the duodenal microbial nitrogen flow =
0.81±0.09).
Dietary protein: Although the content of AA in the residual protein in nylon

bags incubated in rumen varies generally between 0.80 and 0.90, a value of
1.0 is adopted (as in the previous version) to account for microbial contam-
ination.
The digestibility of the UDP in the small intestine (dsi) is considered to be

variable among feedstuffs, raging from 0.60 to 0.95. It was predicted from
the equation:
Faecal N = 4.62 ±0.18×FOM+0.216±0.33×bag UDN+0.6±0.7×NDOM,
This equation splits up the faecal N excretion (CP-DCP taken from

tables) into its three main components; microbial, dietary and endogenous
N, which are related to the FOM, UDN and NDOM intake, respectively. From
this equation the indigestible dietary protein is calculated as: IDN (indi-
gestible dietary protein) = Faecal N–4.62 FOM–9.6×NDOM, and the intes-
tinal digestibility of UDN as: dsi = (UDN-IDN)/UDN.
- Protein requirements As previously commented, unlike the UK , USA and

Germany systems, that use a factorial approach to asses the protein
requirements, the PDI requirements for maintenance and the PDI require-
ments and efficiency of utilization of PDI for different production functions)
were estimated from N balance trials or based on feeding trials.
Within experiments, when PDI intake decreased generally protein

production decreased and efficiency increased both in a curvilinear manner.
The optimal allowance was considered to be the quantity of PDI below which
the protein yield exhibits a decrease in biological significance. Thus, the
proposed requirements correspond to an optimal situation in practice with
well-balanced diets. Nevertheless, increased efficiency can be expected in
the future when the balance between absorbed nutrients will be better
understood with regard to the amino acid profile and to the nature of ener-
gy nutrients.
- Recommended allowances
For the animal:
228
The recommended PDI allowances are in general equal to the PDI require-
ments, with some exceptions. For example PDI allowances are slightly lower
than requirements for dairy cows and goats in the first weeks of lactation
because of their ability to mobilize body reserves that can cover part of milk
production requirements. On the other hand allowances may be greater
than requirements for milking ewes because milk yield and milk protein
content usually respond positively to PDI supply above requirements.
Some positive responses can be obtained by supplementation with

protected amino acids, particularly methionine and lysine, the most likely
to be the first limiting in dairy cows, although more information on animal
amino acid requirements and amino acid composition of UDP would be
needed to safely predict the response.
For rumen microbes: In the PDI system, the balance in the rumen
between degradable N supply and microbial requirements is given by the
difference (PDIN supply – PDIE supply). The INRA system considers the pos-
sibility of covering a small deficit of PDIN with recycled urea coming from
amino acid catabolism. The relative deficit that is acceptable is expressed as
g (PDIN-PDIE) per Feed Unit (FU) intake. This acceptable deficit is in gener-
al higher in animals at low levels of production compared with those at high
level of production (e.g. in beef cows compared to dairy cows) and also high-
er when the supply of amino acids exceeds requirements.
10.3.4. The Hungarian System
The Hungarian protein evaluation system (metabolizable protein, MP) was

introduced in the teaching and practice in 1999. The system is partly based
on the elements of other protein evaluation systems. Ruminal protein
degradability of the feeds mostly used in the feeding practice in Hungary
was measured in situ. Metabolizable protein is the sum of microbial protein
and bypass (UDP) true protein truly digested in the small intestine (SCHMIDT
et al. 2000).
Protein values of feeds. The quantity of microbial protein is mainly

dependent on the energy and the rumen degradable protein (RDP) supplies
for microbes. Feeds differ in their levels of available energy and RDP, so the
quantity of microbial protein synthesized in the rumen can be calculated on
the basis of both energy and RDP supplies. Thus, each feed has two protein
229
values including digestible UDP and digestible true microbial protein pre-
dicted either from the energy or degraded protein (RDP) available for
microbes. The two protein values are: the MPE – metabolizable protein
dependent on energy and the MPN – metabolizable protein dependent on
nitrogen. The formulas to calculate the protein values are as follows:
-MPE, g/kg DM = 0.9(UDP-ADIN×6.25) + 0.16 FOM × 0.8 × 0.8 and
-MPN, g/kg DM = 0.9(UDP-ADIN×6.25) + 0.9 RDP × 0.8 × 0.8, where ADIN

= acid detergent insoluble N and FOM = fermented organic matter.
Acid detergent insoluble nitrogen (ADIN) is considered as unavailable N-

fraction, which is neither degraded nor digested. The digestibility of UDP
excluding ADIN×6.25 is supposed to be 90 percent similar to the MP-system
(AFRC, 1992). Digestibility values calculated in this way, are lower for for-
ages and roughages than for concentrates and are similar to the values of
true digestibility of undegraded protein in the small intestine (dsi) in the PDI
system (INRA, 1989).
Energy available for microbes is expressed as fermented organic mat-

ter (FOM), which equals to digested organic matter (DOM) excluding fer-
mentation products of ensiled feeds, UDP, digestible fat and bypass starch.
Nutrients escaping rumen fermentation (UDP, bypass starch), digestible fat
and the end products of fermentation (organic acids of silage) are not con-
sidered as energy sources for microbes though they are utilized well by
ruminants. Energy values used for ruminant feeding (metabolizable energy,
net energy etc.) do not completely cover the energy requirement of rumen
microbes.
Microbial crude protein (MCP) production is expected to be 160 g/kg

FOM. True protein content of MCP and the digestibility of microbial true
protein is 0.64MCP. The efficiency of utilization of RDP for microbial crude
protein synthesis is considered to be 90 percent for feed RDP and 80 per-
cent for NPN-sources. In summary, the Hungarian MPE value covers the
same concept as PDI (INRA, 1989) or AAT (MADSEN et al., 1995). MPN is sim-
ilar to PDIN (INRA, 1989).
Protein requirements. The protein requirements of ruminants is

expressed as metabolizable protein (MP), which equals net protein require-
ment divided by the efficiency of utilization of metabolizable protein. Net
protein requirement is equal to the quantity of protein depositied or secret-
230
ed. Net protein requirement for maintenance includes endogenous losses in

urine, obligatory fecal metabolic N, and scurf protein (only in cattle). Net
requirements for milk production, daily gain, pregnancy, wool production
are calculated according to the quantities of protein in the product.
For the efficiency of utilization of metabolizable protein for production

the following values are used: milk production 65%, growth 50%, pregnan-
cy 50% and wool production 40%. If 1 of kg milk contains 32 g of protein,
the net MP requirement for the production of 1 kg of milk equals 32/65 =
49 g MP. The efficiency of utilization of MP for growth is variable. It decreas-
es as the age and the weight of ruminants increase. It is assumed that the-
efficiency is higher for young animals combined with high protein deposi-
tion and decreases with age. Efficiency changes from 0.56 to 0.44 between
200 and 500 kg of live weight for e.g. in growing bulls. Metabolizable pro-
tein requirements for maintenance are calculated by the following formulas:
Maintenance MP, g/day = 3.41W0.75. Maintenance requirement for sheep,
in MP, g/day = 2.60W0.75, where W = live weight, kg.
Ration formulation. For ration formulation, both MPE and MPN values
of feeds should be multiplied by the quantity of feeds consumed. The small-
er sum of MPE or MPN value is the MP content of the ration, which should
be compared to the daily requirement. By subtracting MPE from MPN, the
protein balance in the rumen can be calculated. When protein balance is
positiv, there is more degradable protein available than energy. Negative
protein balance indicates the deficiency of nitrogen for rumen microbes.
(Positive protein balance means a negative value of UFP, therefore no feed-
ing of urea is allowed!) It shows the difference between the quantity of
digestible true microbial protein predicted from RDP and energy available
for microbes. For high-yielding dairy cows, producing more than 30 kg of
milk per day, low positive protein balance (+ 100 g) is advised in the rumen.
It seems that microbial protein production is higher per unit of energy avail-
able for microbes at higher energy intake (AFRC, 1992; NRC, 1985). Positive
protein balance supplying more RDP per unit of energy can support micro-
bial protein synthesis of higher efficiency. Feed intake increases as protein
balance in the rumen changes from negative to positive. At the same time,
high positive protein balance (+250-300 g) is to be avoided for dairy cows,
because there are several observations indicating that overfeeding RDP
results in poor reproductive performance. Negative protein balance in the
231
rumen is allowed for dry cows, beef cows and slowly growing animals, as
long as MP requirement is met.
10.3.5. Shortcomings of current European systems
The main shortcoming of current European systems is the lack of sufficient

quantitative information to construct mechanistic equations that accurate-
ly describe the different processes involved in ruminal and host nitrogen
metabolism, such as the degradation rate of both carbohydrates and pro-
tein in the rumen, microbial growth, amino acid composition, digestion and
absorption of microbial and mainly undegraded feed protein or the efficien-
cy of utilization of absorbed amino acids. On the other hand revised systems
are based merely on protein and energy supply with diet, and a better
knowledge is required of the absorbed nutrients that are made available to
the host animal and the metabolic interactions between them. Likewise,
quantitative information and a deeper understanding of feed conversion and
animal response when nutrient supply is lower or higher than the optimum
recommended for specific production levels is necessary.
In relation to feed evaluation, the in situ method used by most sys-

tems to estimate the nitrogen available for microbial growth and dietary pro-
tein by-passing the rumen, is hampered by low precision and reproducibil-
ity, as shown in different Ring tests carried out in UK by the IDWP (AFRC,
1992) and at a European level (MADSEN and HVELPLUND, 1994). In a first ring
test of the IDWP in which thirteen different laboratories in the UK partici-
pated, with no constraints as to methodology, great differences in N disap-
pearance from bags before incubation (dg 0h) were found for a same hay
(from 10 to 44%) and soybean meal (from10 to 36%). In a second ring test,
where a standard procedure was applied in five laboratories, the effective
protein degradability calculated for a 0.08 h-1 rumen outflow rate varied
between 0.60 and 0.89 for barley and from 0.56 to 0.69 for soybean meal.
In the European ring test 23 laboratories from different countries partici-
pated, applying a standardized methodology. Nevertheless dg0h was esti-
mated with a coefficient of variation of 44% for barley and 56% for soybean
meal, and the effective degradability for k= 0.08 h-1 with a coefficient of
variation of 12 and 18%, respectively. A bias of 10 percent units in the esti-
mation of effective crude protein degraded in rumen of soybean meal, would
account for a bias of proximately 50 g/kg feed in the UDP supply, which
232
would be sufficient to support 1 kg of milk production.
Moreover, N disappearance from the bag is not synonymous of pro-

tein degradation, because an important and variable fraction of insoluble
particles may leave the bags without being degraded, which leads to the
overestimation of the quickly degradable and also the effective degradable
protein. As previously commented, the French and Dutch systems try to
take this into account by multiplying by 1.11 the UDP estimate from the in
situ method. By contrast, contamination of bag residues with microbial
protein, which may be quantitatively important in high fibre and high
degradable low protein foods, results in the overestimation of UDP.
The negative effect of diets rich in concentrates on the in situ protein

degradation rate is also well documented, but no system takes into account
the effect of diet composition and, in particular, the effect of the proportion
of forage and concentrate on the rate of protein degradability.
Amino acid composition and digestibility of undegraded protein may

be variable as shown above. The approach of AFRC, which estimates the
true digestibility of UDP considering the ADIN indigestible and a constant
digestibility value of 0.9 for the potentially digestible UDP, is very simplis-
tic. Although for a majority of conventional diets ADIN may be assumed to
be undegradable and indigestible, it is an unreliable index for head-treated
proteins and distillers’ products, in which ADIN content increases during
the process, without being necessarily indigestible.
The greatest element of uncertainty in the estimation of protein reach-

ing the duodenum in all of the new system is probably the quantification of
microbial yield. The first constraint is the unit used to express the energy
available for microbial maintenance and growth. Whereas ME in a good
index of energy available to the host, it does not represents the energy avail-
able for microorganisms because some faecal and urinary energy losses
come from metabolites previously synthesised by microbes, and the
methane losses which are also subtracted from DE to calculate the ME con-
tent of foods, are also a product of energy utilization by microorganisms. In
addition, ME from fat, undegraded protein and volatile fatty acids in the
case of silage is not available to microorganisms. Although this is consid-
ered, at least in part, by the AFRC, INRA and Nordic systems, the FME and
FOM do not reflect precisely the energy available for microorganisms
because the proportion of ME or DMO which is available in the rumen varies
233
with the rate of degradation of organic matter in the rumen and the feed
ruminal retention time.
Another noteworthy aspect is that, although all current systems take

into account the rate of rumen protein degradation and rumen outflow rate
to estimate the nitrogen supply to rumen microorganisms, no attempt is
made to describe the rate of fermentation of organic matter and to account
for effects of intake on the amount of organic matter fermented in rumen.
None of the revised systems consider either the cost of microbial mainte-
nance and protein synthesis or the partitioning of available energy between
microbial maintenance and growth, although the cost of microbial mainte-
nance may vary considerably with diet and level of feeding, which may have
a substantial repercussion on net microbial growth.
Current European systems only take into account the microbial need
for nitrogen, which is very simplistic considering the complex nitrogen
metabolism of microorganisms and the needs for different nitrogen sub-
strates (ammonia, amino acids or peptides) depending on microbial popula-
tion and energy disposal. The repercussion of the protozoa population on
nitrogen turnover in the rumen is not considered and the contribution of
endogenous N to microbial protein synthesis is also poorly understood.
The possible benefits of synchronising energy and nitrogen supply

with diet according to the relative rates at which energy and protein
becomes available to rumen microorganisms is not considered by any of the
current European systems either. Amino acid composition and digestibility
of microbial true protein is considered to be constant without taking into
account the relative proportion of protozoa and bacteria and the possible
variation of microbial protein composition according to dietary conditions.
Apart from the imprecision that the different systems can incur in the
estimation of metabolizable protein supply, for the reasons outlined, the
quantification of post-ruminal metabolism of absorbed amino acids is even
more superficial. As stated by NEWBOLD (1994) “Translating and under-
standing of biochemistry of protein utilisation at tissue and organ level into
appropriate coefficients to convert net essential AA requirements into metab-
olizable essential AA (MEAA) requirements remains a significant challenge as
important perhaps, as further refinements of estimates of MEAA supply”.
However, there is insufficient data at present concerning net requirements
of individual amino acids for different functions, and the quantification of
234
MEAA supply is not easy. Therefore, revised protein evaluation systems do

not take individual amino acids into consideration.
Current systems assume expected efficiencies of utilization of

absorbed protein in conditions in which the supply of protein limits pro-
duction, but the optimal production response is certain to occur when the
ratio of increasing yield to supply is below of the maximum efficiency of uti-
lization of absorbed protein. In fact, the AFRC (1992) revised data from dif-
ferent trials carried out with dairy cows in different UK research centres,
showing that in each centre, the response of milk protein to increasing lev-
els of MP was between 0.21 to 0.27, which is clearly lower than the values
of 0.68 adopted for knl by the different current systems (between 0.64 and
0.80).
The current MP system can be used to predict MP requirements for a

stated level of production but does not define the marginal response to an
increment in protein supply above the level for maximum efficiency of uti-
lization, in terms of yield of saleable products. The positive response of milk
protein yield to MP supply above the optimum dietary MP:ME ratio is well
documented in both ewes and cows. This response may be caused by the
increase in DMI, the mobilisation of body lipids or the use of amino acid as
glucose precursors via neoglucogenesess. Moreover, elevated propionate
absorption may spare amino acids used for glucose production.
It should be noted that current systems do not account for intake

responses to MP and that there is an inadequate understanding of interac-
tions between MP supply and body reserves mobilization and how products
of carbohydrate fermentation may affect the response. There is also a lack
of knowledge of the protein composition of live weight loss of ewes and dairy
cows, and of the influence of the nutritional background and body condition
of an animal in the potential mobilisation of body reserves and its composi-
tion. The accurate prediction not only of energy and protein supply but also
of the absorption of lipogenic and glycogenic precursors is also essential in
the development of feed evaluation systems capable of predicting milk com-
ponent concentrations.
10.3.6. Evaluation system of “absorbed protein” (USA, NRC, 1985).
It classifies feedstuffs in three classes: protein, which degrades fast and
235
completely to ammonia, slowly degradable and utilizable protein, and

undegradadable and even potentially not utilizable protein. Degradation of
utilizable protein is influenced also by outflow rate. Protein recycled with
saliva is considered as a separate nitrogen source. Total nitrogen of feeds
and saliva make up what is referred to as intake protein (IP). Protein
requirement of animals is calculated according to the requirements of main-
tenance and production (growth, milk production, pregnancy) and metabol-
ic feacal losses. There are some values of protein requirement calculated
with the softver NRC (Table X-16).
10.3.7. New aspects introduced in the Cornell (CNCPS) proposal
The Cornell Net Carbohydrate and Protein System (CNCPS) is an integrated

set of equations and transfer coefficients that try to describe the dynamic
processes that occur in the ruminant. Contrary to the rest of systems, the
CNCPS was not designed specifically for diet formulation, but rather it pro-
vides a powerful tool for evaluating cattle diets and detecting possible fac-
tors that may limit its efficient use. It is based on a more mechanistic
approach than previous systems, taking in consideration numerous factors
related to the chemical characteristics of feeds, diet composition and level
of intake determining the supply of nutrients for microbial growth and the
236
host animal.
The model comprises a series of submodels that predict the feed car-
bohydrate and protein availability (SNIFFEN et al. 1992); the fermentation
process and microbial protein synthesis that occur in the rumen (RUSSELL et
al. 1992); cattle requirements (FOX et al. 1992) and amino acids supply and
requirements (O’CONNORS et al. 1993). The ruminal fermentation submodel
was complemented with a new submodel that tries to predict the produc-
tion, absorption, passage and concentration of ruminal VFA and pH (PITT et
al. 1996). Although both of these submodels are innovative, contributing to
a better prediction of absorbed nutrients and their efficiency of utilization,
and in spite of the fact that the requirements submodel also has some novel
contributions allowing a better prediction of cattle dry matter intake and
maintenance and growth requirements, the framework of CNCPS is mainly
based on the submodels of SNIFFEN et al (1992) and RUSSELL et al. (1992.)
The CNCPS considers that bacterial growth is dependent on the

amount of carbohydrates fermented in the rumen and on their rate of fer-
mentation. The amount of carbohydrates fermented in rumen from each
feed depends on its carbohydrate composition, the rate of fermentation of
each carbohydrate fraction (kd) and the rate of passage (kp). The proportion
of each carbohydrate fraction fermented in the rumen would be kd/(kd +
kp), where kd varies with feed and carbohydrate fraction and kp varies with
level of feeding and feed characteristics, particularly their effective NDF con-
tent (eNDF), which depends on total NDF content but also on particle size
and density.
The CNCPS distinguishes between four different carbohydrate frac-

tions with different degradation rates (Table X-17). Total carbohydrate con-
tent (% DM) is calculated as CHO = 100 – CP – fat – ash). Carbohydrates are
then partitioned according to degradation rate in: fraction A, which com-
prises mainly sugars and organic acids and is degraded very quickly; frac-
tion B1, comprising mainly starch and pectin, which have an intermediate
rate of degradation; fraction B2, representing the available cell wall compo-
nents (cellulose and hemicelluloses), which degrade slowly, and Fraction C,
representing the unavailable cell wall. These fractions are computed from
the structural carbohydrates (SC = ash free NDF corrected for associated
protein = ash free NDF–NDFN × 6.25) and non-structural carbohydrates
(NSC = Total CHO–SC), following the VAN SOEST procedure. Fraction C is
237
calculated as the lignin content × 2.4; fraction B2 is calculated as SC – frac-

tion C; fraction B1 is computed as the amount of starch and fraction A by
difference (NSC-starch). It must be pointed out that, according to this
method of calculation of fraction B1 and A, as originally proposed by SNIFFEN
et al (1992), pectin would be included in fraction A, not in fraction B1. Thus
further work is needed to better define NSC fraction.
To estimate the amount of nitrogen degraded in rumen, available for

microbial growth, and the amount reaching the duodenum, the CNCPS fol-
lows a similar approach. Feed protein is partitioned into three main frac-
tions: fraction A, that comprises the NPN (ammonia, nitrate, free amino
acids and peptides) and is instantaneously soluble in the rumen; fraction B,
that comprises the true protein potentially degradable in the rumen; and
fraction C that comprises Maillard products and lignin and tannin-bound
protein, considered as totally undegradable in the rumen and indigestible in
the small intestine. The true fraction is further partitioned into three frac-
tions (B1, B2 and B3) characterised in that they have different rates of
degradation. Fraction B1 includes most globulins and some albumins,
which are rapidly degraded in the rumen; fraction B2 includes most pro-
lamins and glutelins, with intermediate rates of degradations; and fraction
B3 comprising mainly prolamins and cell wall proteins showing a low rate
238
of degradation (Table X-18).
The analytical procedure to quantify these fractions is based in the

solubility of nitrogen in buffer and different detergent solutions following the
procedure of GOERING and VAN SOEST. The rate of degradation of each frac-
tion is determined using enzymatic or in situ methods. Once we know the
proportion and amount of each fraction in the feed, their rate of degradation
(kd) and the rate of passage of feed (kp), the supply of nitrogen available to
the microorganisms and the proportion of feed protein leaving the rumen
undegraded is calculated.
Although the theoretical framework used to estimate the carbohy-

drate available for microbial growth and the amount of nitrogen degraded in
the rumen and reaching the duodenum is physiologically meaningful and
original, there is little current information on several inputs in the model
(mainly the rates of degradation of different CHO and nitrogen fractions and
rate of passage) and current methods used to estimate rates of degradation
are not very reliable.
As previously commented, the main contribution and the core of the

CNCPS system is the rumen fermentation submodel and, in particular, the
approach used to estimate microbial protein synthesis. Current revised sys-
tems establish a simple empirical relationship between available energy and
microbial protein yield, which is a very simplistic approach that does not
239
reflect the complexity of interactions between rumen fermentation and

microbial growth processes occurring in the rumen. Using a mechanistic
approach, the CNCPS rumen fermentation submodel attempts to integrate
some of the factors affecting microbial protein synthesis: rate of fermenta-
tion and passage of carbohydrates, maintenance requirements and maxi-
mum theoretical growth rate of bacteria, form of nitrogen available to
microbes and rumen environment.
The CNCPS divides the ruminal microbial ecosystem into two micro-
bial groups, microbes that ferment nonstructural carbohydrates (NSC; CHO
fraction A and B1) and microbes that ferment structural carbohydrates (SC;
CHO fraction B2). The former bacterial population grows rapidly, is able to
utilise either ammonia or amino acids and peptides as a nitrogen source,
and can produce ammonia. The latter bacterial population grows more
slowly, does not ferment peptides or amino acids and only uses ammonia as
source of nitrogen.
Microbial yield (Y, g bacteria/g CHO) is calculated from a derivation

of the Pirt equation: 1/Y = 1/Ym + M/µ, where Ym represents the theoreti-
cal maximum microbial yield (g bacteria/g of fermented carbohydrate), M
represents the maintenance requirement of the microorganisms (g ferment-
ed carbohydrate/g bacteria per hour) and µ represents the bacterial specif-
ic growth rate, that in steady-state conditions may be substituted by the
rate of carbohydrate fermentation (kd, h-1). Whereas Ym is considered to be
0.4 g for the whole microbial population, NSC and SC bacteria are assigned
different maintenance requirements (0.150 and 0.050 g fermented CHO/g
bacteria, respectively). However, since the rate of degradation of NSC is
much higher than the rate of degradation of SC, microbial yield of NSC is
higher than that of SC bacteria and over the range bacterial specific growth
rate or carbohydrate degradation rates possible in the rumen, microbial
yield can vary by more than threefold.
Another factor considered to influence microbial yield in the CNCPS

is the supply of nitrogen as peptides and rumen content pH. The yield of
NSC bacteria is increased by almost 20% as the ratio of peptides to NSC
plus peptides in rumen liquor increases from 0 to 14%. Diets that are very
rich in concentrates and soluble carbohydrate have been shown to depress
rumen pH, SC bacteria growth and total microbial protein production. The
effect of pH on microbial growth is considered in the CNCPS, by reducing
240
microbial yield by 2.5% for every 1% decrease in effective NDF or NDF sup-
ply as forage below 20%. The flow of true microbial protein to the duodenum
is calculated assuming that 62.5% of bacteria dry matter is CP, and 60% of
this is true protein. The rest would be non-available nitrogen bound to
microbial cell walls (25%) and nitrogen in form of nucleic acids (15%).
FOR FURTHER READINGS
AFRC (1992): Technical Committee on Responses to Nutrients, Report Nº9. Nutritive

Requirements of Ruminant Animals: Protein. Nutr. Abs & Rev., Series B, 62, (12),
787-835, CAB International, Wallingford, Oxon
AFRC (1993): Energy and Protein Requirements of Ruminant. An advisory manual pre-
pared by the AFRC Technical Committee on Responses to Nutrients. CAB
International, Wallingford, UK
ARC (1980): The Nutrient Requirements of Ruminant Livestock. Technical Review,
Farnham Royal: CAB
ARC (1984): The Nutrient Requirements of Ruminant Livestock, Supplement Nº 1. Report
of the Protein Group of the ARC Working party. Farnham Royal: CAB
Bickel, H. and Landis, J. (1987): Present situation of protein evaluation for ruminants in
Switzerland. In: Jarrige, R. and Alderman, G. (Eds). Agriculture. Feed evaluation
and protein requirement systems for ruminants. Commission of the European
Communities. pp 41-45.
CEC (1987): Jarrige, R. and Alderman, G. (Editors). Agriculture. Feed evaluation and pro-
tein requirement systems for ruminants. 331 p. Commission of the European
Communities
Corbett, J.L., Freer, M. and Nolan, J.V. (1987): Present situation of the modern protein sys-
tems: Australia. In: Jarrige, R. and Alderman, G. (Eds). Agriculture. Feed evaluation
and protein requirement systems for ruminants. Commission of the European
Communities. pp 69-79.
CSIRO (1990): Feeding standards for Australian livestock. Ruminants. Standings
Committee on Agridulture and Resource Managemant. Ruminants subcommittee.
CSIRO Publications.
CVB (1991): Veevoedertabel, Centraal Veevoederbureau in Nederland, Lelystad.
De Boer, F. and Bickel, H. (1988): (Eds). Livestock Feed Resources and Feed Evaluation in
Europe. Present situation and future prospects. E.A.A.P. Publication Nº 37. pp 408
Reprinted from Livestock Production Science, 19(1,2), Elsevier. Amsterdam.
DLG (1997): Futerwerttabellen, Wiederkäuer, DLG-Verlag, Frankfurt
Fox, D.G., Sniffen, C.J., O’Connor, J.D., Russell, J.B. and Van Soest, P.J. (1992): A Net
Carbohydrate and Protein System for Evaluating Cattle Diets: III. Cattle
Requirements and Diet Adequacy. J. Anim. Sci. 70, 3578-3596.
Hvelplund, T. and Madsen, J. (1993): Protein systems for ruminants. ICEL. AGR. SCI, 7,
21-36
INRA (1978): Jarrige, R. (ed.), Alimentation des ruminants. INRA Publications, Versailles.
INRA (1988): Jarrige, R. (ed.), Alimentation des bovines, ovins et caprins. INRA
Publications, Versailles.
INRA (1989): Jarrige, R., (Ed.): Ruminant nutrition. Paris, John Libbez.Madsen, J,
Hvelplund, M.R., Weisbjerg, J., Bertilsson, I., Olson, R., Spröndly, O.M., Harstad, H.,
Volden, M., Tuori, T., Varvikko, P., Huhtanen, P. and Olafsson, B.L. (1995): The
AAT/PBV protein evaluation system for ruminants. A revision. Norwegian J.
Agric. Sci. Suppl. 19-37.
Madsen, J. and Hvelplund, T. (1994): Prediction of in situ protein degradability in the
rumen. Results of European ringtest. Liv. Prod. Sci. 39, 201-212.
Newbold, J.R. (1994): Practical application of the metabolizable protein system. In:
Garnsworthy, P.C. and Cole, D.J.A. (Eds), Recent Advances in Animal Nutrition.
Nottinghan. University Press. pp 231-264.
NKJ-NJF (1985): Protein evaluation for Ruminants. Acta Agric. Scand., Suppl., 25, 220 p.
NRC (1985): Ruminant Nitrogen Usage. 138 p. National Academy Press. Washington, D.C.
241
O’Connor, J.D., Sniffen, C.J., Fox, D.G. and Chalupa, W. (1993): A Net Carbohydrate and
Protein System for Evaluating Cattle Diets: IV. Predicting Amino Acid Adequacy. J.
Anim. Sci. 71, 1298-1311.
Ørkov, E.R. and McDonald, I. (1979): The estimation of protein degradability in the rumen
from incubation measurements weighted according to rate of passage. J. Agric. Sci.,
Camb. 92, 499-503.
Pitt, R.E., Van Kessel, J.S., Fox, D.G., Pell, A.N., Barry, M.C. and Van Soest, P.J. (1996):
Prediction of Ruminal Volatile Fatty Acids and pH within the Net Carbohydrate and
Protein System. J. Anim. Sci. 74, 226-224.
Rohr, K. (1987): Present situation of the modern protein systems: Germany. In: Jarrige, R.
and Alderman, G. (Eds). Agriculture. Feed evaluation and protein requirement sys-
tems for ruminants. Commission of the European Communities. pp 3-10.
Russell, J.B., O’Connor, J.D., Fox, D.G., Van Soest, P.J. and Sniffen, C.J., (1992): A Net
Carbohydrate and Protein System for Evaluating Cattle Diets: I. Ruminal
Fermentation. J. Anim. Sci. 70, 3551-3561.
Schmidt, J., Varhegyi, I., Varhegyi, J. and Cenkvari, E. (1999): New Hungarian Protein
evaluation system. Hungarian Agricultural Research, 8(1), 8-11.
Schmidt, J., Varhegyi, I., Varhegyi, J. Turine, Cenkvari, E. (2000): Energy and protein eval-
uation of ruminants, Mezőgazda Publisher, Budapest
Sniffen, C.J., O’Connor, J.D., Van Soest, P.J., Fox, D.G. and Russell, J.B. (1992): A Net
Carbohydrate and Protein System for Evaluating Cattle Diets: II. Carbohydrate and
Protein Availability. J. Anim. Sci. 70, 3562-3577.
Tamminga, S., Van Straalen, W.M., Subnel, A.P.J., Meijer, R.G.M., Steg, A., Wever, C.J.G.
and Blok, M.C. (1994): The Dutch protein system: the DVE/OEB-system. Liv. Prod.
Sci. 40, 139-155.
Veresegyhazy, T., Nagy, A., Fekete, S. and Kutas, F. (1989) In vitro evaluation of protein
degradability in the rumen and digestibility of undegraded protein. Acta Vet. Hung.
37, 103-115.
Vérité, R., Michalet-Doreau, B., Chapoutot, P., Peyraud, J.L. and Poncet, C. (1987):
Révision du système des protéines digestibles dans l’intestin (PDI). Bull Tech CRZV
Theix, INRA, 70, 19-34.
Wallace, R.J. (1983): Hydrolysis of 14C-labeled proteins by rumen microorganisms and by
proteolytic enzymes prepared from rumen bacteria. Br. J. Nut. 50, 345-355.
242
Chapter XI
MINERALS IN THE ANIMAL NUTRITION
©Sandor Gy. Fekete
11.1. Generally about minerals

11.1.1. Justification of the need for minerals
11.1.2. Mineral content of the body and individual organs.
11.1.3. Sources of mineral supply
11.1.4. Absorption and excretion of minerals
11.1.5. Deficiency syndromes, toxicity.
11.2. Homeostasis of minerals
11.3. Effect of feeding on acid-base balance of animal organism.

Cation–anion balance
11.3.1. Theoretical basics
11.3.2. Field of practical application
11.4. Macro-elements
11.5. Micro- or trace elements

Chapter XI: MINERALS IN THE ANIMAL NUTRITION
In the feeds and animal body approximately fifty elements occur. Mineral
composition of the animal body reflects the requirements, too.
11.1. Generally about minerals
11.1.1. Justification of the need for minerals

Mineral content of the animal body is approximately 3 percent. These ele-
ments can be classified into two (or three) groups. Macroelements occur in
higher concentration (>70 mg/kg LW, or according others 50 mg/kg LW)
and these are the calcium (Ca), phosphorus (P), magnesium (Mg), potassi-
um (K), sodium (Na), chlorine (Cl) and sulphur (S) (Table XI-1). They are also
called as major element, those which are required in gram/day quantities
in diet. The other elements are called microminerals. There are classifica-
tions, according to element of the concentration 10 to 200 ppm (mg/kg) are
the meso-elements. Until now from the approximately 40 microelements of
the organism 16 proved to be really essential, these are the iron (Fe), zinc
(Zn), manganese (Mn), copper (Cu), fluorine (F), iodine (I), cobalt (Co), sele-
nium (Se), chromium (Cr), nickel (Ni), molybdenum (Mo), silicon (Si), tin
(Sn), vanadium (V), lead (Pb) and arsenic (As).
Table XI-1: Mineral content of adult mammalian body and blood plasma
*Inorganic phosphorus; ** In the blood serum higher by approx. 12-16%
The main functions of the minerals in the animal organism are as fol-
244
lows: structural, regulatory and contribution to the milk and egg produc-
tion. MINERALS AS BODY CONSTITUENTS. The highest mineral containing tissues
in the organism are the bones and the teeth. The hair, feather and horn for-
mation (horn, nail, claw, hoof etc.) have medium, whereas soft tissues and
faeces have lower mineral content. This does not mean an order of impor-
tance, of course. Generally spoken 25% of the bones is ash, which contains
approximately one third calcium and one fifth phosphorus, but the concen-
tration of magnesium is considerable, too. Besides bones, the second great
mineral storages of the organism are the hair and the horn formations. They
offer also a possibility for monitoring the mineral status. Some of the min-
erals have organ (or tissue) preference for accumulation, for example the
cadmium in the renal cortex, having a biological half time of more than 17
years and in turn, the red blood cells have important iron concentration.
REGULATORY FUNCTION. The majority of mineral takes part in the bio-
chemical-physiological regulation of the organism. This role may be an inde-
pendent or as part of a vitamin (e.g. Co in the vitamin B12), enzyme (e.g. Se
in gluthation-peroxydase) or of hormone (iodine in the thyroid hormones).
The regulatory function can be realized through biochemical reactions or
through influencing the appropriate concentration of body fluids. A trifle
portion of calcium, magnesium and phosphorus, whereas the majority of
sodium, potassium and chloride are present in the electrolytes of body flu-
ids and soft tissues. The role of electrolytes of body fluids (blood, cere-
brospinal fluid etc.) is important in maintaining of acid-base balance and
the osmotic pressure. They do influence the membrane permeability and the
irritability of muscles and nerves. Salts of saliva, gastric juice and rumen
fluid are indispensable for functioning of digestive enzymes and as living
medium for the micro-organisms.
Contribution in the MILK AND EGG PRODUCTION. Cow milk contains 0.5-
0.6% minerals (4-5% in dry matter). Except iron, mammals’ mineral supply
can be assured by giving milk. All of the mineral elements of the milk,
directly, or indirectly (previously stored in the body) derives from the feed or
drinking water. Sodium content of milk comes directly from the ingested
feed, whereas calcium, phosphorus and magnesium partly derive from the
momentary feed intake and partly from the mobilization of body reserves
(bones). In principle, the same is valid for the eggshell formation, too.
11.1.2. Mineral content of the body and individual organs.
245
As it can be seen in Table XI-1 too, the values of mineral concentrations are
usually given not only in fresh (original) but also in dry matter basis. Using
the dry matter basis, as a form of expression, there are no significant dif-
ferences among values of adult individuals of different species. Specific min-
eral composition of the individual organs of mammals is also very similar to
each other. Undernutrition and water deprivation increase the mineral con-
centration of fat-free dry matter. It is worth mentioning that while concen-
tration of sodium, potassium and chloride in fat-free dry matter does not
change during ontogenesis (from the embryo till the adult age), that of cal-
cium, phosphorus and magnesium in the foetus and newborn are only
approx. 50% of the adult values. Blood concentration of macroelements
(mostly that of calcium, magnesium, sodium, potassium and chloride) is
maintained, almost independently from the nutritional supply, in narrow
limits by the neurohormonal regulation. In case of the essential microele-
ments this regulation is not so strict; consequently the blood concentration
may reflect the actual status.
11.1.3. Sources of mineral supply
The most important mineral sources of the production animals are the for-
ages and the concentrates (grains and seeds). Animal products like bone
meal may also have important mineral content, but in many countries they
are not authorized. Geological supplements (limestone, rock phosphates
and rock salt) are also important. Contribution of the drinking water (except
some elements, like iodine) to the mineral supply is negligible. Soil (mud)
contamination of green forages and crops may also provide minerals.
Mineral content of feedstuffs depends upon the species and vegeta-
tion (phenological) period of the plant, the type and composition of the soil,
climate and the applied agricultural engineering (fertilization, irrigation
etc.). While the mineral content of the vegetative parts may change within
wide ranges in relation to the environmental factors, that of the germinative
parts (grains, seeds) is rather constant (KÁDÁR, 1992). Relative deviation of
the microelement concentrations is much higher than that of the macroele-
ments.
11.1.4. Absorption and excretion of minerals
The main sites of absorption are the small intestine and the cranial section
of the colon; microelements are mostly in form of ions. In ruminants, there
is absorption (e.g. Mg) across the rumen wall, too. Part of minerals, excret-
ed into the intestinal lumen by the digestive juices, can be re-absorbed. The
246
main route of excretion may change according to the animal species and the
feed composition. For example, while ruminants eliminate calcium and
phosphorus mainly by the faeces, the monogastrics do it by the urine.
Availability of minerals. While evaluating a feed mixture, besides the
absolute mineral content, the data of utilization in the body (percentage of
absorption) are also indispensable. In case of the minerals the determina-
tion of the apparent digestion coefficient has no meaning, because in the
faeces the non-absorbed and the endogenous mineral are mixed. There is
no difference between the feed-originated and the endogenous (from saliva
and digestive juices) minerals. Part of the endogenous mineral may re-
absorbed; the remaining portion gives the endogenous (metabolic) mineral
content of the faeces. In ruminant, the endogenous portion may give more
than the half of the faeces mineral content. On the contrary, in mono-
gastrics the degree of endogenous mineral excretion is low. Availability of
minerals can be measured using balance trial or by applying labelled min-
erals. The mineral supply is considered as good, if the balance is zero at the
adult and a positive value at the growing animals. In some special physio-
logical states, the negative balance may be normal (e.g. lactational osteo-
porosis).
11.1.5. Deficiency syndromes, toxicity.
Long-lasting deficiency of minerals results in declining production, repro-
ductive failures and impaired immune system. In case of some elements
(Cu, Co and Se) the “therapeutic index” is small and 4-5 times of the
requirement may be toxic. Thus, in case of minerals, besides the minimum
requirement the maximum tolerable level should also be known.
11.2. Homeostasis of minerals

The conception has originally been developed for the microelements, but
except the structural role, it can be applied to the macroelements, too (FIG-
URE XI-1).
247
FIGURE XI-1: Outline of mineral homeostasis ©Sandor Gy. Fekete
248
Mineral, ingested by the feed is only partially absorbed. The absorbed

portion may or may not create a reserve. If there is a depot (e.g. Cu), the
given mineral should not be provided every day. On the contrary, if no stor-
age forming happens (e.g. Na); the continuous daily supply must be
assured. Mineral, present in blood or body fluids, may participate in the
osmo- and acid-base regulation or may contribute to the synthesis of bio-
logically active substances. The main routes of excretion are the re-excre-
tion into the gut lumen, elimination by bile, urine, saliva, sweat and milk.
Considering the above described, according to their homeostatic con-
trol, minerals can be divided into three main groups. For example, the iron
homeostasis in the organism is regulated unipolar (exclusively) at the level
of absorption. Practically 100% is the absorption in case of the sodium and
the regulation is also unipolar, but at the level of excretion. In other words,
the sodium absorption basically is independent from the needs of organism;
if there is an excess, every route of excretion (faeces, urine, saliva, sweet) is
used, proportionally to the measure of surplus. There is a sodium excretion
through the milk, too, but this concentration is fixed. Elements of the third
group have a bipolar regulation, namely there is regulation both at the level
of absorption and excretion. It means that absorption varies within the ele-
ment-specific ranges. The elimination is realized by the different endoge-
nous excretion into the gut lumen (digestive juices, bile: Zn, Mn, Cu and Ni)
and by the changeable urinary excretion (Se, Cr, Mo and Si). Owing to this
homeostatic control, the concentration of minerals in the organism remains
within ranges, sufficient to fulfil the physiological function, but does not
reach a possibly damaging accumulation. As a cumulative effect of the
homeostatic regulation, the mineral content of organism does not follow lin-
early the dietary intake, avoiding a possible poisoning (e.g. Cu and Zn).
Absorption and utilization of minerals are strongly influenced by the
other feed constituents/ingredients (protein, fat, carbohydrates and espe-
cially fibre) and by the interrelated, mineral-to-mineral INTERACTIONS. The
interaction can be facilitating (chelating by an amino acids may enhance
absorption of some microelements) or inhibiting (phytic acid bond decreas-
es the phosphorus availability). Mineral-to-mineral interaction may act in
the soil, in the plant and also in the animal organism (rumen, gut lumen
and tissues).
249
In all animal species occur the sodium-potassium and the calcium-

phosphorus interactions. The latter means that the excess of phosphorus
decreases the calcium absorption. The copper-molybdenum-sulphur inter-
action is characteristic of ruminants, because the site of it is the rumen.
Basically, the copper absorption decreases under the influence of sulphur;
the excess molybdenum reinforces this effect (FIGURE XI-3). The most com-
mon is the secondary copper deficiency, induced by the excess of molybde-
num and/or sulphur. On the other hand, too low molybdenum and sulphur
concentrations may lead to copper-poisoning in the susceptible sheep.
FIGURE XI-2: Copper-molybdenum-sulphur interaction
11.3. Effect of feeding on acid-base balance of animal organ-

ism. Cation-anion balance
11.3.1. Theoretical basics

The potential of a nutrient can be evaluated by its electrical charge. That of
alkaline character are the Na, K, Ca, Mg, carboxylate, malate, citrate,
fumarate, glutamate, Lys and NH4, in cationic form and in turn, that of
acidifying effect are the Cl, S, P, Met, in form of anion.
EARTH-ALKALINOGENIC ALKALINOGENICITY (EAA). Hungarian researchers
250
(MAREK, WELLMANN and URBANYI) during the late twenties, early thirties
searched for the causes of rickets, studied the relationship of feeding and
acid base balance. During metabolism both acidic and alkaline metabolites
emerge in the organism, proportionally to the intensity of processes. The
acidic compounds are predominant (Table XI-2).
Table XI-2. Some important acid and base producing process of the metabolism
* e.g. ketosis; ** e.g. lactacidosis
Continuously producing acids can be counterbalanced, if the ingest-

ed feed is of alkaline character, specifically in the measure of the degree of
metabolic intensity. In formula, EAA = CaO+MgO-P2O5, meq/100 g DM,
should be a positive value. In term of definition, the EAA of a feedstuff (or a
feed mixture) is a value, which shows that the joint amount of alkaline cal-
cium oxide and magnesium oxide is larger or smaller than the acidic phos-
phoric pentoxyde, expressed in milliequivalent. The recommended numbers
in the ration for weaned animals is +25, for growing +10 and for adults +5.
For example, the EAA-value of corn grain is -5.3 g/kg, therefore for an adult
pig to achieve the optimum value, +10.3 EAA supplementation is required.
The EAA-value of limestone (CaCO3) is 2.04, thus the necessary addition is
10.3/2.0=5.1 gram. The reason of using oxides, instead of elements, is only
of technical origin, having made the mineral determination from the ash of
the feeds and animal body.
251
Alkalinogenic alkalinogenicity (AA) compared the joint amount of

potassium and sodium to the joint amount of chlorine and sulphur. Sum of
EAA and AA gives the total alkalinity (or acidity, if negative) of the feedstuff.
This type of calculation used to be applied in Middle Europe till the late six-
ties, but after that it has been replaced by the simple calculation of the
required amount of calcium, phosphorus and sodium. Nevertheless, at the
same time, the concept experienced it renaissance in North America
(PATIENCE et al.1987).
Namely, the DIETARY UNDETERMINED ANION (dUA) is a modern form of
expression of the same principle.
dUA=(Na++K++Ca2++Mg2+)-(Cl-+H2PO4-+HPO42-+SO42-)
The name derives from the fact that analytical determination of the above
„fixed” cations and anions allows to indirectly measure of the net concen-
tration of metabolizable ions, such as lactate, bicarbonate and acetate,
because the diet must be electroneutral. Thus, dUA in fact is an estimate of
the quantity of acid-consuming metabolizable anions in the diet.
To demonstrate that in the influence of dUA on production parame-
ters the primary is the summarized ratio of cations and anions and not the
effect of individual ions, patience and WOLYNETZ (1990) carried out three tri-
als. In Trial I groups of weaned piglets were fed on isoenergetic and isopro-
teic, corn-soybean diet. In the experimental diets part of the neutral CaCO3
has been changes by the acidifying CaCl2. Results are presented in Table
XI-3 and
Table XI-3: Influence of the dUA value of the feed mixture on the physiological and pro-
duction parameters
The BASIS EXCESS (BE) is the basis surplus in the blood, given mostly
by the bicarbonate (HCO3-) and haemoglobin content. The numerical value
of basis excess is equal to the amount of acid able to re-neutralize one litre
of blood to pH 7.4. Its normal value is 0±2.5 mmol/l. The BE (mmol/l) can
252
be calculated from data of alcalic (+) acidic (-) feed minerals (mmol/kg DM):
BE=2[Ca]+2[Mg]+[K]+[Na]-2[P]-2[Met]-[Cl].
FIGURE XI-3: Effect of dietary undetermined anion (dUA, mEq/kg) on the

average daily gain (ADG, g) in growing pig
Data shows that CaCl2 supplementation decreased both feed intake

and average daily gain. During discussing this result, it occurred that owing
to the corn and soybean ingredients, the potassium concentration in feed
mixture was continuously high. In Trial 2 part of soybean has been replaced
by corn gluten. The tendency of results remained the same at this lower
potassium level. During Trial 1 and 2 the blood chloride level increases,
although it is strongly regulated homeostatically. To decide that the results
are related to the acidifying effect or to the high chloride concentration of
the diet, in a high chloride group additional sodium bicarbonate were given
to achieve the control dUA value. The high chloride but similar to control
dUA diet resulted in similar parameters as in control. This proved that the
data did not derive from the specific effect of the chloride, but the created
lower dUA value. During the 28-day-long trials no adaptation could be
observed, the differences in average daily gain between treatments did not
change.
For ruminants, to express of the acidifying or alkalizing character of
the feed mixture, the DIETARY CATION ANION BALANCE (DCAB) have been devel-
oped: DCAB, mEq/kg DM = [(44Na+26K)-(28Cl+63S)]/100 g DM. Recently,
the nomenclature changed to dietary cation-anion difference (DCAD:S) and
the simplified formula, when sulphur is avoided, DCAD (CHAN et al. 2005).
For horses the DCAD:S is used, even if the S in the abbreviation is omitted
253
(GRAHAM-THIERS. 2005). It worth mentioning, that this formula cannot be

applied if a strong non-volatile acid (e.g. Virtanen-type of silage) is present.
For breeding sow the formula of dietary Cation Anion Balaence includes the
sulphur-containing amino acids, too: dCAB, meq/kg DM=
=(43Na+26K+50Ca+82Mg) – (28Cl+59P+13Met+Cys. For poultry, the sim-
plified, but still of good predictive value DIETARY ELECTROLYTE BALANCE (dEB)
is proposed (MONGIN. 1981): dEB = Na++K+-Cl-. In the practice, it is used for
pig and cattle, too. For dogs the CATION ANION RATIO (CAR=KAV in German)
supposed an average methionine intake, is as follows (SCHUKNECTH, 1991),
where Met and Cys as SO4-sources play a role: CAR, mmol/kg
DM=2Ca+2Mg+Na+K)–(2P+2Met+2Cys+Cl).
For clinical and experimental purposes the following blood parame-
ters are also used for evaluating the acid-base balance of the animal: ANION
GAP (Na+K)-(HCO3+Cl), net strong ion difference (SID), total weak acid con-
centration (ATOT), partial pressure of carbon dioxyde (PCO2), total protein,

albumin and lactate concentration (LEITH, 1991).
11.3.2. Field of practical application

As it is clear from the above described, feeding of slightly alkaline ration has
positive effect on the voluntary feed intake, average daily gain, feed conver-
sion efficiency and even on the milkfat production. In growing poultry the
acid base balance influences also response to heat stress and occurrence of
leg deformities. BORGES et al. (2003) compared the effect of feed mixtures of
different dietary Electrolyte Balance (dEB=mEq Na+K-Cl/kg) on perform-
ance and immune status of chickens exposed to thermoneutral or heat
stress. As a summary, higher dEB (201, 220 and 250) in the thermoneutral
environment improved feed intake, weight gain, feed efficiency and lympho-
cyte percentage, parallel to a decrease in heterophyl percentage, while com-
pared to lower (0, 40 and 140) or higher (340) dEB groups. In heat stress
rooms the results had higher deviation, but the tendency was the same. As
it is generally accepted, the optimum value for growing-finishing pig is 155
mEq/kg and for growing chick 250 mEq/kg dEB. Eggshell quality is also
influenced by the cation-anion balance of ingested feed. Namely, the egg
production and shell quality can be ameliorated by alkaline salts of sodium
and potassium bicarbonate. Acidogenic anions of calcium like monobasic
phosphate, chloride, sulphate and dibasic phosphate have adverse effect on
the mentioned parameters (KESHAVARZ, 1994).
254
However, there are special physiological situations, when application

of an acidogenic feed mixture is benevolent. One of these periods is the week
2 to 5 of piglets (before and after weaning); when in their stomach the
achlorhydria (lack of sufficient hydrochloric acid production) is physiologi-
cal. Lack of low pH in stomach allows the survival of diarrhoea-causing bac-
teria, like E. coli. In order to counterbalance this situation, but without the
great disturbances of acid-base balance, instead of HCl, organic acids are
proposed (e.g. malic acid, fumaric acid) or betain (which is basically a long-
release HCl) around the weaning time. Possible acidosis can be avoided by
keeping the dUA-value of the feed within the range of 250-300 mmol/kg
DM.
Dietary cation-anion balance may influence the incidence of milk
fever in dairy cows. Even an unfavourable calcium and phosphorus supply
during the dry period is unable to provoke milk fever, if the acid-base bal-
ance of the organism facilitates bone mobilization. For the prevention of
milk fever, the DCAD should be lowered below -50 (within the range of -50
to -150). To achieve these values, dry period’s feedstuffs should be selected
(avoidance of alfalfa and other leguminous forage is advisable), sulphuric or
hydrochloric acid or NH4Cl-supplementation can be given at the end of dry
period. Table XI-4 and Table XI-5 show the DCAD of some typical dairy cow
feedstuffs.
Table XI-4: Typical values of European grass and corn silage (after ROTHERT, 1998)
255
Table XI-5: Typical values of some typical USA cattle feedstuffs
*In the EU is not authorized to use!!!!
Some types of urinary calculi, mostly the struvite (magnesium-ammo-

nium phosphate) cannot precipitate in acidic pH. During the prevention of
urolits formation in dogs, cats and intensively grain-fed beef bulls and rams
(wethers) the supplementation of ration by CaCl2, NH4Cl, methionine or
glutamic acid hydrochloride is successful. In case of breeding sows, the goal
of urine acidifying is the reduction of total and Gram negative bacterial
count in the periparturient period. As described by DOBENECKER (1999), the
pH of the urine can be estimated as follows:
pHsow urine= =6.19+0.0031×dCAD+3×10–6×dCAD2.
To achieve a significant reduction of bacterial counts in urine of sows,
a reduction of urinary pH below 6 appears to be necessary.
There is a strong relationship between the acid-base balance and
amino acid metabolism of organism. Changes of acid-base balance may
modify the metabolic pathway of certain amino acids. Degradation of amino
acids (e.g. Lys, Arg, His, Met, Cys) means an acid loading to the organism.
While supplementing by sodium bicarbonate (NaHCO3) the lysine and argi-
nine requirement decreases: “lysine sparing effect”. The possible mode of
action is a down-regulation of kidney arginase activity (see nutrigenomics in
Chapter XVII), degrading lysine and arginine. Therefore the significance of
dUA (or dEB) has been accentuated by the use of synthetic amino acids in
chicken and pig diets.
256
11.4. Macroelements.
From the minerals of living organism, this is the calcium, phosphorus and
magnesium, which are present in the highest concentration.
CALCIUM (CA). Essentiality of calcium in the animal body is known
since more than hundred years. The structural role of calcium is the con-
tribution to the bone, teeth and eggshell formation. However, the bones are
also an active players of the calcium metabolism by providing tissues and
body fluid by calcium and phosphorus. Outside the skeleton, approximate-
ly 1% of the total calcium is present, namely in the extracellular fluid, in soft
tissues and membrane structures, taking part also in vital functions.
Calcium is indispensable in blood clotting, in the functioning of skeletal and
heart muscle and in maintaining membranes’ integrity and in regulating the
calpain-calpastatin group of protease enzymes of muscle protein synthesis
and degradation. Calcium is part of many important enzymes, like trypsine,
chymotrypsine and by means of the calmodulin, modifies the activity of a
series of other enzymes.
Its absorption, distribution in the animal body and excretion are
influenced by several factors. Calcium is absorbed by active transport, facil-
itating by the mucosal calcium-binding protein (CaBP). The synthesis of the
latter is induced by the active vitamin D. There is also a passive and vita-
min D independent absorption of calcium ions (horse, rabbit and guinea pig
do not need vitamin D for their calcium metabolism). The main site of the
active process is the proximal segment of the duodenum, while the passive
absorption occurs in the lower section of the small intestine. The routes of
excretion are the urine and the faeces (predominant in ruminants).
The calcium content of feedstuffs varies within wide ranges. It is gen-
erally true that the calcium concentration in cereals is low (0.1 to 0.3%), in
green forages medium (0.31 to 0.36%) and in leguminous green plants high
(1.2 to 1.7%, Calcium supplementation can be made by using the following
compounds (in parentheses the element content): calcium carbonate or
ground limestone (36%), dolomitic limestone or dolomite (22% Ca 10% Mg),
oyster shell (35% Ca and 0.3% Mg), calcium sulphate or gypsum (29%, but
low availability!), dicalcium phosphate (25 to 28% Ca and 18 to 21% P) and
defluorinated rock phosphate (32% Ca and 18% P).
The perorally ingested calcium practically is not toxic. Assumed an
257
appropriate phosphorus supply, the following calcium concentration are

considered as easy to tolerate: ruminants 2.0%, swine 1.0%, poultry (grow-
ing) 1.2%, laying hen 4.0%, horse and rabbit 2.0%. One of the most com-
mon deficiency syndromes (combined with phosphorus and vitamin D defi-
ciency) is the rickets (rachitis) and its variants (osteomalacia and osteo-
porosis). The typical sign of the rickets is the “rosary” on the osteochondral
part of ribs caused by the lack of mineralization and cartilage overgrowth.
In intensively concentrate-fed beef cattle the osteochondrosis of the tarsal
joint may cause the heel bone (os tarsale) to break off together with the
Achilles tendon. The milk fever of dairy cow and the lactation tetany of
bitches (puerperal eclipse) are treated elsewhere. The severe calcium defi-
ciency in sheep (fed on solely on grain) may result in osteodystrophia fibrosa
and in cats and dog, fed on lean meat, without calcium supplementation,
secondary, alimentary hyperparathyroidism develops (“all meat syn-
drome”).
PHOSPHORUS (P). Although phosphorus is widely spread in the nature,
it cannot be found in free form. In rocks and almost in all body fluid can be
found in form of orthophosphoric acid. Phosphorus is one of the elements
in the body of the most important physiological roles. It takes part in the
bone and teeth formation; contribute to building up the DNA and RNA,
AMP, ADP, ATP and several co-enzymes. The compounds of phosphorus are
important in maintaining the acid-base balance of the body fluids.
Absorption of phosphorus occurs from the duodenum, mostly in form
of orthophosphate. The efficiency of absorption is influenced by a series of
different factors: source of phosphorus, calcium-to-phosphorus ratio, pH in
the gut lumen, vitamin D-status, lactose and lipid intake, and last but not
least, from the concentration of other minerals, like magnesium, alumini-
um, iron, potassium and manganese. Animals respond to increase in phos-
phorus requirement by increasing intestinal absorption via an energy-
dependent process, thus higher body needs improves its absorption. In
monogastrics, active vitamin D is involved in stimulating P absorption
across the intestine. Quantitatively, most of the absorbed phosphorus is
retained in the bones and teeth. In case of a deficiency, the mobilization
from the bones helps in maintaining the physiological phosphorus concen-
tration of blood plasma. Blood level of calcium and phosphorus is regulat-
ed by the parathormone, thyreocalcitonin and active vitamin D. The blood
phosphorus concentration generally moves in opposite direction than that
258
of the calcium, except the milk fever of dairy cows, when both are decreased.
Excess phosphorus in monogastrics is excreted by the urine, in the rumi-
nant mostly by the faeces, but the small urinary excretion is an indicator
for phosphorus status.
Phosphorus content of feedstuffs is quite different. Phosphorus level
in plants depends upon its species, vegetation phase and fertilization of the
soil. In turn, the bioavailability of phosphorus in the feeds shows a great
deviation, too, depending on the chemical structure, species, age and phys-
iological state of the consuming animal. Phytate phosphorus is poorly avail-
able for monogastrics, especially for poultry, but by the application of com-
mercial phytase enzyme practically solves this problem. Moreover, use of
this decreases the need of inorganic phosphorus supplementation, reducing
the phosphorus emission in the environment.
Feed mixtures of the majority of animal species require phosphorus
supplementation. Phosphorus supplementation can be made by using the
following compounds: dicalcium phosphate (25 to 28% Ca and 18 to 21%
P), defluorinated rock phosphate (32% Ca and 18% P), and phosphoric acid
(31.6% P). monobasic sodium phosphate (22.4% P) and sodium tripolyphos-
phate (30.85% P). Phosphorus supplements are generally unpalatable, thus
the ad libitum offering is not an optimal solution (NRC, 2001). Where the
authorization makes possible their use, steamed bone meal (29% Ca, 14%
P and 0.6% Mg), poultry excrement and guano can also be applied. The
bioavailability of the above phosphorus sources is significantly different; the
highest is of the monocalcium phosphate and the lowest of gypsum. These
phosphorus supplements, according to their provenience, may contain toxic
elements (F, Cd, Hg, As, V, etc.). The maximum tolerable concentration of
these elements is legally regulated and controlled (see Chapter 2). The bio-
logical values of phosphorus supplements means the efficiency of absorp-
tion and retention in tissues (in the first place, bone). For measuring the
biological values, the study of absorption, the measurement of retention, the
analysis of bone ash, evaluation of the mineralization and the measurement
of blood alkali phosphatase activity are in use.
Phosphorus not only one of the important elements in the organism
itself, but also influences the availability of other minerals. For example, the
majority of animal species (except laying hen) requires a calcium-to-phos-
phorus ratio of 1.5-2 to 1 in their feed mixture. Horse, rabbit and ruminants
(with the exception of the dry cow) easily tolerate the wider ration, unless
259
they have no real phosphorus deficiency. Actual or related to the calcium

excess of phosphorus may facilitate the formation of urinary calculi, caus-
es eggshell fragility and growth depression, decreased feed utilization and
even higher mortality at broiler chicken. To prevent the milk fever, during
dry period the recommendations should be followed (see Chapter 25).
Phosphorus deficiency in dairy cow may lead to alimentary infertility or in
severe cases acute hypophospataemia in beef cow. If phosphorus deficiency
is combined with selenium, copper and energy, may lead to depress the
ability of red blood cell to produce ATP, their cell wall become rigid lack of
sufficient sodium pump and the postparturient haemoglobinuria may devel-
op in fresh cows. This ailment may occur together with the ketosis. For con-
trolling the calcium and phosphorus status, the balance calculation (real
feed composition and ingested amounts) proved to be the most reliable. The
blood parameters are less sensitive.
MAGNESIUM (MG), together with the calcium and phosphorus, partici-
pates in the formation of bones and in the setting of the neuromuscular irri-
tability. Magnesium is an activator for alkali phospatase, arginase and
kinases enzymes. Magnesium has an important role in the immune func-
tions, too, being involved in lymphoblast transformation and antibody syn-
thesis. In the plants it is a part of the leaf-green (chlorophyll). The main site
of absorption in monogastric is the small intestine, in the ruminants from
the reticulo-rumen and after the recent data, from the large intestine, too.
The efficiency of absorption is about 20 to 30%. The animal practically can-
not regulate Mg absorption and because it is excreted mostly by the urine,
the magnesium homeostasis is achieved by the kidney. Mg reserves are lim-
ited; 2/3 of magnesium is in the skeleton, where the calcium-to magnesium
ratio is 50 to 1.
Magnesium concentration of grasses is low (1 to 2 g/kg DM); on the
contrary, in the other green forages, especially in the leguminous green
plant higher (3 to 4 g/kg DM). There is a magnesium excretion by the milk
(0.12 to 0.15 g Mg/kg), consequently the requirement of lactating dairy cow
is 2.0 mg/kg DM, that of for heifers and dry cows 1.6 g/kg DM. Except some
special geographical regions (see Chapter XXV), and the occurrence of mag-
nesium deficiency (“grass tetany”) in dairy cow and breeding ewes is rare.
On the contrary, calves, fed on inappropriate milk replacer, magnesium
deficiency develops, resulting in hyperirritability, convulsions, muscle
twitching, special posture, then coma and death. Magnesium supplementa-
260
tion can be made by adding magnesium oxide or magnesium sulphate.

Excess Mg intake favours the formation of struvite urolits in cats.
SULPHUR (S) is taken up by plants largely as sulphate anion and to a lesser
extent as gaseous sulphur dioxide. These are reduced to organic sulphur
compounds such as cysteine, which is a constituent of many proteins.
Sulphur is lost by bacterial action on dead organic matter, because most of
the sulphur is converted to sulphide. Sulphides and sulphates can be inter-
converted by many species of bacteria. These organism are responsible for
the production of sulphuric acid to decrease the pH of soils. Sulphur of inor-
ganic bond is absorbed from the ileum. Important proportion is excreted by
the faeces in form of sulphate and by saliva, in form of rodanate. In the ani-
mal organism the approximately 0.15% sulphur is an important constituent
of methionine, cystine, thiamine, biotin, taurine, glutathione, mucopolysac-
charides, for example in the chondroitin sulphate of cartilage and in metal-
lothioneins. (The latter are proteins of small molecular weight in the ente-
rocytes, liver and kidneys, having great affinity to metals, like zinc, copper
and cadmium.) The sulphur content of the wool is 2.7 to 3.3% (up to 4%).
Generally there is sufficient sulphur in the forages, corn silage being an
exception; furthermore, in plants of crucifera its concentration is even high.
Monogastrics require sulphur in form of methionine and cystine, nev-
ertheless, inorganic sulphate has some sparing effect on these amino acids.
Sulphur requirement can be given in dry matter, which is for ruminants
approximately 0.2% of dry matter, above 0.4% it may be toxic, resulting in
polyencephalomalacia. Needs can be given related to the intake nitrogen.
For lactating dairy cow the desirable sulphur-to-nitrogen ratio is 1 to 10.
Lack of sulphur decreases milk production, because methionine is the first
limiting amino acid of the milk synthesis. Sheep requires a ratio of 1 to 7,
explained by the wool production.
Sulphur supply can be given by sulphur-rich plants, methionine-OH-
analogue, sodium or magnesium sulphate or even by sulphur powder.
Ruminant tissues are not able to synthesize methionine, thiamine and
biotin; therefore they have to get it either from the feed or rumen microbes.
Deficient S supply may limit production in sheep, when they are fed on diets
containing NPN and in such a situation is recommended sodium sulphate
supplementation, in an amount of 11-12% of the urea. Sulphur deficiency
not only decrease fleece production but also causes poor wool characteris-
tics, similar to the severe copper deficiency (lack in crimp). Sulphur powder
261
is one of the antidote of copper poisoning of ruminants, mostly sheep. As it

mentioned above, sulphur may combine with molybdenum in the rumen to
form tetrathiomolybdates, which reduce the absorption of copper.
Monogastric animals require the sulphur supply in form of methionine, cys-
tine and chondroitine sulphate.
SODIUM, POTASSIUM and CHLORINE. Occurrence and function of this
three elements is fully interwoven, therefore their presentation follows
together. Their main physiological function is the maintenance of osmotic
and acid base balance of the organism. Practically 100% of sodium and
potassium is absorbed from the rumen and small intestine, then secreted
in saliva and completely reabsorbed in the small and large intestine.
Consequently, their metabolism is closely/strictly linked to that of water.
Any fall in extracellular sodium concentration results in ACTH release,
which in turn, stimulates aldosteron production of the adrenal gland, help-
ing in sodium and water conservation. Extreme extracellular space, on the
contrary, releases the atrial natriuretic peptide from myocardial cells to
inhibit renal reabsorption of sodium. Sodium and potassium concentrations
establish the action potential of cell membranes. Chloride is a co-factor to
the functioning of α-amylase, and part of the gastric hydrochloric acid.
Sodium can almost be found in feedstuffs of animal origin, while
potassium is high in plants. Chloride concentration of feedstuffs falls
between the sodium and potassium. Top of sugar beet (with leaves) and
plants, grown at seaside, as well as in lick (on a saline soil) have a consid-
erable sodium content, too (Table XI-7). Alfalfa is especially poor in sodium
and the alfalfa silage or haylage, having lost plant extracellular fluid, even
poorer.
Table XI-7: Sodium content of the most important forages, g/kg DM (TOLGYESI, 1965)
*Sedge (Carex), bulrush (Scirpus, Typha), reed (Phragmites), bent-grass (Juncus)
Practically chloride is ingested in form of sodium and potassium chlo-

ride. Chloride ions are present in higher concentration in extracellular fluid
262
than in cell interior and tend to diffuse along concentration gradient.

Generally the chloride requirement is not calculated, because it is covered,
unless there is no sodium deficiency, realizing the supplementation in form
of common salt (NaCl). The hard working and intensively sweating horse
and the cow with displaced abomasum being the most important excep-
tions, when the chloride losses are higher, than that of sodium. The defi-
ciency of chloride may lead to alkalosis caused by an excess of bicarbonate,
causing the clinical signs of depression. In other cases recommendations
are given in common salt. The regular sodium chloride supplementation of
monogastric animals’ diet is 0.3 to 0.6% in air-dry matter. Poultry, espe-
cially young turkey is very susceptible to the salt poisoning, but the inci-
dence with the prohibition of the use of fishmeal in production animal feed-
ing is rare. Under stress condition (e.g. transport) the corticosteroids
increases potassium excretion, which in turn, if long-lasting, may cause
diarrhoea and dehydration.
In dairy cow the sodium supply may be the limiting factor of the milk
production. Every litre of cow milk contains 0.63 gram of sodium; to assure
it, cow should receive 1.7 gram common salt. The daily maintenance
requirement of cattle is 30 gram of common salt. The overwhole requirement
of the lactating dairy cow is 0.18% of sodium (0.46% of NaCl) in dry matter,
which generally is not covered by feeding only forages, therefore the sup-
plementation is indispensable. Salt intake is strongly regulated; conse-
quently the supplementation can be given by offering common salt in moult
or rock (licking) salt ad libitum. The daily common salt requirement of grow-
ing lamb is 6 to 7, that of the adult sheep 7 to14 grams. Given the regulat-
ed salt consumption, common salt is suitable to be the carrier of microele-
ments or medicines (“trace mineralized or medicated salt”). The sodium sta-
tus of lactating dairy cow can be evaluated both on the basis of saliva, urine
and faeces sodium concentration (Table XI-8).
Table XI-8: Monitoring of the sodium status of lactating dairy cow
263
An important sign of the sodium deficiency also the sodium-to-potassium

ratio in saliva, normally being approximately 20 and 0.5 to 1 in sodium-defi-
cient animals.
The majority of cattle feeds contain sufficient potassium, even for lac-
tating dairy cow during heat stress. Sodium-to-potassium ration influence
the function of ovaries. If the amount of potassium surpasses the tenfold of
sodium, the incidence of cystic ovaries increases. This can occur if the pas-
ture or cropland is irrigated with the manure and urine of cows and or
swine. Secretory diarrhoea, caused by bacterial toxin, leads to a depletion
of body potassium. In this cases, perorally applied potassium bicarbonate
may be helpful. In Quarter Horses an autosomal codominant genetic dis-
eases is known, causing disturbed sodium-potassium transport across the
skeletal muscle cell, resulting in hyperkalaemia and skeletal muscular
symptoms,
11.5. Micro- or trace elements

Their concentration in the animal body is below 150 ppm in the fat-free dry
matter, but their biological significance is immense.
IRON (FE). Approximately half the body iron can be found in the
haemoglobin and 3 to 7% in the myoglobin. There is some iron in the liver,
spleen and bone marrow, in form of the mobilizable water soluble
haemosiderin. Exchange rate of the iron content of the bone marrow is fast.
During absorption, iron first binds to a specific iron-binding receptor of the
brush border and transported into the enterocyte cytosol, where its trans-
port to the blood transferrin or being trapped by the enterocytes ferritin, is
regulated by the iron status of the organism. The transport protein of iron
in the blood is the transferrin. Except bleeding and other forms of blood
losses, practically there is no iron excretion. Besides the formation of
haemoglobin and myoglobin, the iron has manifold task in the organism. It
is part of the prosthetic groups of haemoproteids (in the citochroms of the
respiratory chain, peroxydase and katalase), as well as of different flavopro-
teids (e.g. succinyl dehydrogenase). The above compounds participate in
oxido-reductive processes.
Piglet and calf anaemia is the most typical deficiency syndrome of the
iron deficiency. Development of piglet anaemia is almost obligatory, because
the suckled sow milk cannot cover the increased needs of the intensive
264
breeds. Hepatic reserves of piglet are insufficient and in indoors-keeping

there are no adventitious sources, viz. soil and faeces of dam. Low iron con-
tent of sow milk means a certain protection against bacterial mastitis,
depriving pathogen bacteria from the essential iron. Wild piglets, in turn,
have a lower growth rate, furthermore they get supplementary iron either
through the dam’s faeces or soil intake. Ways of prevention are the applica-
tion of complex-bond iron injection, like iron dextran or by peroral dosage
of iron oxalate or fumarate. Parenteral iron supplementation may cause
immunosuppressant (iron toxicity (MÉZES et al. 1984). Some exotic birds
(mynahs and toucans) cannot regulate the iron absorption and considerable
amount of iron may accumulate in their liver, (haemochromatosis) causing
anorexia and lethargy.
ZINC (ZN). Most of the body zinc is part of metalloenzymes and acts
catalytically as a Lewis acid. Zinc stabilizes insulin and reinforces the link
between retina and uvea in the eye. In the mammalian organism more than
eighty zinc metalloenzyme and twenty zinc-activated enzymes have been
identified. In the zinc metalloenzymes zinc has a direct functional role, for
example in the carbonic-anhydrase, alcohol dehydrogenase, DNA poly-
merase, carboxypeptidase A, B and C, RNA polymerase and superoxyde dis-
mutase. Zinc is an essential component of the superoxyde dismutase
(ZnCu-SOD) enzyme. The cytosol form of the latter contains as a dimmer
one copper and one zinc ion. This enzyme is of paramount importance in
immune response, producing hydrogen peroxide and oxygen from the
hyperoxydes, indispensable for natural killer cell activity. Many hormone,
like testosterone, prolactin and somatomedin are zinc-dependent. Zinc is
crucial in the immune functions, because zinc is essential in the T-cell life
cycle (influencing apoptosis) and killer cell activity (see also Chapter XVII).
In the organism zinc is present in all of the body fluids; in the mam-
mals blood plasma contains 90 to 100 g/ml, bile 106 to 171, while in sem-
inal fluid 668 to 716 can be found. Females require approximately 20 to
30% less zinc supply than males, because their storage capacity is better
and there is no need for the seminal fluid production. Zinc deficiency of
males results in olygospermia. Zinc is absorbed from the small intestine, by
means of a large protein molecule, using active transport (see homeostatic
control of minerals). There is also a passive transport through proteins of
small molecular mass, like metallothionein of enterocytes. Ligand formation
may both inhibits (phytates) and stimulate (histidine) zinc absorption.
265
Presence of citrate, picolinic acid of milk and the essential fatty acids
enhance its absorption. Zinc is re-excreted into the gut lumen by bile and
pancreatic fluid (enteropancreatic circulation). The real storage capacity of
the organism (in the bones and liver) is small.
Cereal grains have a low (8 to 12 mg/kg), while brans and extracted
oilseeds a high (30 to 80 mg/kg) zinc concentration. Insufficient zinc sup-
ply decreases its tissue concentration and the activity of some zinc-depend-
ent enzymes, like alkali phosphatase, deoxy-timidin kinase, SOD, but this
is not general and inevitable.
The most common clinical sign is the parakeratosis, which can be
generalized (swine) or local (calf, some dog breeds). Parakeratosis means
lesions of the superficial layers of the epidermis with excessive keratinisa-
tion. As a consequence, skin becomes scaly and can crack owing to fissures;
slight pruritus (itching) may occur. In human parakeratosis is combined
with damages of gut epithel (parakeratosis entheropathica). Lack of zinc in
females decreases conception rate; in male impaired spermatogenesis devel-
ops with worsening sperm quality and preservation capability. Hair and
hoofs are also altered. Zinc concentration of hair reflects the status. In defi-
ciency the mucous membrane of the buccal cavity becomes hyperplastic
and the surface of taste bud also damaged. Consequently, troubles in taste
sensation (dysgeusia, hypogeusia), with or without altered smell detection
(dysosmia, hyposmia) can be found, combined with excessive salivation. In
the brain the altered endogen opioid distribution causes anorexia.
Sensitivity of rods in retina decreases. Lack of zinc decreases protein syn-
thesis and especially the methionine retention in tissues. Cross-chains of
collagen fibres weaken. There is a trouble in the metabolism of essential
fatty acids in testicles. Disturbed cholesterol metabolism stability of bio-
membranes decreases.
Ruminants’ zinc deficiency is characterized by local (muzzle, around
the eyes and joints) parakeratosis, excessive salivation, disturbed develop-
ment of hoof horn and bacterial infiltration of the mucous membrane in the
buccal cavity. In case of joint deficiency of zinc and phosphorus the carote-
neevitamin A transformation fails and the zinc deficiency decreases the syn-
thesis of vitamin A transport protein-(retinol binding protein=RBP). There is
genetically determined poor zinc absorption, describes in Black Pied and
Dutch-Friesian cattle. Calves develop scaly thickened skin over the neck
and shoulder, and similarly to the acrodermatitits of bull terriers, they are
266
susceptible to any infections. In pigs zinc deficiency results in generalized

parakeratosis, with erythema, seborrhoea, hyperkeratinisation of skin and
growth depression. Severe cases may cause thymus atrophy, testis degen-
eration, anorexia and farrowing troubles. In growing poultry the arthritis-
like swelling of the hock joint is typical, occasionally with ulceration of sole.
Rodents: the sexual cycle of hamsters get upset in lack of zinc. Rats begin
to practice coprophagy. Severe zinc deficiency is per se teratogenic (like lack
of biotin) in mice and rats. In guinea pigs the dermatitis is the leading clin-
ical sign. In dog and cat the zinc deficiency could be induced experimental-
ly. Naturally it is more common in dogs. In Beagles of marginal zinc status
occurs may happen that in the acute phase of infections even acute zinc
and secondary vitamin A deficiency develops, because the effect of lympho-
cyte endogen mediator (LEM) is so effective in directing zinc (and iron) from
the body fluid to the liver. In growing dogs of genetically predisposed breeds
(Malamut, Alaskian husky, and some German shepherd line), especially fed
on high calcium diet, hyperkeratosis develops all over the body: muzzle, on
the pads, on the interdigital and subaxillar skin. Besides anorexia and
growth depression and general weakness can be observed. Since the cause
basically a defect in zinc absorption, continuous supplementation is
required.
MANGANESE. The two main functions in the organism of manganese

are the cartilage formation (being part of the glucosyl transferase enzyme)
and the promotion of development and functioning of the ovaries.
Manganese is part of the Mn-dependent superoxyde dismutase. MnSOD
and CuZnSOD partly may replace each other’s function. The absorption of
manganese, especially from alkaline soils, is low (1 to 4%), the presence of
calcium further decreases its absorption. Monocotyledon plants (grasses
and “sauer” grasses) contain considerable amount of manganese; dicotyle-
dons (included the leguminous plants) have very low manganese concen-
tration (TÖLGYESI, 1969). Absorbed manganese is stored in bones, liver, pan-
creas and kidneys, but the capacity of organism for retention is limited.
Main routes of excretion are the bile and the large intestinal glands into the
gut lumen. Considering the very low absorption rate (no more than 4%), TÖL-
GYESI (1990) proposed to use as an endogenous indicator (see Chapter VII)
in digestibility trial.
Manganese deficiency in gilts results in anoestrus, in boars in dis-
267
turbed sperm production. Inappropriate supply of pregnant sows may cause

ataxia and bone deformities of newborn piglets. Heifers and cows on man-
ganese deficient diet may suffer in alimentary sterility, owing to ovarial trou-
bles, clinically they are slower to exhibit oestrus, or they have “silent heats”.
Deficient feeding of pregnant cows results in higher mortality of female foe-
tuses and the newborn; mostly bull calves had swollen joints from the trou-
bles of cartilage development. For controlling the manganese status, the
analysis of pigmented hair is applicable; the acceptable lower limit is 7 ppm.
Ovarian functions of queens also are very susceptible to the manganese
deficiency. In poultry the deficient feeding may lead to perosis and altered
egg hatchability.
COPPER. The absorption of copper is 5-10%, but excess sulphur and

molybdenum decrease its availability. The main site of absorption is the
small intestine, helping by the enterocyte copper-binding protein, a metal-
lothionein. . Ingestion of soil (Fe content) as leaf contaminant may partly be
responsible for seasonal drop in copper absorption. The majority of body
copper can be found in the liver, bones, muscles and skin. The most impor-
tant among them is the liver, which even in the foetus has an important
storage capacity. In the organism copper is bond to proteins. One of the lat-
ter is the blood’s coeruloplasmin, which catalyzing the oxidation of Fe2+,
making it suitable for the haemoglobin synthesis. This is the reason of the
phenomenon that copper deficiency leads to a secondary anaemia. Copper
is an essential part of at least ten major metalloenzymes, like cytochrome
oxydase, copper-dependent superoxyde dismutase (SOD) and polyphenyl-
oxydase. In copper deficiency the structure and pigmentation of hair and
wool are altered, there is a trouble in the development of foetal nervous sys-
tem, resulting in congenital ataxia, as well as on skeletal system deformities
and fragility can be found. These damages derived from the decreased activ-
ity of copper-dependent enzymes: polyphenyloxidase – pigment formation,
citochrom oxydase – respiratory chain, monoamino oxydase – in lysine
dezamination for the cross links of proteins in the conjunctive tissue.
Copper deficiency can be waited while feeding plants, grown in sandy
soil, or in case of inhibiting interaction, by calcium, molybdenum or/and
sulphur excess. On the contrary, in soils, low in molybdenum and/or sul-
phur, the copper concentration of plant may increase in a dangerous extent.
As mentioned, dicotyledons contain more copper than monocotyledons. The
268
optimal botanical composition of meadow hay would be two third grasses

and one third dicotyledons (TÖLGYESI, 1969). If the ratio of dicotyledon is
higher, the danger of manganese, if lower, the possibility of copper defi-
ciency occurs.
The earliest sings of sheep copper deficiency is the loss of wool crimp
(steely wool). Sheep may exhibit swayback or enzootic ataxia, caused by a
impaired myelin synthesis in the nervous system. Other signs of ruminants’
copper deficiency are the anaemia, diarrhoea, depigmentation of hair or
wool, developing unpigmented circles around the eyes, congenital ataxia of
lambs, kids and calves, fertility problems in heifers and cows, occasionally
nymphomania. Copper concentration in blood and liver, as well as the
coeruloplasmin level in blood decreases. The recommended supplementa-
tion is 2 gram copper sulphate per cattle, or chelated injection monthly. The
best indicator of copper status is the hair; the acceptable lower limit is 6
ppm. Low (less than 0.3 to 0.5 IU/mg haemoglobin) SOD activity of red
blood cells indicates a prolonged deficiency.
In the Wilson-disease of man there is a copper deposition in the
liver, brain, kidneys and eyes, causing severe local damage. The cause is the
inherited lack of a transport protein capable of eliminating copper, fixed by
hepatic metallothionein. It has been described in some dog breeds (e.g.
Beddlington terrier), too. The Menkes syndrome is a sex-linked, inherited
trouble of the copper metabolism, while the copper intake of cells is dis-
turbed. In sheep chronic copper poisoning develop after long-term inges-
tion of feeds of higher copper content (e.g. growing pig feed with the copper
sulphate additive). When the copper quantity in the liver surpasses the stor-
age capacity of metallothionein molecules, the copper gets into the periph-
eral blood circulation, causing haemolysis, jaundice and occasionally death.
FLUORINE. Free fluorine is rare in the nature, but fluoride can be found
in several chemical compounds. Fluorine is a constant and natural compo-
nent of bones and teeth. Trace quantities of ingested fluoride minimizes the
incidence of caries and the senile bone demineralization. Fluoride absorp-
tion basically is a passive process, beginning in the stomach (in ruminants
in the rumen) and ended in the small intestine. Rate and extent of absorp-
tion depend on the source of fluoride, for example from the sodium fluoride
the absorption is more than 90%, that from phosphates approximately only
50%.
269
Three third of the blood fluoride is in the plasma, where its concen-
tration is basically regulated by kidneys and skeletal system. Continuous
ingestion even of small quantities of fluoride causes its accumulation in
teeth and bones, which manifest in morphological damage only above a
threshold concentration. In dairy cow only a small portion is excreted by the
milk. On the contrary, the placenta does not mean a real barrier; thereby
the fluoride concentration of the bones in the newborn calf is proportional
to that of the dam’s blood plasma. Notwithstanding, the calf health is dan-
gered neither by the placental transport nor the milk intake. The same is
true for milk from the point of view of human health concerns.
Phosphorus rock mostly contains fluorapatite (Ca10F2[PO4]6}, which
is the raw material of the production of phosphorus fertilizers and phos-
phorus feed supplements. With the exception of industrial pollution, the
chronic fluorosis of man and animal is seldom caused by the food/feed,
because the intake capacity of plants (except the tea and camellia) is limit-
ed. Plants in the pasture have a F content of 2 to 16, the cereal grains 1 to
3 mg/kg DM. The highest F level of bone ash cannot be more than 1.5
mg/kg DM. The main sources of the animals fluorine intake is the phos-
phorus supplement. Since fluorine overdosage may endanger animal
health, the fluorine concentration is regulated both in the phosphorus sup-
plements and in the final feed mixture (see Chapter II). Excess fluorine may
accumulate in skeletal system, causing bone weakness, lameness and mot-
tled, eroded and stained teeth.
IODINE (I). High physiological significance of iodine is given by its con-

tribution to the formation of thyroid hormones. The iodine of the seas has
an important role in natural cycle. The iodine concentration of freshwaters
follows the iodine content of soils. Regions with iodine deficiency have less
than 2 μg/litre iodine. Iodine level of European drinking water shows a
great deviation (10 to 50 μg/litre). The concentration of 3 μg/litre of iodine
is considered as optimal. The iodine content of forages basically depends
upon the iodine concentration of soil and generally it averages in 1 mg/kg
DM. On the contrary, feed plants from iodine deficient regions have gener-
ally only approximately 10 μg/kg DM.
In monogastric animals iodine is absorbed from the small intestine
and it is excreted by the urine, saliva and milk. In ruminants iodine is
absorbed from the rumen and reticulum, but is re-excreted into the abo-
270
masum and finally from the small intestine almost the whole amount of
ingested iodine gets into the blood. The majority of blood iodine is taken up
by the thyroid gland, the remaining part distributes in the body fluids,
included the milk. Two third of the total body iodine content (15 to 2ö mg)
can be found in the thyroid gland. The normal iodine concentration of milk
(5 ?g/l), according to the iodine content of the ingested feed and drinking
water, shows a considerable fluctuation (5 to 694 μg/l), thereby it is suit-
able for controlling the supply of animals.
In severe iodine deficiency the foetal and postnatal growth and devel-
opment pathologically slow down. As a consequence, goitre, cretinism???,
reproduction troubles and foetal and postnatal mortality can be observed,
summarizing called IDD (iodine deficit disorders). Iodine deficiency may be
aggravated by the presence of thyreostatic (goitrogenic) substances, like
cations of the hard water, the nitrite and nitrate content of the water (KAC-
SKOVICS, 1985) and the antinutritional factors of the crucifera plants (see
Chapter VIII), in addition the arsenic and the rubidium (secondary alimen-
tary iodine deficiency). In prematurely born mammals the iodine regulating
mechanism is immature, thus the newborn requires higher iodine intake.
Iodine deficiency of production animals is characterized by abor-
tion, stillbirth, congenital goitre and lack of viability and fertility problems
(low conception rate, retained placenta) in dams. The most typical is the
birth of newborn lambs, calves and piglets with congenital goitre and myx-
oedema (BOKORI, 1958). The histological findings of the thyroid gland are
abnormal. Colloid depletion, cell proliferation and cell enlargement, fol-
lowed by invagination and finally collapse and infiltration of follicles. In case
of iodine deficiency goitre develops in fish, too. Iodine requirement of for
non-lactating cattle 0.33 mg/kg dietary DM and 0.45 mg iodine/kg DM for
the lactating dairy cow (NRC, 2001). Inorganic iodine sources (iodide of cal-
cium, potassium and sodium) are unstable; therefore in licking salt even the
pentacalcium orthoperiodate and the ethylenediamine dihydroiodine (EDDI)
are incorporated. The daily intake of 2.5 to 3.0 gram of iodine by dry cows
may prove harmful for the foetus. Besides the milk, the urinary iodine con-
centration is also suitable for controlling the status, using the iodine to cre-
atinine value. Iodine poisoning. Sheep are less susceptible to the iodine
toxicity than cattle. Horses are extremely susceptible to iodine toxicity.
Foals of mares, fed on high iodine concentration diet during pregnancy,
large goitre may develop. Foals generally die before birth of shortly there-
271
after. As clinical signs, the limb abnormalities, long hair and general weak-
ness are present. Excess iodine intake may cause the feline hyperthy-
roidismus in cat, characterized by an extremely high blood T3 and T4 level.
SELENIUM. The highest selenium concentration in the animal organism

can be found in the liver and kidneys; the extremely high, toxic intake accu-
mulates mostly in hair and horny formations, substituting the sulphur in
the proteins. A small portion is exhaled, giving the breath garlic smell. As
active part of the glutathione peroxydase enzyme (GSH-Px), the main phys-
iological role of the selenium is the defence of biomembranes in the organ-
ism. Its effect in protecting double bonds of biomembranes, is supplement-
ed by the action of vitamin E and in some extent, by the carotenoids and
ascorbic acid and riboflavin. Many Se-dependent deiodinase enzymes has
been identified, therefore, selenium indirectly affect basic metabolic rate
and a series of processes influenced by active thyroid hormones.
The lipidperoxidation caused cell damages have different clinical and
pathological manifestation according to the species and age of the animal.
Italic printing outlines syndromes, which can be prevented by giving only
selenium; the other should be treated with both selenium and vitamin E,
and some can be cured only by using vitamin E.
I. REPRODUCTION TROUBLES: embryo degeneration (rat, hen, turkey, cow,
ewe, damage in embryonic vascular system), sterility (male rat, guinea pig,
golden hamster, dog, cock), decrease in egg production and hatchability
(hen).
II. MULTIPLE-ORGAN AILMENTS: liver, blood, brain, capillaries, pancreas:
liver necrosis (rat, swine, beef cattle), pancreas fibrosis (chicken, mouse),
haemolysis of red blood cells (rat, chicken, newborn human baby), decrease
in serum albumin concentration (poultry), anaemia (monkey), encephaloma-
lacia (chicken), exsudathiv diathesis (chicken, turkey), degeneration of kid-
ney tubuli (rat, mouse, monkey, mink), ceroid steatitis in fat tissue (mink,
pig, chicken), depigmentation of incisors (rat), loss of hair or feather (rat,
horse, pig, chicken). The pancreas atrophy of young chickens finally leads
to pancreatic fibrosis, with subnormal production of digestive enzymes.
Exsudathiv diathesis of growing poultry is a generalized oedema of breast,
wing and neck area, caused by an increased permeability of capillaries.
Fluids under the ventral skin give greenish-blue discolouration.
III. ALIMENTARY MYOPHATHIES: Type A myopathy (rabbit, guinea pig,
272
monkey, duck, mouse, and mink), Type B (cardio) myopathy (lamb, calf, kid,
and pig), Type C cardio- and gizzard myopathy (turkey) and Type D myopa-
thy (chicken).
However, the selenium and essential fatty acids cannot replace vita-
min E in its immunostimulant effect. Lack of selenium in pregnant ewe diet
will cause the typical congenital “white muscle disease” or nutritional
muscular dystrophy of lambs. Denomination derives from the white stria-
tion in the muscles, especially those in the thigh and shoulder. Not only are
the skeletal, but also cardiac and respiratory muscles affected. Heart mus-
cle typically contains white formations. Abnormalities in the electrocardio-
gram develop early. The basic lesions are hyaline degeneration followed by
coagulative necrosis and calcification. Individual muscle fibres may be
oedematous and swollen, with loss of cross striations and haemorrhage is
often present. The VESD (Vitamin E Selenium Deficiency) syndrome occurs
in pigs, including myopathy of the hind legs and m. longissimus dorsi, liver
necrosis (hepatosis dietetica), multberry heart disease (bleedings, haemor-
rhage and necrotic lesions in heart muscle, showing red mottled appear-
ance) and occasionally oesophageal gastric ulcer. In dairy cow the selenium
and/or vitamin E deficiency increases the incidence of non-infective reten-
tion of foetal membranes (retained placenta) and the number of mastitis.
Large dosage of sodium selenite injection at the end of dry period may help
in preventing the mentioned troubles. For monitoring the cow’s selenium
status, both the gluthathion peroxydase activity of the red blood cells and
the selenium content of the pigmented hair (0.25 ppm being the critical
lower limit) are suitable.
Selenium toxicity. Overdosage of selenium (more than 2 ppm) or
ingestion of seleniferous plant on the pasture may cause poisoning; the clin-
ical signs in ruminants (and rarely in horses) are erosions of the joints of
long bones, liver cirrhosis, anaemia and ascites (“alkali disease”). Ingestion
of seleniferous plants for months results in the chronic “blind staggers dis-
ease”, frequently leading to death. For the pig’s selenium toxicity may cause
on the coronet of hoof clinical signs, similar to the mouth-and foot disease.
Growing chicken rather resistant to selenium poisoning, but 8 pp in the diet
has already reduced growth rate. In laying hens, feeding of diet of more than
5 ppm selenium results in decreased hatchability of eggs and malformations
of the embryo.
273
COBALT is the main component of the vitamin B12. or cyanocobalamin,

which cannot be found in plant feedstuffs. The only known physiological
role of cobalt in mammals is as a constituent of vitamin B12, which in turn
is the coenzyme of two key enzymes of metabolism. If the cobalt supply is
sufficient, well-functioning rumen microflora may provide ruminants with
vitamin B12; in this case they do not require exogen intake. For the efficient
vitamin B12. synthesis the rumen fluid should contain at least 20 μg/l
cobalt. In the caecum of herbivorous and omnivorous monogastric animals
the bacterial flora synthesizes the vitamin, too. Nevertheless, except the cae-
cotrophy of rabbit, the utilization is low. Lack of cobalt results in the “ovine
white liver syndrome” (OWLS).
NICKEL. Notwithstanding the fact that the nickel is an essential ele-

ment, the incidence of deficiency syndromes in average feeding conditions
rather rare. The observed signs of manifested nickel deficiency are the
decrease of the haematocrit value, damage of the endoplasmatic reticulum
in the liver, reproduction troubles and high perinatal mortality. Enzyme
functions, like pancreatic amylase, bacterial urease are also altered.
Described clinical signed have been provoked by special nickel depletion
diet, thus in recommendations for the practice generally there are no
requirement values.
However, the nickel is a prosthetic part of several metalloenzymes in
ruminal and intestinal bacteria. Consequently, the intake of supplementary
nickel may improve the function of these bacteria, which can have some
advantage for the host animal, too (ReGIUSNÉ, 1991).
CHROMIUM. The trivalent form of chromium is part of the glucose tol-

erance factor (GTF), which in turn, is indispensable for the proper func-
tioning of insulin receptors. GTF, in the presence of sufficient insulin, help
the glucose uptake of cells. Its deficiency may cause hyperglycaemia.
Organic chromium (in form of chromium picolinate and nicotinate) is sug-
gested to modulate immune response (MOWAT, 1994). Its application
increased lean meat percentage in growing-fattening pigs. On the other
hand, the inorganic salts of chromium are carcinogenic and toxic.
MOLYBDENUM is an integrated part of the xantin oxydase enzyme,

indispensable in the nucleic acid and peroxyde metabolism. Besides the
274
xantin-oxydase, it can be found in many plant and animal enzymes (alde-

hid-oxydase, sulphit-oxydase), and the deficient intake may lead to dam-
ages both in plants and animals. Salts of molybdenum stimulate the cellu-
lose activity of rumen bacteria. Owing to the dependence of soil conditions
and to several interactions (see copper, sulphur etc.), the molybdenum con-
tent of plant varies within wide ranges. It is a mobile element; its concen-
tration even in the generative parts of plants may change/differ by several
magnitudes (KÁDÁR and FEKETE, 1992) according to the soil Mo concentra-
tion. Consequently, plants grown on peat soils may contain high amount of
molybdenum and their ingestion results in chronic diarrhoea. Toxicity: the
maximal tolerable dietary concentration for cattle is estimated to be 10
mg/kg diet (NRC, 2001).
SILICON is present as structural element in the mucopolysaccharides,

establishing cross-chains between the polypeptide and polysaccharide
chains. Consequently, most of the silicon in the animal organism can be
found in the epithelial and conjunctive tissue, as well as in the growth
plate/zone of bones. Lack of silicon may cause disturbance in skeletal
growth. Inhalation of silica-containing mineral dust induces silicosis as an
occupational diseases.
OTHER MICROELEMENTS. Tin (Sn). Its exact function is unknown, but

the essentiality is proved in rat model. Vanadium (V) is required for opti-
mum growth of rats and calves in a concentration of 0.1 ppm. In some
invertebrate animals, like tunicates, the protein-bond vanadium con-
tributes in the oxygen transport as part of the green blood cells, called
vanadocytes. In mammals may help the incorporation of phosphorus into
bone. Rock phosphate may have a high vanadium concentration, capable to
affect the quality of white of egg (albumen). Arsenic (As). Essentiality was
first demonstrated in rats. Animals fed on a very low (0.03 mg/kg DM)
arsenic diet, the growth rate and haematocrit value decreased an
splenomegaly developed. ANKE(1977) kept female dwarf goat on semisyn-
thetic arsenic deficient diet and reported about reproduction troubles and
damages in foetal development. In higher concentration arsenic is a defi-
nitely toxic element. Very severe deficiency of lead (Pb), developed by feed-
ing carefully purified diet, may result in anaemia, not modifying the very
toxic effect of higher lead intake. Boron (B) is a rare element, in the Earth
275
crust its concentration is only 3 mg/kg. In poultry, independently from the

vitamin D, may help the maturation of chondrocytes. In women of post-
menopausal period there is an oestrogen-like effect of boron compounds.
FOR FURTHER READING
Borges, S.A., Fischer da Silva, A.V., Ariki, J., Hooge, D.M. and Cummings, K.R. (2003):
Dietary Electrolyte Balance for Broiler Chickens Exposed to Thermoneutral or Heat
Stress Environment. Poult. Sci. 82, 428-435.
Bowen, H.J.M. (1966): Trace elements in biochemistry. Academic Press. London and New
York
Keshavarz, K. (1994): Laying hens respond differently to high dietary levels of phosphorus
in monobasic and dibasic calcium phosphate. Poult. Sci. 73, 687-703.
Leith, D.E. (1991): The new acid-base: power and simplicity, in Proc. Am. Coll. Vet. Intern.
Med. 611-617.
Marek, J. and Wellmann, O. (1932): Die Rachitis. Biochemischer Teil [Rickets. Biochemical
part – in German] Fischer Verlag, Jena
NRC: Mineral tolerance of animals. Second rev. Ed. The National Academies Press.
Washington, D.C., www.nap.edu
Underwood, E.J. ( 1981 and later editions): The mineral nutrition of livestock. CAB.
Farnham Royal. Slough, England
Tolgyesi, G. (1969): Trace element content of plants and its agricultural respects (in
Hungarian) Mezogazda Kiado. Budapest
276
Chapter XII
VITAMINS IN FEEDING AND NUTRTITION
12.1. Generally about vitamins

12.1.1. Relationship of nutrition and diseases
12.1.2. How many vitamins should be given ?
12.1.3. Stability of vitamins
12.1.4. Relationship of vitamin requirements with the
function of rumen and intestinal flora
12.1.5. How to express the vitamin requirement
12.1.6. Methods for monitoring the vitamin status
12.1.7. Recently discovered physiological role of some
vitamins
12.2. Overview of the nutritional significance

of vitamins
12.2.1. Fat- soluble vitamins
12.2.2. Water soluble vitamins
Chapter XII: VITAMINS - FUNCTION- DEFICIENCY
12.1. Generally about vitamins
12.1.1. Relationship of nutrition and diseases

There is a lack of satisfactory information concerning the relationship of
nutritional status and susceptibility to infectious diseases in domestic ani-
mals. According to the generally accepted opinion the overfeeding makes
organism susceptible to viral infections. On the contrary, good body condi-
tion has many advantages against bacterial and parasitical infections. It
worth also mentioning that the related data rather derive from epidemiolog-
ic survey than from designed animal trials. Nevertheless, results of well-
planed trials show that protein deficiency weakens the immune response
against virus, bacterial and fungal infections. Similarly, it is known that
mineral and vitamin deficiency (e.g. selenium, vitamin A and E) can be
linked with a decreased general resistance to diseases. In dogs, the effect of
obesity in impairing immune status has also been demonstrated. The genet-
ic and environmental (basically: nutritional) determination of some meta-
bolic diseases can be well interpreted only parallel to each others.
In case of haemophilia and progeria the generic determination is of
100% and their manifestation cannot be influenced by feeding. People with
phenylketonuria, the production of phenylalanine hydroxilase, indispensa-
ble in the tyrosine synthesis, fail. By balancing the phenylalanine-tyrosine
intake, the status of concerned individuals can be improved. Pernicious
anaemia and osteoporosis are mostly of genetic determination. However, by
adapting the diet (vitamin B12, calcium etc.) the signs of clinical manifesta-
tion can be alleviated. The other extremes are the protein-energy malnutri-
tion (PEM) or the scurvy, which can be totally controlled by the feeding.
Atherosclerosis and Parkinson’s disease are decisively of genetic, on the
contrary, pellagra and rickets of nutritional determination, but the role of
278
other factor cannot be underestimated.
12.1.2. How many vitamins should be given ?

The recommended vitamin concentrations in feed (or the daily allowances)
depend largely on the aim of vitamin supplementation (FIGURE XII-1).
FIGURE XII-1: Factors influencing vitamin recommendations (after La Roche, modified)
Analysing FIGURE XII-2, the lowest recommended vitamin concen-

tration is able to prevent clinically manifestation of deficiency syndromes.
The prevention of subclinical signs requires a little more. Optimum vitamin
recommendation can never be one value, but a range. Within the range, a
diverse series of different factors will determine the actual allowances, (sym-
bolizing by the arrow) included the genotype of animal, its keeping and hus-
bandry, the incidentally present infectious and parasitic diseases, stress
status, environmental temperature and the related voluntary feed intake
and the goal of supplementation. Dosages above the optimum range may be
advantageous (e.g. immunostimulant effect of vitamin C and E) and disad-
vantageous, causing hypervitaminosis, even poisoning. Production and bio-
logical goal-oriented/determined vitamin requirements can clearly be illus-
trated on the example of vitamin E (FIGURE XII-2).
279
FIGURE XII-2: Vitamin E recommendations according to the expected biological effect
The first, basic recommendation aimed the prevention of clinical

signs, like myodegeneration etc. If the feed mixture contains more lipids
(carnivore diet, increased energy density in poultry), the requirement for the
natural antioxidant, the vitamin E also increases. The above described can
be clearly explained by the complex defence mechanism of the organism
against the peroxide loading. This principle partly remains valid even after
slaughtering and to develop good storage capability of carcass by avoiding
the post mortem rancidity of carcass fat, the vitamin E need are even high-
er. In order to achieve the immunostimulant effectiveness of tocopherols, at
least tenfold of the basic requirement should be given.
12.1.3. Stability of vitamins
The stability of vitamins depends on a large scale on the water content of
environment, or to be more precise, on the value of water activity (for details
see Chapter XIV). FIGURE XII-3 demonstrates the degradation of ascorbic
acid in orange juice and wheat flour as a function of water activity.
280
FIGURE XII-3: Degradation of ascorbic acid in function of the water activity of the envi-
ronment
The of stability of the individual vitamins depend on the different pH

conditions, the influence of air, oxygen and light. Data are related to pure
chemical substances, or to commercial vitamin preparations actually.
According to the used production technology, the 2 values maybe substan-
tially different. For example, β-carotene and ascorbic acids are very sensi-
tive to the oxygen and light, but after microcapsulation (coating) by ethyl-
cellulose or gelatine (both), or by presenting in a stable, but biologically
available chemical compound (L-ascorbyl-2-popyphosphate) they are quite
resistant.
Some example the ascorbic acid is so sensitive, that only in acidic
media can remain undegraded. Vitamin B6 shows a high resistance against
many destroying factors. (It worth mentioning that related data of scientific
literature are many times contradictory, the present compilation used the
most commonly accepted sources.) While preparing diets for laboratory ani-
mals and before feeding a heat treatment is necessary, the destructive effect
of sterilization should be taken into account and sensitive vitamins should
be mixed in a higher concentration.
12.1.4. Relationship of vitamin requirements with the function of

rumen and intestinal flora
As a general rule can be stated that the more developed and complex the
ruminal/intestinal microflora is, the more the host animal is independent
from the exogenous vitamin intake. Bacteria are able to synthesise a series
281
of vitamins (vitamin K, vitamin B group etc.), which is totally available for

the adult ruminants and for the rabbit (caecotrophy). Nevertheless, changes
of rumen pH (high-grain diet) or use of peroral antibiotics may significantly
alter vitamin-synthesizing ability of bacteria.
12.1.5. How to express the vitamin requirement

In case of ruminants and companion animals it is possibly to give the indi-
vidual daily need of an average size animal or related to kg LW, but with the
use of TMR (total mixed ratio) the recommendation can be given on dry mat-
ter basis, too. For monogastrics, included preruminant calf, the most fre-
quently the recommendations are given in air-dry matter basis. Scientific
vitamin allowances (NRC, ARC, DLG, INRA, CODEX FABULARIS HUNGAR-
ICUS) generally give the minimum requirement, sufficient to prevent clini-
cal deficiency syndromes with safety. Commercial companies (Hoffman-La
Roche, Bayer, BASF, Rhone-Poulenc etc.) for all domestic animal species
and production categories and the AAFCO (American Association of Feed
Chemistry Officials) for dogs and cats give higher values, an assumed opti-
mum level.
12.1.6. Methods for monitoring the vitamin status
It can be lain down as an absolute rule that single measurement of vitamin

concentrations in blood and/or tissue is not sensitive enough to detect a
slight deficiency. More reliable results can be collected by measuring the
changes of vitamin concentration in body storage organ after the application
of a known amount of vitamin. For example the RDR (relative dose
response), in case of the vitamin A, when blood concentration is measured
before and after vitamin A application. Changes in blood level are related to
the vitamin A content, the saturation of the liver. In other cases the activi-
ty of enzyme is measured, which is dependent on the presence of the given
vitamin (e.g. glutathione reductase activity for riboflavin) or is related to the
activity of the measured enzyme activity (e.g. vitamin E and superoxide
dimutase). The special physiological function can also be a good indicator of
supply, for example the prothrombine time (see blood clotting) for vitamin
K. In many cases the parallel use of several methods is necessary. The par-
ticular methods by vitamins are given in 14.2. Table XII-1 present an exam-
ple for the diagnossis (NRC, 1986)
282
283
12.1.7. Recently discovered physiological role of some vitamins
In case of many vitamins turned out that behind the strictly spoken and
already known function, in higher dosages, they have special pharmacolog-
ical or preventive effects. For example vitamin E and certain carotinoids
including β-carotene were capable of stimulating cellular immune function.
Plasma tocopherol level has been shown to be inversely related to gastroin-
testinal cancer risks. Ascorbic acid, given in huge amounts is a nitrosation
inhibitor, because reduced formation of nitrosamines, which are potential-
ly carcinogenic. From the wide diversity of examples, the immunostimulant
effect of carotinoids is presented in more details.
Role of immune response basically is implied in the protection against
infectious agents and foreign protein as well as against tumour formation.
GREEN and MELLANDY have already found in 1930 that in vitamin A deficient
rats large amount of β-carotene prevented the infections of ear, urinary
bladder and intestine. The necessary carotene dosage was higher than that
of optimal growth. CLAUSEN (1931) observed carotene ingestion facilitate
recovery in children of infectious inner ear inflammation. At that time, it
could not exclude that part of carotene effect derives from its vitamin A pro-
vision. Researches of the since then elapsed period of time proved that
carotinoids of nine or more double bonds are able to neutralize reactive free
radicals and by this mean, they have a vitamin A independent immunos-
timulant effect. Anyhow, in comparison with carotinoids, the antioxidant
effect of the vitamin A in itself is negligible.
During the immune response (especially in its beginning phase) some
groups of white blood cells destroy the intruder bacteria by means of reac-
tive free radicals, like peroxides and nitrogen monoxide. Although in the
above mentioned function the free radicals are indispensable, in other time
and in other place their presence in tissues and blood may be dangerous
and damaging. Namely, if during immune response the amount of released
free radicals is overwhelming or they are not neutralized properly, the white
blood cells and the neighbouring tissues may be damaged. Neutrophyl gran-
ulocytes participate in the struggle against bacteria. If neutrophyl granulo-
cytes are incubated with bacteria in the presence of β-carotene, their killer
activity is efficient and they are protected against free radicals. In lack of β-
carotene granulocytes may be damaged by their own oxidative products.
The released free radicals may cause chromosome injury, too.
284
Cells of specific immune response are also susceptible to the lipid-

peroxides. It has been known since more than twenty years that both vita-
min A and carotene intake is capable of preventing stress-induced thymus
atrophy. Ingestion of both vitamins increased the number of lymphocytes in
the circulation and stimulated the rejection of grafts in mice. Moreover,
mice thymocytes proved to be more resistant against radioactive radiation,
if the β-carotene and vitamin A supply was abundant; the cytotoxic T-cells
showed intensive proliferation.
Antigens are presented to T- and B-lymphocytes by macrophages. The
recognition of antigen takes place on the macrophage’s membrane; during
this process macrophages produce several chemical mediators (interferon,
tumour necrosis factor, prostaglandins, and interleukin-1). Oxidative
metabolites of inflammative events may damage the membrane receptors of
the macrophages. Application of β-carotene or canthaxantine was able to
counterbalance this harmful effect. Role of T-helper cells includes the facil-
itating of secretion (specific antibody production) by B-lymphocytes. Two-
week-long ingestion of high dosage of β-carotene (180 mg/day vs. 5 mg/day)
significantly increase the number of T-helper cells in human.
To elucidate the independent carotinoid effect from the role as a vita-
min A precursor, rats were fed on β-carotene and canthaxantine containing
diets. The latter compound is effective against the free radicals, but cannot
serve as a vitamin A precursor. At the end of the 20-week long trial, it could
be stated, that on mitogen stimulation by concanavalin, phytohaemagglu-
tinin A and lipopolisaccharide, the T- and B-cells of carotinoid-pretreated
animals reacted more intensively (BENDICH and SAPIRO, 1986).
As above already mentioned above, a series of evidences indicate that
the ingested carotinoids decrease the prevalence of different tumours.
Comparable epidemiological investigations involving vitamin A, have not
found a similar relationship. Three cell types of the immune system take
part in destroying tumour cells: macrophages, the natural killer cells (NK)
and cytotoxic T-cells. Both α- and β-carotene were capable to increase the
activity of these components of the immune defence in cell culture. It was
demonstrated in golden hamster, that the production of tumour necrosis
factor (TNF) may be enhanced by feeding a-, β-carotene or canthaxanthine
TOMITA et al. (1987) fed mice, injected by tumour cells, on placebo or
120 β-carotene daily. From the lymph nodes of both groups lymphocytes
have been taken, mixed with fresh tumour cells and injected into healthy
285
mice. After the application of both “cell mixture”, liver tumour developed in
the healthy mice. However, the size of tumours, caused by the “cell mix-
ture”, containing the tumour cells and the carotene-pre-treated lympho-
cytes, proved to be tenfold smaller, than in the other group, with the ploace-
bo-pretreated lymphocytes. Thus, the previous β-carotene feeding enhanced
the tumour cell destroying capability of lymphocytes. Later the trial has
been repeated using canthaxanthine and asthaxanthine, as well as algae
extract instead of the β-carotene, receiving similar results, even the meas-
ured citotoxic activity of T-cells increased, too. These data proved that the
effect can be attributed to the carotinoids and not to the vitamin A precur-
sor role.
Anticancerogenic effect of carotinoids can be explained by the bind-
ing/neutralization of the reactive free radicals. Carotinoids are able to pro-
tect cell membranes and DNA against the damaging effect of free radicals,
help in maintain membrane fluidity and the integrity of membrane recep-
tors. At the same time, they stimulate the production of immunomodulant
prostaglandins and leukotriens, as well as that of the tumour necrosis fac-
tor. Moreover, dietary carotinoids increase the activity of cytotoxic T-cells,
macrophages and natural killer cells against emerging tumour cells.
12.2. Overview of the nutritional significance of vitamins
12.2.1. Fat- soluble vitamins
VITAMIN A (retinol, axeroftol) fulfils its main physiological function in the

mechanism of sight, in the bone formation and in the protection of epithels,
mucous membranes, skin and germ epithel. In some way, vitamin A is a
precursor to a group of hormones, the retinoic acids. Once it is converted to
one of the retinoic acids, it will affect gene expression. Namely, four nuclear
retinoic acid receptor proteins have been identified in vitamin A target tis-
sues. After having bound to the retinoic acid receptor, it initiates transcrip-
tion and translation of certain genes, influencing thyroid hormone and vita-
min D receptor proteins, as well as the leptin gene expression (see obesity).
In considerable quantities it can be found in cod-liver oil, viscera
(especially in liver), in egg yolk and in milk. Its deficient intake in poultry
may cause a keratinisation of mouth and oesophageal mucosa, with accu-
mulation of dead cells around gland ducts, which should be differentiate
286
from some forms of Fowl Pox. Risk of secondary infection of the affected
areas increases. Parallel to the above described, or independently, visceral
gout develops with tophus formation and urate crystal deposition in the
abdominal cavity, underneath the pleura and on the surface of kidney and
heart. It should be differentiate from the viral avian nephritis (MÁNDOKI et al.
2005). Lack in dog causes dermatitis. Deficient supply of pregnant sows
results in birth of newborn piglets with hydrocephalus, malformation of the
eye, mouth and cleft palate. It is worth mentioning that vitamin A hypervi-
taminosis during the gestation may also lead to the malformation of off-
spring. In calves the disturbed reabsorption of the cerebrospinal fluid leads
to a papiloedema, which in turn causes blindness. Cerebrospinal fluid (CSF)
is continuously produced by the numerous microvilli of choroid plexus. CSF
normally is absorbed through arachnoid villi, small nodules that extend
from the arachnoid into cranial venous sinuses as function as one-way
valves. In hypervitaminosis, the arachnoid villi become clogged with fibrob-
last and are unable to release the fluid. On the cornea ulcus may develop.
Slight vitamin A deficiency causes the night blindness (nyctalopia) by defi-
cient building of rhodopsin. Typical deficiency syndrome of birds of prey
(typically in captivity) the prolepsis of the third eye lid. Vitamin A deficien-
cy of fish manifests in ascites, exophthalmia and haemorrhages in the kid-
ney.
One IU vitamin A equals to 0.344 µg crystalline vitamin A acetate.
Requirement of monogastric animals is 2,000 to 10,000 IU/kg feed on air-
dry matter. Lactating cows requires 110 IU/kg LW (NRC, 2001). Vitamin A
is sensitive to oxygen, light and acids, the lipoxygenase of some leguminous
seed (e.g. soybean) may degrade it. To control the animal’s status, the meas-
urement of its concentration is one of the choices. Since the differences in
vitamin A concentration of the individual liver lobes significantly differ, it
can be recommended rather in cadaver, during necropsy. The relative dose
response (RDR) much more appropriate procedure makes possible a much
more accurate evaluation. It include the measurement of the basal vitamin
A concentration of blood, then after giving standard quantity of vitamin A
injection the changed blood concentration should be analysed. The higher
is the blood concentration after the vitamin A injection; the better is the
vitamin A supply of the organism, because the storages (liver) are saturat-
ed.
Overdosage (10 to 1000 times) of the recommendation is harmful,
287
even toxic. Too much liver ingestion may cause vitamin A toxicity (typically
in cats). Clinical sings include exostosis on cervical vertebrae and limbs,
general disturbance in skeletal development and increased pressure of cere-
brospinal fluid. In poultry, the vitamin A overdosage causes growth depres-
sion and fatty liver. In mammals, included human, the higher vitamin
intake may prove teratogenic. A special form a vitamin A toxicity is the
hyena disease (mostly combined with adeno virus infection), when the
administration of huge amount of vitamin A for calves may induce prema-
ture closure of the physes, predominantly in the hind limbs. The manifes-
tation is that the skeletal growth will disturbed and the hind legs are spec-
tacularly shorter than the front legs.
β-CAROTENE is the previtamin of vitamin A of molecular weight of 537.
Its main physiological function is the protection of membranes and the
stimulation of progesterone synthesis. In species, where the main site of
carotene to vitamin A transformation the gut wall, practically no carotene
can be found in blood plasma (rodents, swine, sheep, goat and poultry);
while in species (cattle and deer), where the main site of transformation is
the liver, the carotene concentration of blood is important. In the blood (and
milk) of Jersey and Guernsey cattle breeds the β-carotene concentration is
higher than in other breeds, the explanation is unknown. Horse and rabbit
are intermediary from this respect. In poultry, coccidiosis severely disturbs
carotene metabolism.
The main sources of carotene are the green forages and some tuber.
In this way, in green alfalfa (50 mg/kg), grass (40 to 100 mg/kg), whole corn
plant (10 to 12 mg/kg), alfalfa hay (10 to 40 mg/kg), grass hay( 5 to 20
mg/kg), alfalfa haylage (20 mg/kg), corn silage (4 to 5 mg/kg), alfalfa meal
(60 to 120 mg/kg), carrot (20 to 130 mg/kg), pumpkin (5 to 20 mg/kg, relat-
ed to the raw material (“as fed”).
The most important deficiency syndromes are the disturbed concep-
tion of heifers and cows (ovarial troubles), as well as the unviability of the
newborn calves. Daily requirement of lactating dairy cow is 300 to 600 mg
and this is independent from the vitamin A supply, because vitamin A can-
not replace the reproductive function of β-carotene. Except cattle and deer,
vitamin A and β-carotene are interchangeable, 1 IU vitamin A equals to 0.6
µg ß-carotene. The latter is sensitive to light, oxygen and acids and the soya
lipoxigenase (antivitamin A) degrades it. To control the supply, balance cal-
culation of the daily ration can be made, but the blood serum of cows is an
288
excellent indicator, 5.6 µmol/l (=300 µg/100 ml) being the lower critical
limit. Hypervitaminosis cannot be develops perorally, because if the blood
concentration attains the 22 µmol/litre, there is no absorption from the gut
(“carotene blockade”).
It new, pharmacological characteristics are the antioxidant,
immunostimulant and anticancerogenic effect. In this latter functions
other, plant and aquatic carotinoids may replace: lutein (grass, alfalfa),
zeaxanthine (corn grain), apocarotene (orange), canthaxanthine
(chanterelle), capsantine (paprika), violaxanthine (pumpkin), asthaxanthine
(crabs), licopine (tomato) and echinenone (sea star).
VITAMIN D in nature occurs in form of the ergosterol-originated ergo-
calciferol (D2) and 7-dehydro-colesterol-originated cholecalciferol (D3). Its
molecular weight is 397 (D2) and 385 (D3). One IU equals to 0.025 µg crys-
talline vitamin D. It is a precursor of the active 1,25-(OH)2-cholecalciferol,
produced by hydrolization in the kidneys and liver. Its main physiological
function is the promoting of synthesis of calcium-binding protein (CaBP) in
gut mucosa, which help in calcium absorption (except horse, rabbit and
guinea pig). Action of vitamin active vitamin D (calcitriol) is also indispen-
sable in mineralization of chondrocytes. In this function the presence of
vitamin C and K may enhance the calcium deposition (see cage layer fatigue
of hens and human osteoporosis). Vitamin D deficiency is also involved in
the pathogenesis of tibial dyschondroplasia of poultry.
Ergocalciferol can be found in the cod-liver oil and in marine sponges;
sources of cholecalciferol is besides the cod-liver oil the fat of warm-blood-
ed animals. Some plants (Solanum malacoxylon, Cestrum diurnum, Trisetum
flavescens) may have a high active vitamin D content (a glycoside form of
1,25-(OH)2-vitamin D), which can cause poisoning of grazing animals
(chronic wasting disease or enteque seco).
The typical deficiency signs derive from the lack of ossification and
overgrowth of cartilage: “ricketsy rosary” on the ribs of growing poultry, pig
and dog at the bone-cartilage border, “S”-shaped breastbone in poultry,
bented legs and crooked spine bone (pig, dog, calf), haemorrhage and frac-
ture of femur-head in breeding sow, deformities of the pelvic bones of poor-
ly supplied breeding gilts. In dairy cow vitamin D deficiency may contribute
to the development of milk fever. Recommended concentration for mono-
gastric animals is 100 to 300 IU/kg air dry feed and 30 IU/kg LW for dairy
cows. It is sensitive to light, oxygen and acids; raw soybean contains an
289
antivitamin D.
For evaluation of supply, the calculation of feed and body calcium and
phosphorus state, concentration of alkali phosphatase (AP) and hydrox-
yproline in blood and the ratio of plasma 1,25-24,25-(OH)2-D3 (active-inac-
tive form). After ingestion of ten- to twenty fold of the requirement may leas
to hypervitaminosis, which in turn results in calcinosis. The latter mani-
fests in a calcium deposition in soft tissues all over the organism, in this
way for example in the ovaries, causing infertility in rabbits.
Into the VITAMIN E GROUP belong eight, similar to each other structured
tocopherols. The a-tocopherol has the highest biological activity. Main phys-
iological function of tocopherols, together with selenium, riboflavin,
carotenoides and vitamin C, the protection of organism from the damages
of free radicals. Germ of cereal grains, most of the oilseed and green plants
have important amount of vitamin E.
Deficiency syndromes of vitamin E can be interpreted as a free-radi-
cal poisoning, because all over the body the extra- and intracellular mem-
branes (mitochondria, Golgi apparatus, lysosomes, peroxysomes, surface
receptors etc.), as well as the cell wall are damaged. The predilection points
and the deriving clinical signs are species dependent. In calves and lambs
(“white muscle disease”) the cardyomyopathy is typical, in form of Zenker-
type or waxy necrosis of the fibres; in ducks the damage of leg musculature
causes the so-called “seal posture” and an atrophy of gizzard muscles can
be found. In mink, marine mammals and cat a pansteatitis (“yellow fat dis-
eases”) develops, in chicken encephalomalacia (“crazy chick diseases”: cere-
bellum hyperplasia and oedema can be observed, typically at the age of 2-4
weeks. The cerebellum softens (malacia means softening). On the base of
clinical signs, it should be differentiated from the congenital manganese
deficiency and from some forms of pasteurellosis. Exsudative diathesis also
may be observed in growing poultry. Fragility of red blood cells generally
increases. The VESD (vitamin E and selenium deficiency) of growing-finish-
ing pig leads to bleedings in myocardium (“mulberry heart disease”), necrot-
ic spots in liver, oesophageal gastric ulcer and myodegeneration, in form of
pale, soft and exsudative (PSE) muscles, especially in the hind quarter (m.
longissimus dorrsi, m. gastrocnemius and semimembranaceus). The latter
clinically results in shaky gait. (It cannot be confused with the post-mortem
PSE -meat, caused by hidden, strong stress before slaughtering.)
In dairy cow the incidence of mastitis, the somatic cell count of milk
290
and non-infective placenta retention increases. In males of many species

degeneration of testis may develop. In rodent resorption sterility has been
observed, given the synonym of tocopherols as anti-sterility vitamin. In fish
a ceroid pigment deposition can be observed in the liver, kidneys and
spleen, besides ascites and pericardial oedema develop.
Recommendation for monogastric animals generally are 10 to 40
mg/kg of air-dry feed, for dry cows and heifers fed stored forages during the
last 60 days of gestation require approximately 1.6 IU supplemental vitamin
E/kg LW (or 80 IU/kg of DM), for lactating 0.8 IU/kg LW (approximately 20
IU/kg of DM), having calculating with the reduction of mastitis (NRC, 2001).
If the feed mixture has higher lipid content or the storage characteristics of
carcass should be improved, five- to tenfold, if the immune system is aimed
to be stimulated, twenty- hundredfold higher dosage is required. Vitamin E
is not a too sensitive vitamin, but against air, light and heat is not resist-
ant. In bean (Phaseolus vulgaris) there is an antivitamin E and pigment of
gossypol seed also decreases the tocopherol level of cows, decreasing the
herd’s conception rate
For monitoring the supply, the measurement of blood vitamin E level
and the activity of superoxide dimutase are appropriate. Hypervitaminosis
is very uncommon, only after long-lasting ingestion of high-tocopherol feed
(containing more than 1000-2000 IU/kg air-dry feed). Clinical signs of vita-
min E hypervitaminosis are not specific. A special feature of vitamin E, that
short-term application of mega-dosage has immunostimulant effect, fur-
thermore help in alleviating damaging effects of mycotoxins.
VITAMIN K or anti-haemorrhagic vitamin occurs in nature in form of
phytonadion (K1) and phytomenadion (K2); its synthetic variant is the
menadion (K3). The main physiological function of vitamin K is the contri-
bution in the blood clotting by helping the synthesis of prothrombin (factor
II) and other blood clotting compounds (factor VII, IX and X) in the liver.
Vitamin K is common in green plants and microbes of rumen and gut are
also capable of its synthesis.
Deficiency signs are linked with the trouble of blood clotting. Affected
poultry has an anaemic appearance, the comb, balls are pale, at the same
time, all over the body haemorrhage and petechia in the subcutaneous con-
junctive tissue can be seen. For the latter, it should be differentiate for the
Newcastle disease and from some mycotoxicoses. In rabbit does, pregnant
and lactating at the same time, bleeding can occur in the placenta. In
291
women, some types of premature birth are related to vitamin K deficiency.

Most of the requirement is covered by the microbial synthesis. Thus,
adult healthy ruminant does not need external supply. The recommenda-
tions for monogastric feed mixtures are within the range of 2 to 10 mg/kg.
Vitamin K is sensitive to light and bases. In the stuffy sweet clover emerg-
ing dicoumarol and the rat poison warfarin have antivitamin K effects, like
the sweet clover or melilot (Melilotus officinalis). Originally, sweet clover con-
tains only a harmless chemical coumarol, which gives the plant its charac-
teristic odour. During heating or spoilage of sweet clover silage of hay, the
latter compound is converted into the toxic dicoumarol. This toxin prevents
blood clotting; the first signs after a 14-day-long of ingestion are stiffness,
lameness, dull attitude, and haematoma in tissues manifesting in swellings
beneath the skin and uncontrolled bleeding may lead to death. The problem
is more common in ruminants than in horses. The toxin production in sweet
clover results from poorly managed ensilage or in hay put up too wet or
allowed to draw dampness when bales or loose hay remain in contact with
moist ground, The problem is that the stem of sweet clover is relatively thick
and retains moisture after the small leaves have dried. Since suspected
sweet clover has to be fed for about 2 weeks before chronic bleeding occurs,
an intermittent feeding with alfalfa or other dicoumarol-free forage may be
one of the solutions.
Drugs, damaging gut flora, like sulphonamides, antibiotics, some
cocidiostat may cause deficiency, especially in poultry. Long-term use of
nonsteroidal anti-inflammatory drugs (e.g. phenylbutazone) makes horses
susceptible to vitamin K deficiency and a possible coumarin poisoning. Even
an overdosage of thousand-fold could not provoke a hypervitaminosis. As
new, pharmacological feature of vitamin K that has a preventive effect on
the human postmenopausal osteoporosis. It should be borne in mind that
although natural vitamin K1 and K2 are fat soluble, the synthetic vitamin
K3 is water-soluble and practically there is no depot formation in the organ-
ism. Consequently, the every day supply is indispensable, if supplementa-
tion is necessary.
12.2.2. Water soluble vitamins
VITAMIN C chemically is ascorbic acid and its main physiological func-

tions rely in the formation of bones and teeth, cartilage, conjunctive tissue
292
and in iron transport in synergism with the folic acid. Main sources of vita-
min C are the green forages, fresh vegetables, fruits, potato, carrot, and
some animal product. Deficiency syndrome occurs only in species lacking
L-gulonolacton oxydase enzyme that converts L-gulonolactone to ascorbic
acid. The list of these animals includes man, monkeys, guinea pig, red-vent-
ed bulbul bird, fruit-eater bats, fish, butterfly, crabs, shrimp and krill
(CHEEKE, 1999). There is a mutant pig strain, too. In affected growing ani-
mals troubles of ossification and bone development occur, which in turn,
cause a shortening of femur and atrophy of linked muscles resulting in
prone position. Scurvy (gingival haemorrhage, loose teeth and leucopoenia)
is a general sign both for young and adult. In laying hen the endogenous
renal synthesis may prove insufficient and tibial dyschondroplasia (osteope-
nia or “cage layer fatigue”) develops, showing a poor mineralization of
growth plate and an overproduction of chondrocytes. Thus, vitamin D effect
requires the presence of vitamin C. In fish retracted head, scoliosis and lor-
dosis can be observed, typically the fracture and dislocation of vertebrae is
near to the caudal fin, accompanied by a general haemorrhage all over the
body.
Guinea pig requires 5 mg/kg LW daily, in the feed of fishes the rec-
ommended concentration is 500 mg/kg. To counterbalance the heat stress
of poultry, 100 to 200 mg can be added to one kg of feed mixture. According
some authors, is can be used in carnivores to acidify the urine. Ascorbic
acid is a very degradable compound, especially in humid environment and
in the presence of metal ions. It can be maintained its activity only in acidic
milieu or in microcapsulated form. It is stable as of L-ascorbyl-2-polyphos-
phatate, consequently, in fish feed this is the suitable supplementation.
Pharmacological property of ascorbic acid that in megadoses stimulates the
immune system and may be anticancerogenic.
VITAMIN B1, the thiamine or aneurine is the firstly discovered vitamin.
Its main function in organism is in the carbohydrate metabolism, namely
carrying acetyl-Co-A into the tricarboxyl cycle. In brans, germs of grains,
yeast, milk and egg yolk (see JAMES CLAVELL: King Rat novel) can be found in
higher concentration. In case of a thiamine deficiency, the cardiovascular
and the nervous system are primarily affected. In human beriberi, in mink,
cat, fox and marine mammals Chastek paralysis, in many other species
(birds, lamb, calf etc.) general weakness, paleness (pallor), cyanosis,
polyneuropathy, causing opisthothonus (head drawn back over neck),
293
aggravate in lamb and calf by cerebro-corticalis necrosis (CCN) (FIGURE

XII-4).
Living animals show medial strabismus (cross-eyed) and may often be blind
(cortical blindness). From the point of view of differ-
ential diagnosis, the hypomagnesaemia (calf), con-
genital ataxia (lamb, kid) and listeriosis lamb should
be mentioned. In felidae (lion, kitten) an opisthot
honus (“stargazer-disease”) and instability are pre-
dominant (animal easily fall or roll aside. In fish con-
vulsions and atrophy of muscles are characteristic.
FIGURE XII-4: Typical posture of chick with thiamin deficiency
For monogastric animals the recommendation is 1 to 3 mg/kg air-dry feed.
Adult, healthy ruminant has no needs, for calves, growing and feeder lamb
(especially if it is on high-grain ration) the supplementation is necessary.
Thiamine is sensitive to light, humidity, heat and alkali. In raw body of
many fish species (Table XXIX-3) there is a thiaminase enzyme, which
degrade it; in bracken fern and there is an antimetabolit of it. In poultry
some, coccidiostats or medicine (e.g. amprolium), disturbing or killing gut
flora, may cause thiamine deficiency. For controlling the state the transke-
tolase activity of red blood cells and lymphocytes is appropriate. For human
purposes (polyneuropathy) fat-soluble thiamine-analogues have been devel-
oped.
VITAMIN B2 (riboflavin or lactoflavin) takes part in the respiratory
chain (as part of flavin adenine dinucleotid (FAD) and in fat synthesis as
flavin mono*nucleotide (FMN). Considerable amount can be found in yeast,
milk, meat and liver. Deficiency syndromes primarily affect skin and nerv-
ous system. Growing pigs exhibit dry, desquamative dermatitis with loss of
hair on the whole body. In young chicken (1 to 3 week-old) the myelin
degeneration of peripheral nerves cause the “curled toe disease”. Fish body
receives dark pigmentation, on the eyes there is a corneal vascularisation
and haemorrhage, thereby animals become photophobic and move inco-
ordinated. In horses the lack of riboflavin may result in uveitis (or irido-
cyclochoroiditis, which is an inflammation of the iris, the ciliary body and
the choroid). In the majority of cases the cause is the insufficient absorp-
tion of the microbial riboflavin synthesized in the hind gut. It is called also
as periodic ophtalmia and moonblindness. It should be emphasized that the
riboflavin deficiency is not the only possible cause of equine uveitis.
294
Requirement of monogastric animals is 4 to 10 mg/kg air-dry feed.

Riboflavin is sensitive to alkali, light and UV-radiation. Riboflavin status
can be evaluated on the basis of glutathione reductase activity of erythro-
cytes.
VITAMIN B6 (pyridoxine, pyridoxal, pyridoxamine, or adermin) displays
its main biochemical action in the protein and amino acid metabolism
(desamination). Pyridoxine is also required for the conversion of tryptophan
to 5-hydroxytryptamine (or serotonin), a neurotransmitter. It is in important
concentration in red meat, spawn, cod-liver oil, yeast, egg yolk, viscera
(liver), cereals (except corn grain) and green plants. Deficiency syndrome of
pyridoxine in rodents is the acrodynia of tail, in growing chicken the eyelid
oedema and rough, brightless plumage are characteristics. In growing pig
growth depression and generalized exsudative dermatitis occurs, which can
be turn into purulent dermatitis, owing to secondary infections. Affected
dogs show lachrymation and ascites; in mink sterility develops in both
sexes. Hatchability of hen’s egg decreases owing to an embryo mortality in
week 2 of incubation. Fishes show epileptiform seizures, hyperirritability
and ascites. The requirement is 2 to 8 mg/kg air-dry feed. In human medi-
cine, it is prohibited to give vitamin B6 for vitamin B6 in case of Parkinson-
disease, because of possible neurotransmitter interaction. is sensitive to the
light and alkali. Pyridoxine status can be evaluated either by L-tryptophan
loading or by the aspartate aminotransferase (AST) and alanin amino-trans-
ferase (ALT) activity of red blood cells. As a pharmacological effect of vitamin
B6 it was found that in extra dosage may alleviate the premenstrual syn-
drome (PMS).
VITAMIN B12 (syn. cyanocobalamin, animal protein factor or APF). Its
main biochemical function is the contribution in the production of labile
methyl-groups. Thereby, vitamin B12, together with the folic acid, has a role
in nucleic acid synthesis. It is essential in methionine synthesis from homo-
cysteine and in the choline synthesis from ethanol-amine. The activity of
hydroxy-methyl-tetrahydro folate dehydrogenase and that of methylmalonyl
coenzyme A isomerase decreases in lack of cyanocobalamin. Even in the
pathogenesis of human diseases is important, that vitamin B12 is a coen-
zyme of methionine synthethase, which recycles methyl groups via homo-
cysteine, in this way promotes methionine synthesis. In lack of vitamin B12
(and folic acid) deficiency, the blood homocysteine level increases, this
means an increased susceptibility to myocardial infarction and atheroscle-
295
rosis in human. Another key enzyme of vitamin B12, especially for rumi-
nants, is the methylmalonyl coenzyme A mutase, which helps propionate
metabolism via succinate and the tricarboxylic acid cycle. Deficient activity
of this enzyme causes an accumulation of methylmalonil coenzyme A, which
in turn, inhibits the β-oxydation of fatty acids. This is the reason, why the
presence of methylmalonic acid in urine is an indicator of vitamin B12 defi-
ciency (SUTTLE and JONES, 2005).
Vitamin B12 practically occurs only in animal tissues (liver, kidney,
egg yolk), in addition in yeasts and some microbes can be found. It worth
mentioning that for the absorption man and swine requires a so-called
intrinsic factor. A special deficiency syndrome of vitamin B12 deficiency the
anaemia perniciosa; in pigs and carnivores normocytaer, in fishes macrocy-
taer anaemia develop. In young animals there is a growth depression, the
nitrogen retention decreases and the feed utilization worsens. These pigs
have thin body and longer legs, their coat is dishevelled (the so-called “dog-
like fattening pig”) and they are susceptible to parasites (ascariasis and
manges). In calves axons are degenerated; manifested clinical signs should
be marked off from the hypomagnesaemic status. Eggs hatchability
decreases, embryonic mortality falls to the last week of incubation. Dead
embryos show myodegeneration.
Recommended concentration in monogastric animals’ feed is 10 to 50
µg, for lactating dairy cow, according to the milk yield, zero to 300 µg daily.
Alkaline medium, light and UV radiation degrade it. Antivitamin B12 of soy-
bean binds it and prevents absorption from the intestine. The status of
organism can be evaluated by the vitamin B12 level in the blood and/or
blood and urine methylmalonic acid concentration (for metabolic pathway,
see above).
The main biochemical role of the NIACIN (syn. nicotinic acid, nicotinic
acid amid, pellagra preventive or PP factor, vitamin B5) is to take part in the
intermediary metabolism as part of nicotinic acid-adenin-dinucleotid (NAD)
and nicotinic acid-adenin-dinucleotid-phosphate (NADP). Except the cat,
the other species are capable to synthesize niacin from tryptophan. It can
be found in higher amounts in liver, meats, cereals (except corn grain), but
the bioavailability of latter is low.
In human the characteristic syndrome of niacin deficiency is the pel-
lagra (inflammation of skin, mucous membrane of buccal cavity, tongue,
stomach and intestine and nervous troubles). In dog and fox the dark, blue
296
pigmentation of the edges and tip of the tongue (“black tongue disease”) is
typical. Localized, dry and desquamative dermatitis develops on the ears,
dorsal surface of neck and back of growing-finishing pig with a coarse hair-
coat and on the head and axillar part (“armpit”) of the wings in poultry,
which, especially in poultry, can be accompanied by stomatitis and
oesophagitis. In fishes lesions and ulcers may develop in the hind gut.
Niacin deficiency is one of the causes of perosis in poultry and turkey.
Damages of nervous system is shown by ataxia and epileptic seizures (dog)
and jerking swimming (trout).
Monogastric animals require 15 to 80 mg/kg feed dry matter, lactat-
ing dairy cow on high-grain ration 1000 to 6000 mg daily. It is a quite resist-
ant vitamin, it is not sensitive against oxygen, light and heat. The
acetylpyridine and indolacetic acid of corn grain have antiniacin feature
(corn-inhibitor or anti PP factor).
It is a new, pharmacological effect of the niacin that in high dosage (6
g/cow/day) decreases the triglyceride and non-esterified fatty acid (NEFA)
concentration of the blood, contributing to the prevention of fatty liver. The
explanation of this benevolent effect is that niacin decreases the cAMP con-
centration of fat tissue, inhibiting in this way the adrenalin-stimulated lipol-
ysis. Megadoses of niacin (at least 3 g/day/person) significantly decreased
the cholesterol and β-lipoprotein-cholesterol (syn. [a]lipoprotein) concentra-
tion in blood, helping in prevention of atherosclerosis.
PANTOTHENIC ACID (syn. chicken antidermatitis factor, B3), as con-
stituent of coenzyme A, takes part in the intermediary metabolism. It con-
tributes to the mucopolysaccharide and acetyl-choline synthesis. By means
of his acetyl group, it has a role in the detoxification, too. Panthothenic acid
can be detected in higher concentration in animal tissues, egg yolk, yeasts
and green plants.
Generally characteristic deficiency signs are the growth depression,
damages of skin and nervous system, decreased antibody production and
disturbed adrenal gland activity. As clinical manifestation, in poultry the
rough plumage is faded (depigmented), especially the wings. At the corner
of beak crust can be seen, eyelids are sticked together by brownish exsu-
datum and on the dorsal surface of foot circumscribed ulcers develop
(descendent dermatitis). In rodents depigmentation (greying) can be
observed, if deficient intake is long-term. Growing pigs have peculiar loco-
motor disturbances, particularly on the hind legs (goose stepping,
297
Paradenmarsch), derived from the ailment of peripheral nerves. (FIGURE

XII-5a and FIGURE XII-5b) Carnivores exhibits convulsions and later paral-
ysis; necropsy reveals bleedings in the cortex of adrenal glands. On the gill
of fish waxy deposition, later necrosis develops, leading to death.
Only monogastric animals require supplementation, namely 6 to 20
mg/kg feed. Panthothenic acid is resistant to oxygen and light, but heat,
bases and acids degrade it. To control the supply, the coenzyme A concen-
tration of red blood cells or hepatocytes can be measured.
FIGURE XII-5a and FIGURE V-5b: Locomotob trouble at swine
The main biochemical functions of BIOTIN (VITAMIN H) ARE the contri-

bution in the active CO2 transport, in the fatty acid synthesis and in gluco-
neogenesis. It can be found in higher concentration in vegetables, fruits,
rice bran, milk, yeast (!), egg yolk and fungi. Lack of biotin in poultry results
in ascending, sloughy dermatitits, beginning on the balls of the feet and ris-
ing up to the balls, combs, eyelid and angle of the mouth maybe it ought to
be differentiated from the vitamin B6, and panthothenic acid deficiency).
Most frequently in turkey, perosis develops, which should be differentiated
from the niacin, folic acid and choline deficiency. In fox and dogs there is a
depigmentation of hair and a dry, desquamative dermatitits, which in case
of mink and cat leads to a fall of hair, too. Alterations are symmetrical and
generally they begin at the root/base of the tail and progresses backward
and forwards. In breeding sows the litter size decreases and their soles
become ragged having ulceration and from the pain their gait is sneaking.
(FIGURE XII-6)
298
FIGURE XII-6: “Sneaker- sow” syndrome
In broiler chicken Fatty Liver and Kidney Disease may develop. Biotin
deficiency of fishes can be characterized by the fragility of erythrocytes,
muscle atrophy, spastic convulsions and lesions of skin.
Monogastric animals require 50 to 300 µg/kg air-dry feed, which in
case of overwhelming corn and wheat feeding may increase by 20 to 50%.
Feeding approximately 20 mg supplemental biotin daily improved hoof
health parameters in dairy cows (NRC, 2001). Biotin is one of the most sta-
ble vitamin; it is sensitive only against strong heat and peroxides. Avidin of
raw egg white may bind biotin in the gut lumen preventing it s absorption.
For controlling the supply, the measurement of blood pyruvate carboxylase
activity and the calculation of plasma fatty acid ratio (C16:1-C18:0) are suit-
able. It is a special feature of biotin, that its single deficit (per se) is terato-
genic in mice.
FOLIC ACID or FOLATE (pteroyl-glutamic acid, vitamin Bc), in form of
tetrahydro-folic acid, takes part in the synthesis of purins and pirimidins,
nucleic acids, and (together with the vitamin B12) contributes to the forma-
tion of methionine, serine and choline. Cereal germs, green plants, meat,
milk and viscera are important sources of folic acid.
General deficiency syndromes are the growth depression, damages of
skin and gastro-intestinal epithel. In birds cervical paralysis can be
observed, manifesting in neck twisting and wing waving, finally the birds
gaze at the ground. This is particularly characteristic for turkey poults. In
pig and fish macrocytaer, in birds, carnivores and rabbit microcytaer
anaemia, in addition leuco- and thrombopenia develop. Minks exhibit fatty
liver; at breeding sow the litter size decreases. Deficient folic acid supply is
299
teratogenic to the bird embryos: the malformation of beak and femur can be
found.
Recommendation for monogastric animals is 0.6 to 2 mg/kg feed. Part
of dietary supplemental folate (0.2 to 4 mg folate/kg LW) may escape rumi-
nal degradation, and possibly by methionine-sparing effect, increases milk
production and milk protein. Folic acid is rather sensitive vitamin, but is
resists to air and heat. For the evaluation of states the activity of dihydro-
folate reductase of hepatocytes, that of serine-hydromethyl transferase in
blood and the urinary concentration of formino-tetrahydro folate are appro-
priate. Spina bifida of newborn is related to the deficient folic acid supply of
pregnant women, as well as the incidence of human cervical cancer.
The biochemical importance CHOLINE (choline chloride, B4) derives
from its role as a methyl donator and lipotrop agent (latter function is effec-
tive even as in part of the betain). Choline is an element of acetyl choline,
phosphatidil choline in the lecithin and in this way takes part in formation
of biological membranes. Larger amounts can be found in yeast, fish meal,
distiller and brewer products and soybean. The choline content of corn
grain is especially small.
Lack of choline decreases the sows’ litter size, which is more
expressed in case of concomitant methionine deficiency. Deficient supply
may cause perosis of growing poultry and fatty liver in laying hen. For
monogastric animals the recommendation is 200 to 1200 mg/kg feed.
Choline is a stable vitamin, it is sensitive only to the oxygen. To control the
supply, the incorporation of 14C into phosphatidil etanolamine and phos-
phatidil choline, after injection of 14C-ethanolamine. Hypervitaminosis may
develop, characterized by nervous troubles, anaemia and growth depres-
sion. High dosage of rumen protected choline helps high-yielding dairy cow
to prevent fatty liver and increases milk production, too.
VITAMIN U (S-methyl-methionine-sulphonium salts, anti-ulcus vita-
min, or cabagin) is an efficient methyl donor and supports the regeneration
of epithels and mucous membranes. Its sources are green plants (especial-
ly the cabbage) and fruits. Its deficiency sign in human is the peptic ulcer
in the stomach; or rather it helps in curing the subclinical cases. The exact
needs are not known. It degrades under the influence of light and alkali.
Hypervitaminosis practically does not occur. In small concentration it may
increase the feed intake in growing pig.
300
PANGAMINIC ACID (vitamin B15) is of strong lipotrop character. It can be

found in paprika seed, liver, brewer’s yeast and molasses. Its deficiency may
cause fatty liver in mink. Physiological needs are not known. Bases destroy
it, but it is resistant to acids. A special characteristic of pangamic acid that
in pharmacological dosage is a vasodilatator compound.
OTHER VITAMIN-LIKE SUBSTANCES are the inositol (hexa-hydroxi-cyclo-
hexan), the essential fatty acids (linolic acid, C18:2; linolenic acid, C18:3;
eicosatrienic acid, C20:3; arachidonic acid or eicosatetraen acid, C20:4;
timnodonic acid or eicosapentaenic acid, EPA, C22:5 and the clupanodonic
acid or docosahexaenic acid, DHA, C22:6. (EPA and DHA are called as
PUFA=polyunsaturated fatty acids), the taurin (amino-sulphonic acid), the
L-carnitine (quaternary amine), the orotic acids (B13), and the glucose
tolerance factor, GTF.
Essential fatty acids are precursors of prostaglandins and
leukotriens; they take part in formation of biomembranes and in water and
fat metabolism. L-carnitine facilitates the transport of long-chained fatty
acids into the mitochondria, especially in liver, muscles and heart. Inositol
or myo-inositol is found in feedstuffs as a component of phytate. It has
lipotrop character it may help minimize triglyceride accumulation in the
liver of lactating dairy cow. It is essential for calves and laboratory rodents
fed on semisynthethic diet. Essential fatty acids are in oils of grains and
seeds, as well as in animal fats of low melting point; taurin in animal tis-
sues, especially in marine molluscs and fish. L-carnitine sources are the
viscera and the meat.
Lack of essential fatty acids contributes to the pathogenesis of human
atherosclerosis, generally skin alterations and reproductive troubles are
characteristic to its deficient supply. Aquatic organisms (fish, crab, and
shrimp) especially require essential fatty acids. Lack of inositol causes
growth depression in rodents, fishes exhibit stomach atony and skin
lesions. Taurin deficiency in cats reflects in central retina degeneration,
dilatation cardiomyopathy, troubles in growth and reproduction, as well as
impaired immune functions. L-carnitine is essential to the growth of meal-
worm (larva of Tenebrio molitor beetle); in human newborn babies fatty liver
and mitochondria damages may develop in L-carnitine deficiency.
Their exact requirement data are not known, except linolic acid (1 to
1.5% in poultry feeds) and taurin (100 mg/kg cat feeds). Essential fatty
acids are very sensitive compounds, any slight oxidation destroys them.
301
Chapter XII: VITAMINS - FUNCTION- DEFICiENCY
Evaluation of essential fatty acid status is possible by measuring the

linoleate, arachidonate and eicosatrienate concentration of liver lipids.
Ingestion of to much essential fatty acids may cause vitamin E deficiency,
because it increases the requirement to tocopherols.
Tioctic acid is essential for some microorganism. The orotic acid
(B13) is an intermediary of pyrimidin metabolism and it enhances the
growth of rodents. The letrile (B17) or amygdaline is a β-cyanoglycoside,
like found in fruit kernels like apricot. In vitro it was capable of destroying
cancer cell. Vitamin H3 or gerovityl (chemically procain-hydrocloride or
novocaine) is able to alleviate the clinical signs of aging. Coenzim Q is the
common name of ubuquinons; it has tocopherols-sparing effect. Lack of
pyrroloquinolin-quinon (PQQ) may cause reproductive troubles and der-
matitis in mice. This is the most recently discovered vitamin-like substance.
It is higher concentration in fermented soybean (natto), in parsley, green
tea, green pepper, kiwi and the fruit of papaya.
FOR FURTHER READING
Ammerman, CB., Baker, D.H. and LEwis, A.J. (1995): Bioavailability of

nutrients for animals. Amino acids, minerals, and vitamins.
Academic Press. New York- London- Sydney.
Kolb, E. (1998): Evoluation and application of vitamind at domestic animals. Hofmann- La
Roche. Grenzach-Whyhlen.
McDowell, L.R. (1989): Vitamins in animal nutrition. Comparative aspects to human nutri
tion. Academic Press Inc. New York- London- Tokyo.
NRC (1987): Vitamin tolerance of animals. National Academy Press. Washington, DC
302
Chapter XIII
THE INTESTINAL AND RUMINAL FLORA

13.1.1. Historical overview
13.1.2. Methods of flora investigation
13.1.3. Composition of the intestinal flora
13.1.4. Connection of autochton flora and intestinal mucosa
13.1.5. Physiological role of intestinal flora
13.1.6. Defence systems of intestinal ecosystem
13.1.7. Intestinal immune defence system (GALT)
13.1.8. Interaction of intestinal flora and immune system
13.1.9. The peristalsis
13.1.10. Other components of the intestinal defence system
13.1.11. Upset of the balance of intestinal ecosystem and
flora (dysbiosis)
13.2.1. Ecosystem in the rumen
13.2.2. Rumen bacteria
13.2.3. Protozoa
13.2.4. Anaerobic fungi
13.2.5. Interactions of rumen microbes
13.2.6. Phenomenon of “attachment”
CHAPTER XIII: INTESTINAL AND RUMINAL FLORA AND FAUNA
SMITH (1965) compared the composition of intestinal flora of a series of ani-

mals (monkey, dog, cat, cattle, sheep, horse, pig, rabbit, guinea pig, rat,
mouse, golden hamster, gerbil, poultry, duck, tortoise, bullfrog, cod, finch
and cockroach) and has found that with the exception of rabbit, in which E.
coli is not a natural inhabitant, the intestinal tract of all homoeothermic
animals is basically populated by the same microorganisms. However, the
ratio of individual strains, depending on the peculiarities of digestive tract
and the feeding, considerable difference occurred. To concentrate on the
monogastrics, the main representatives of the intestinal tract are the strict
anaerobic bacterium species of Bacteroides, Eubacterium, Peptostrepto-coc-
cus, Bifidobacterium, Endosporus, Plectridium and Clostridium genus and the
facultative anaerobic species of Enterobacteriacae, Streptococcus and
Lactobacillus. In small number yeasts are also present. The concentration of
microbes in the different section of the gut is not the same: 104to 105 in the
duodenum and jejunum, 105 to 108 in the ileum and finally 109to 1011 in
the caecum and colon, related to 1 gram gut content.
The PHYSIOLOGICAL ROLE OF THE INTESTINAL FLORA is the synthesis of vita-
mins, degradation and modification of some endogenous compounds, fer-
mentation of non-absorbed nutrients, changing of pH and redox potential,
breaking down xenobiotics; formation the ratio of individual intestinal sec-
tions, determination of the villus density on surface, stimulation of the
regeneration of epithelial cells, maintenance or acceleration of peristalsis,
energy supply of hind gut mucosal cell, finally initiating and maintaining
the necessary stimulus for the intestinal immune system, the gut associat-
ed lymphoid tissue (GALT). In case of necessity, the suppression and extru-
sion of emerging pathogens is possible. On the basis of the above described
304
the intestinal flora of monogastric animals should be considered as an

ecosystem, which stands in integrant relation with the host organism.
13.1.1. Historical overview

Intestinal flora of the monogastric animals consists of approximately 1016
microbes; this number is tenfold more than the cells of body. The metabol-
ic activity of intestinal flora has a similar magnitude as the liver. As PASTEUR-
recognized in the XIXth century that the function of intestinal flora is indis-
pensable to the normal life of mammals, in 1906 MECSNYIKOV has already
raised the idea of intervention, by drinking yogurt. While discovery of path-
ogenic agents of the important infectious diseases drew the interest on the
specific treatment by sulphonamides and antibiotics, the achievements of
the last decades medical researches focused the attention on the intestinal
flora-host interaction. Importance of this field has only been highlighted by
the restriction of antibiotics application, as well as the opportunity of genet-
ic engineering of intestinal bacteria. The modern nomenclature uses com-
mensal) bacteria, instead of non-pathogenic. These classifications are rela-
tive, because many saprophytic bacteria proved to be able to cause infec-
tious metabolic trouble. Members of this group can display their damaging
effect if the general resistance of host organism falls. For these microbes the
“facultative pathogenic” and the “opportunist” names have been introduced.
Parallel to the development of bacteriology, a new science evolved, which
studies the interactions of intestinal flora and the host organism. The eco-
logic approach and the application of idea “ecosystem” onto the intestinal
flora made possible the better understanding of a series of pathological
processes and the development of efficient medical treatments.
13.1.2. Methods of flora investigation

Development of bacteriological techniques allowed the more accurate char-
acterization of intestinal bacteria. From this respect especially important is
the perfection of anaerobic techniques. As a further development, the com-
parison of animals with sterile (axenic), regulated (gnotobiotic) and natural
(holoxenic) intestinal flora occurred. These types of investigations may be
direct (study of gut content and/or faeces) or indirect. The latter means that
a specific substrate is applied perorally and the released metabolites are
analysed. As example, the lactulose test (hydrogen production), the cholyl-
14C-glycin test (CO -release) and the D-glucose test (hydrogen release) may
2
305
be mention, which are suitable to evaluate the extent and quality of colo-
nization in the hind gut.
13.1.3. Composition of the intestinal flora

The intestinal tract of newborn is sterile. The COLONISATION (ESTABLISHMENT) is
fast and important and till the 48th hours may achieve the adult values.
However, the composition of this flora is more simple, consisting mainly of
Enterobacteriacea, Streptococci and some absolute anaerobic species, like
Bacteroides and Bifidobacterium spp. Colonisation is selective, but its exact
mechanism is unknown. Composition of intestinal flora in suckling animal
basically depends on its environment and only some weeks after weaning
has the adult values
To illustrate the quantitative conditions, it worth mentioning that in
the gastrointestinal tract of man live 1014 microbes/g gut content, includ-
ing virus, bacteriophags, bacteria, mycoplasma and protozoa. In the mono-
gastric animals this value is even higher. Forty percent of the faeces are
microbe body. In one ml of hindgut content the number of bacteria is ten-
fold more than the erythrocyte number in one ml blood. Intestinal flora of
monogastrics consists of approximately 400 microbe species and a complete
bacteriological investigation of one faeces sample would last a year. The
majority of component of intestinal flora (up to 99%) is anaerobic and the
classical sampling and cultivation techniques are not suitable for their
detection.
Definition of the “normal” intestinal flora is not simple; it is called also
as indigenous, endogenous, resident, physiological and saprophyte??? flora,
too. Their common feature is that they are permanently present in the
ecosystem and are capable to proliferate in the conditions of anaerobiosis.
Composition of intestinal flora changes in physiological conditions, too:
depending on species, breed, individual and mainly the feeding; neverthe-
less, in the rear section the microbe population is rather constant. Stress
stands in interrelationship with the intestinal flora: stressors may influence
the composition of flora, and in turn, the composition of flora may mean
stressor for the host organism.
GASTRIC FLORA: although yeasts are low in number, but their biological
significance is great. In some herbivores (horse, rabbit) and in the omnivo-
rous pig from time to time Lactobacilli can be found. Bacterium concentra-
tion of duodenum and jejunum is low (104-5/g gut content) and mainly con-
306
sists of aerobic-facultative anaerobic species, like Streptococci. Others, espe-

cially if the sampling happened with the biopsy of mucous membrane, have
found an important number (106-7/g) of strict anaerobic species, too. The
bacterium concentration in ileum (105-8/g gut content) is higher than in
jejunum and the strict anaerobic species (e.g. Bacteroides) are dominant,
but occasionally Streptococci and Enterobacteriacea can also be found.
Owing to the function of ileocaecal vale, the composition and importance of
colonic flora substantially differ from the previous intestinal sections.
Bacterium concentration attains the 109-11/g gut content value, in which
the strict anaerobic species are dominant and their portion is hundred-
thousand times than that of facultative anaerobic bacteria. Caecal flora
roughly reflects those of the colon, the bacterial concentration is 1010-11/g
gut content. The microbes of content and those of attached to the caecal
wall should be treated as two individual compartments. Generally the strict
anaerobic (Bacteroides, Eubacterium, Peptostreptococcus, Bifidobacterium,
Endosporus, Plactridium, Clostridium) are dominant, but a small number of
facultative anaerobic bacteria (Enterobacteriacea, Streptococcus), as well as
yeasts and flagellate can be detected.
To study the flora of gut content is much more simple than investi-
gation of attached to the wall bacteria, hence most of our knowledge relat-
ed to the microbes of the gut content. Based on the number and significance
of microbes, the main groups are the following. Dominant species (109-11/g
content): Bacteroides, Eubacterium, Clostridium, Plectridium, Endosporus
and Peptostreptococcus. The secondary dominant (106-8/g: E. coli and
Streptococcus. In the group of “others” (below 106/g concentration), there
are permanent (Lactobacilli) and occasional flora component
(Pseudomonas). In other nomenclature, the first two groups are called pri-
mary and this latter as secondary flora. The number of a given group
expresses also in the magnitude of the exerted physiological effect.
13.1.4. Connection of autochton flora and intestinal mucosa

Most of the statements, concerning the intestinal flora, derive from the
investigation of faeces or gut content. Nevertheless, the pathology considers
the capability of bacteria to attach to the intestinal mucosa as an important
component of pathogenicity. Tools of attachment are surface formations,
called fimbria. Electro-microscopic studies revealed that the members of
intestinal flora are linked both to each others and the intestinal mucosa by
307
means of filaments. Members of the indigenous flora establish themselves

in special niche, using microcins and bacteriocins, these antibiotic-like sub-
stances against the concurrent microbes. This is a species-related charac-
teristic is after many authors is a fundamental feature of auchtoton species.
It is a striking new recognition that the genotype of host animal by produc-
ing endogenous substrates may have a more decisive effect on the makeup
of bacterial population, than the exogenous nutrients (GACSKIN, 2001). Other
bacteria are only embedded into the intestinal mucosa. Some of the
researchers consider only the epithelial flora as endogenous; others include
both the luminal and the crypt-colonized bacteria.
13.1.5. Physiological role of intestinal flora

It is evident that the 1014 microbes and the produced enzymes have an
important influence on the host organism, which is as follows.
NUTRITIONAL-METABOLIC EFFECTS. Synthesis of vitamin K and group B (in
particular vitamin B12), deconjugation and dehydroxylation of bile salts,
degradation of cholesterol, hydrolysis and transformation of bile pigments,
splitting of urea and ammonia release, deconjugation of sexual steroids, fer-
mentation of the non-absorbed carbohydrates (cellulose, pectin, starch etc.)
with the release of organic acids, carbon dioxide and hydrogen; digestion of
feed lipids by the bacterial lipase and degradation of proteins and amino
acids, like tryptophan, modification of pH, decrease of oxido-reductive
potential and modification of the metabolism of some drugs (antibiotics, dig-
italis, imipramin, L-DOPA etc.)
EFFECT ON MORPHOLOGICAL AND PHYSIOLOGICAL STATE OF INTESTINAL TRACT.
The flora modifies the ratio of the different sections of the intestinal tract:
the caecum of axenic animals is dilated and the wall is loose/flabby. By
introducing exogenous flora, these anomalies will cease. There is more
intestinal villus in a given surface, in bacterial flora is present. The cell
regeneration (the mitotic index) in axenic animals is slower, because of the
lack of nutritive volatile fatty acids, produced by the caecal and colonic
strict anaerobic flora.
As GENERAL PHYSIOLOGICAL EFFECT, in animals with normal intestinal
flora, the speed of blood circulation, the erythrocyte number are higher, the
emptying time of stomach faster and even some behavioural elements are
different, compared with the axenic counterpartners. The auchtoton
microflora has an ANTIMICROBIAL EFFECT, being an integral part of the defence
308
system of the host organism. By means of its mass, complexity and the
epithelial membrane connection it is a real barrier against the establish-
ment, attachment and proliferation of pathogenic microbes. On the other
hand, it has an indispensable role in the development and maturation of
intestinal immune system (GALT), too.
13.1.6. Defence systems of intestinal ecosystem

Although by means of the feed different exogenous strains arrive, the intes-
tinal flora show a high stability. This compositional stability is owed to a
regulatory-defence mechanism, which consist of microbial (active) and
intestinal (passive) elements. From the side of the intestinal wall the good
vascularisation, the production of digestive juice and the presence of lym-
phoid tissue worth mentioning. The harmonic coexistence of host animal
and intestinal flora, which means the least antigen and toxin loading for the
host organism, is the state of eubiosis (SZIGETI, 1990).
The above mentioned active and passive elements are in a complex
interaction (synergism or antagonism) with each other, with the com-
pounds, present in the intestinal lumen, even with the host organism as a
whole. Long-term existence of ecosystem testifies that the totalities of inter-
actions are for the benefit of each participant. This complex defence mech-
anism suppresses the potentially pathogenic bacteria, stabilizing the intes-
tinal flora and preventing its removal.
WAKSMAN (1945) has already raised the idea that the normal intestin-
al flora may exert an antagonist effect against the pathogenic bacteria. Later
this hypothesis has been justified by mouse and guinea pig experiments.
Thus, in the gastro intestinal tract exists a real microbial barrier, prevent-
ing or decreasing the establishment and multiplication of exogenous bacte-
ria (“COLONIZATION RESISTANCE”). For example, if axenic mice are perorally
treated by Shigella flexnerii culture, these bacteria will establish and prolif-
erate in the intestinal tract. The same bacteria, given for holoxenic mice,
according to the speed of peristalsis, during 35 to 36 hours, are completely
excreted. If axenic child receives Lactobacillus casei, it will establish during
24 hours. If then, representatives of normal, combined human flora (Str.
faecalis, Bifidobacterium and Bacteroides spp.) are applied, the L. casei will
be expulsed within 72 hours. It proves that even the settled, but not indige-
nous bacteria are eliminated by the intestinal ecosystem.
They are TWO MAIN TYPES OF BARRIER EFFECTS: the drastic, which elimi-
309
nates the exogenous strain completely, and the permissive, which makes
possible the establishment of exogenous bacteria, but inhibiting their pro-
liferation, their number remains low. In the majority of cases, barrier effect
derives from a complicated interaction of strict anaerobic strains. Barrier
effect consists of many factors, namely a mechanical closure of mucous
membrane, production and excretion of inhibitory metabolites (bacteri-
ocins), changing of the pH, modifying the redox potential, enzymatic action,
competition for the same nutrient and “extraction” of plasmids from the
exogenous bacteria by conjugation, putting them into the state of bacte-
riostasis
Besides the described antimicrobial antagonist effects, auchtoton
flora inhibits the translocation of external bacteria through the mucous
membrane and lamina propria into the lymph vessels and furthermore, it
inactivates bacterial toxins. Microbial barrier inhibits not only the estab-
lishment and proliferation of exogenous bacteria, but also suppress endoge-
nous, potentially pathogenic bacteria. The described defence mechanisms
cannot be separated from the function of intestinal immune system, the
GALT.
13.1.7. Intestinal immune defence system (GALT)

It is obvious the existence of an immune system at the level of gastroin-
testinal tract, if considering the continuous and important antigenous load
by means of infectious agents, parasites and feed. It is known since the
results of EHRLICH (1891) that immune response can be triggered through
the digestive system, too. The intestinal mucous membrane is a real lym-
phoid organ, all types of the cell-mediated immune response are present,
include B- and T-lymphocytes, neutrophyls, macrophags and plasma cells.
The lamina propria is the most plasmacyte-rich tissue in the organism, their
average concentration in the intestinal fluid is 450,000. Along the whole
length of the intestinal tract the immuno-competent cells are present both
in the mucosa and the submucosa, but in some species they form agglom-
erations (Peyer’s-patches and Ampulla ilei in rabbit). The microfold cells (M
cells) overlaying the follicles of Peyer’s patches have a role in antigen sam-
pling for the whole immune system. Commensal bacteria help by stimulat-
ing lymphoid follicle development.
The majority of data relate to the humoral immune response, namely
to the production of secretory IgA, the most important immunoglobulin on
310
the mucous membrane. IgA is synthesized in form of dimers in the plasma

cells, then links to the enterocyte-produced secretory unit, on the surface of
the mucous membrane for a real antibody painting).
IgA molecules are capable of inhibiting the attachment of bacteria to
the enterocytes, they takes part is the virus neutralization and in elimina-
tion of liquid antigens. The mucosal antibodies provide protection basically
by preventing the adherence of bacteria or toxin to the epithelial cells, which
process is called as IMMUNE EXCLUSION. However, their role in antibacterial
defence is contradictory, because lack of IgA did not lead to a really higher
incidence of intestinal infections. In the gut content are also present local
and blood-derived IgG, IgM and IgE molecules. Cellular immune function in
gut are not completely elucidated, but it is a fact that in case of its lower
function the incidence of bacterial, viral or fungal intestinal infections
increases. Of course, the function of macrophages, facilitating the bacteri-
olysis and phagocytosis, is not negligible, too.
13.1.8. Interaction of intestinal flora and immune system

Axenic animals are more susceptive both for intestinal and for systemic
infections. The comparison with the holoxenic animal revealed that the
cause of impaired immune function is the lack of normal intestinal flora. In
the axenic animals the lymphoid tissue underdeveloped, the level of natural
antibodies is lower and the polynucleo-granulated neutrophyls are less
active. Thus, the intestinal flora is the main stimulus of the development
and maturation of the intestinal immune system. The presence of flora
increases the measure of Peyer’s-patches and enhances the proliferation of
IgA-producing plasma cells. IgA deficit, in turn, inversely influences the
quantity and composition of the intestinal flora. In animals, after thymus
extirpation the intestinal bacteria frequently intrude into the bloodstream
and establish in organs.
It is clearly demonstrated that the microbial and immunological bar-
rier act in synergism to prevent the establishment and proliferation of path-
ogenic agents. For example, the resistance of axenic mice against Vibrio
cholerae is higher, when before the experimental infection receive not only
parenteral vaccination, but also peroral a mixture of natural flora compo-
nents.
311
13.1.9. The peristalsis

Peristalsis has an important influence on the attachment of proliferation of
bacteria, especially in the small intestine. In physiological circumstances
the colonisation is in inverse relation with the motricity of gut wall and the
possibility of attachment is higher in section of stasis (e.g. in the caecum),
than in the fast moving duodenum and jejunum. In pathological situation
the stasis of gut content facilitates the attachment and proliferation of
undesirable bacteria and they even may intrude into the bloodstream. For
example, the fibre-deficient diet of pregnant sows predisposes for the metri-
tis-mastitis-agalactiae, [MMA] syndrome. Thus, the peristalsis controls the
intestinal flora, and microbes, in turn, stimulate motricity of gut wall.
Really, the transit is slower in each gut section of axenic animals.
13.1.10. Other components of the intestinal defence system

Saliva contains a series of antimicrobial substances, like lysozyme, comple-
ment, lactoferrin, peroxidase, defensins and even antibodies. Acidity of gas-
tric juice assures an important protection against the feed bacteria; even it
keeps practically sterile the first section of the small intestine, too. Bile
acids have also some antibacterial feature. Mucin, produced on the mucous
membrane means physical and biochemical barrier against the attachment
of exogenous bacteria. Different discharges, excreted into the gut lumen,
like lysosime, lactoferrin peroxydase have antimicrobial effect, too. Under
the conditions of large gut the produced short-chain fatty acids (butyric acid
and lactic acids, produced by obligate anaerobes, are extremely toxic to fac-
ultative anaerobes, particularly to the species of the family Entero-bacteri-
aceae. Continuous regeneration (2 days on an average) of epithelial cells and
enterocytes also hinders the colonization of bacteria.
13.1.11. Upset of the balance of intestinal ecosystem and flora (dys-

biosis)
Defence mechanisms of the intestinal ecosystem form a dynamic steady
state, which can easily be upset. Each component is a potential cause of
disturbance, which is called dysbiosis. First group of causes are related to
the anatomical and physiological state of the host organism: the congenital
or required immune deficiency, malnutrition, premature birth and serious
diseases may contribute to the upset of balance between the intestinal flora
and the organism. The result is the same in case of anatomical, histological
312
or functional abnormalities of the gastrointestinal tract: viral infections,

enteritis, tumours, diverticulum, fistula, stenosis, achlorhydria and any
other factor, decreasing motility and motricity of gut wall.
Exogenous factors are also important, for example exogenous infec-
tions, abrupt and significant change in feeding, stress, even psychological
(increased blood cortisol level) may disturb intestinal ecosystem. From the
iatrogenic factors the gastrointestinal operation, the anti-inflammatory
therapy and the antibiotic treatment worth mentioning. It has been experi-
mentally proved that peroral application of antibiotics or other antimicrobial
compounds may upset the balance of intestinal flora. Damage of auchtoton
flora involves serious consequences: destroying of bacterium strains of
physical and biological barrier, emptying of binding sites on mucous mem-
brane and selection of antibiotics-resistant strains. Some of the antibiotics
may have an immunosuppressive characteristic, too. Holoxenic mice are
rather resistant against the colonization of perorally give E. coli, but after
treatment of antibiotics, having no effect on coli, the colonization will hap-
pen. Infective dosage of Salmonella enteritidis can be decreased up to hun-
dred thousandth part by using previously streptomycin treatment.
It has to be emphasized that not only the broad-spectre antibiotics
are capable to upset the balance of intestinal flora, because the naming
came from the therapy. For example the clindamycin is considered as a
selective antibiotic, having effect only on cocci and Gram positive bacilli.
However, it is very effective against the strict anaerobic barrier-forming cae-
cum bacteria and in this way may significantly decrease the Fusobacterium
and Bacteroides concentration of faeces. If SPF mice perorally receive peni-
cillin, the strict anaerobic flora of their caecum completely collapses, on the
contrary, the Gram negative Enterobacteriaceae proliferate and by translo-
cation may even results in bacteriaemia.
It has generally been accepted that perorally applied antibiotics are
able to drastically disturb the intestinal flora. For example, the ampicillin is
used to produce diarrhoea in pharmaceutical researches. At the same time,
it was a surprise that parenteral application of gentamycin, ampicillin or
cefpiramid caused the disappearance of strict anaerobic (Bacteroides,
Bifidobacterium, Veilonella, Clostridium) and facultative anaerobic
(Enterobacteriacea, Streptococci and Staphylococci) flora and the prolifera-
tion of yeast in the gut of human babies.
Exogenous stressors of management may lead to the changes in the
313
intestinal flora, mainly resulting in a fall of the anaerobic members. The

concentration of coliform bacteria will be higher after a decrease in amount
of fatty acids produced by the obligate anaerobic flora. From pathological
aspect the disturbance in the ecosystem of intestinal flora and host organ-
ism has two consequences. First, while some functions of the flora patho-
logically intensify, others exactly disappear. Secondly, owing to the weak-
ened or lacking barrier function allows facultative can pathogenic bacteria
to establish and to proliferate. If they achieve a population size, their viru-
lence can be manifested. This number of bacteria (this population size) is
called by ecomicrobiology the “critical threshold of pathogenic population”.
The occurring clinical and pathological signs basically depend on the inten-
sity of effect caused the disturbance and that which species are favoured by
the factors of new environment.
13.2.1. Ecosystem in the rumen

The composition of ruminal microbe population is under the control of
existing ecosystem. Rumen environment is peculiar and are different from
any other anaerobic systems. One of the most important factors is, that
owing to the host animal’s metabolism, the temperature is constant
(homothermia). The more or less continuous water and feed supply allows
the production of considerable amount of acids. Notwithstanding, the pH is
rather stablet (within 6 to 7), because the arising acid partly are absorbed
across the rumen wall and the remaining part is neutralized by the buffers
of saliva. For the same reasons the osmotic pressure remains close to the
isotonic. Ion concentration is regulated by dilution, absorption and passage.
Non-degraded solid feed particle, ground in small pieces by rumination,
pass also into the hind digestive sections. Since end products and residues
do not accumulate, but leave, the factors of microbial environment change
only within narrow ranges. More or less, these conditions are characteristic
for the hindgut fermentation, too.
The majority of rumen microbes is strict (obligatory) anaerobic.
However, they have a certain resistance to oxygen, supposed that the fer-
mentation is active enough to eliminate oxygen and to keep the electric
potential of media within the physiological -250 and -450 mV. Part of the
oxygen gets in by the feed and drinking water, but there is also diffusion
314
through the rumen wall. This normal amount of oxygen is quickly metabo-
lized and serves an electron donator. Excess oxygen in rumen may intoxi-
cate anaerobic microorganisms. However, the presence of any oxygen is dis-
advantageous to the anaerobic metabolism, included the methane produc-
tion. At the same time, methanogenic bacteria are important components of
the normal rumen flora and their role is important in assuring the balance
of fermentation processes. Activity of methanogenic bacteria depend on
many other factors, included the quantity and quality of arising products in
rumen. Oxygen (and generally electron donor) resistance of rumen ecosys-
tem can be stimulated by the provision of readily degradable carbohydrate
substrates. This is the reason, why a well supplied rumen is not sensitive
to the oxygen, arriving by an open fistula. On the contrary, having sub-
stantially lower substrate concentration, the in vitro rumen cultures have
significantly less oxygen resistance.
Continuous elimination of end product is extremely important for the
ecological equilibrium. This is the reason, why the pH drops in the silage
and the magnitude of cell synthesis is negligible. The base of the compari-
son is that both systems are anaerobic and the substrates are similar or the
same. Nevertheless, the silage reminds to a badly buffered culture, charac-
terized by the proliferation of one type of microbe, which struggle with the
other microorganisms for the dominance and for the control of substrate. In
ideal ensiling conditions the lactobacilli are in predominance and decreas-
ing the pH prevent the functioning of the other microbe groups. In the silage
there is no cellulose degradation, in the normal rumen this is of outstand-
ing characteristic. While in the growth of microbes is trifling, this is consid-
erable in the rumen, considerably contributing to the protein supply of host
organism.
Which is optimal for preparing good silage, it is disadvantages for the
rumen fermentation. Without considerable fibre degradation, rumen
microbes cannot really utilize the whole energy content of the substrate. On
the other hand, in during ensiling, the production of large amount of organ-
ic acid occurs on the account of microbial protein synthesis. However, pro-
duction of organic acids in rumen is inevitable during the release of sub-
strate energy content; the synthesized ATP and the released C are built in
the microbes. Owing to the continuous leaving/and or elimination of end
products and the buffer capacity of saliva the pH in the rumen is relatively
constant. It means that in spite of the continuous acid production, the pH
315
does not drop significantly, not favouring acidophil bacteria. Feeding ordi-
nary feed mixture (i.e. mostly forages), the role of lactobacilli is negligible.
On the contrary, giving ration rich in readily fermentable carbohydrates
(sugars, starch etc.), the importance of lactic acid bacteria increases.
Efficiency of rumen energy transformation (viz. substrateÞmicrobe body)
depends upon the dominant representatives of the flora and their acid-pro-
ducing capacity. The main factor during the selection of rumen microbes is
their maximal biochemical performance. In other words, bacteria of highest
degree of efficiency will multiplicate and dominate in the rumen. Close to
the neutral pH of rumen offers opportunity for survival for a wide diversity
of microbe species, and the formation of feeding or supplementing each
other groups. Continuous buffering of the rumen content is especially
favourable for the acid-sensitive fibrolytic bacteria.
13.2.2. Rumen bacteria

Steadiness of the ruminal environment and the regular substrate supply by
the feed intake establish favourable opportunity for an active fermentation.
Since similar conditions are dominating in the whole rumen, niche forma-
tion does not occur, otherwise so characteristic for other anaerobic circum-
stances (e.g. in soil or sludge). Moreover, the number of species is limited by
the continuous exchange of the rumen content (turnover), because the slow
growing and proliferating strains are washed out by the passage.
Although the majority of rumen bacteria is obligate anaerobic, some
facultative anaerobic occur, too. Under normal conditions they are insignif-
icant, but during some functional troubles of the rumen they may become
important. Number of rumen bacteria generally is determined by direct
counting. Bacteria of small body size (16 109/ml) give the half of the rumi-
nal biomass, but their metabolic activity is even of more important than
their proportion. Generally, metabolic activity exhibits an inverse relation-
ship with the body size. Microbes of large body size are not negligible, too,
notwithstanding that they give only a smaller portion of biomass. The most
important representatives are the Selenomonas (108/ml)), Oscillospira flag-
ellates (106/ml)), protozoa and phycomyceta. Group of ciliate-flagella proto-
zoa, like Entodinium (3Î105/ml), Dasytricha and Diplodinium (3104/ml),
Isotricha and Epidinium (1.1 104/ml) contains the lower number of microbes
present, but by their huge body size they give approximately 40% of the
ruminal biomass. Their metabolic activity is not proportional with their
316
mass, even their role is unknown. Anaerobic fungi have been detected in the
rumen content of grazing cattle and sheep. They are mostly attached to the
rumen wall and the feed particles, contributing basically in the fibre degra-
dation.FIGURE XIII-1
FIGURE XIII-1: Typical selenomonas bacterium
Group of small bacteria is a diversified population, consisting of pri-

mary carbohydrate degrading bacteria, breaking down cellulose, hemicellu-
lose, pectin, starch and sugars, as well as microbes, using up carbohydrates
and its derivates (cellulodextrins, pentoses, glucose, lactic acid, succinate,
formic acid and hydrogen) in their own metabolism. They should be treated
separately, because the used substrate, the produced metabolites and the
requirement of growth may differ. Many of the strains are rather specialized,
concerning their nutrient requirements, but the latter are assured by the
general rumen fermentation. The majority of species has already been iso-
lated and characterized. Some groups and species from the most important
rumen bacteria (afterVAN SOEST, 1994) are the structural carbohydrate fer-
menters (Ruminococcus albus, R. flavafaciens, Fibrobacter succinogenes,
Butyribibrio fibriosolvens and Eubacterium cellosolvens), the pectinolytic
species (Succinivibrio dextrinosolvens and Lachnospira multiparus), the non-
structural carbohydrate fermenters (Bacteroides ruminicola, B. amylophilus,
Selenomonas ruminantium, Streptococcus bovis, Succinomonas amylolytica,
Eubacterium limosum and Megasphaera elsdeni), the lipolytic species (e.g.
Anaerovibrio lipolytica), the proteolytic species (Peptostreptococci spp. and
317
Clostridia spp.), the organic acid fermenters (Megasphaera elsdeni and

Veilonella alcalescens) and the hydrogen utilizers, like Methanobacterium
ruminantioum and Vibrio succinogenes (see also Table XXV-1, too). Besides
the fundamental building compounds (volatile fatty acids, carbon dioxide,
certain vitamins, ammonia etc.) microbes require branched-chained fatty
acids as essential nutrient. Most of the rumen bacteria need B-vitamins,
ammonia, carbon dioxide, acetic acid, valeriate, isobutyrate, isovaleriate
and 2-methyl butyrate. Moreover, one third of the strains require haem, cal-
cium, magnesium, sodium, potassium, phosphorus, cobalt, sulphur, iron,
molybdenum, selenium, nickel, manganese and zinc.
Bacteroides genus has the highest number of species, included
starch-degrading and cellulolytic species. Notwithstanding that they are
capable of utilizing ammonia and peptide, many of them cannot be culti-
vated in pure culture, having the single N sources as free amino acids.
Furthermore, if nitrogen is provided mostly by ammonia, they need
branched-chain fatty acids for the amino acid synthesis. Members of
Bacteroides genus require carbon dioxide as a growth factor. On the con-
trary, for the cellulolytic, methane-producing and for some starch degrad-
ing species the ammonia is an appropriate nitrogen source. However, some
other species of this group have strong proteolytic activity and prefer as N-
sources peptides and amino acids.
Some Ruminococcus strain splits cellulose and hemicellulose, but for
the majority of species the primary energy sources are the cellulose-derivate
glucose and the hemicellulose-derivate pentoses (e.g. xilan), which are
excreted into the environment and the bacteria staying in the proximity can
use these compounds. On the other hand, the latter species have an impor-
tant need for isovalarate, 2-methyl-butyrate and/or isobutyrate. There are
strains, specialized to the use only one substrate as energy source; anoth-
er is more flexible from this respect. Being a considerable overlapping in the
functioning of the strains, the disappearance of a group of microbes gener-
ally does not modify the rumen function as a whole. More than half of the
bacterial population uses bacterial metabolites as source of energy. This
group has a high importance from the point of view of the passing bacteri-
al yield. Members of this group have a close interaction with the primary
carbohydrate utilizers, provide them with growth factors and influence the
variety of produced metabolites and end products into the maximal efficacy
of the whole ecosystem.
318
Establishment of calf rumen with bacteria (INOCULATIO) and the ques-

tion of transmission of the rumen bacteria and protozoa from one ruminant
into another have not still completely elucidated. The background of the
possibility of transmission is, that although rumen bacteria are obligate
anaerobic, they are pretty resistant against the oxygen. In this way, by
means of feed, saliva and possibly air, the microbes can be transmitted from
animal to animal. Isolated raising up prevent faunation of calf developing
rumen, but the bacterial fermentation may start normally.
13.2.3. Protozoa
Notwithstanding that protozoa give an important portion of biomass in aver-
age rumen content, their role in the rumen metabolism and in the fermen-
tation processes are not completely clear, despite that in Chapter XXV some
recent data are presented. One of the reasons is that it is extremely difficult
to cultivate protozoa completely bacterium-free. Protozoon-free rumen can
be achieved by fasting, applying certain chemicals or isolated rearing of calf.
Under experimental conditions, the weight gain of defaunated feeder cattle
proved to be higher than the control. Phenomenon can be explained by the
uptake of bacterial protein by the protozoa, decreasing thereby the total out-
put performance of the rumen.
Exclusive roughage feeding may also cause defaunation of the rumen.
Dafaunation hardly influence the rumen fermentation as a whole, but the
ratio and balance between metabolites and end products will change. For
example, in the presence of protozoa, high-grain feeding cause the produc-
tion of butyric acid; lack of protozoa favours the release of propionic acid.
This shift in the produced organic acid can be explained by the biochemical
characteristic of protozoa, which using starch and sugar substrate predom-
inantly produce formic acid and butyric acid. The most important protozoon
groups (after VAN SOEST, 1994) are the Holotrich (Isotricha and Dasytricha
genera) and the Entodiniomorph (Entodinia, Epidinium, Ophryoscolex,
Diplodinium, Eudiplodinium and Polyplastron genera (see also Table XXV-2).
In the defaunated rumen the number of bacteria increases, justifying
those protozoa is competing with bacteria for energy sources, moreover,
occasionally they consume them. The Entodinia genus of protozoa worth
mentioning for the outstanding acid resistance and for the fastest growth
rate. It can be experienced during in vitro cultivation that protozoa cannot
survive if they are inoculated further (onward) within than 24-48 hours.
319
Whereas the complete turnover time of the rumen fluid is 10 to 20 hours,

therefore only the attached to solid feed particle protozoa are able to remain.
Protozoa are capable of producing a considerable amount of ammo-
nia, forming an ecological niche with the task of providing bacteria with this
nitrogen source. Protozoa degrade part of the bacterial and feed-originated
protein, and they contain the 10 to 40% of the total ruminal nitrogen. They
use protein both as nitrogen and as energy source. The other energy-pro-
viding compounds are sugars (Holotrichia), starch and probably cellulose
and hemicellulose, too (in Entodiniomorph genus). Starch is an important
energy source for many groups, like members of Epidinium and
Ophryoscolex genus. The Holotrichs use sugar and other, soluble feed com-
ponents, while Entodiniomorphs (FIGURE XIII-2) depend on solid feed con-
stituents, like cloroplasts, fibre and bacteria. The main products of the pro-
tozoal fermentation activity are the acetate, butyrate and hydrogen. In the
presence of large amount of starch the lactic acid production is important,
too. Arising of propionic acid is trifling. Reports about the methane produc-
tion could be discussed, because it may a product of the symbiotic bacte-
ria. Many protozoa ingest and utilize bacteria; it is not yet completely eluci-
dated whether the symbiosis with bacteria (adhesion or inclusion) is essen-
tial for the protozoon. Anyway, it is similar to the evolution theory of
Eucaryotes. On the other hand, it cannot be excluded that some groups of
protozoa are able to degrade the vegetable cell wall on their own.
FIGURE XIII-2: Typical rumen protozoon (Entodinium)
Role of protozoa in the rumen ecosystem

The majority of protozoa cannot breakdown the fibre, even there are species
320
consuming bacteria. Therefore, the main benefit of their presence is not the
production of volatile fatty acids, but the preservation and bypassing unsat-
urated fatty acid into the abomasum and the continuous mixing of the
rumen content. Finally, the defaunation does not influence either the nitro-
gen metabolism of the whole rumen or the ruminal degradation processes.
Namely, from one hand, by consuming bacteria, they decrease the degree of
ruminal energetic efficiency (double transformation), from the other hand
they positively influence the ratio of emerging volatile fatty acids. It is strik-
ingly true in case of high-grain fed systems, when the extinction of ciliates
caused unfavourable close acetate to propionate ratio. It is supposed that
between the ciliate protozoa and methanogenic-cellulolytic bacteria,
attached on their surface, there is a real symbiosis.
13.2.4. Anaerobic fungi

The concentration of fungi in the rumen content is approximately 104/ml.
Their staying time is relatively short and they represent only a small portion
of the biomass. The most important fungi are the members of
Neocallomastix, the Caecomyces (formerly Sphaeromona), the Pyromyces
(formerly Pyromonas) and the Orpinomyces (polycentric fungi) genera. Fungi
produce fruiting body (sporangium), which in turn, getting mature, release
into environment swimming zoosspora. The latter are capable of attacking
rough fibre particle and developing sporangia and rhizoid filaments. The
main role of fungi is the facilitating of breakdown of coarse-fibred feed parts,
because fungal filaments are capable to penetrate lignified cell wall, too and
bring to place of possible action their cellulase complex. According to recent
data, inoculation of rumen with fungi may improve fibre degradation. Fungi
get in the faeces and external world embedded in cysts, resistant to the aer-
obic circumstances. Similarly to bacteria, fungi are producing volatile fatty
acids and gases; in traces ethanol and lactic acid are also released.
13.2.5. Interactions of rumen microbes

Composition of ruminal flora and fauna, depending mostly on feeding, may
vary between wide ranges. However, they can be classified into two groups:
those, which degrade feed component and those, living on the metabolites,
produced by the first members of the first group. The importance of the sec-
ond group is huge, because beyond the elimination of end products they
provide essential compounds for the members of first group.
321
Notwithstanding, the interaction between the two groups is not negligible,

because bacteria of different substrates reciprocally contribute to cover the
energy and nitrogen requirements of each other. For example, the survival
of exclusively cellulolytic Ruminococcus albus is supported, if it is incubat-
ed together with Bacteroides ruminocola, which is capable for degrading
both carbohydrates and proteins.
The interdependence of rumen microbes is called SYNTROPHY, which is
more, than the symbiosis, having the co-existence of two species for the
mutual benefit. In the syntrophy, many of microbial species are classified in
a feeding group, being specialized for the utilization of a special substrate,
for example fibre. In the latter syntrophic groups the cellulolytic, the hemi-
cellulolytic and the secondary fermenters (including methanogens) can take
advantage. In other words, syntrophic relations among species involve
shared metabolism and cross-feeding. The mutual benefit of two syntroph-
ic groups is the CONSORTIA, see above the example. This interaction allows a
series of further influence on the distribution and sharing of fermentation
metabolites.
Interactions modify the composition of really appearing/emerging end
products in the rumen. For example, the cellulolytic Bacteroides succino-
genes, through some metabolites, produces succinate, then acetate and
formic acid. The Selenomonas ruminantium, in turn, is able to use the aris-
ing cellulose metabolites and the succinate. Consequently, in the simulta-
neous presence of the two bacteria only propionic acid, acetic acid and car-
bon dioxide are produced. On the other hand, the latter is an essential
growth factor for the Bacteroides succinogenes.
13.2.6. Phenomenon of “attachment”

Many of the rumen bacteria are able to attach to the solid feed particle, even
to the ruminal wall, improving the chances of survival in the ecosystem.
(This phenomenon can be compared to the attachment of intestinal bacte-
ria, improving their pathogenicity.) By means of the attachment microbes
not only avoid the washing out with the liquid phase, but also they have
access to the substrate of the given feed particle. Tools of attachment are
glycocalix filaments. Cell wall degrading bacteria corrode cavities by their
enzymes on the surface of feed. Many of them digest hemicellulose, while
releasing pentosan fragments, in order to obtain access to their main sub-
strate, the cellulose. There is no free cellulase activity in the rumen content,
322
but, on the contrary, hemicellulose activity can be detected. This is the rea-
son for the fact that successful breakdown of cellulose is possible only in
case of a previous adhesion to the feed particle.
FOR FURTHER READING
Broudiscou, L. and Jouany, J.P. (1995): Reassessing the manipulation of protein synthe
sis by rumen microbes. Reprod. Nutr. Dev, 35, 517-535.
Ewing, WN. and Cole, D.J.A (1994): The living gut. An introduction to micro-organisms in
nutrition. Context. Dungannon.(N. Ireland)
Gaskin, H.R. (2001): Intestinal Bacteria and Their Influence on Swine Growth. In: Lewis,
A.J. and Southern, L.L. (2001): Swine Nutrition. 2nd ed. CRC Press. Boca Raton, London,
New York, Washington, D.C.p. 545-602.
Hirsh, D.C., MacLachlan, N.J. and Walker, R.L. (2004): Veterinary Microbiology. Blackwell
Publishing. Ames, Iowa
Russel J.B. (2002): Rumen microbiology and irs role in ruminant nutrition. Cornell. Ithaca,
NY
Tannock, G.W. (1999): Probiotics. A critical rewiew. Horizon Scientific Press. Wymondham
Van Soest, P.J. (1994): Nutritional ecology of ruminants. Cornell Univ. Press. Ithaca-
London
323
Chapter XIV
DETERIORATION OF FEEDS. RANCIDITY.

FEED MICROBIOLOGY AND MYCOLOGY

14.1.1. Rancidity of fats
14.1.2. Detection of rancidity
14.1.3. Prevention of rancidity
14.1.4. Veterinary aspects of feed rancidit
14. 2. Microbiological deterioration and mycotoxins contamina

tion of feeds
14.2.1. Microbiological decay
14.2.2. Mycotoxin-producing feed spoilage
Chapter XIV: FEED DETERIORATION

14.1.1. Rancidity of fats
The value and hygienic status of feeds are influenced by several factors. The
heat damage of proteins has been treated in Chapter 4.2. Here the oxidative
damage of lipids and spoilages of bacterial and fungal origin are delineated.
The rancidity of fats and oils begins with a hydrolytic or oxidative break-
down. Hydrolytic degradation of lipids to glycerine and fatty acids is caused
by the lipase of bacteria and fungi, facilitated by the presence of catalytic
metal cations, heat and light. The rancidity, produced by the described
hydrolytic process significantly decreases the organoleptic characteristics
and the preference, but it is not toxic and the changes in nutritive value are
trifling. Nevertheless, the “flavour threshold” of these compounds (capric,
lauric and myristic acids) is low, therefore the importance of hydrolytic ran-
cidity in case of milk powders and milk replacer is not negligible. The pro-
duced acids have a distinct soapy flavour, which is why hydrolytic rancidi-
ty is often referred to as “soapy rancidity”. The oxidative rancidity sometimes
referred to as autoxidation; it is not stopped by lowering the temperature of
feed or food storage. The cause for that is because the autoxidation is a
reaction with low activation energy.
The rancidity, resulting from the oxidative breakdown, produces not
only unpleasant smell, taste and abnormal colouration, but also the energy
value and the concentration of essential fatty acids and a series of biologi-
cally important substances are decreased. Oxygen directly attacks double
bonds: the oxygen substitutes hydrogen and very reactive free radical is
generated. The latter is transformed by entering of molecular oxygen, into
fatty acid peroxide, then through hydrogen uptake into fatty acids. The
extremely aggressive free radicals (ROO*, RO*, HOO*) means an oxidative
destruction of the enzymes, membrane structures and other lipid sub-
326
stances in feed and in organism, included the fat-soluble vitamins, too.

Oxidation of free fatty acids and the triglycerides is catalyzed by heat, light,
metal ions (especially the copper, iron and manganese) and the lipoxydase
enzymes of plant. The oxidation is concerned primarily with the unsaturat-
ed fatty acids, of which oleic (C18:1), linoleic (C18:2) and linolenic (C18:3)
acids are the most abundant. The first product of oxidation is an interme-
diate, which is itself odourless, but which breaks down to smaller molecules
which do produce the unpleasant taste. Thus, during the so-called autoxi-
dation, first hydroperoxids, then aldehydes, organic acids and polymerisation
compounds are produced. Consequently, if the peroxide value is observed for
long enough it will be seen to reach a maximum and then finally decline.
This occurs when the rate of decomposition of peroxides of hydroperoxids
into smaller volatile compounds outstrips their rate of formation.
In autoxidation the three main unsaturated acids are oxidised at dif-
ferent rates, namely linoleic acid is oxidised 64 times faster than oleic acid
and linolenic acid is oxidised 100 times faster than oleic acid. The produc-
tion of the mentioned intermediate can occur by one of three mechanisms:
the classical free radical mechanism, which can operate in the dark or a
photo-oxidation mechanism, which is initiated by exposure to light and
finally, by lipoxygenase route. As summarised above, the classical free rad-
ical route depends on the production of free radicals from lipid molecules by
their interaction with oxygen in the presence of catalysts, for example the
initiation may occur by the action of external energy sources such as heat,
light, high energy radiation or by chemical initiation involving metal ions (e.
g. copper ions) or metalloproteins such as haem of blood or muscle. The free
radical produced in the initiation steps can then react to form a lipidperoxy
radical, which, in turn can react further to give the hydroxyperoxyde. The
second reaction of the PROPAGATION steps provides at the same time a further
free radical, making it a self-propagating chain process. Course of oxidation
shows an initial phase or induction period, during the oxidation goes slow-
ly and at a uniform rate, followed by the phase of rapidly accelerating rate
of oxidation. The latter can be slowed down by adding antioxidant. The
photo-oxidation route is producing singlet oxygen (1O2*) from the triplet
oxygen (3O2). Singlet oxygen reacts 1500 times faster with double bonds,
producing hydroperoxids. Singlet oxygen is produced by light in the pres-
ence of sensitizers, like chlorophyll, myoglobin, erythrosine, riboflavin or
heavy metal ions. There is a Type I and Type II form of photo-oxidations,
327
producing different hydroperoxids. Type II photo-oxidation can be prevent-

ed by β-carotene and α-tocopherol which act as a quencher of singlet oxy-
gen. The lipoxygenase route is very similar to the autoxidation, but accord-
ing to the origin of enzyme, is very specific in substrate. These enzymes
chiefly prefer free fatty acid substrates, but there are lipoxygenase isoen-
zymes which can react with triglycerides, too.
It should be bore in mind that each of these three routes leads to
hydroperoxids which are relatively involatile and do not produce aftertaste.
There is a conversion of the hydroperoxids into the final flavour compo-
nents, because lipid hydroperoxids are very unstable and break down to an
alkoxy free radical. The mechanism of cleavage of alkoxy free radicals pro-
duces aldehydes. Parallel to the range of hydroperoxids available, there are
a great many aldehydes which can be produced. Aldehydes are responsible
for the pungent off-flavour and unpleasant smell (perfume or keton rancid-
ity), which are described as ranging from fresh, sweet through milky, citrus
and sweet oily. Formation of alcohols is very similar to that of aldehydes.
They are believed to contribute to the taste in the same manner as aldehy-
des, but in a milder way. Besides aldehydes and alcohols, there is also a
production of hydrocarbons. The polymerisation products cause the hot,
scratchy taste of the rancid fats. Hydrolytic rancidity, while the glyceride
molecule, under the action of heat and moisture, may break down to keto
acids, which lose carbon dioxide readily and methyl ketones, lactones and
esters may be formed primarily. Then the arising free oleic, linoleic and
linolenic acids undergo more rapid autoxidation. Methyl ketones contribute
a piercing sweet fruitiness, ranging from C3, pungent, sweet, through C7
blue cheesy to C11 fatty sweet. The aliphatic acids contribute to the taste
by being sour, fruity, cheesy or animal-like. Their contribution ranges from
C2 vinegary, C3 sour, Swiss cheesy, C4 sweaty cheesy, C8 goat cheesy, C9
paraffin to C14-C18 with very little odour. Water content of feed will create
a very great impact of the taste, reinforcing it. If the water content is high
enough, micro-organism can also produce methyl ketones.
It worth mentioning the third type of rancidity (non common), occurs
when fungal attacks on feeds, in the presence of limited amounts of oxygen
and water, leads to liberation of short chain saturated fatty acids. The lat-
ter then undergo a β-oxidation to produce a range of homologous even-car-
boned methyl ketones and aliphatic alcohols. Moreover, in the presence of
limited amount of water and oxygen, moulds (e.g. Eurotium amstelodami)
328
can interact with the lauric fat, first liberating short-chain fatty acids, which
are then subjected to a β-oxidation, yielding two homologous series of com-
pounds, namely methyl ketones and aliphatic alcohols, both with odd car-
bon chain (see milk replacers, based on cocoa powder or palm oil).
Measurement of ketonic rancidity is possible by analysing moisture, lipase
activity and free fatty acid contents.
14.1.2. Detection of rancidity

To detect rancidity, the peroxide value (number), the acid number and the
anisidin value (previously also: aldehyde or benzidin number) are applied.
High peroxide value is characteristic for the initial phase of the oxidative
process; the other parameters inform about the secondary oxidation prod-
ucts, as well as the damage of waxes. Nevertheless, the first products
formed are the (hydro)peroxides, therefore the most frequently the PEROXIDE
VALUE (PV) and the ACID NUMBER is used in the practice. The principle of the
methods is as follows. After cold, petroleum ether extraction of the lipids

from the sample, the fat concentration is determined by drying, the perox-
ide value by iodometric titration in glacial acetic acid-chlorophorm medium
and the acid number by acidi-alkalimetric titration in diethyletheric medi-
um. Peroxide value is equals to the iodine, separated by the peroxides of
1000 gram fat, i.e the amount of the 1 Mol concentration sodium-thiosul-
phate titration solutions (in ml). On the contrary, the acid number is relat-
ed to 1 gram of fat and it is equal to the amount of potassium hydroxide (in
mg), necessary for the neutralisation of the free fatty acids present. In other
words, the amount of free fatty acid that has developed in a feedstuff or mix-
ture is conveniently determined by direct titration of the extracted fat
against alkali: direct titration method. The Hungarian Feed Codex (2000)
published guideline to the evaluation of the degree of feed rancidity (Table
XIV-1) is.
329
Table XIV-1: Guidelines for the evaluation of the rancidity of fats
--------------------------------------------------------------------------------------------------------------
Peroxide value Qualifications Acid number
--------------------------------------------------------------------------------------------------------------
0-15 fresh 0-35
16-25 slightly stale 36-45
26-40 stale 46-55
41-50 very stale 56-60
51-60 starting rancidity 61-70
60< rancid 70<
--------------------------------------------------------------------------------------------------------------
The Hungarian Feed Codex (1990) published data concerning the maximum
allowed acid number, peroxide and paraanisidin values of commercial feeds
(Table XIV-2). The anisidin value (AV) is defined as 100 times of absorbance
of solution resulting from the reaction of 1 gram of fat of oil/fat in 100 ml
of a mixture of solvent and p-anisidin, measured at 350 nm. The test esti-
mates the level of aldehydes, principally 2-alkenals present. The AV test is
particularly useful for abused oils with low PVs such as frying oils.
Table XIV-2: Maximum allowed rancidity quality parameters of commercial feeds

--------------------------------------------------------------------------------------------------------------
Feed (raw material) Acid number Peroxide value Paraanisidin value
--------------------------------------------------------------------------------------------------------------
Milling by-products 50 25 -
Extr. solvent meals 40 25 -
Plant phosphatide 45 10 -
Starch by-products 50 25 -
Distiller’s feeds 50 25 -
Feeds of animal origin 50 25 -
Fat premix of milk replacers 5 8 25
Feed mixtures 50 25 -
Fish feeds 40 20 -
--------------------------------------------------------------------------------------------------------------
In delicate cases the AV is often used in conjunction with the PV to

calculate the “total oxidation value” or TOTOX VALUE: totox value=2PV+AV. It
is important to emphasize that each parameter is related to the fat content.
Consequently, the decision about the use of the feed or feed mixture
depends upon the fat concentration, too. Nevertheless, the given parameters
provide important basis to the evaluation of the potential animal (and
human) health hazard. The totox value is considered important in that it
combines evidence about the past history of the lipid, in the AV, with that
of the present state in the PV. The thiobarbituric acid (TBA) test can also be
carried out on whole feeds and may pick up oxidation damage to substances
330
other than the extractable triglyceride fats themselves. The test relates to
the level of aldehydes present in lipid, reacting specifically with malonalde-
hyde to give a red chromogen, which may then be determined spectropho-
tometrically. For the purposes of extension service, the Kreis test or rancid-
ity index has the advantage that it is rapid and gives an indication of incip-
ient rancidity, too. The test involves the production of a red colour when
phloroglucinol reacts with oxidised fat in acid solution. The Kreis-colour
reflects the concentration of epoxy aldehydes or their acetals, reported in
red units on the Lovibond scale. For evaluating rancidity, there are also
physical (e.g. infrared spectroscopy) and chromatographic (e.g. liquid col-
umn chromatography, GC, HPLC etc.) methods available for measuring
thermally labile peroxides, hydroperoxids and both volatile and non-volatile
secondary oxidation products.
Peroxidation is accelerated in the presence of salt and even faster if
salted feed or raw materials are frozen. It should be kept in mind that the
rate of rancidity is higher at about -5oC than it is at 0oC. The reason is that
the formation of ice crystals causes the reactants remaining in solution to
become more concentrated and thus stimulate reactions go faster. While the
temperature is reduced below minus 5oC, the lower temperature can dom-
inate the reaction rate, even though the reactants have got even more con-
centrated. These aspects have special concern during storage of petfood raw
materials (liver, fish, MBM, rendered fat, deep-frozen poultry broth etc.).
14.1.3. Prevention of rancidity

Rancidity of lipids can be prevented by supplementing feeds with antioxi-
dants. Antioxidants may be “any substance which is capable of delaying,
retarding or preventing the development of food rancidity of other flavour
deterioration due to oxidation” (Antioxidants in Food Regulation, 1978). The
major justification for using an antioxidant is one of the needs: it can extend
the shelf-life of a feed, reducing wastage and complaints; it may reduce
nutritional losses (especially fat-soluble vitamins and biotin are prone to
oxidation) and improve the technological possibilities of the feed industry of
using added fats and oils.
The general formula of a compound having antioxidant property is
“AH”; they provide hydrogen for the peroxide radical:
ROO* +AH_______ROOH + A*
(peroxide radical) (hydroperoxid)
331
It means that the free radical of the fatty acid turns back to the orig-
inal fatty acid, and the free radical of antioxidant (A*) arises. The latter
either has no reaction with the oxygen or this reaction is very slow, there-
fore the peroxidation will stop (“chain-breaking”). Most of the vegetable oils
contains some natural antioxidant, (e.g. tocopherols), but the animal fat
much less. This is the reason of the recent proposal, that the autoxidation
of carcass and meat during preservation may be prevented by feeding of
huge amount of vitamin E. There are also preventive antioxidants, which
act by reducing the rate of chain initiation. Metal inactivators, which coor-
dinate with metal ions capable of catalysing chain initiation, include citric,
phosphoric acid, ascorbic acid and EDTA. Absorption of UV radiation (e.g.
by carbon black, phenyl salicylate and hydroxybenzophenon) may also
inhibit forming radicals. Synergism: by mixing of two stabilisers has a much
better effect than either of the stabilisers alone. If a chain-breaking and a
preventive antioxidant are mixed both initiation and propagation are sup-
pressed (e.g. tocopherols+ascorbic acids+phospholipids). However, not all
combinations of antioxidants display synergism.
From the synthetic compounds the phenols (butylated hydroxi-
anysole, the BHA, butylated hydroxytoluene, the BHT, gallates, like doda-
cyl-3,4,5-trihydroxybenzoate and lauryl-3,4,5-trihydroxybenzoate and
mono-tert-butylhydroquinone, the TBHQ) and the cyclic amines (trimetil-
etoxy-dihydro-quinolin, the etoxiquine or EMQ= etoxy-methylquinolin).
Tocopherols and carotenoids are efficient natural antioxidants. Antioxidant
can be added either to the fat preparation or to the feed mixture (in the
majority of cases in both), but what is extremely important that in order of
prevention. Thus, rancid feed cannot be “improved” by mixing antioxidant,
only in some cases to decrease its harmful effect. The generally used con-
centration of the common antioxidants (BHT, EMQ and BHA) is 150 mg/kg
feed. Notwithstanding, an antioxidant cannot improve the taste of poor
quality feeds and cannot improve an oil or fat in which oxidative rancidity
has already developed, they cannot prevent hydrolytic and ketonic rancidi-
ty and they cannot prevent microbial decay. Although phenols have a quite
marked antimicrobial activity, but it is difficult to employ this activity of
BHA or BHT are practically insoluble in water. In watery media the vitamin
C itself, in fats and oils the fat-soluble derivate of vitamin C, the ascorbyl
palmitate can be applied effectively.
The tocopherols are widely distributed throughout the plant tissues
332
and have been called natural antioxidants. They are present in significant
quantities in all vegetable oils; fats derived from animals and fish contain
virtually no tocopherol. Gallates are also effective antioxidant for both ani-
mal fats and vegetable oils. The TBHQ (t-butylhydroquinone) has excellent
antioxidant properties, but it is not authorized in the EU. Its so-called
“carry-through” property, i.e. the capability to survive technological
processes, like pelleting, extrusion, is outstanding. The future ideal antiox-
idant maybe a high molecular weight polymer with phenolic side-chains
having structures of BHA or its derivates. These compounds are designed to
pass through the animal and human gut without being absorbed. Anyway,
the local regulations should be checked before application.
14.1.4. Veterinary aspects of feed rancidity.

The original vitamin A and E content will be destructed in the rancid feeds,
which should be considered during preservation and storage of these feeds.
The taste and smell of the rancid fats and oils is so unpleasant that fre-
quently feed refusal occurs. Part of the peroxides is absorbed from the intes-
tine and exerts its damaging effect in the organism, too. During excessive or
long-lasting heat treatment toxic polymerisation products may develop in
feed lipids. Feeding of these fats in a concentration of 10 to 20%, resulted
in anorexia, loss of hair, different organic ailments and death of experimen-
tal rodents. When chickens are fed on this type of lipids, growth depression
and oedema (primarily in the abdominal cavity and pericardium) can be
observed (“chickens’ toxic fat disease”). Laying hens react on rancid feed
intake by stopping the egg laying and an increase of aspecific mortality,
owed to the impaired general defence mechanism. Testicular tissue of grow-
ing cockerels is especially susceptible to the effect of the ingested rancid
feed. Monomer oxidation products (aldehydes, ketones) are less harmful,
they mostly cause organoleptic failures. FEKETE et al. (2006) added natural-
ly rancid meat-and-bone meal, with and without antioxidant supplementa-
tion to the growing rats's feed mixture. According to the histopathology, giv-
ing of rancid feed mixtures caused disappearance of glycogen from the
hepatocytes of liver, accompanied by a slight centrolobular fatty infiltration.
The addition of antioxidant failed to prevent these histological signs.
Peroxydes showed immunosuppressive character like lymphocyte depletion
in spleen, decreased measure of Malpighi bodies and decreased number of
lymphoblasts (FIGURE XIV-1 and FIGURE XIV-2) altered spermiogenesis
333
(FIGURE XIV-3 and FIGURE XIV-4). Antioxidant mixture's protective effect

seemed to be negligible in this respect. In a subsequent experiment with
growing chickens, chemically pure hydrogen peroxide supplementation has
been used (FEKETE et al.2008). Owing to the water-soluble hydrogen perox-
ide ingestion the degeneration and partly necrosis of cells in tubular epithe-
lia of kidneys, showing lyses, shrinking and detachment were found in the
treated bird. One fifth of the chickens of the highest treatment (3 ml/kg
H202 watery solutions) showed partial depletion of lymphocytes from the
lymphoid follicles of their bursa Fabricii. The lymphoid tissue and the
parenchyma organs, showed significant alterations even in the clinically
healthy animals.
From the point of view of warranty (guarantee) decisive is that the
rancidity of feed alone may be qualified as “significant failure” or not. Based
on several precedence cases, the statement of VÁRNAGY (1979) may serve as
a guideline: ”The peroxide number above 20 to 30 is the trustworthy sign of
the harmful oxidative process; the acid number of 40 to 50 (or higher) is also
sign of faulty performance, basically because it shows a significant drop of
feeding value of the batch by the decrease of energy concentration, by the
destruction of vitamin content and many time by the decreased preference.”
14. 2. Microbiological deterioration and mycotoxins contam-

ination of feeds
14.2.1. Microbiological decay
Microorganism are generally present on the feed; if their number becomes
high by means of contamination or by multiplication of undesirable bacter-
ial populations, the quality of feed may decrease or even a total spoilage
develops. Moulds may produce mycotoxins, which in turn, excreted into the
feed medium, threaten the animal health, and moreover, through the food
chain they may risk human health, too. Some of the yeasts and moulds are
capable of entering and proliferating in the organism, causing mycosis (e. g.
Candida-enteritis in piglets and calves). Feed components may serve as cul-
ture medium for pathogenic bacteria (salmonella, clostridia, staphylococci
etc.), which in organism, but often already in the feed, can produce toxins.
Occurrence of high number of apathogenic bacteria may also produce
health damage. The presence of their aspecific toxins and detrimental cell
wall residues in gut lumen results in subclinical inflammation, thickening
of the mucous membrane and thereby in decreased absorption of nutrients.
334
The real pathological significance of a microbial contamination is influenced

by the environmental temperature, the water activity, the pH, the partial
pressure of oxygen, the actual chemical composition of the feed, the cir-
cumstances of harvesting-preservation-storage trio and the preharvesting
events (e. g. insect or bird damage
The most important is the water activity (aw), which is the main lim-
iting factor of microbial growth. Water activity is an index, a proportional
number, showing the ratio of the water vapour pressure around the sub-
strate (feed or food) and the saturation steam (or water vapour) pressure of
the clear water, at a given temperature: aw = substrate steam (or water
vapour) pressure/steam pressure of water. The chemically pure water has
a water activity value of 1.0. The water activity is equal to the 1/100 part of
relative humidity in and around the substrate (aw= relative humidity/100),
at a given temperature. In other words, water activity shows the available
water for bacterial biochemical reactions. Because of their different physi-
cal structure and chemical composition, individual feeds of the same water
activity may have different water content (Table XIV-3).For example at tem-
perature of 21oC, corn grain has 15% and soybean seed 16% water content
at the same (0.72) water activity. Higher environmental temperature
increases the water activity. Table XIV-3 shows, among others, that to
achieve a same water activity of 0.85, corn grain requires 18.0% water con-
tent, while the dehulled sunflower only 9.0% water content. It mains, that
the latter is much more susceptible to bacterial and fungal spoilage.
Table XIV-3.: Water activity and water content of different grains and seeds at different rel-
ative air humidity, on 27.5oC
--------------------------------------------------------------------------------------------------------------
Relative aw Corn and Rice Soybean Sunflower
humidity, % wheat standard polished whole grain dehulled
--------------------------------------------------------------------------------------------------------------
65 0.65 12.5-13.5 12.5 14.0 12.5 8.5 5.0
70 0.70 13.5-14.5 13.5 15.0 13.0 9.5 6.0
75 0.75 14.5-15.5 14.5 15.5 14.0 10.5 7.0
80 0.80 15.5-16.5 15.0 16.5 16.0 11.5 7.0
85 0.85 18.0-18.5 16.5 17.5 18.0 13.5 9.0
--------------------------------------------------------------------------------------------------------------
Feed contaminating microorganisms have different water activity
335
requirement. Some xerophile moulds (e. g. xerophile Aspergillus spp.) may

already start to develop at a water activity of 0.62 to 0.65; most of the
moulds (Penicillium genus, other Aspergillus spp.) require medium (aw=0.80
to 0.91) and the hygrophilous Mucor spp. may develop only above a water
activity of 0.95. The minimal required water activity value for the develop-
ment of the individual microorganism groups is summarized in Table XIV-4
(SZIGETI, 1995)
.
Table XIV-4: Water activity (aw) values, required for the microbial growth
--------------------------------------------------------------------------------------------
Group of microbes aw
--------------------------------------------------------------------------------------------
Gram negative bacterium 1.00-0.95
Bacillaceae family 0.95-0.91
Yeasts 0.91-0.88
Moulds 0.88-0.80
Xerophile moulds 0.75-0.65
Osmophile moulds 0.65-0.60
--------------------------------------------------------------------------------------------
Data clearly demonstrate that bacteria require the highest and the
osmophile moulds the lowest water activity for multiplication. Feed dam-
ages occur already on the crop field by the attack of so-called “cropfield
moulds” (Fusarium, Alternaria, Cladosporium, Stachybotris etc.) and the
ergot. After harvesting, the circumstances do not favour development of
these moulds any more. At that time can start their activity of the “storage
or granary moulds”, having a need only for a lower water activity (e. g.
Penicillium, Aspergillus, xerophile Aspergillus spp.).
Some of the moulds cannot be classified definitely, because for exam-
ple member of the Fusarium genus may continue their growth even in the
silo, if the feed is not dry enough at the beginning of storage. On the con-
trary, in tropical conditions, cereal and corn grain and cottonseed may be
infected by Aspergillus flavus, if owing to insect or bird damage the seed-
coat (testa) injured. Microorganisms of lower water requirement will produce
water during their functioning, which in turn, facilitates and supports the
proliferation of moulds or even bacteria of higher water needs. According to
their required water activity, the microorganism may be ordered in a so-
called METABIOTIC SERIES or SEQUENCE).
Thus, during the deterioration the microflora of feeds significantly
changes. According to the feedstuff and the climatic circumstances,
spoilage is caused by a characteristic group of microorganisms. In moder-
336
ate climatic zones in the crop field the Fusarium genus, during the storage
the Penicillium genus are common. On the contrary, in animal cadavers bac-
terial development may already start on the pasture or cropfield. As a con-
sequence, in raw feed ingredients of animal origin, salmonella or coli infec-
tion may occur. On carbohydrate-rich substrates, like corn silage, yeasts
can proliferate.
From the microbiological aspect, saprophyte, indicator and pathogen-
ic bacteria can be distinguished. In the everyday practice, most of the analy-
ses are aimed at the determination of the quantitative status of the indica-
tor bacteria and as the pathogenic bacteria concern, only the registration of
the presence of Salmonella spp. is common. The Escerichia coli bacteria are
the signs of faeces impurity and their high concentration suggests the case
of a recent contamination. Unfortunately, the faeces might contain patho-
genic bacteria (e. g. the spores of Bacillus anthracis), viruses and parasites,
too; for the isolation of the latter the routine methods are not applicable.
14.2.2. Mycotoxin-producing feed spoilage

Feed damage without the production of mycotoxins may develop either
under the influence of atoxic mould species or by means of toxic moulds
strains, unless the environmental circumstances (e. g. the chemical compo-
sition of the substrate, water activity, temperature, humidity, the presence
of other microbes, the so-called accompanying flora, allow the synthesis of
toxic substance. Owing to the bacterial activity, first the concentration of
readily fermentable carbohydrates, like sugars and starch, decreases, then
the degradation and transformation of proteins and lipids follow. Dry mat-
ter and energy concentration of mouldy feedstuffs may decrease by 10 to
30% and the vitamin E and β-carotene content by 30 to 70%. On the high-
protein raw materials, like extracted soybean or sunflower, the amount of
free nitrogen grows and parallel to that, the pH of the substrate increases.
On the contrary, in the high-starch feedstuffs (e. g. cereals) the acidity and
the alkaline-binding capacity augment, parallel to the starch breakdown.
For the high-fibre stuffs an elevation of water content is typical, which may
cause a virtual increase in crude protein level. Unsaturated fatty acids
quickly undergo (during 2-3 day) the peroxide phase of oxidation and in the
following phases the growth of the acid number can be observed. The latter
phenomenon is in close correlation with the strong lipase activity of many
mould species.
337
Many of the feed-spoiling mould species excrete such secondary

metabolites (mycotoxins), which are toxic for the vertebrate animals. To dis-
tinguish from the mycosis, these syndromes are called mycotoxicosis.
Mycotoxicosis cases can be classified into three main groups.
a) Primary acute form. Clinical signs: hepatitis, haemorrhages, nephri-
tis, necrosis of the mucous membrane of mouth and gut, death. Combining
of these symptoms depends on the individual character of the mycotoxins.
b) Primary chronic form. Clinical signs: reproductive troubles, growth
depression, “ill thrisfft” of feed and worsening of the quality of final product
(e. g. meat). Their incidence in the practice is more common than that of the
acute forms. The recognition of chronic forms is not easy, because the oth-
erwise specific signs can be covered by the manifestation of other ailments,
metabolic troubles or deficiency syndromes.
c) Secondary mycotoxin disease. It may occur after the frequent inges-
tion of very low quantities of mycotoxins, decreasing the natural defence
mechanisms of animal. Mycotoxins primarily alter cell-mediated immune
response; however, in case of aflatoxins the antibody production may also
be depressed. At the same time, the phagocytosis and the production of
complement is altered, too. As a consequence, animals get more susceptible
to infectious and parasitic diseases.
The amount of toxic dosage, according to the toxin, the species, age
and sex of the animals, moves within wide ranges. For example, the LD50-
value of the zearalenon (F–2 toxin) is very high, that of ochratoxin A very low
(6 mg/kg in piglet and 3.5 mg/kg in one-day chicken). From the point of
view of incidence and economic impact the two most important mycotoxins
groups are the aflatoxins and the trichothecenes. As an illustration, it’s
worth mentioning that the screening of the average North-American corn in
the eightieth have found 2 ppm T–2 toxin; at the same time 4 ppm is effi-
cient enough to decrease the weight gain of poultry. In 1984 in Middle-
European climatic conditions the T–2 toxin concentrations of corn grain
samples fall in the range of 0.05-50 ppm. After ingesting of these corn
batches, feed refusal, vomiting and damage of the intestinal mucosa have
been observed.
The most characteristic feature of the zearalenon (F–2 toxin), pro-
duced similarly to the trichothecenes, by Fusarium spp., is a hyperoestro-
genism: embryonic mortality, foetal absorption and mummification, pseu-
do-oestrus of female animals are common. In male the sperm quality
338
decreases and in pig herds the “oestrogenic syndrome” develops, including

the birth of newborn piglets with swollen vulva, mammary gland and many
times having spread legs. TRICHOTHECENES. The T–2 toxin and its metabolites
(HT–2 toxin, deoxynivalenol, the DON, nivalenol and the fusarenon-X)
inhibit protein synthesis at cell level, which in turn, causes alteration of cor-
pora lutea in the ovaries, resulting in insufficient progesterone synthesis. In
the lymphoid tissues depletion of T- and B-lymphocytes can be observed,
manifesting in immunosuppression. By its characteristic effect, DON is also
called as “vomitoxin”. Ochratoxin A and B cause primarily kidney damage;
aflatoxins (aflatoxin B1, B2, G1, G2) may induce liver cancer. Patulin is of
antibiotic character and may occur in badly fermented silages, too.
Nowadays the incidence of stachybotricosis, which is a result of satratoxin-
poisoning, causing bleedings all over the body, is uncommon. Fumonisins
are the causing factors of the swine lung oedema and the encephalomalacia
in horses.
T–2 toxin addition to the growing rabbits’ feed caused lymphocyte
depletion in spleen, bone marrow and ampulla ilei ( FEKETE et al. 1989) (FIG-
URE XIX-5 and FIGURE XIV-6)
Simultaneous ingestion of otherwise very low concentration of myco-
toxins and other undesirable substances may exponentially reinforces the
potential of each others’ harmful character. As it was proven in fish model
for the liver carcinogenic effect of PCBs (polychlorinated biphenyls) and afla-
toxin B1; in turn, fusarium acid or 5-butyl-cholic acid (produced by
Fusarium moulds from the tryptophan in feed) redoubles the growth-depres-
sive influence of DON in pigs. Rumen microflora and fauna is able to
degrade part of the mycotoxins, but the efficacy depends upon the rumen
outflow rate, too (HUSZENICZA et al. 2000). Some of the mycotoxins in feed
mixture can be detoxified or neutralized by different treatments or supple-
mentation, but unfortunately they are very resistant chemical structures,
too. On the contrary, to prevent mould growth, mould inhibitors, like organ-
ic acids (propionic, sorbic, benzoic and acetic) and salts of organic acids
(calcium propionate and potassium sorbate) can be added to the feeds. The
mycotoxins, already present, can be bound by using mycotoxins-binders
(e.g. clay-derivates, like sodium bentonite etc.) is partly possible.
Significance of the mycotoxins contamination of the feeds is empha-
sized by the fact that by means of eating of animal products, the health of
human consumer is also endangered (KOVÁCS, 1998). Otherwise, mycotoxins
339
CHAPTER XIV: FEED DETERIORATION
can enter the human organism by the flour, bread, coffee, tea, bier, egg
meat and dairy products. SZUTS et al. (1997) described troubles in sexual
maturation of fusarium toxin exposed (wheat bran of the muesli) girls.
Generally, in the milling by-products, included bran, the concentration of
mycotoxins becomes higher, than in the flour. KOVACS et al. (1994) and BATA
et al. (1996) determined ochratoxin A from human blood and breast milk
samples, which is so much the more dangerous, being the organism of
embryo and infants by far more susceptible than that of the adults.
REGULATION
Table XIV-4: Maximum permitted concentration of mycotoxins contamination in feeds
--------------------------------------------------------------------------------------------------------------
Mycotoxin Feed Maximum allowed
concentration, mg/kg
--------------------------------------------------------------------------------------------------------------
Aflatoxin B1 Raw materials 0.05
Concentrates and supplements for non-lactating
cattle, sheep and goat 0.05
Concentrates for milking animals !!!
Poultry and pig feed, except young animals 0.02
Poultry and pig supplements, except young animals 0.03
Other feeds and supplements 0.01
Deoxynivalenol
(DON, vomitoxin) Feeds for pig, goose, duck, turkey and pets 0.4
Zearalenon
(F–2 toxin) Raw materials 10
Non-breeding ruminants 2.0
Feeds for broiler chicken, turkey, goose, duck,
laying hen, other birds and growing-finishing pig 0.5
Feeds for breeding and replacement cattle, sheep,
swine, turkey, guinea-fowl, goose and duck 0.08
*T-2, HT-2, nivalenol
DAS (diacetoxy-
scirpenol) Feeds for adult ruminants 2.0
ruminant concentrate, over 5 kg daily intake 1.0
Feeds for broiler poultry 0.5
Feeds for laying poultry and pig 0.3
Ochratoxin A Feeds for egg-producing birds 0.01
Feeds for adult ruminants 0.1
Other feeds 0.025
Ergot Whole cereal grains (rye, triticale) 1000
--------------------------------------------------------------------------------------------------------------
!!! zero tolerance
* Mode of action and toxicity of the listed trichothecene-structured toxin practically are
the same, consequently the summarized value should be considered while evaluating.
Legend to Table XIV-5

“-“ no data available
* on the basis of the 43/2003. (IV. 26.) FVM departmental order of the Agricultural and
Rural Development Ministry (Hungary)
340
341
Notwithstanding that nowadays only the (sole) detection of aflatoxin

B1 means automatically as faulty performance in feed commerce of the
European Union, the trends are that the maximum permitted concentration
will be prescribed for more and more individual mycotoxins. As an approx-
imation, data of the previous Hungarian Feed Codex (1990) may be helpful
and informative (Table XIV-4).
Followed the future tendencies, the Veterinary Medicinal Committee
of the Hungarian Academy of Sciences accepted and published the stand-
point of the special commission of the Hungarian Feed Codex Committee
(MATRAI, RAFFAI and VARGA, 2003). This declaration reinforced the regulation,
related to the aflatoxin B1 and elaborated for the fusarium toxins a toxic
and a so-called depressive (recognisable subclinical effects) limits (Table
XIV-5). For example, below the threshold of 840 ppb of feed the deoxyni-
valenol (DON) is not capable of altering the performance, blood biochemistry
and immune status of weanling piglets (ACCENSI et al. 2006).
The interpretation of the fusarium toxin limits are the following.
Ingestion of feed, containing fusarium toxin in a concentration no higher
than the “depressive” level, does not mean hazard to the given group of ani-
mals, the occasionally occurring losses in production or the possible health
problems cannot be connected with the consumption of the feed. Even the
long-term use of this feed is safe and tolerable. Toxin exposure through
feed, having more toxin concentration than the toxic limit, can be consid-
ered as a direct animal health risk and the feeding and economic value of
the given feed is strongly decreased. Mycotoxin concentrations within the
range of depressive and toxic limits mean decreased value of feed and risk
of use, too. The ruminal and intestinal flora may destruct many of the myco-
toxins, and there is a natural inhibition of the intestinal absorption, too
(PIZZAMIGLIO et al. 2006). The real manifestation of these effects depends on
a series of other factors, like the other components of the feed (essential
nutrients, toxin adsorbents, vitamin E level, peroxides etc.), immune status
of the animals and the environmental conditions.
FOR FURTHER READING
Accensi, F., Pinton, P., Callu, P., et al. (2006): Ingestion of low doses of deoxynivalenol
does not affect haematological, biochemical, or immune responses of piglets. J.
Anim. Sci. 84, 1935-1942.
Hungarian Feed Codex. FM-MMI. Budapest, 1990.
Kovacs, F. (1998): Mycotoxins in food chain (in Hungarian), Hung. Acad. Sci. Budapest
342
FIGURE XIV-1: Detail from a control spleen. Intact Malpighi body of large zones of dark
black lymphocytes and grey lymphoblasts. H-E. painting, 200 x enlargements. Bar= 50
(left)
FIGURE XIV-2: Detail from spleen of a peroxide-fed rat spleen. Decreased measure of
Malpighi body caused by the decreased number of lymphocyts and lymphoblasts of
Malpighi body. H-E. staining, 200x enlargement.
FIGURE XIV-3: Detail from a control testis. Developing germ cells of different phase of mat-
uration in the seminiferous tubulus and a lot of elong dark stained spermiocytes in the
proximity of the lumen. H-E. Pd staining 200 x enlargement. Bar=50 μ (left)
FIGURE XIV-4: Detail from a testis of peroxide-fed rat Lack'
seminiferous tubulus. H-E staining 200 x enlargement. Bar=50 μ (right)
FIGURE XVI-5: Lympocythe depletion in raqbbit bone marrow (left)

FIGURE XVI-6: Lymphocyte depletion in rabbit spleen (right
343
014-015 közötti színes.indd 343 2008.09.03. 15:04:31

Chapter XV
FEED ADDITIVES AND

PERFORMANCE PROMOTERS

15.2.1. Hormones
15.2.2. Repartitioning agents or metabolic modifier

15.4.1. Acidification of feed or drinking water after weaning
15.4.2. Enzymes
15.4.3. Application of plant extrats
15.4.4. Probiotics in poultry, pig and ruminant nutrition
15.4.5. Feeding of organic minerals
15.4.6. Prebiotic
15.4.7. “Fat burners” for more lean meats
15.4.8. Peroral passive immunization
15.4.9. Miscellaneous
15.4.10. Stress –Immune status – Performance enhancers
15.4.11. Nucleotides and their role in nutrition
Chapter XV: FEED ADDITIVES - PERFORMANCE PROMOTERS
To properly supply the humanity, the use of food of animal origin cannot be
given up. At the same time, it should be economically based. While using
performance promoters (syn. performance-enhancing substances), the prof-
it emerges from the increase of the average daily gain, milk, egg, wool pro-
duction and/or improvement of the feed efficiency. The measure of improve-
ment, when repeating trials, showed significant deviation, which can be
explained by the genotype-environment interaction. In the European Union
the use of hormone preparations, antibiotics or antimicrobial compounds for
performance promotion is not allowed since 2003. However, the knowledge of
the mode of action may be useful for each practical veterinarian. On the
other hand, in a lot of country of the Worlds (included the United States and
Canada), many of the compounds are in legal use. Only for using in swine
nutrition 12 antibiotics (apramycin, bacitrin methylene disalicylate, baci-
tracin zinc, bambermycins, chlortetracycline, lincomycin, neomycin, oxytet-
racycline, penicillin, tiamulin, tylosin and virginiamycin) and 5 chemothe-
rapeutics (arsanilic acid, carbadox, roxarsone, sulfamethazine and sulfathi-
azole) are approved (FDA, 2000).
BRAUDE (1981) summarized the results of his more than 100 copper
sulphate pig trials. The used concentration was 1000 mg/kg of feed and the
main parameter to collect the average daily gain. Points of the Gauss-curve
represent the result of an individual trial of more than hundred growing-fin-
ishing pigs (FIGURE XV-1).
346
FIGURE XV-1: Effect of copper sulphate supplementation upon the weight gain of grow
ingfinishing pigs
It can be seen in the figure that in the majority of cases the copper
sulphate supplementation improved the average daily gain by 4 to 14
(mean; 9.1%) percent units. In a small number of trials (in the territory over
the ±2 SD) extremely high positive answer (+37.1%) has been received, but
it also occurred in some experiments that the gain of the experimental
groups proved to be lower (-15.6%). The conclusion is that the effect of
growth promoters is influenced by genetic and environmental factors.
Moreover, between the two latter there may have been an interaction.
Among the environmental factors, influencing the effect of a growth pro-
moter the components of the outside physical world, the feeding, keeping,
the background genotype and the social effects (hierarchy fight, stress state)
should be mentioned.
Considering the above described which are the circumstances, when
a considerable result can be expected from the use of performance promot-
ers? Well, the better is the quality of feeding and the lower is the stress state
of animals, the application of performance promoter gives the smallest pos-
itive effect. The antibiotic response is greater in a dirty environment than is
a clean one. It means that the inadequacy of feeding, the harmful effect of
environmental stressors, is possible to counterbalance by using of perform-
ance promoting feed additives. On the contrary, no positive effect can be
expected from the application of copper sulphate or carbadox, if the grow-
ing-finishing pigs are in climatised chamber and fed on the best feed mix-
ture.
This statement is especially true for the probiotics and the nutriceu-
347
ticals. For example, the feeding of probiotics for gestating-lactating sows

may influence the mortality of suckling piglets, but the extent of improve-
ment is changeable. If the original mortality is anyhow low (below 9%), the
feeding of probiotics has no effect. If the piglet mortality is above 9%, in the
offspring of probiotics fed sows less baby pig will die.
Most of the growth promoters have an indirect effect, namely they
influence the composition of ruminal and intestinal flora and help in the
maintenance of that new, advantageous status. This is the reason why in
the companion animals of short intestinal tract these compounds have tri-
fling effect and in germ-free laboratory animals they are virtually ineffective.
The potential hazards of using antibiotics are
linked to the possibility of developing of an
antibiotic resistance. However, plasmid, coding
resistance can be transferred in human patho-
genic bacteria, too (FIGURE XV-2). This is the so-
called infective drug resistance (WATANABE, 1963).
FIGURE XV-2: Infective drug resistance
This discovery inspired the SWANN-report (Report of the Joint
Committee on the use of antibiotics in animal husbandry and veterinary med-
icine. HMSO, 1969), which declared that an antibiotics may be used either
for medicinal or for performance promoting purposes. The used compound
cannot be absorbed from the digestive tracts, its degradation in the outside
environment and soil should be fast and it cannot have allergenic side
effect. To protect the consumer, between the ingestion of performance pro-
moter and the consumption (slaughter, use etc.) a so-called withdrawal time
has been inserted. It should be mentioned that in the real life the infective
resistance occurs rarely, but still exists, see the Italian vancomycine case.
In the United States the Food and Drug Administration (FDA) must approve
a feed additive before it can be used in the practice. About the authorized
products and compounds the current “Feed Additive Compendium” (2000)
provides data, giving the generic name e.g. carbadox), chemical name (e.g.
methyl 3-[2-quinoxalinylmethylene] carbazate-N,N-dioxide), allowed species
(swine) and concentration 15-25 g/ton), FDA status (feed mill licence
required), warning statement (e.g. 10 weeks withdrawal time) and the trade
name Mecadox 10).
The classical performance promoters of the production animals can
be classified in two groups, viz. the direct effect on the metabolism, like ana-
348
CHAPTER XV: FEED ADDITIVES - PERFORMANCE PROMOTERS
bolic compounds and the group of substances acting through influencing

the ruminal and intestinal flora.
15.2.1. Hormones and their derivates support the anabolic processes of

the organism. Function of that can be found at the STEROIDS. Although the
medical-veterinarian evaluation of these compounds is contradictory, their
use in Europe is not authorized. A synthetic compound, similar to the F-2
toxin (Ralgro) is also used in the United States to enhance protein retention
of growing cattle.
Application of GROWTH HORMONE (GH), growth hormone-releasing hor-
mone or somatomedin pushes the homeorrhetic control towards lipolysis
and protein synthesis. By their use increases the milk production of sows
and cows and enhances the lean meat gain of feeder animals. Using sever-
al consecutive lactational periods, fat reserves of milking animals may be
exhausted, decreasing the conception rate. However, use of growth hormone
hides no hazard for the consumers; its application in Europe is not author-
ized. Regulation of poultry fat metabolism is different to the mammalian
one; the role of glucagon is not the same (RUDAS and SCANES, 1983). Thereby
the use of growth hormone or repartioning agents is not successful, more-
over, the bird can get fat (SZAKALL et al. 2001). Improved weight gain has
been observed while applying of THYREOSTATIC COMPOUNDS (e. g. ammonium-
perchlorate), inhibiting the hormone production of the thyroid gland,
decreasing the basal metabolic rate.
15.2.2. Repartitioning agents or metabolic modifier

THE BETA-2-AGONISTS. By feeding of these compounds, there is a shift between
the fat and protein synthesis and while consuming the same amount of feed
more protein and less fat will be produced. The beta-2-agonists have been
found effective in cattle, pig and rabbit, data concerning poultry are contra-
dictory. Since being catecholamines, their structure of these compounds
(ractopamin, cimaterol, clenbuterol, salbutamol, isoproterenol and L-
644,969) is similar to that of adrenalin (epinephrine) and dopamine, they
enhance lipolysis and protein retention of tissues. The prerequisite of the
positive effect is an optimal protein and amino acid supply. Their peroral
349
application should be continuous. As a summarized result, the fat content

of carcass decreases by 10 to 12% and that of the protein increases by 9 to
10%. The proposed withdrawal time before slaughtering is 24 to 48 hours,
which cannot be longer, because an intensive fat synthesis begins. The rac-
topamine is approved in the USA by the FDA in 2000.
Components of the ruminal and intestinal flora and fauna of a healthy ani-
mal live in balance with each others. The upset of this balance causes diges-
tion troubles, even health problems. The positive effect of many growth pro-
moters (antibiotics and antimicrobial compounds) is realized by the estab-
lishment and stabilization of a new microflora and fauna. This new ecosys-
tem is more advantageous for the host organism, because by reducing the
state of so-called physiological inflammation that is manifest constitutively
in the intestine and explains the high degree of cell and mucus turnover via
epithelial and goblet cell responses to cytokines, expressed by lamina pro-
pria cells. As a consequence, the absorption of nutrients increases, more
vitamin and fewer toxins are produced. The measure and energetic costs of
bacterial mucin degradation may also diminish. Many of intestinal bacteria
are capable of producing amines via decarboxylation of amino acids and
breakdown of polyamines (see Chapter VIII). In antibiotic treated rats the
concentration of both intestinal and urinary histamine decreased (WILSON,
1954). Typical representative of this additive group are the ionophor antibi-
otics (e.g. monensin, narasin, salinomycine etc.). Besides the described
effects in the monogastrics, in the rumen they are able to selectively kill
Gram-positive bacteria, included methanobacteria, improving in this way
the utilization of roughages in ruminants.
In monogastric animals the use of antibiotics (bacitracin, flavomycine,
virginiamycin etc.) was common. For growing-finishing pigs the application
of copper sulphate, zinc oxide (in high concentration), carbadox and
olaquindox used to be very effective. Because the last two substances are
synthetic antimicrobial compounds, long (approximately 10 weeks) with-
drawal time was necessary. Before the real application of any growth
promoter, the actual and local legal regulation should be considered.
350
THE FORMER EU-REGULATION (GRUNE BROSCHÜRE, 1994) the preven-

tive and nutritive biologically active substances used to classify into three
groups (first the generic and in parentheses the trade mark name).
I. Performance promoting antibiotics: avilamycin (Manus premix),
avoparcin (Avotan R-100 medicated premix) flavophospholypol (Flavomycin
80), sodium monensin, sodium salinomycin, spiramycin, tylosin phosphate
(Tylan performance promoting medicated premix), virginiamycin
(Virginiamycin premix, Stafac), zinc bacitracin (Fermin-10 medicated pre-
mix)
II. Other growth promoters: carbadox (Getroxel-25), olaquindox
(Bayonox).
III. Preventive compounds
histomonadosis: dimetridasol, ipronidasol, nifursol, ronidasol
coccidiosis: amprolium (Amprolium medicated premix), amprolium-
ethopabate, (Amprolium plus medicated premix) aprinocid, decoquinate,
diclasuril, dinitolimid, DOT, halofuginon (Stenorol), sodium lasalocid
(Avatec-Roche), maduramycin ammonium (Cygro medicated premix),
methychlorpindol, methychlorpindol+methylbensoquat, monensin sodium
(Elancoban medicated premix and Rumensin premix for ruminants),
narasin (Monteban 100 medicated premix), narasin/nicarbasin, robenidin
(Cycostat premix), salinomycin sodium (Sacox-120; Salocin-120 for pigs).
The maduramycin, the sodium lasalocid, the sodium monensin, the
salinomycin sodium should be kept far from horses because of the danger
of caecal dysbacteriosis and profuse diarrhoea. For rabbit and dog the
narasin, for one-day chicken the carbadox may cause poisoning.
At the same time, it is interesting, that right now a series of growth
promoter antibiotics (avilamycin, efrotomycin, salinomycin, nosiheptide and
thiopeptin) have not received the FDA authorization for use in pig feeds.
GROWTH PROMOTERS INFLUENCING THE RUMEN FUNCTION
a) Antibiotics: monensin, lasalocid, salinomycin, narasin, lisocellin

(ionophores), avoparcin (glycopeptid) transform the rumen flora by killing
Gram-positive bacteria, thereby decreasing the number methane and car-
bon dioxide producing bacteria. By this indirect inhibition of gas produc-
tion, they spare energy for the host animal. These compounds have origi-
nally been coccidiostats and the effect on coccidia and rumen bacteria is
similar to each other: they disturb the cation transport of the cell mem-
brane. Owing to the physiological regulatory mechanisms the voluntary dry
351
matter intake generally decreases and the economic benefit realizes in an

improved feed utilization. According some data, the ionophor antibiotics act
not only by inhibiting the methane production and promoting the propionic
acid production, but also decrease the basal metabolism and increase the
ratio of rumen undegradable protein (RUP or UDP). Their use is not gener-
ally advised for lactating dairy cow, because the reduced acetic acid con-
centration in rumen may result in a drop of milkfat concentration.
Notwithstanding, the incidence of ketosis may be reduced by this way. In
the EU, the use of monensin sodium (Rumensin premix) used to be author-
ized in the concentration of 125 (up to 250 kg LW) and of 250 mg/kg DM in
older age. The intake of monensin is dangerous for horses, rabbits and car-
nivores and it is incompatible with the tiamulin.
b) Direct inhibition of the methane production
Each compound, which inhibits the formation of methane from
hydrogen and formiate, achieves the same final effect, described above. For
example do the direct hydrogen attracting nitrates, sulphates and the long-
chained unsaturated fatty acids. The latter are also efficient in decreasing
the number of methane-producing bacteria. It is worth mentioning the chlo-
ral hydrate derivates, too (e. g. hemi-acetal chlorate starch, HCS), which by
killing methanobacteria and by adsorbing the hydrogen and vitamin B12
improves the feed utilization.
c) Antibiotics of other mode of action, like flavophospholipids, tylosin
and avoparcin may contribute to the stabilization of an effective rumen
microflora and fauna, correcting fermentation processes (FEBEL, 1995) and
decreasing the incidence of liver abscesses. It should be emphasized that
the peroral use of broad-spectre antibiotics is not advisable, because the
useful members of the microbe population may perish, too.
d) By feeding rumen buffers (NaHCO3, sodium sesquicarbonate and
MgO) or ion-exchanger clays (zeolite, bentonit or fuller’s earth) the harmful
side-effects of the high-grain feeding can be counterbalanced. By binding
and releasing ammonia, they may improve the nitrogen utilisation, too.
e) Others. It means also as an influencing of rumen function, when
protected protein, amino acid or lipids are fed. By applying nonil-phenol-
ethoxylate the number of protozoa decreases: defaunation. The orothic acid
may increase the ammonia uptake of rumen bacteria. Antibloating agents,
like poloxalene, incorporated commonly is salt block, may be useful during
grazing period. The parasympatico-mimetic compounds (salivation inducer)
352
of mould origin (e.g. slaframine) contribute to the stabilization of the rumen

pH, while feeding high-grain ration.
In order of the safer animal and especially human feeding the replacing of
performance promoter antibiotics and antimicrobial compounds in the feeds
and foods is necessary. For this purpose the application of functional feeds
and foods (or nutriceuticals) is continuously increasing. Thus, certain nutri-
ents may have pharmacological effects while given in higher doses than the
established nutritional requirement. These raw materials, preparations,
feeds, foods are commercialized, because they are supposed to have advan-
tageous effects on performance and/or health. Positive effects include aspe-
cific (e.g. increasing vitality), specific and directed on one of the body organ
systems (e. g. stimulation of the immune system) and specific, systemic (e.g.
slimming diet for human and companion animals or the increase of the per-
centage of lean body mass in case of slaughtering animals). The following
materials are considered as nutriceuticals: amino acids, minerals, vitamins,
organic acids, linseed and fish oil, conjugated linolenic acids (CLA), plant
extracts (e.g. garlic, oregano, rosemary, ginkgo, ginseng, grape seed), which
are generally accepted as safe (GRAS) and they are prescription-free.
Supporting of health status and prevention of diseases by means of these
preparations may also be valuable but it is difficult to evaluate their effica-
cy and many times they turn to be ineffective. Regulatory authorities treat
nutriceuticals rather as food or feed and the strict authorization rules are
not applied. The Hungarian Feed Codex (Codex Pabularis Hungaricus) in
2000 proposed the „feeds of special nutritional goal” denomination. The
main reason of using nutriceuticals, especially in the European Union, is to
replace the performance promoting effect of prohibited feed additive antibi-
otics.
The main nutriceuticals (nutricines, functional feeds) in the animal
feeding are as follows:
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15.4.1. Acidification of feed and/or drinking water after weaning

One of the causes of the digestive troubles of weaning piglet is that the pH
of the feed mixture is above 6.5. The genuine hydrochloride acid production
of piglets at that time is insufficient to set the desirable pH of 3 to 4.
Considering the above described, the acidification of drinking water or/and
the feed mixture is suggested by using organic acids, betain-HCl or lactic
acid producing bacterial cultures. Organic acids (malonic acid, fumaric
acid, citric acid) counterbalance high alkalinocity of feeds and decrease inci-
dence of diarrhoea. The non-condensated (hydrolysable) tannic acid is use-
ful in treating clinical signs of enteritis (e.g. FARMATAN® from Tanin
Sevnica d.d. and Matischa Ltd.). The previously fermented, liquid feed of
weaning pigs cannot have a pH below 4.8, unless feed refusal occurs.
15.4.2. Enzymes
Some components of the monogastric feed mixtures has a low digestibility,
even some of them is indigestible for these group of animals. The reason for
that is the occurrence of non starch carbohydrates (NSC) and/or antinutri-
354
tive substances in high concentration. The use of enzyme offers solution for
this problem. The improved digestibility means at the same time more
absorption and less nitrogen and phosphorus emission, which is increas-
ingly more important for the protection of environment. Let us overview the
most important compounds, decreasing utilization of feed ingredients.
CEREALS COMPONENT TO DEGRADE. For the digestion of fibre fractions and
other non-starch carbohydrates pig and poultry do not produce efficient
enzymes. The majority of these cell wall components are mostly carbohy-
drates, but proteins and phenolic acids (polyphenol-ethers and ferulic acids)
are present, too. The carbohydrate subunits form cellulose micro-filaments,
which are surrounded by non-cellulose polysaccharides. The cellulose
chains of cellulose are built up by linking glucose subunits, which chains
are fixed by intra- and intercellular hydrogen-bonds, making resistant to
the digestive enzymes of mammals and birds.
The beta-glucans consist of glucose subunits of 1,4-β-bond. The structure
is supplemented by 1,3 β side-bond. This is the main difference, compared
to the cellulose, and owing to this structure, this polymer can form a vis-
cous solution. The concentration of beta-glucans is the highest in the bar-
ley and oats grains.
The arabinoxylans (or pentosanes) consist of a xylopiranosyl chain,
linked by beta-1,4-bonds, supplemented by 1,2- and 1,3-arabinofuranosyl.
The latter decreases the formation of hydrogen bridges; hence the com-
pound becomes water-soluble, forming a very viscous solution. The individ-
ual cereals differ from each other in the measure of arabinofuranosyl sub-
stitution, which determines their water solubility. Arabinoxylans occur in
high concentration in wheat, rye and triticale grains. Linking by the ferulic
acid, they form a complex of high molecular mass. On the contrary, ara-
binogalactans form complexes of low molecular mass, which in turn, may
bind proteins. Uronic acid can also been isolated from the cereal grains. At
the same time, the beta-glucan and arabinoxylan content of the cereal
grains, depending on the plant genotype and the circumstances of the pro-
duction, may vary within wide ranges (Table XV-1).
355
Table XV-1: Carbohydrate components of cereals fibre, % of DM (after HESSELMAN, 1989)
* as part of the cellulose ** as part of the hemicellulose *** as part of the arabi-
noxylan (pentosane)
The more viscous is the gut content (digesta), the lower is the blood
cholesterol level in poultry. These data have not only of comparative patho-
logical value, but indirectly suggest that the absorption of nutrients from
viscous gut content is lower. Addition of beta-glucanase enzymes decreases
viscosity.
NUTRITIONAL ROLE OF NON-STRUCTURAL CARBOHYDRATES. The ingested beta-
glucans and arabinoxylans, arriving into the stomach and intestine, dis-
solve, increasing the viscosity of gut content. It is supposed that the lower
nutritional value of rye, barley and wheat, compared to the corn grain can
be explained by this effect. The degree of viscosity is in close positive corre-
lation with the concentration of high molecular mass (>500,000 Da) carbo-
hydrates. The described phenomenon primarily is valid for the poultry,
because the viscosity of swine gut content is much lower.
Viscosity affects the efficacy of digestion by several means: it decreas-
es the diffusion of indigenous digestive enzymes in the gut content; the peri-
stalsis slows down, which in turn, negatively influences the appetite. In the
viscous gut content the degradation and absorption of nutrients diminish.
The slower peristalsis allows the proliferation of microbes and the bacterial
count of the small intestine increases. Undigested beta-glucans bind water
and excreted by the faeces, making bedding wet and smelly. Compounds,
causing viscosity, can be extracted from the grains using HCl-KCl solution.
Viscosity of this extract shows a direct relationship with the production
parameters of broiler chickens. By adding beta-glucanases to the feed mix-
tures, the differences between chickens, fed by barley of different viscosity,
will disappear (Table XV-2).
356
Table XV-2: Performance of broiler chicken fed low and high viscosity barley, with and-
without enzyme supplementation (after CAMBELL et al. 1989)
Supplementation of wheat-based chicken feeds by pentosanes

increases the AME value by 10 percent unit and the digestibility of protein
by 0.5 percent unit. Joint use of pentosanase and beta-glucanase is really
effective in decreasing the viscosity of gut content. The necessary enzymes
(endo-1,4-beta-xylanase, beta-xylosidase, beta-L-arabino-furanosidase,
endo-1,3:1,4-beta-glucanase, endo-1,4-beta-glucanase, beta-glucosidase
and beta-glucan-solubilase) are present in the nature as bacterial, yeast
and fungal products.
The enzyme supplementation gives better results in the performance
if the animals are young. Poultry in the first three weeks of age is especial-
ly susceptible to the growth depressing effect of beta-glucans and ara-
binixylans. Parallel to the advancement of age, the relative dry matter intake
capacity increases, the benevolent members of the gut flora proliferates and
the whole gastro-intestinal tract becomes mature. However, using a barley-
based diet, the pentosanase and beta-glucanase supplementation positive-
ly influenced the performance and the faeces consistency in older birds, too.
SPECIES CHARACTERISTICS OF SWINE CONCERNING THE USE OF THE ABOVE

ENZYMES. Since the dry matter content of the pig small intestinal content is
approximately the half of that of the poultry, the compounds, capable of

causing viscosity, will be diluted. In the ileum, due to bacterial activity,
there is a digestion of beta-glucanases. Strong acidity of the swine stomach
may also weaken the activity of added enzymes. Nevertheless, enzyme sup-
plementation may have positive effects using barley-based diet, especially in
weaned piglets.
357
Summarizing, the most important cause of the performance improve-

ment, owing to the enzyme supplementation is the decrease in the viscosi-
ty of the gut content. Beta-glucanases and arabinoxylanases relieve enclo-
sure of nutrients by the cell wall components; thereto the glucose
monomers of the degraded beta-glucans provide energy. Availability of pen-
tosan monomer is lower.
The alfa-amylase is used to improve starch digestion, because other-
wise in the young poultry, rabbit and foal, 10 to 40% of the corn starch gets
undigested in the caecum. Pigs generally produces sufficient amount of
amylase, but in case of early weaning or in liquid feeding system the addi-
tion proves to be efficient.
Alpha-galactosidase supplementation serves to improve the digestibil-
ity of oligosaccharides. Alpha-galactosides derive from the raffinose, sta-
chiose and verbascose and if getting undigested in the caecum, they under-
go microbial fermentation. The latter produces intestinal gases, which
means energy losses. The concentration of the mentioned compounds is
high in leguminous seeds (Table XV-3), besides this table, the lupine seed
worth mentioning.
Table XV-3: Oligosaccharide and fibre content of leguminous seeds, % (after CHEEKE, 1998)
PHYTASE. Addition of phytase enzyme to the feed improves the avail-

ability of phosphorus in cereals, bound by phytic acid. Consequently, the
mineral phosphate supplementation may be decreased, minimizing the
phosphorus loading of the environment by the lower emission. At the same
time, in human and companion animals, the phytate as a chelating agent
that binds iron and other metals, may serve to reduce the amount of free
radical production that occurs in the large intestine. Free iron forms free
radicals from lipids (lipid peroxidation). In addition to binding free radical
forming iron, phytate binds Zn, which normally induces DNA synthesis and
cell proliferation. So by binding Zn phytate reduces synthesis and cell
358
proliferation in the colonic wall, yet another anticarcinogenic effect of phy-

tate emerges. By bioengineering, the heat and acid tolerance of phytase can
be improved by manipulating of yeast; KIM et al. (2006) prepared single and
multiple mutants, using for the expression the Pichia pastoris yeast, of the
commonly used Aspergilus niger. In this way the pH optimum of PhyA phy-
tase shifted from pH 5.5 to more acidic side. As a result, the activity of
enzyme hardly decreases in the stomach, which was confirmed in animal
feed trial, showing a greater hydrolysis of soy phytate at pH 3.5.
SPECIAL FEATURES OF RUMINANTS. In one hand, the rumen may be con-
sidered as a disadvantage, because simple compounds, like sugars, some
cell wall components otherwise available in the small intestine for absorp-
tion, will be degraded. On the other hand, preparation of different industri-
al processes (bypass or protected proteins, amino acids, lipids, vitamins)
may escape rumen fermentation, absorbed from the small intestine and can
exert their new biological effect. As typical examples, the positive reproduc-
tive influence of rumen protected methionine and the lipotrop effect of pro-
tected niacin and choline in preventing fatty liver in cow. By mixing of
fibrolytic enzymes to the diet of periparturient cow, the adaptation of rumi-
nal flora and fauna can be supported.
The TASTE OF INDUSTRIAL DOG AND CAT FOODS is improved by hydrolysed
animal viscera (“digest”). For the production of digest, protease, lipase and
amylase enzymes are applied; as a result of the process peptides, free amino
acids, monoglycerids, free fatty acids, oligosaccharides and sugars emerge
(e.g. SPF-Diana products). The total of these compounds gives a taste,
which will be modified by the original raw materials of the mixture, the pos-
sible Maillard-reaction of proteins and of the occasionally added synthetic
preparation.
15.4.3. Application of plant extrats

Plant extracts are called also “botanical feed additives”; that of antimicrobial
activity have the name of “plant antibacterial” or “phytobiotics”, too.
359
Table XV-4: Active components and characteristics of the most important plant extracts
The majority of plant extracts is present in form of essential, volatile

oils. According to the biological characteristics of their active substances,
they can be applied as antioxidant, antimicrobial, fungicide, coccidiostatic
and/or immunostimulant -modulator feed additive. Table XV-4 displays the
most frequently used plants and their major features. Above the enumerat-
ed functions, some of them are simply natural food/feed preservative or col-
orant substances. For example, in case of rosemary clear evidence suggests
that the extract is less effective than the better antioxidants currently used,
but it is possible that identification of the active principle, such as the triter-
pene carnosol, may lead to synthetic manufacturing routes enabling pro-
duction of a new antioxidant. In this case, of course, it will be necessary to
prove the safety of such products, as it cannot be assumed that they mean
any hazards simply because they occur in nature. The antioxidant effect can
be detected in vivo, too: oil of thyme, fed by mice, was able to protect polyun-
saturated fatty acids and remove free radicals, delaying aging process (DEAN
et al. 1993). Some of them is used to improve liver function or as a prepa-
ration increasing the tolerance against (myco)toxins.
360
Although until now only on cell cultures and in experimental animals

is proved, the results are very promising, received by applying plant extracts
against free radicals and tumour formation. For example, the diallyl-sul-
phide of the garlic, the D-limonen of lemon and grapefruit, the epigallo-cat-
echin of the tea, the glycirrhetic acid of the sweet-root, the genistein of the
soybean seed, polyphenols (especially the resveratrol of the grape seed). The
knowledge of the active substances (Table XV-5) offers an opportunity for
making useful combinations.
Table XV-5: Biologically active substances and their concentration in the plants
Concerning the antimicrobial effect the comparison of minimal

inhibitory concentration (MIC) give an objective picture (Table XV-6), but
there are preparations of anticoccidial activity, too. The antimicrobial effect,
among others, is attributed to the short chain fatty acids (SCFA) and medi-
um chain fatty acids (MCFA) of plant extracts. (The latter have 6 to 12 car-
bon atoms.)
361
Table XV-6: The minimal inhibitory concentration (MIC) value of plant oils and olaquin
dox against food/feed microbes
*strong, synthetic antimicrobial compound
The MECHANISM OF ACTION of both fatty acid families is similar: non-

associated fatty acids enter microbes’ cytoplasm, change the pH and
exhaust the buffering mechanisms. Moreover, anions of the medium chain
fatty acids bind protons, produced by the respiratory chain (“uncoupling
effect”). The emerged non-associated fatty acid enter the protoplasm (see
above described effect), the proton will remain on the cell membrane, dis-
turbing the permeability and the ATP synthesis. In lack of energy, the lysine
and glutamic acid uptake of microbes decrease, finally the cell membrane
tears (open). All these damages quickly result in death of the bacteria.
Plant extracts are proposed for performance promoting and for the
prevention of diarrhoea, but they may also stimulate general resistance of
poultry, pig and calf. Dairy cows may be fed on such a mixture (nettle, car-
away seed, chamomile and dandelion), which improves the processing char-
acters (e. g, heat resistance, gelatinization capacity etc.) of milk. At the same
time, in case of ruminants, the use of plant extracts should be with pre-
caution, because some components of volatile oils, notably alcohols such as
linalool and terpineol are capable of reducing fermentive efficiency of
rumen, having adverse effect on cellulolytic bacteria (HAY and WATERMAN,
1993).
Exact composition of commercial product, owing to patent causes,
generally is unknown, which may raise authorization and quality control
concerns. For example, the producer (Belgian Vitamex) of the SPE’ for grow-
ing finishing pigs and the Galli d’Or for broiler chickens simply declares that
the active substances are medium chain fatty acids of 6, 8 and 10 carbon
362
number, deriving from a tropical plant. In most of the countries nutriceuti-

cals are classified in the category of GRAS (generally recognized as safe)
products and they are prescription-free. Above the mention product, are
available the Aldasyl (allycin active substance, producer: Europharma), the
Orego-Stim (oregano extract, produced by United Animal Health) and chest-
nut extract with hydrolysable tannins (Tanin Sevnica d.d.; Matischa Ltd).
CHESTNUT-STEVIA EXTRACT AND ITS ROLE IN NUTRITION

As the actual EU-regulations do not allow the use of antibiotics and antimi-
crobial agents as performance enhancer, the role of natural product is
becoming more and more important. One of these natural products is the
Farmatan product family. This extract, obtained from the natural chestnut
(Castanea sativa MILL) wood gained extract contains a series of estheric and
glycosidic tannins (elagitannins and other vegetable polyphenols, mostly
hydrolysable, but also condensed tannins). Tannins are astringent, bitter
plant polyphenols that either bind and precipitate or shrink proteins. The
terms tannin is widely applied to any large polyphenolic compound con-
taining sufficient hydroxyl and other suitable groups (mostly carboxyl) to
form strong complexes with proteins and other macromolecules. Tannins
have a molecular masses ranging from 500 to over 3000, and are generally
classified in hydrolysable and condensed tannins. Hydrolysable tannins
have a carbohydrate (e.g. D-glucose) in the centre. The hydroxyl groups of
the carbohydrates are totally or partially esterifies with phenolic groups
such as gallic acid in gallotannins or ellagic acid in ellagitannins.
Hydrolysable tannins are hydrolysed by weak acids or bases, resulting in
carbohydrate and phenolic acids. Condensed tannins (syn. proanthocyani-
dins) are polymers of 2 to 50 or more flavonoid units that are joined by car-
bon-carbon bonds, which are not susceptible to being cleaved by hydroly-
sis.
Tannins occur in the leaves and bark of many plants, tea, wine, fruits,
smoked foods, beer, citrus, fruit juices, some condiments, legumes and
chocolate.
The chestnut wood extract Farmatan is effective in preventing and
treating digestive troubles of animals, especially in the weaning period and
in stress situation. After coming into contact with tannins, the mucous
membrane of the gut becomes more resistant against pathogenic bacteria.
Tannins in the rumen slow down the protein degradation and increase the
363
bypass value of highly degradable dietary proteins, such as soybean or sun-

flower. As a result, the amount of amino acids, absorbable in the small
intestine increases, improving the protein supply of ruminants. Unlike con-
densed tannins that bind proteins through C-C binding, cannot be digested
and are mostly excreted in the faeces, hydrolysable tannins hold the protein
only between pH 4 and 7 and will be hydrolysed under the acidic conditions
of stomach, thus providing amino acids for the guest animal. GONZALES et al.
(2002) demonstrated that chestnut tannins are more efficient in protecting
soybean meal from in vitro degradation by rumen bacteria than are tannins
from quebracho and acacia.
The extract of species belonging to the genus Stevia (active com-
pound: steviol glycoside from the leaf, in the majority of cases Stevia rebau-
diana or candy leaf, sweetleaf (German nama: Süsskraut, Hungarian name:
jázmin pakóca) is a sweetener, that improves the acceptance of feed mixture
and lacks the possible diarrhoea-stimulating effect of sugars. Stevia has an
up to 300 times sweeter taste than sucrose and together with the wood sug-
ars, it positively influences the appetance of the supplemented feed mixture.
Therefore this composition (Farmatan®, Farmatan Plus® – sensory feed
additive from Tanin Sevnica d.d. and Farmitannin® from Matischa Ltd.) is
not only able to prevent and cure digestive disturbances and prevent coloni-
sation of the gut by pathogenic bacteria („immuno-facilitation”), but it also
optimizes feed intake.
Its inhibitory effect is selectively much more potent against patho-
genic bacteria, like E. coli and Clostridia, because the product deprives these
bacteria from the essential zinc and iron. Owing to this additive effect the
product is recommended for usealso in stress situations and, as a conse-
quence, it will also improve the performance of animals. According to the
opinion of the European Safety Authority (EFSA, 2004) Farmatan is totally
harmless. The results of other investigations suggest that it even improves
the insulin sensitivity.
Farmatan is also a very effective antioxidant, protecting the organism
among others immune cells, against the aggressive free radicals, and pre-
venting the DNA damage of leukocytes (SALOBIR, 2007). Therefore, the level
of antioxidants can be increased with Farmatan. In this way, the obtained
food product would get functional qualities and be protected from spoiling,
which is especially important for foods containing of higher levels of polyun-
saturated fatty acids (e.g. egg, milk and meat enriched with n-3 fatty acids).
364
15.4. Probiotics in poultry, pig and ruminant nutrition

Disease-preventive effect of the intestinal flora bears many names, for
example bacterium competition, bacterial interference and competitive
exclusion (CE). The latter will be used in the following. The early observa-
tion in this field was done by BLAXTER (1979): if with Salmonella typhimuri-
um or Salmonella gallinarum infected rats and chicks were fed on wheat
bran, the survival rate of animals increased. Namely, the ingested bran
enhanced the proliferation of Aerobacter species, producing enterobactin, a
salmonella resistance factor.
One of the most important fields of the application of the competitive
exclusion is the struggle against the poultry salmonellosis. The first experi-
ences showed that one-day-chickens became protected against S. infantis
and S. typhimurium if they were “treated” perorally with gut content of adult
birds. The total immunity develops during approximately 32 hours; howev-
er, the competitive antagonism is not able to prevent intestinal colonization
of the salmonellas, if the exposure is too high. At the same time, the colo-
nized chicken excreted significantly less salmonella than the control birds.
Both chicken and turkey intestinal flora have a certain protective effect at
the other species. Competitive antagonism can be induced by feeding sug-
ars (lactose, mannose) transforming in the gut into organic acids. The latter
are partly of antibacterial character, partly cover the binding sites if the
intestinal mucosa (see Prebiotics unit).
Botulisms are commonly caused by the ingestion of the pre-formed
toxin, produced by Clostridia spp. However, it has been proved in model tri-
als with mice and rats, that enterotoxaemic botulism may develop with the
proliferation of Clostridium botulinum and Cl. perfringens, unless a presence
of a normal intestinal flora. In broiler chicken the Cl. perfringens may cause
necrotic enteritis. VOROS et al. (2003) explain the first described in Hungary
FOAL GRASS SICKNESS by a proliferation of Cl. botulinum in the gut and the
absorption of its C-toxin. In rabbit with carbohydrate overload of the hind

gut, the Cl. spiroforme is capable of kill the animal by the produced iota
toxin (see Chapter XXIII).
Protective effect of competitive exclusion derives primarily from the
prevention of pathogenic bacteria to adhere the mucosa of the gastroin-
testinal canal. Notwithstanding, production of volatile fatty acids, bacteri-
365
ocins and other toxic metabolites, competition for nutrients and decreasing
of the redox potential may play a role, too. Lactobacilli are also capable of
adhering to the mucous membrane of the mouth and the epithelium of the
crop. Once settled, the protective bacterial flora is rather stable. Heat stress
and lack of drinking water disturb initial colonization in young animals.
Later, systemic infection by mycoplasma, viruses or coccidia may cause
competitive antagonism collapse.
According to the modern conception, the probiotics are living microor-
ganism of bioregulative character, applied perorally (SZIGETI, 1991). The pro-
biotics effects include the production of lactic acid, protection of mucous
membranes by forming biofilm, competitive exclusion of pathogenic germs
and production of vitamins and other growth factors.
The most important probiotic organisms are as follows: Bacillus sub-
tilis var. licheniformis and toyoi, Lactobacillus acidophilus, L. bifidus, L.
casei, L. lactis, Streptococcus bulgaricus, Str. faecium and Str. thermophilus
bacteria and the Saccharomyces cerevisiae yeast. In Europe is available
under the name of Lactiferm the freeze-dried form of Streptococcus faecium
M74 in milk powder-dextran carrier. Lactosacc contains Lactobacillus aci-
dophilus, Streptococcus faecium and Saccharomyces cervisiae. The water
soluble Entrodex consists of vitamins, electrolytes and Enterococcus faeci-
um. Spores of B. toyoi in limestone carrier (Toyocerin) and the Paciflor are
also on the market. The applied microorganisms are able to inhibit the pro-
liferation and colonization of pathogenic bacteria (E.coli, salmonella,
clostridia, and treponema). The first goal of using these preparations is the
prevention of diarrhoea of poultry, pig and calves, but the improvement of
weight gain and feed utilization are also expected.
A strain of the bier yeast Saccharomyces cerevisiae alone (Yea-Sacc®)
from Alltech or combined with lactobacilli is capable of stimulating rumen
function. There are promising investigations to apply lactobacillus cultures
(e. g. Broilac) in the exemption of poultry flocks from salmonella.
In the United States the FDA requires feed manufactures to use the
term “direct-fed microbials” instead of probiotics. The definition of direct-fed
microbials is as follows: “a source of live (viable), naturally occurring micro
oorganisms” (PEDELTON, 1992). They are considered both by the FDA and the
Association of American Feed Control Officials (AAFCO, 2000) as feed addi-
tives as generally recognised as safe (GRAS). (Author’s remark: similar to
the European qualified presumption of Safety, QPS, see FIGURE XVIII-1).
366
Nowadays 42 different microorganisms are recognised GRAS and are autho-

rised to used as direct-fed microbials: Aspergillus niger, A. oryzae, Bacillus
coagulans, B. lentus, B. lichenformis, B. pumilus, B. subtilis, Bacteroides
amphophilus, B. capillosus, B. ruminocola, B. suis, Bifidobacterium adoles-
centis, B. animalis, B. bifidum, B. infantis, B. longum, B. thermophilum,
Enterococcus (or previously: Streptococcus) cremoris, Enterococcus diacety-
lactis, E. faecium, E. intermedius, E. lactis, E. thermophilus Lactobacillus aci-
dophilus, L. brevis, L. bulgaricus, L. casei, L. cellobiosus, L. curvatus, L. del-
bruckii, L. fermentum, L. helveticus, L. lactis, L. plantarum, L. enterii,
Leuconostoc mesenteroides, Pediococcus acidilactici, P. cerevisiae, P. pen-
tosaceus, Propionibacterium freudenreichii, P. shermanii and Saccharomyces
cerevisiae.
15.4.5. Feeding of organic minerals

Organic metals, bound bond to proteins, amino acids, or nucleoproteids of
yeasts can readily be absorbed, transported across the placental barrier
than that of the inorganic forms. In some cases the accumulation in a par-
ticular tissue (e.g. horse hoof) can also be achieved in this way. The most
important are in this group the glycine, methionine, lysine, histidine or
caseinate chelates of zinc, iron, manganese and copper, as well as the nick-
el-histidine chelate. Yeasts, enriched by zinc, selenium and chromium are
also available. The chelated form of minerals are able to increase the tissue
concentration in a targeted way, improving the special physiological func-
tion (fertility, male reproduction, immune response, blood composition, hoof
integrity etc.) in a very effective way. Especially the yeast-based minerals
proved to be highly profitable in the feeding praxis (e.g. Sel-Plex® and
Bioplex® of Se, Fe, Cu, Mn and Zn from Alltech).
15.4.6. Prebiotics
The prebiotics are indigestible, carbohydrate-type compounds, which help
the proliferation of useful members of intestinal flora and inhibits at the
same time the attachment and proliferation of pathogenic germs. Already in
1990, PUSZTAI et al. proved that even plant lectines may exert a probiotics-
like effect. In their experiment they added lectines and/or glycoconjugates
of plant origin into the feed of suckling piglets. The mentioned compounds
inhibited the adhesion of pathogenic intestinal bacteria, either by competi-
tive exclusion (antagonism) or by modifying the expression of surface anti-
367
gens in the mucous membrane. Phosphorylated glucomannan of yeast cells,

the MOS (mannan oligosaccharides) also competes with pathogenic microbes
for the binding sites (the carbohydrate moieties of glycoproteins on the sur-
face of the host cells) of the mucous membrane and decreasing their colo-
nization opportunities by the developed “carbohydrate layer”. (The most fre-
quent bacterial adhesion factors are the Type I fimbriae on the surface of
Gram-negative bacteria, having great affinity for mannose-containing glyco-
proteins on the surface cells.)
Moreover, the pre-treated yeast or the synthetic mannan oligosaccha-
rides attach to the surface of the pathogenic bacteria too and this complex
is excreted by the faeces. MOS is a growth promoter of the intestinal villi
and has both locally and systematically an immunostimulant effect. Thus,
the MOS compounds, having lectin-like feature in recognizing binding site
of gut mucosa. Jerusalem artichoke (Helianthus tuberosus), sugar beet pulp
and yeat are rich sources of MOS. By using MOS-containing products like
Bio-Mos® (Alltech), the blocked mucosal binding sites are not available for
the pathogen bacteria and the state of eubiosis can be maintained for a long
time. The general consequence is an improvement of immune status, which
results in an increase of immunoglobulin level in dam’s milk, the prevention
of diarrhoea and promoting growth in young animals (e.g. piglets).
The fructo-olygosaccharides (FOS) and the transgalacto-olygosaccha-
rides (TGOS) are short-chained (2 to 10 units) polysaccharides with β-(2-1)
oxidic bond and with a terminal glucose unit found naturally in cereal
grains and leguminous seeds. It can be industrially produced from the
pectin of sugar beet or apple, or by the partial hydrolysation of the inulin,
found in high concentration in Jerusalem artichoke, artichoke, chicory, or
in purple coneflower (Echinacea purpurea) or by means of an enzymatic
treatment (transgalactosidation) of the sucrose. Their positive effect is that
they can serve as nutrient for the useful members (e. g. lactobacilli, bifido-
cacteria) of the intestinal flora. Beneficial effect reflects in the intestinal
morphology, too: the villus height and the villus to crypt ratio increases in
the treated pigs.
Preparations, containing both probiotics and prebiotics, usually are
called SYMBIOTICS OR SYNBIOTICS. While designing these products, the choice
of appropriate bacterial count is of great importance, because the exagger-
ated microbial bile acid deconjugation in gut lumen may cause growth
depression. Many of the protein degradation products (ammonia, biogenic
368
amines, branched-chain fatty acids, indols, phenols and sulphur-contain-

ing compounds) of the hind gut have an unpleasant odour. By feeding pre-
biotics (mostly MOS and FOS together), the production of undesirable com-
pounds can be suppressed, which is of paramount importance in case of
indoor-kept dogs and cats.
15.4.7. “Fat burners” for more lean meat

The L-carnitine received its name from the Latin word “carne”, meaning
meat, because it was isolated first from this tissue. L-carnitine as a tri-
methylated compound, which is synthesized mainly from the lysine. L-car-
nitine may be conditionally essential for neonates and parenterally fed
patients. Its main role is to help the long-chain fatty acids inside the mito-
chondria for being burned. To increase the lean meat production, the pro-
posed dosage for monogastric animals is 50 to 200 mg/kg air-dry feed. In
the experiment with rats FEKETE et al. (2001) had no effect; in broiler chick-
en trial they found only a slight improvement of the production parameters.
According some data, the supplementation of pregnant sows’ fed by L-car-
nitine may increase the number of muscle fibres in the newborn piglets.
The trivalent (organic) chromium has been first isolated from human
mother milk, in form of chromium picolinate; the commercial preparation is
the relative compound, the chromium nicotinate. When during the yeast
production, inorganic chromium salts are given to the biomass, the yeast
cells incorporate it in organic bond. This type of chromium-enriched yeast
may also used as a feed additive. Trivalent chromium is part of the glucose
tolerance factor (GTF) and by that means improve insulin function. The fat-
burner effect is partly explained by an indirect negative feed-back of insulin,
partly by an independent mechanism (NRC, 1997). The proposed dosage for
growing-finishing pig is 2 to 5 mg/kg feed mixture. In the experiment of
FEKETE et al. (2001) with conventional rats the chromium nicotinate sup-
plementation improved voluntary feed intake and weight gain, but paradox-
ically most of the increment was fat. While giving chromium picolinate for
breeding sow, the ovulation rate and litter size increased, because chromi-
um picolinate potentates the action of insulin. Organic chromium, L-carni-
tine and vitamin A supplementation of the obese dog and cat feed facilitates
the slimming process and help in preventing the hepatic lipidosis. For hors-
es and cattle (mostly calves) the organic chromium supposed to have
immunostimulant effect.
369
Betain (trimethyl glycine is also a potential performance enhancer. It

may serve as methyl donor and proved to be effective in poultry, fish and
swine, improving nitrogen and energy retention. As betain-hydrochloric,
help in the post weaning hypochlorhydria of piglets. Some chemical forms
of the conjugated linoleic acids (CLA) seemed to increase lean meat produc-
tion and improvement of fat quality and firmness in slaughtering pig. In a
very experimental phase, there are data about the transitory immunisation
against leptin in order to improve average daily gain and protein accretion
in growing-finishing pig.
15.4.8. Peroral passive immunization

Dried blood plasma. The specially dried blood plasma (SDBP) of previously
immunized adult animals, besides the albumin and globulin fractions, con-
tains the immunoglobulins against the pathogenic bacteria, too. These
preparations are fed by piglets, some days before and after weaning, to
decrease colonization of harmful microbes (e.g. entheropathogenic coli and
salmonella) in the intestinal tract.
Egg-yolk antigen (IgY). Antibodies, produced by laying hens after tar-
geted immunization get into the egg yolk. The glycoprotein-type
immunoglobulin yolk (IgY) undergoes freeze-drying and is commercialized in
form of microcapsules. The preparation is fed by weaning piglets to prevent
diarrhoea, caused the pathogenic enterobacteria.
15.4.9. Miscellaneous
Glycocomponent of the originally plant (Yucca shidigera) extract sarsaponin
is capable of binding ammonia both in the gut and in the environment. This
reaction indirectly may improve animal performance
Among the poly-unsaturated fatty acids (PUFA), the timnodonic acid
(EPA=eicosapentaenoic acid, C20:5) and the clupanodonic acid (docosa-
hexaenoic acid, C22:6) are called omega-3-fatty acids. They can practically
be found only in fish oils and animal fats. Previously they were supposed to
be able to decrease fat deposition in the abdominal fat pad of broiler chick-
ens; the related experimental evidences are contradictory. Their function is
more general, they are responsible for the integrity of cell membranes. The
proposed level of supplementation in breeding swine (boar and sow) feed is
1%, in poultry feed mixtures 1 to 2%. The expected positive effects are the
improved sperm quality, increased litter size and weight, healthier state of
370
the retina and brain tissue in the newborn animals and a decline in the
intensity of inflammation processes. Laying hen, fed on omega-3-fatty acid-
containing diet, have an improved feed utilization, higher number of eggs
and better hatchability of breeding eggs. In the later case the roosters
should receive the same feed. In flocks, producing eating eggs, only high
quality, and refined fish oil can be used, to avoid the fishy taste and smell
of eggs. Higher dosages (5%) of omega-3-fatty acids are able to counterbal-
ance harmful effects of Eimeria tenella infection.
While supplementing of dog and cat diets by omega-3-fatty acids, the
optimum ratio of omega-3-fatty acids to omega-6-fatty acids in their diets is
1 to 5-10. If the occurrence of omega-6-fatty acids is higher, the inflamma-
tive processes prevail. The expected benevolent effects are similar to that in
pigs. It worth mentioning the balanced development of the young animals’
brain and nervous system, a decrease in the incidence of the inflammatory
phenomena, like atopic dermatitis and improved wound healing.
15.4.10. Stress – Immune status – Performance enhancers

Modern, industrialized animal production, even besides a good and profes-
sional management, means an important stress to the animals. The
response on stressors of the organism consists of three periods, the adren-
alin (epinephrine) and noradrenalin (norepinephrin) dominated, relatively
short alarm reaction, the ACTH and cortisol-controlled resistance and the
final exhaustion, depression phases. The cortisolinaemia, among others,
causes immunosuppression, too. The latter may be aggravated by some feed
mycotoxins, like T-2 toxin. Mycotoxins decrease the absorption of vitamins,
too. Considering the above described, those feed additives are more and
more preferred, which support the natural resistance systems of the ani-
mals and help the prevention of diseases and subclinical ailments. In this
way animals will be able to produce according to their genetic capacity. As
some of the most important substances of this type, the increased addition
of vitamin E, vitamin C, selenium (mostly in organic bond, e.g. Sel-Plex®
from Alltech), toxin adsorbents and/or binder (e.g. Mycosorb® from Alltech),
immunostimulant and immunomodulant preparations (e.g. nucleotids).
15.4.11. Nucleotides and their role in nutrition

The impact of nucleotides in cellular biology is dominating more and more
the research in natural sciences and medicine during the last decades. The
371
predominant role of these molecules in cells is primarily in gene duplication

and protein synthesis, but more recently the key role of small RNA mole-
cules in gene regulation has been uncovered. This means that nucleotides
are not only involved in all processes related with the proliferation and func-
tionality of cells but also have major impact on specific regulatory mecha-
nisms triggering the expression of genes within cells. This fundamental
importance explains the effects of nucleotides on health, cell maintenance,
immunity and growth regulation of living organisms. The universal use of
nucleotides for the storage of the genetic information as well as for efficient
gene regulation and expression in multicellular organisms and protozoan
highlights their evolutionary grown importance for every living organism. It
moreover explains the apparently widely different effects that have been
observed when nucleotides are included in the diets of farmed animals.
Structure and function. A nucleotide consists of a base (either a purine
such as adenine, or a pyrimidine, such as cytosine) linked to a sugar (either
ribose or deoxyribose) and up to three phosphate residues. Nucleotides as
the basic building blocks of DNA and RNA are especially important in tis-
sues with rapid turnover like for example, gut and immune system.
Moreover they are metabolic regulators involved in energy transfer, „acid”
handling, synthesis and breakdown of large molecules.
Nucleotides are not considered „essential” nutrients as they can be
synthesised in most cells of the body. However, during certain times of high
demand for nucleotides, the cellular synthesis may not be able to meet this
increased demand. This mostly occurs for example, during periods of rapid
cell turnover e.g. in young animals or in animals challenged by bacterial,
viral or parasite infections. At these times, the body needs to rely more on
dietary sources of nucleotides to ensure the unhindered supply of these
molecules. Therefore, nucleotides must be are regarded as „conditionally
essential” nutrients. Polynucleotides (DNA and RNA) contain up to several
1000 nucleotides and oligonucleotides consist of only some nucleotides
(RUDOLF, 1994).
Here it seems to be useful to clarify the nomenclature of the yeast cell
wall components accepted in natural science and medicine (FLASCHENTRÄGER,
1951). Nucleic acids are the protein-free elements of the cell nucleus. Purine
and pyrimidine nucleotides are parts of the nucleic acids. The nucleopro-
372
teins consist of nucleic acids associated with an alkaline protein compound

(protamine). Nucleosides are decomposed parts of mono- and polynu-
cleotides.
Many yeast components have immuno-stimulatory effects (GLAWISCHNIG
and PEDIT, 1992; VAN BUREN et al. 1983; KULKARNI et al. 1986; COFFEY, 1988;
RUDOLPH et al. 1984), showing that lymphocytes and enterocytes require
purine and pyrimidine for growth. The absence of nucleotides tends to
cause an arrest in the cell cycle of T-helper cells between G0-G1 as well as
S-phases (WALKER, 1994). T-helper cells and T-lymphocytes need purine and
pyrimidine bases for maintaining their immunological function. Nucleotide
supplementation of feed improved survival of BALB/c mice intravenously
injected with Staphylococcus aureus or Candida albicans (VAN BUREN et al.
1994).
Polynucleotides and RNA in vitro stimulated T-helper cell-mediated B-
cell antibody responsiveness using murine spleen cells and human periph-
eral blood mononuclear cells (JYONOUCHI, 1994). Moreover, the IL-2 produc-
tion and the natural killer activity is also elevated (CARVER, 1994). In these
experiments mononucleotides were not tor less effective.
The immuno-modulation is especially useful in order to improve the
antigen production after vaccination against a given disease. The best
results (high and persistent antibody titres) can be expected if the
nucleotides were applied before, during or even after vaccination (KOEPPEL,
1992).
To test the effects of nucleotide supplementation on crypt villus differ-
entiation in vitro, human cancer cell lines (Caco-2) and non-malignant
rodent cell lines (IEC-6) were used. Cell differentiation was measured by
means of brush-border enzyme activities. Nucleotides supplementation ge-
nerally enhanced enterocyte gene expression in IEC-6 cells; in case of the
carcinoma Caco-2 cells this effect became obvious under dietary stress only,
i.e. while using glutamine subducted media (SANDERSON and HE, 1994).
UAUY and STRINGEL (1988) as well as UAUY et al. (1990) found that the
intestinal mucosa and the concomitant enzyme activities are more devel-
oped in nucleotide-fed rats. As a consequence, the DNA levels in enterocytes
determined by cell counts of these cells and the villus height, calculated
from the crypt-villus ratio are both improved.
Hydrolysation and extraction of yeast (especially the brewers yeast,
(Saccharomyces cerevisiae) help to increase the nutritional availability of
373
valuable components such as proteins, amino acids and nucleotides. The

processes used are heating, swelling, cracking, drying, occasional applica-
tion of enzymes, autolysis and finally grinding. The result is a simple yeast
extract containing plenty biologically active compounds with high immuno-
logic impact. First of all the glucans or mannanoglucans should be men-
tioned. These are present in the yeast cell wall and have an immunostimu-
latory effect. In the given natural configuration and binding form, the glu-
comannanes are able to aggregate with adhesive parts on the surface of sev-
eral Enterobacteria species in the intestinal lumen but not in the stomach
and thereby they may block or inhibit their microbial activity. Other com-
pounds of yeast such as nucleic acids, purines and pyrimidines have
immunostimulatory effects; moreover they enhance enzyme activity and
intestinal secretion. Summarizing, different integral parts of yeast are
accepted as being nutrients having stimulating effects on the gut flora.
Some oligonucleotides inhibit the deamination of undesirable intestinal
microbes, therefore sparing amino acids in the intestinal tract and simulta-
neously decreasing the ammonia production in the gut. Other components
of yeast cell are supposed to influence and optimize the enzyme functions of
gut mucosa. Consequently, yeast hydrolysates are valuable components of
dietetic mixtures to improve the immune status of and promoting eubiosis
in the intestinal flora.
Besides this positive effect on the colonisation with beneficial microflo-
ra, there are data about enhanced liver regeneration after D-galactosamine-
induced injury (OGOSHI et al. 1988) and stimulated fatty acid metabolism
upon nucleotide application with the diet. The latter may prevent lipid accu-
mulation in organs and blood (CARVER, 1994). In babies fed nucleotide-sup-
plemented diets, the ratio of indigenous gut flora (especially bifidobacteria)
is improved (UAUY, 1994). Mother’s and dam’s milk (except cow’s milk) are
excellent sources of nucleotides (BARNESS, 1994).
Dietary nucleotides are reported to play a decisive role in the mainte-
nance of immune responses. Exogenous nucleotides supplied in the diet
contribute to the pool of nucleotides available to stimulated leukocytes,
which show a rapid turnover (short shelf life) and therefore increased
nucleotide requirements. (KULKARNI et al. 1994). Induction of lymphocyte
proliferation is accompanied by a drastic increase of the intracellular
nucleotide pools and the expression of a large number of transmembrane
nucloside transporters (SMITH et al. 1989).
374
By this regulatory way nucleotides support the organism to overcome

infections, digestive challenges and metabolic disturbances. The presence of
nucleotides in the lumen (with exception of adenosine) prevents ischemia,
decrease leukocyte extravasations as well as protein and nitric oxide pro-
duction (BUSATAMANTE et al. 1994). All this leads to the conclusion that
nucleotides enhance beneficial drug effects and prevent potential immuno-
suppressive side effects (e.g. mycotoxins).
In studies with burn and surgical post-trauma cancer patients, intra-
venous and enteric nucleotide supplementation reduced postoperative com-
plications (VAN BUREN et al. 1994). The incidence and severity of post-radia-
tion gut injuries decreased, but the lesions developed more rapidly if exoge-
nous nucleotides were provided. In a recent broiler chicken trial in Hungary,
applied water soluble nucleotides, higher villi with less dense lymphocyte
infiltration have been found (SZABO, GLAVITS, MATISA and FEKETE, 2008) . UAUY
et al. (1993) supposed that excessive nucleotide ingestion may lead to the
production of free oxygen radicals after conversion of hypoxanthine to urate
catalyzed by xanthine oxidase in the gut wall. In contrary to this, SALOBIR et
al (2005) found that feeding piglets with nucleotides prevented the damage
of DNA of lymphocytes by oxygen radicals after feeding linseed oil with high
PUFA content. The same effect was observed in leukocytes of chicken fed
with a diet containing mycotoxins. (FRANKIC et al, 2006)
Today nucleotides must be considered as semi-essential nutrients,
under some physiological and/or pathological conditions even as essential
nutrients. Consequently, the different nucleotide-containing cell extracts of
yeast may have immuno-stimulatory effects and a certain role in the for-
mation of functionally active lymphatic tissues and epithelia of mucous
membranes. Moreover there are feed additives available on the market, con-
taining hydrolysed yeast or yeast extracts and including free (purified)
mono-, oligo- and polynucleotides. These are effective in poultry, pig, fish
and crustacean production exploiting the beneficial effects of nucleotides in
livestock and aquaculture (e.g. Nu-Pro® – Alltech, Ascogen® – Chemoforma
and Matischa, Nucleoforce® – Bioiberica etc.).
375
FOR FURTHER READING
AAFCO (Association of American Feed Control Officials) (2000): Official Publication. POB.
478. Oxford, IN 47971.
Arnaud, A., Fontana, L., Angulo, A.J., Gil, A. and López-Pedrosa, J. M. (2003): Exogenous
nucleosides alter the intracellular nucleotide pool in hepatic cell cultures.
Implications in cell proliferation and function. Clin. Nutr. 22, 391-399.
Rodriges-Serrano, F., Marchal, J.A., Rios, A., Martinez-Amat, A., Boulaiz, H., Prados, J.,
Perán, M., Caba, O., Carillo, E., Hita, F. and Aránega, A. (2007): Exogenous nucleo-
sides modulate proliferation of rat intestinal epithelial IEC-6 cell. J. Nutr. 137879-
884.
Barness, L.A. (1994): Dietary sources of nucleotides – From breast milk to weaning. J. Nutr.
124, 128S-130S.
Barness, L.A. (1994): Dietary sources of nucleotides – From breast milk to weaning. J. Nutr.
124, 128S-130S.
Blaxter, K. (1079): The role of nutrition in animal disease. Vet. Rec. 104:595-598.
Brown, D.L., Scott, J.T., DePeters, E.J. and Baldwin, R.L. (1989): Influence of somettribove,
USAN 8Recombinant Methionyl bovine somatotropin) on the body composition of lac-
tating dairy cattle. J. Nutr. 119:633-638.
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(1994): Dietary nucleotides: effects on the gastrointestinal system in swine. J. Nutr.
124, 149S-156S.
Carver, J.D. (1994): Dietary nucleotides: cellular immune, intestinal and hepatic system
effects. J. Nutr. 124, 144S-148S.
Cheeke, P.R. (1998): Natural toxicants in feeds, forages, and poisonous plants. Interstate
Publ. Inc. Danville, IL..
Feed Additive Compendium from (2000):Volume 38, Miller Publishing Co. Minnetonka, MN
in cooperation with The Animal Health Institute, Alexandria, Virginia
Fekete, S. (1998): Complex study of biological effect of Farmatan, a feed and drinking water
additive in the broiler chicken and rabbit rearing (In German with Croatian summa-
ry). Krmiva (Zagreb), 1998. 5, 235-251. .
Fekete, S. and Hullar, I. (1994): New ways in the growth promotion of monogastrics: admin-
istration of enzymes and plants extracts. Compilatory account. (in Hungarian, with
English summary). Magyar Állatorvosok Lapja, 49, 489-492.
Fekete, S., Szakáll, I., Kósa, E., Andrásofszky, E., Fodor, K., Hidas, A. and Tozser, J. (2001):
J.: Alteration of body composition in rats: effect of organic chromium and L-carnitine.
Acta Vet. Hung. 49, 385-398.
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nucleotides in reduction of DNA damage induced by T-2 toxin and deoxynivalenol in
chicken leukocytes. Food and Chem. Toxicol. 44, 1838-1844. Gonzales, S., Pabón,
M.L. and Carulla, J. (2002): Effects of tannins on in vitro ammonia release and dry
matter degradation of soybean meal. Arch. Latinam. Prod. Anim. 10(2), 97-101.
Gonzales, S., Pabón, M.L. and Carulla, J. (2002): Effects of tannins on in vitro ammonia
release and dry matter degradation of soybean meal. Arch. Latinam. Prod. Anim.
10(2), 97-101.
Hay, R.K.M. and Waterman, P.G., eds. (1993): Volatile oil crops: their bio;ogy, biochemistry
and production. Longman Scientific & Technical. Harlow, Essex
http://www.alltech.com
http://www.bioberica.com/dig_mejora_prod4, _prod5.asp
http://www.chemoforma.com
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138S-143S.
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A.H.J., Daly, C.B., Welch, R. and Lei, X,G, (2006): Shifting the pH of Aspergilus niger
PhyA phytase to match the stomach pH enhances its effecteveness as an animal feed
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Koeppel, P. – Chemoforma (1992): Immuno-modulation. Poultry Intern.April, pp. 58-62.

Kulkarni A.D., Rudolph F.B. and, Van Buren L.A. (1994): The role of dietary sources of
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testinal tract of piglets. Asian-Aust. J. Anim. Sci. 18, 1353-1362.
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Washington, D.C.
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(1988): Effect of total parenteral nutrition with nucleoside and nucleotide mixture on
D-galactosamine-induced liver injury in rats. J. Parenter. Enter. Nutr. 12, 53-57.
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avian adipose tissue. Poult. Sci. 62. 1838-1845.
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(2007): Effect of natural extract of chestnut wood on digestibility, performance traits,
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Walker, W.A. (1994): Nucleotids and nutrition: role as dietary supplement. J. Nutr. 124,
121S-123S.
377
is a tannin extract from Sweet Chestnut Wood (Castanea Sativa
Mill.). Natural antibiotic. For all kind of animals. It is health promotor,
environment improver and performance enhancer.
Modified products of Sweet Chestnut extract:

- Farmatan plus: Sweet Chestnut extract and natural sweetener
- for young animals and pets;
- Acidad Dry: Sweet Chestnut extract and different acids
- for pigs, rabbits and broilers;
Certificates:
Producer:
Tanin Sevnica d.d.
® Hermanova cesta 1
8290 Sevnica, Slovenia
tel.: + 386 7 81 64 455
tel.: + 386 7 81 64 425
fax: + 386 7 81 64 445
e-mail: tajnistvoj@tanin.si
www.tanin.si
015-016 közötti színes.indd 379 2008.09.03. 15:08:58

JAV_16_VND_FSGy1:MINTAÚJ.qxd 2008.09.03. 15:11 Page 381
Chapter XVI
FEEDING–REPRODUCTION RELATIONSHIP

16.1.1. Nutrient supply and puberty
16.1.2. Ovulation rate
16.1.3. Feeding, fertilisation and embryonic survival
16.1.4. Feeding and foetal growth
16.1.5. Mammary development and colostrum production
16.1.6. Feeding and newborn viability
16.1.7. Feeding and parturition to rereconception interval
16.1.8. Feeding and postpartum hormonal changes
16.1.9. Role of mycotoxins and heavy metals on the reproduc
tive functions
16.2. Endocrine and metabolic factors of reproductive functions

of dairy cows
16.2.1. Negative energy balance (NEB) in postpartum dairy
cows
16.2.2. Puerperal endocrine changes
16.2.3. Resumption of cyclic ovarian function during the neg
ative energy balance
16.2.4. Signalling the metabolic state: metabolic hormones
in regulation of ovarian function
16.2.5. Factors influencing fertility in postpartum dairy cows
16.2.6. Bacterial complications of uterine involution
16.2.7. The effect of circulating progesterone level in already
cyclic cows
16.2.8. Post-ovulatory rise of progesterone in inseminated
cows.
16.2.9. The quality of oocytes
Chapter XVI: FEEDING - REPRODUCTION

Introduction
The professional opinions are not univocal concerning the earliest develop-
mental phase of the animals, whose nutrition may already have an influ-
ence upon the later reproductive performance. During the foetal period the
energy deficit has no irreservible effect on the development of primordial
germ-cells. The primordial follicles may not be susceptible to the low toxin
effects, either. On the contrary, in pregnant ewes a moderate undernutrition
during the last six weeks of pregnancy and during the lactation already sig-
nificantly alters the ovulation rate of the offspring’s in adulthood. Excessive
consumption of beef meat by women during pregnancy may alter a male’s
in utero testicular development and compromise his future reproductive
capacity having significantly lower sperm than that of men whose mothers
ate less beef (SWAN et al. 2007). The basic concept of the above phenomenon
is the „foetal programming“ which means that maternal stimuli during the
foetal development has long-lasting impacts on progeny postnatal growth
and physiology (BARKER et al. 1993). The uncertainty extends only on the
long-term relationship of feeding and reproduction, because the short-term
and mid-term effects of nutrition on the reproduction there are sufficient
amount of information. Therefore, in this subchapter only the reproductive
effects of the feeding as a whole, that of the energy and protein supply will
be discussed and the role of minerals, vitamins, mycotoxins and heavy met-
als will be treated only marginally.
16.1.1. Nutrient supply and puberty

16.1.1.1. Females. A nutritional supply of 50 to 70% of the real needs delays
the sexual maturation in cattle, deer, ewe, goat, gilt, rabbit and rat (SEKI et
al. 1997). This effect is the more expressed the younger the animals are. In
the seasonal breeders an incorrect grazing technology can lead to insuffi-
382
cient nutrient supply and the onset of the puberty may be postponed by a
whole year. Less severe lack in nutrient supply delays the onset of puberty,
the re-assumption of the cyclic ovarian activity after the first calving occurs
later and the chances of reconception are bad (HUSZENICZA et al. 1989). The
physiological ranges are broad, this is why the live weight at the puberty of
cattles, even in the same ruminant breed, is within wide ranges. From the
major chemical components this is the fat concentration which has a defin-
itive influence on the sexual maturation. As described in the cases of
human, rabbit doe and gilt, the fat percentage should achieve a “threshold
value” to enable the onset of the first oestrus (see the importance of leptin,
too).
The experiences sometimes are contradictory, because the influence
of energy supply can be modified by the protein intake. For example, a rel-
atively high protein (16% crude protein) diet may assure the onset of the
puberty in term even given only a small daily ration. According to rodent
and gilt studies the tyrosine supplementation could advance the sexual
maturation. The possible explanation is that the tyrosine may play the role
of the precursor of neurotransmitters like dopamine and noradrenalin. The
low protein content (less than 8% of the dry matter) of tropical pastures
delays growth rate and sexual maturation of the there grazing ruminants.
Most of the mineral and vitamin undernutrition postpone the onset of first
oestrus through the retardation of growth.
16.1.1.2. Males. The relationship of feeding, growth (average daily gain) and
sexual maturation is more simple and direct in males than in the females.
Males raised on a high nutrition plan have a higher growth rate and will
attain puberty early and with a high live weight. Similarly to the females,
the more intensive the feeding influence on reproduction the younger the
male animals are; therefore, the suckling age is the most susceptible peri-
od. In growing rats fed on 55% of the ad libitum level Leydig cell atrophy was
found in the testes. A reduction a sperm production and concentration has
been observed in rams fed on 75% of the maintenance ration for 5 to 6
weeks. At the same time, overweight of breeding boars results in leg weak-
ness and reduction in the libido. On day of services 1.3 MJ ME (approxi-
mately 100 gram mixed feed) surplus should be given above the mainte-
nance energy and protein supply, otherwise the sperm production but not
the quality may significantly drop. Sperm production and quality of buck
rabbits raised with a supply of 70% of the energy and protein requirements
383
did not change; but the libido decreased (FODOR et al. 2002). Zinc has a
direct role in the spermatogenesis, besides it is indispensable to the vitamin
A uptake of the spermatozoa. As the vitamin A concerns, it is important
alone in achieving the puberty, the maintenance of libido and the integrity
of germ epithelium in the testes.
16.1.1.3. Feeding factors of sexual maturation. It is generally accepted that
in the sexually mature females the poor energy supply inhibits the pulsatile
gonadotropin releasing hormone (GnRH) from the hypothalamus and there-
by the consecutive luteinizing hormone (LH) release in the pituitary gland.
High GnRH and pituitary gonadotropin blood concentration cause the break
of pulsatile GnRH release. Undernutrition suppresses the secretion of LH
and follicle stimulating hormone (FSH). This phenomenon can be elicited in
some days. In ovariectomized ewes the pulsatile LH release can be main-
tained by parenteral applied mixture of dextrose and amino acids exactly
the same as by an abundant feed supply. Thus GnRH-producing neurons
are highly responsive to the nutritional status of the animal. In the trans-
mission of the effect of energy providing nutrients the role of insulin may be
important, because it is linked to the receptors of those brain areas (nucle-
us arcuatus and eminentia mediana) that participate in the regulation of
GnRH secretion. In case of the protein alone the role of amino acids may
have a role, which is necessary to the synthesis of GnRH neurotransmitters.
The undernutrition exerts its detrimental effect on GnRH secretion and
therefore on the sexual maturation directly and immediately by the nutrient
deficiency as well as long-term, indirectly by the retardation of bodily growth
and development and by decreasing the responsiveness to the steroids of
gonads.
16.1.2. Ovulation rate

16.1.2.1. Effect of feeding level. The, under practical circumstances occur-
ring fluctuations in the nutrient supply have a significant influence on the
ovulation rate in case of sheep and goat; this effect is less expressed in pig
and negligible or does not exist in rabbit. Long-term effect of feeding could
reflect in the body condition. For example ewes kept on mountain or tropi-
cal pastures frequently have less than 10% total fat in their carcass and
thereby the ovulation rate remains below one. By abundant feeding their fat
content can be increased up to 25% causing the doubling of ovulation rate.
The influence of body condition is considerable in case of the prolific sheep
384
breeds of the flatland: if the total body fat is 20%, the ovulation rate is 2.3;
while measuring 30% fat, it is 3.4. The jerky, excess nutrient intake (the so-
called flushing) prove to be efficient only in case ewes of medium and poor
condition and it means 0.2 to 0.4 ovulated egg-cell surplus on an average.
The long-term effect of nutrient supply is more expressed in case of adult
ewes than in case of tegs. This tendency is the opposite in case of sows and
gilts. In this way by increasing the daily energy supply from 21 MJ ME to
34 MJ ME resulted in 1.5 higher ovulation rates in gilts. Different nutrient
supply during the first lactation of gilts will change the live weight but has
no influence on the subsequent ovulation rate. Similarly, flushing between
the weaning and reconception has no influence the number of ovulated egg-
cell. On the contrary, the daily higher ration during the oestrus conse-
quently increases the ovulation rate in a small extent.
16.1.2.2. Effect of the individual nutrients. In case of the pig the is a lot of
studies were carried out concerning not only the influence of energy supply
on the ovulation rate, but also concerning the possible effects of protein
intake, lysine concentration or fat level, but no increased ovulation rate
occurred while compared to animals fed on a concentrate of average com-
position. Although it is well known that the mineral and vitamin deficien-
cies deteriorate reproductive performance, but evidences about the effect on
the ovulation rate frequently are indirect. For example, on the improving
effect of zinc supplementation can be concluded from the increase of the
number of newborn lambs. Supplementation of a corn-soybean based preg-
nant sow concentrate by 300 µg/kg biotin result in an increase of litter size
at birth. The causative role of the zinc theoretically can be accounted for the
activation of the enzymes of steroid synthesis and its presence in the steroid
receptors; the biotin, in turn, activates the metabolism (so-called inner
flushing).
16.1.2.3. Possible regulatory mechanisms. The higher ovulation rate of ewes
of good or rather improving condition is the result of the previous oestrus
cycle, while more big (pre-ovulatory) follicles developed during the late cor-
pus luteum (CL) phase. The increase of the follicle dimensions is accompa-
nied by an elevation in the gonadotropin secretion, but the blood concen-
tration does not consequently reflect the hormone production. In gilts on a
higher ration during the four days before the oestrus, parallel to the
improved ovulation rate the blood concentration of FSH and LH are
increased, too. The effects of acute changes in dietary intake on ovarian
385
activity can be correlated with changes in circulating concentrations of

metabolic hormones including INSULIN, IGF-I, GH and LEPTIN (AMSTRONG et al.
2002). Exogenous insulin significantly improved the ovulation rate in beef
heifers, ewes and gilts. This phenomenon can be explained by an increased
GnRH secretion but also by the stimulation progesterone production of the
CL cells. Within 15 minutes after the insulin application the oestrogen con-
centration in the ovarium vena increased, too. Besides this direct ovarian
effect the insulin may act upon the ovaria indirectly, by means of the growth
hormone (GH) and the insulin-like growth factor I (IGF-I). The later effect is
made possible by the strong affinity IGF-I receptors of the granulosa cells
(LUCY, 2002). At a cellular level, physiological concentrations of insulin and
IGF-I interact to stimulate oestradiol production by the granulosa cell.
Conversely, leptin inhibits FSH-stimulated oestradiol production by the
granulosa cells and LH-stimulated androstenedione production by the theca
cells. Ovulation rate of gilts can be improved by GH application. On the con-
trary, GH application before the puberty delayed the development of ovaria
and decrease the proportion of gilts on heat (TROTTIER and JOHNSTON, 2003).
The KiSS-1 gene, which produces a protein (KISSPEPTIN) in the hypothala-
mus, stimulates gonadotropin secretion by acting directly on GnRH neu-
rons (SMITH et al. 2006). According to an other possible explanation the feed-
ing effects are mediated by the enhanced hepatic steroid metabolism, mod-
erating the inhibitory steroid feed-back on the gonadotropin secretion.
16.1.3. Feeding, fertilisation and embryonic survival

While studying the effect of feeding on embryonic survival the generally
used presumption is that the conception rate even in very varying nutri-
tional conditions is very high, except in animals fed on phytoestrogen-con-
taining plants. In the practice, during evaluating the effectiveness of supple-
mentations (e.g. beta-carotene to improve the sperm or semen index = num-
bers of inseminations per successful conception; or selenium in order to
shorten the calving to calving interval) the conception rate and the embry-
onic mortality are not separated. It is well-known that selenium deficiency
during the implantation increases the embryonic mortality. Selenium sup-
plementation improves the conception rate by facilitating the sperm-cell
transport and helping in the development of a favourable uterine medium.
The described effect is especially well-marked using organic selenium sup-
ply (e.g. Sel-Plex®). In superovulated beef cows the conception rate
386
improved from 40% to 100% after the selenium supplementation, but with-
out effect in non-superovulated cows. One of the reasons of the bad results
in the course of the first insemination of high-yielding dairy cow is the pos-
itive rumen N-balance (excess of rumen undegradable protein, RDP) and the
consecutive ammonia release. The later and its metabolites are harmful to
the gametes and/or the embryo (FERGUSON, 2005).
Long-lasting undernutrition inevitable damages the growth and sur-
vival of embryo in cattle, sheep and pig. From the point of view of embryon-
ic survival the nutrient supply of days just before and after the ovulation are
important and the feeding during the early pregnancy does not mean.
Excess nutrient intake during the early pregnancy expressly may increase
the embryonic mortality by 25 to 30%. The background mechanisms can be
concluded from the fact that the effect of excess nutrient supply on embry-
onic mortality can be counterbalanced by exogenous progesterone applica-
tion. Excess rations may stimulate the hepatic circulation and thereby the
metabolic break-down of the progesterone, which may decrease the blood
progesterone level below the critical to the survival of embryo. It is rein-
forced by several evidences that high nutrional plan during the early preg-
nancy is accompanied by a low progesterone level of the circulating blood.
In most of the species the early embryonic mortality occurs in the pre-
implantation period during the fast elongation phase of the trophoblast. On
the basis of this it is supposed that the lack of progesterone modifies and/or
decreases the synthesis of trophoblast proteins (included the IGF-binding
protein) and/or endometrium secretory proteins. By means of exogenous
progesterone application the amount of endometrium secretory protein can
be increased. The endometrium and trophoblast proteins are necessary for
the “mother-foetus biochemical dialogue”, which is indispensable to the
survival of embryo. The feeding as a whole and the individual nutrients may
stimulate the synthesis of the mentioned proteins directly or through
changing the blood progesterone level. For example the riboflavin supply of
gilts 4 days before and 4 days after the oestrus decisively influences the sur-
vival of embryo. If preventing the usual drop in the blood folic acid level dur-
ing the same period may shorten the weaning-to-reconception period.
16.1.4 Feeding and foetal growth

The retention efficiency of individual nutrients into the foetus is the start-
ing-point of the factorial methods predicting the nutrient requirements of
387
the foetal growth. For example beef cows utilize the ME with a 13% and the
metabolizable protein (MP) with a 50% of efficiency for gestation products.
There is a great range of variety in this field owing to the different genotype
(lean pig, highly prolific, multiple-pregnant ewes, goat and deer). According
to several authors the pregnancy by itself (per se) increases the amount of
small intestinal available protein. Nevertheless, it is important to underline
that the feeding standards and recommendations are useful tools in calcu-
lating the daily ration of the pregnant animals, but they do not aim for that
the nutrient supply should be every day as high as that of the needs of
mother and dam. That is precisely why the feeding allowances can be eval-
uated on the basis of the viability of offspring at birth.
16.1.4.1. Effect of feeding level. Amongst the production animals the cow is
one of the larger, therefore needs of her single, relatively small can easily be
satisfied. The calves birth weights did not decrease if the beef cows of twins
are fed on a daily ration providing 1.5-times the energy of the maintenance
requirement during the pregnancy. The relative needs of gestation are low
even in female of small body weight like the rabbit doe. Nevertheless, if
using an intensive reproductive technology, while the breeding occurs on
day 1 to 2 after kindling, besides inappropriate nutrient supply the high lac-
tational needs cause the regression of CLs and the ending of the pregnan-
cy. If the used concentrate is satisfactory (15.4 MJ DE/kg air-dry matter
and 14.8 gram digestible crude protein per MJ DE) the lactation will not be
disadvantageous to the pregnancy and to the foetal growth. However, under
such circumstances the milk production will abruptly drop on week 4 of the
contemporary suckling making the accretion of ingested nutrients into the
pregnant uterus.
Concerning the accurate feeding of pregnant gilts and sows it is
important to emphasize that even a high amount of excess nutrient can
hardly increase the birth weight of piglets. On the other hand, high-fibre
diets during gestation may increase litter size by means of the beneficial
dietetic effects of the fibre fractions (RU and BAO, 2004; PAPOCSI et al. 2007).
Fat or oil supplementation (7.5 to 15%) of both gestational and lactational
diets of breeding sows may also increase litter size and newborn mortality.
Besides the foetal growth the needs of sow’s body weight gain should be cov-
ered, which will be mobilized later to assure the lactation requirements. The
pregnant ewes are less sensitive to the changes in nutrient supply. Although
mild (30 to 50% of maintenance requirement) undernutrition during the
388
first 30 to 40 days of pregnancy decreased foetal growth by 10%, but it

could be compensated till the lambing. Reactions of the middle gestation
depend upon the condition of animals. Mild undernutrition of ewes high
condition score (BCS of 4 on a 1 to 5 scale) stimulated placental growth and
thereby the expected birth weight of lambs. On the contrary, undernutrition
of pregnant tegs and ewes of low condition score (BCS of 2 on a 1 to 5 scale)
has no positive consequences. Effect of late pregnancy feeding on the birth
weight of lambs depends upon the genotype, the actual state of body
reserves and the ration energy concentration-dependent partial kc-value
(utilization of ME for foetal growth). The negative effect of undernutrition on
the foetus may be counterbalanced to certain degree by the mobilization of
the maternal tissue.
16.1.4.2. Effect of individual nutrients. If the energy deficiency of high-preg-
nancy of ewes is accompanied by a lack of protein the decrease in the birth
weight of lambs is greater. The amino acid requirement of the mentioned
period cannot be covered by bacterial protein and especially in breeds of
high prolificacy the histidine and cystine became limiting. In case of one foe-
tus even the additional energy supply in form of glucose could help unless
sufficient methionine is not available. Although the calcium retention into
the foetus is important, minor failures of the nutrition can satisfactorily cor-
rected by the mobilization from the mother’s organism. The plentiful calci-
um supply of foetus is helped in many species the parathormon-like peptide
(PTH-like peptide) isolated recently from lamb foetus and from the placen-
ta.Calcium mobilization from bones during lactation is inevitable in the
dairy cattle, ewe and goat. Factors of this mobilization can gradually be acti-
vated by the feeding during gestation, in a way which assures the preven-
tion of the hypocalcaemic state. Old animals are more prone to the parturi-
tion hypocalcaemia because of the reduced expression of 1.25 dihydro-
cholecalciferol (DHCC) receptors (see also Chapter XXV). Worldwide experi-
ences from practical cases show the importance of zinc and iodine supply
during pregnancy.
16.1.5. Mammary development and colostrum production

In ruminants the lobulo-alveolar cell system develops mainly during the
gestation, therefore an undernutrition during late pregnancy decreases the
growth of the mammary gland and the future colostrum production.
Additional feeding of rumen undegradable protein (RUP) during late preg-
389
nancy of ewes may improve the colostrum supply of the newborn lambs. The
colostrum energy concentration can be increased by the (protected) fat sup-
plementation of the concentrates for high-pregnant ewes and sows. In accu-
rately fed ewes the colostrum accumulates in the mammary gland during
the last some days of gestation, assuring the appropriate amount (approxi-
mately 50 ml/kg LW) for the newborn lambs just after birth. Undernutrition
not only decreases the quantity of produced colostrum but also delays the
onset of the milk production. This shows that the question is not only the
lack of nutrient, but the regular drop in the mediated progesterone level
before partirution may delay.
The mammogenesis and the expected milk production are highly sen-
sitive to the excess in nutrient supply during the period before the puberty.
The regulation of this phenomenon is complex, but the consecutive decrease
in plasma GH concentration has the main role. Finally, this is the GH-
dependent IGF-I, which is responsible for the mammogenesis. (For details
see the Raising of dairy heifer section in Chapter XXV).
16.1.6. Feeding and newborn viability

Owing to its effects on the foetal metabolism, growth and the colostrum pro-
duction dam’s nutrition has a key role in determining the newborn viabili-
ty. Undernutrition decreases the blood supply and glucose uptake of the
uterus leading to the hypoglycaemia and hypoxia of the foetus. Latter is
accompanied by hormonal changes in the foetus: there is a decrease in
insulin concentration and an increase in the blood level of GH, ACTH and
corticosteroids, which in turn, modify the effect of maternal undernutrion
on the foetal growth and development. In rodents, rabbit and sheep, espe-
cially if the birth is in a cold environment, the brown fat produced extra heat
is indispensable for the survival of offspring. If the nutrient intake is insuf-
ficient during the pregnancy less brown fat tissue is produced and owing to
the described hormonal changes its heat production will be inhibited, too.
Although lipids are the main energy source (60 to 70%) for the new-
born lamb, the role of hepatic and muscular glycogen (15%) is not negligi-
ble, either. The undernutrition of ewes decreases the foetal glycogen reten-
tion; conversely, foetal insulin and IGF-I stimulate it. The glycogen is the
main energy source for the newborn piglet. The effect of the fat supplemen-
tation of pregnant sows’ diet (10%) has positive but inconsistent effect on
the newborn mortality: it increased the glycogen content of the liver by 36
390
to 46%, its effect on survival rate is not such an expressed one (KEMP and
SOEDE, 2003).
Besides its effects on the foetal metabolism and energy reserves the
mother’s undernutrition has an essential influence on the birth weight and
thereby on the survival rate. The big birth weight is especially important in
case of newborn piglet and lamb. Namely parallel to the decrease in birth
weight a relative increase in the ratio of the body surface (potential heat
losses) and body weight (potential heat production) occur. That makes new-
born susceptible to hypothermia in cold and dehydration in hot environ-
ment. The situation is only aggravated by the underdeveloped wool coat of
lamb caused by the maternal undernutrition. Notwithstanding the ewe’s
undernutrition has no effect beyond the newborn mortality in lambs,
because the ability of lamb tissue for the hyperplasia is already present and
there is a possibility for the compensatory growth. Conversely, hyperplastic
capacity of foetal muscles discontinues on day 85 to 95 days of pregnancy.
In piglets, undernourished until this moment, the number of m. semi-
tendineus fibres significantly decreases even inside the same litter. An extra
L-carnitine supply of sows during the last third of pregnancy may increase
the leg musculature of piglets.
Under practical circumstances it is not uncommon that newborn
calves, lambs, incidentally obtain the first colostrum intake delayed, mak-
ing newborn susceptible to the otherwise only slight nutrient supply of the
dam. Extra selenium (5 mg sodium selenite) and/or dl-tocopherol (1 gram
tocopherol acetate) supplementation of sow on day 100 of the pregnancy
intramuscular resulted in an increased immunoglobulin supply of newborn
(KEMP and SOEDE, 2003). Organic chromium supplementation of pregnant
cows may improve the immune status of newborn calves (MOWAT, 1997).)
16.1.7. Feeding and parturition to rereconception interval

The shortage of calving to reconception period is one of the most important
fields of improving the reproductive performance. In the re-assumption of
the cyclic ovarium functions the nutrition has a key role. In high-yielding
dairy cow the nutrient supply during the dry period has already influence
on the above period, namely if the nutrient supply is abundant, the onset of
the cycle delays (HUSZENICZA, 1987). Deers are seasonal breeder. If the lac-
tational weight losses cannot be supplemented on a poor nutrition the cyclic
ovarium activity delays and could be postponed even till the next breeding
391
season (year). Under unfavourable circumstances a similar situation may

occur in case of the ewe and goat, too. In case of female that are less sensi-
tive to the changes of photoperiod like cattle and pig the feeding should sup-
port the re-assumption of the normal cyclic ovarium activity as early as pos-
sible. Although the deficiency of some minerals (Ca, P, Cu, Fe and I) may
cause postpartum anoestrus, but most of the problems are due to the lack
of energy and the troubles in the protein supply.
The energy supply is reflected in the body condition which in turn,
has a close correlation with the cyclic ovarium function (DEROUEN et al.
1994). The energy deficiency during lactation acts through the hypothala-
mus and ovaria. This state can be aggravated by a lack of protein (FEKETE et
al. 1996), of some minerals (REGIUSNE-MOCSENYI, 1988), as well as of some
vitamins, like -carotene (PETHES et al. 1985). In dams the milk production
generally enjoys a priority against the reproduction. Data of in vivo body
composition measurement showed that the shortest postpartum anovulato-
ry period could be found in ewes that accumulated the highest amount of
fat during the first month of lactation. In dairy cows there is a significant
correlation between both the cumulative and average energy balance and
the length of postpartum anoestrus. Poor condition and negative energy bal-
ance decrease the blood progesterone and non-esterified fatty acid concen-
trations.
The interconnection of condition, body composition and fertility can
be explained by the different homeorrhetic regulation of the milk production
and the reproduction (BAUMAN and CURRIE, 1980). Rumen functions as a
whole and its supply with undegraded protein act on the reproduction also
through changing the body composition. A decrease by 1 body condition
score (BCS) is accompanied by the using of 56 to 87 kg of body fat. In the
majority of dairy cows the postpartum negative energy balance may last up
till the day 70 of lactation. The time of the first ovulation shows a positive
correlation with the degree of energy deficit of week 2 to 3. Numerically it
means that on the average of the first 20 day energy deficit of 1 MJ falls 0.7
day delay in ovulation time, i.e. each gram fat mobilization corresponds to
a fraction day postponement. Described correlations can be modified by
rumen degradation of the intake protein. For example while feeding cows on
a high RUP diet in order to support milk production it may stimulate fat
mobilization and rise in the energy deficit (FEKETE et al. 1996). Too much
RDP may delays the reconception, too by altering the composition and pH
392
of the uterus secretion. Although the prerequisite of a short calving to

reconception period the onset of the oestrus as early as possible and the
reassumption of the cyclic ovarian function, the latter is influenced by a
series of further factors, too.
In beef cows the suckling may postpone postpartum anoestrus up to
90 days (HUSZENICZA et al. 1988) in this way it is the main cause of the bad
reproductive parameters. Undernutrition not only prolongs elapsed time to
the first ovulation but increases the individual deviation that makes uni-
form feeding technology impossible.
In case of breeding sows the shortage of the weaning to reconception
period is important, too, because the failure of cyclic ovarian function is one
of the most important culling considerations in the practice. An undernu-
trition during the lactation (less than 45 MJ energy intake per day) will
increase the weaning to the first oestrus period. This phenomenon is more
expressed in gilts and so much the more the younger were at the time of the
first fertilization. Feed mixtures of average energy concentration (13.2 MJ
DE/kg) can be fed ad libitum, but keeping on a feed of high energy concen-
tration (e.g. 15.6 MJ/kg, owing to fat supplementation) ad libitum will pro-
long the weaning to reconception time. The danger of feeding lactating sows
ad libitum is that animals of good condition and thereby of bad appetite will
loose more than the physiological. First lactation sow should be fed ad libi-
tum if only it has a backfat thickness no more than 20 cm at farrowing. (one
mm in backfat thickness is equivalent to 3 kg body fat.) In this situation the
first new oestrus will occur on day 6 to 9 after the weaning and this onset
shows a correlation with the energy intake during lactation.
16.1.8. Feeding and postpartum hormonal changes

The nutrition achieves the re-assumption of the cyclic sexual function by
means of timing and quantitatively modifying hormonal changes of the late
pregnancy and early lactation. The number of GnRH receptors in the
hypophysis does not change but GnRH release from the hypothalamus may
vary. Differently according to the species the suckling or rather the fre-
quency of suckling, by means of the opioid peptides of the hypothalamus
(e.g. β-endorfin) inhibits the pulsatile LH release of hypophysis. Under the
circumstances of practical pig production the suckling totally suppresses
the ovarian function. In deer, beef cow and ewes the suckling, except it coin-
cides with the seasonal anoestrus, solely postpones the ovulation. The feed-
393
ing modifies the described neuroendocrine events, the most frequently it is

the frequency of the pulsatile LH release which is commonly influenced. The
feeding lengthening the parturition to reconception interval acts by the inhi-
bition of the GnRH secretion in the hypothalamus (HESS et al. 2005).
Identification of the information pathway to the GnRH neurons nowadays is
a research priority. The blood IGF-I concentration reflects reliably the post-
partum body condition of the beef cow.
16.1.9. Role of mycotoxins and heavy metals on the reproductive func-

tions
Several mycotoxins, the secondary metabolic products of the microscopic
moulds could alter the reproductive functions. The F–2 toxin (zearalenon)
produced by Fusarium species causes pseudo-oestrus, embryonic mortality
and absorption, in newborn piglets the oestrogen syndrome and in males
the inhibition of the spermatogenesis. The trichotecene-structured T–2
toxin and its metabolites through the inhibition of the protein synthesis at
cell level decrease the progesterone production (FIGURE XVI-1) causing the
prolongation or absence of the cycle in the monogastric animals (FEKETE and
HUSZENICZA, 1993).
FIGURE XVIII-1: Effect of 30-day long pre-feeding of low T–2 toxin (0.19 mg/kg feed) dosage
on the GnRH injection induced progesterone profile in virgin rabbit does
394
Healthy rumen microflora on a balanced diet (ratio of concentrate to

roughages) is able to break down the trichotecene-structured mycotoxins.
On a low-fibre high-concentrate diet the pH falls, the microflora undergoes
transformation and the outflow rate of rumen content accelerates, thereby
mycotoxins can get the abomasum and duodenum intact, unchanged. In
such a case even an extremely low mycotoxin intake may cause the cycle be
upset (FIGURE XVI-2; HUSZENICZA et al. 2000).
FIGURE XVIII-2: Effect of rumen acidosis and T–2 toxin intake upon the blood progesterone
concentration: the minimum protesterone level (3 nmol/l) required for the ovulation
occurred in the toxin-loaded heifers (below) 2 days later (8.3 days instead of 6.3 days) than
in the control (above)
Among the heavy metals the mercury, the cadmium and the lead
could damage both ovarium and testes tissue (BERSENYI et al. 2003). Similar
alterations (FIGURE XIV-3 and XIV-4) were found in rats’ testes after per-
oxide loading (FEKETE et al. 2006).
395
16.2. Endocrine and metabolic factors of reproductive func-

tions of dairy cows
© Gyula Huszenicza
n the last few decades a continuous increase has been recorded in the
I average milk yield of dairy cattle population kept in intensive production

systems all over the world. However, the incidence of metabolic mal-
functions related to the postpartum (pp) negative energy balance has also
increased in the early weeks of lactation predisposing the affected cows for
several puerperal and ovarian malfunctions. Simultaneously, dramatic
decrease was seen in reproductive performance. All these unfavourable ten-
dencies are based on the known discrepancy between the dry matter (main-
ly energy) intake and the requirements of milk production. The energy
demand of high-producing dairy cow’s shifts abruptly after calving as the
daily milk yield rapidly increases and the ensuing negative energy balance
extends over a period of 10-12 weeks. In principle it is a physiological phe-
nomenon, which may, however, result in disorders in metabolism (fat accu-
mulation in hepatocytes, ketosis) and antimicrobial self-defence mecha-
nisms. Also it postpones the first ovulation and oestrus. In the first 3-4
weeks after calving the negative energy balance is highly correlated with
both milk yield and the length of time before the first ovulation (pp acyclia).
Because the number of ovulatory cycles preceding the first insemination (AI)
has been shown to influence the conception rate, the length of pp acyclia
provides an important parameter for assessing the effect of negative energy
balance on reproductive performance. Also, the poorer than expected qual-
ity of oocytes and embryos, and the diminished character of post-ovulato-
ry/post-AI progesterone (P4) rise have been thought to decrease the first
service conception rate.
16.2.1 Negative energy balance (NEB) in postpartum dairy cows

THE INCREASED LIPID MOBILIZATION AND ITS CONSEQUENCES
During the period of NEB high-producing cows must mobilize their body
reserves, first of all their subcutaneous lipid stores (increased or forced lipid
mobilization). Generally, the animals overfed before calving [body condition
score (BCS) at calving: >3.75] are the most endangered (fat cow syndrome).
The rapid reduction in live weight (LW) and BCS are the most common clin-
396
ical consequence of forced lipid mobilisation: dairy cows often lose over 60%
of their body fat during early lactation. As one of the earliest changes a
sharp elevation is seen in the levels of non-esterified fatty acids (NEFA).
Simultaneously lipids (mainly triacylglycerols) are accumulated in the liver
(fatty liver disease). From moderate to severe forms of fatty liver (fat content:
³20 %) may result in well-defined disorders in hepatocellular functions (glu-
coneogenesis, cholesterol, bile acid and bilirubin metabolism, synthesis of
certain apolipoproteins and the 25-hydroxylation of cholecalciferol).
Concentrations of total bilirubin and total bile acids in plasma correlate
positively with degree of hepatocellular fatty infiltration. Simultaneously
hypoalbuminaemia, evidently reduced levels of very low-density lipoproteins
and low-density lipoproteins. Furthermore, lower than normal concentra-
tions of all the lipoprotein-transported substances (total cholesterol, triacyl-
glycerols, b-carotene and tocopherols) are detected in the peripheral blood.
Although the membrane-damaging effect of hepatocellular fatty infiltration
is not conspicuous, moderate elevation in serum activities of aspartate
aminotransferase, alkaline phosphatase and lactate dehyd-rogenase is usu-
ally observed. Also production of bOH-butyrate and other ketone bodies
may increase (hyperketonaemia). When the hepatocellular NEFA uptake ele-
vates these fatty acids are either esterified to triacylglycerols, or converted
into acetyl-CoA. Whether this acetyl-CoA is introduced into the Kreb’s cycle
depends on the availability of oxaloacetate, which is predominantly deliv-
ered from gluconeogenic precursors such as propionate, pyruvate, glycerol,
or certain amino acids. If the supply of these precursors is inadequate, as
can occur frequently during NEB, the availability of oxaloacetate for intro-
ducing acetyl-CoA into the Kreb’s cycle decreases. The excess acetyl-CoA is
than used for ketogenesis and the subclinical or clinical forms of ketosis
develop. Inadequate supply with gluconeogenic precursors seems to be the
core event in this process. The intrahepatic detoxifying capacity of ruminal
ammonia (and endotoxin) is also diminished, but it results in clinical symp-
toms of hepatic coma with the subsequent death of animal only exception-
ally, in the most severe cases. From economic point of view it is more impor-
tant, however, that these metabolic changes predispose the cow for milk
fever, retained foetal membrane, acute putrid endometritis, ovarian mal-
functions and displaced abomasums.
This predisposition for bacterial disorders in the puerperal period is
based on impairments of the immune system. Polymorph nuclear leukocytes
397
are the most important cellular elements of the host’s antimicrobial self-
defence mechanism. The peri- and post-parturient metabolic changes,
mainly the high plasma levels of NEFA and β-OH-butyrate impair their
migration and phagocyte activity, enhancing the susceptibility of mammary
gland to invading pathogens. On day 1-3 after calving the elevated (>1.00
mmol/l) β-OH-butyrate levels predispose the cows for mastitis in the sub-
sequent 4 weeks. The same factors predispose the cow also for bacterial
complications of uterine involution (acute putrid endometritis).
16.2.2 Puerperal endocrine changes

ENDOCRINOLOGY OF THE NEGATIVE ENDOCRINE BALANCE
In early weeks of lactation the NEB are characterized by marked endocrine
changes, known as important factors in regulation of forced lipid mobilisa-
tion. Besides the involvement of catecholamines, elevated glucagon and
decreased insulin concentrations are reported to occur, whereas the growth
hormone (GH) levels remain unchanged or perhaps slightly increase.
Simultaneously the GH-induced hepatic insulin-like growth factor-1 (IGF-1)
production and the glucagon-induced insulin responsiveness of pancreatic
β-islets are diminished and certain tissues loose their insulin sensitivity. All
these endocrine events can shift the metabolism from anabolic to catabolic
direction. Tissues try to fit their current local energy metabolism to this new
catabolic state through increasing the capacity of an inactivating path of
thyroid hormones. As a result, although the thyreotropin releasing hormone
(or thyrotrophic) induced thyroxin (T4) response is only slightly altered,
markedly decreased T4 and 3,3’,5-triiodothyronine (T3) levels and elevated
rT3 concentrations are observed in the peripheral blood (PETHES et al. 1985).
Also the adrenocortical function including cortisol production is increased.
Recently the NEB-related endocrine research has been focused on the
role of leptin. This newly identified cytokine like protein hormone secreted
mainly by the adipose tissue is believed to act through hypothalamic nerve
centres in mediation of neuroendocrine responses to energy supply or dep-
rivation. It may signal nutritional status perhaps also for the peripheral
organs. Leptin is one of the primary agents communicating information
about the level of peripheral energy stores to hypothalamic regions that con-
trol feeding behaviour, metabolism, and endocrine function so as to main-
tain energy homeostasis. Insulin, glucocorticoids and endotoxin exposure
may increase its gene expression and/or plasma level, whereas leptin can
directly inhibit cortisol synthesis by adrenal cells. Thus leptin and cortisol
398
interact in a negative feedback loop. Leptin liberation from adipocytes is

down-regulated by adrenergic stimulation and it is also supposed to inter-
act with growth hormone and IGF-I secretion. In ruminants, as in other
species, leptin concentrations vary with changes in body weight and tria-
cylglycerol percent of body fat. Plasma leptin content was found to be a good
indicator of body fatness (BCS) in peripartum dairy cows as well, especially
when 3 weeks before calving measured.
16.2.3. Resumption of cyclic ovarian function during the negative

energy balance
RESUMPTION OF REGULAR FOLLICULAR GROWTH. ONSET OF OVARIAN CYCLICITY

In almost the entire non-suckling dairy cows the FSH concentrations in
plasma increase to peak values on d 4-5 after calving. This first FSH peak
is followed immediately by the initiation of the first pp follicular wave pro-
ducing the first dominant [>9 mm] follicle. Subsequently the regular forma-
tion of new FSH waves followed by the growing of new follicular cohorts and
producing new dominant follicle- is reported to proceed despite the average
NEB of -31 MJ/day during the first 3-week period after calving. It appears
that the initiation of follicular waves in early pp cows is unperturbed by
NEB and occurs in response to the re-establishment of periodic FSH surges
synchronized only by the end of gestation and parturition. During the NEB
in early weeks of lactation LH, but not FSH appears to be deficient.
Therefore regular onset of FSH dependent follicular growth seems to be
insensitive to NEB.
Three patterns of pp follicular development based on the fate of the first
dominant follicle (DF) have been described (1) ovulation of the first-wave DF;
(2) development of a first-wave anovulatory DF followed by (several other)
additional waves of follicular development before the first ovulation; (3)
development of a first-wave DF that becomes cystic. It has been widely
accepted that the EB is one of the most important factors influencing the
duration of this acyclic period: in non-suckling dairy cows the first pp. ovu-
lation takes place on about the day 10 after the nadir of NEB).
Relationship between NEB and follicular dynamics in pp. dairy cows. It
was demonstrated that the number of class 3 (10-15 mm in diameter) - but
not of the classes 1 (3-5 mm) and 2 (6-9 mm) - follicles increased with more
positive energy balance (EB) before day 25 after calving. This observation
suggests that as cows improve in EB, the movement of smaller follicles into
399
larger size classes is enhanced. During the first pp follicular wave (d 8-14
after calving) in cows receiving three levels of dietary fat, the number of
class 1 and class 2 follicles was not correlated with EB during either the 1st
or 2nd wk pp, regardless of diet. The development of a DF in pp dairy cows
is tolerant to NEB. However, several studies demonstrated that the ultimate
diameter and E2 production of DF are influenced by metabolic factors: both
the size of DF and the E2 level in plasma increased after EB improved rela-
tive to its most negative level.
16.2.4 Signalling the metabolic state: metabolic hormones in regula-

tion of ovarian function.
Effect of negative energy balance on the time of the first pp ovulation was
confirmed a relatively long time ago. The restricted energy intake did not
alter the pituitary GnRH receptor density in pp cows but dietary energy
restrictions were followed by both decreased and increased (WHISNANT et al.
1985) responsiveness to exogenous GnRH. Loss of pulsatile LH secretion
was shown to result from prolonged inadequacy of energy supply in both of
suckling beef cows, and non-suckling dairy cows. Up to now it has been
widely accepted that the re-establishment of a pulsatile LH secretion pat-
tern conductive to preovulatory follicular development and function is a key
event in the return of ovarian cyclicity in pp dairy cows experiencing NEB.
It appears that pp recovery of pituitary LH content and responsiveness to
GnRH is complete by day 10 after calving in dairy cows, and available evi-
dence across species suggests a predominantly hypothalamic locus for the
primary effect of decreased energy intake. In early weeks of lactation the
reduced activity of the GnRH pulse generator in pp dairy cows is expressed
as reduced pulsatile LH support of follicular steroid genesis necessary for
induction of a preovulatory like LH surge and subsequent ovulation.
However, a seemingly low LH pulse frequency (2 pulses per 6 h) is appar-
ently adequate to sustain the morphological development of dominant folli-
cle by the 2nd wk pp. This observation is consistent with the growth and
differentiation of competent DF-s during the mid-luteal phase of the bovine
oestrous cycle when the LH pulse frequency is low.
Studies investigating the potential metabolic signals for hypothalamus-
anterior pituitary-ovary (HPO) axis have been focused primarily on blood
metabolites (NEFA, glucose) and metabolic hormones (insulin to GH ratio,
insulin, IGF-I) known to fluctuate during altered states of energy metabo-
400
lism. It is generally accepted nowadays that any mechanism such as meta-

bolic status coupled with HPO has to involve ultimately hormonal compo-
nents. Many authors confirmed the physiological importance of plasma
insulin to GH ratio and the day of the EB nadir, indicate that these hor-
monal differences in the intermediate pp period may influence the first wave
of follicular function. Insulin has been shown to stimulate follicular cells
and small increases pp could have important effects during the very early
triacylglycerolses of follicular development. Circulating concentrations of
IGF-I and one of its binding proteins (IGFBP-2) in the peri-parturient peri-
od were good indicators of the capacity of energy-restricted cows to resume
cycling after calving. (Furthermore, an increased insulin to GH ratio follow-
ing parturition may be conductive to greater hepatic IGF-I production,
resulting in increased amounts of this growth factor earlier after calving.
During the first 2 weeks pp higher circulating IGF-I concentration can be
detected in cows developing E2-active, ovulatory first-wave DF than in those
with E2-inactive anovulatory first-wave dominant follicle. In cows the circu-
lating IGF-I concentrations correlated with IGF-I levels in the follicular fluid
of large follicles. Both insulin and IGF-I are known to stimulate the in vitro
steroid genesis and proliferation of bovine thecal and granulose cell cul-
tures. Likewise, cows that ovulated within 35 days postpartum presented
higher IGF-I concentrations as well as higher glucose and insulin and lower
NEFA and beta-OH-butyrate concentrations (HUSZENICZA et al. 2001). Under
in vitro conditions in bovine theca cells the IGF-I increased the number of
LH-binding sites and enhance the LH-induced production of androstene-
dione and progesterone. This finding confirms the in vivo observations
demonstrating an apparent relationship between the steroidogenic activity
of first-wave DF and circulating levels of IGF-I.
In cattle the T4 was a much weaker (i.e., increase to 1.3-fold) inducer
of theca cell P4 production than was LH (i.e., increases to four- to nine fold)
and its effect was only evident at hyperthyroid levels (i.e., 100 ng/ml, but
not 30 ng/ml); T3 had no effect on granulosa and theca cell P4 production
in this study. Association between low E2, negative energy balance, and low
T3 was also found in young distance runner women. In contrast, diameters
of dominant follicles from multiparous Brahman cows were not affected by
induced hypothyroidism.
Through affecting the hypothalamic GnRH and pituitary gonadotrop
secretions the leptin has also been reported to influence the genital func-
401
tions in rodents, primates and recently also in farm mammals. Experiences

have confirmed that this hormone may be one of the indicators of nutri-
tional status that allows reproductive processes to proceed. Results in
rodents, non-human primates and also in porcine and ovine models suggest
that the suppression of GnRH/LH during fasting is mediated by central
action of leptin in the pituitary or in the brain. However, pulsatile leptin
secretion was independent of LH secretion in lambs. Treatment with leptin
accelerates puberty in normal mice at doses that do not change body
weight, and suggested that actions of leptin to regulate neuroendocrine and
reproductive function in normal mice are not secondary to effects on ener-
gy balance. Low leptin induced markedly increased production of neu-
ropeptid Y (NPY) in hypothalamus, and increased NPY inhibited the gonadal
axis in lab rodents and also in ruminants, so it have postulated that
impaired reproductive function in adverse metabolic condition such as fast-
ing could be due to excessive hypothalamic NPY release. Several observa-
tions suggest that leptin is an important biochemical message between fat
stores and the reproductive axis. Central (intracerebro-ventricular) admin-
istration of leptin increased LH secretion in fasting cows and ewes, but not
in control fed animals indicating that metabolic state is an important factor
in modulating the response of HPO axis to leptin. Leptin receptors have
been detected also in gonads. Under in vitro conditions the supraphysiolog-
ical dose of leptin has week inhibitory effects on gonadotropin- and/or IGF-
I-induced steroidogenesis of theca and granulosa cells. In conclusion, not
only the hypo-, but also the hyperleptinaemia can be supposed to have a neg-
ative influence on reproduction in domestic mammals including cattle.
In periparturient dairy cows, changes in the plasma concentration of
leptin have been measured during the period from 35 d before to 56 d after
parturition. The plasma concentration of leptin was highest during late
pregnancy and declined by ~50% after parturition. Leptin concentrations
started to decrease 20 days before parturition. The plasma leptin concen-
tration remained depressed during early lactation despite a gradual
improvement in energy balance. Corresponding changes occurred in the
abundance of leptin mRNA in subcutaneous white adipose tissue. The pp
reduction in plasma leptin was due to the NEB because plasma leptin
remained high in cows not milked after parturition. The plasma concentra-
tion of leptin was positively correlated with plasma concentrations of insulin
and glucose, and negatively correlated with plasma concentrations of GH
402
and NEFA. However, wide variety of postpartum tendencies was reported to

occur. The postpartum leptin reduction is likely to be due in part to the neg-
ative energy balance because plasma leptin remained high in cows not
milked after parturition. Regression analyses of plasma leptin during the
pre- and postpartum period in lean and fat cows was carried out: while no
association was found between leptin levels and BCS after parturition in
lean cows, which was maintained in fat cows during the same period.
Overall, our results suggest interactions between the effects of BCS, parity
and lactation on peripartum leptin regulation.
In pp cows data regarding leptin and reproductive performance are lim-
ited. In 20 high-producing Holstein cows kept under model conditions the
plasma leptin concentrations declined after parturition and reached their
nadir on 10.1 ± 2.2 days after calving, then they increased and became sta-
ble near the time of the first ovulation on 25.9 ± 2.0 days. The interval from
calving to first ovulation correlated significantly with the interval from par-
turition to leptin nadir, but it was not in correlation with the prepartum,
pre- and/or postovulatory leptin values. Although there was a lack of rela-
tionship between leptin and first postpartum luteal activity, higher leptin
concentrations were associated with shorter intervals to first observed
oestrus.
16.2.5. Factors influencing fertility in postpartum dairy cows

The detrimental effects of NEB in early lactation appear to be manifested
also as reduced fertility during the pp breeding period. In normal dairy herd
situations direct assessment of energy balance in individual cows is not
possible, but changes in BCS provide an indirect measure. With more
extensive loss of BCS, the reduction in conception rate becomes greater.
Cows losing one unit or more BCS (5-point scale) during early lactation are
at greatest risk for low fertility with conception rates of 17 to 38 %. Recent
studies indicated that cows with marked losses in BCS (>1.25 unit) were
only half as likely to conceive at first AI as cows with more modest loss and
that conception rate increases 10% for every unit increase in BCS. A recent
large survey study found that cows with a BCS of 3.0 at the first AI were
most likely to become pregnant.
DELAYED ONSET OF OVARIAN CYCLICITY. Although reduced fertility as a conse-

quence of NEB in early lactation may be explained by prolonged acyclicity
in 30-36% of cows our understanding of the linkage between NEB and sub-
403
optimum conception rates in ovulatory cyclic cows remains rather specula-

tive. One important link between NEB and lower fertility appears to be
through the aforementioned effects on the timing of first postpartum ovula-
tion. A positive association between the early commencement of ovulatory
cycles and improved conception rate to AI is well documented. Cows
remaining anovulatory for >35-50 days of lactation were significantly less
likely to become pregnant during lactation and would be culled. Puerperal
diseases with intensive endotoxin and cytokine release
Experimental administration of endotoxin can impair both the basal
LH pulsatility and the formation of preovulatory LH peak in ruminants, and
may induce early luteolysis of premature corpora lutea in ruminants. So in
the early weeks of lactation certain inflammatory diseases (e.g. acute putrid
endometritis and severe cases of mastitis) with intensive endotoxin and/or
cytokine release are thought to interact with the effect of NEB, postponing
the time of the first pp ovulation, inducing anovulatory cysts, and influenc-
ing the cyclic ovarian function thereafter.
16.2.6. Bacterial complications of uterine involution

While accepting the general benefit of an early return to ovarian cycles on
fertility, we should be mindful of the interaction with uterine health.
Multiparous cows ovulating before 21 days postpartum exhibited poorer
reproductive performance than those ovulating later in association with a
high incidence of persistent corpora lutea (CLP). CLP presumably result
from uterine infection and their incidence in dairy cows has increased in
recent years.
16.2.7. The effect of circulating progesterone level in already cyclic

cows
Pre-insemination period. Another important link between NEB and fertility
is the carryover effects on plasma P4 concentrations. During the first two or
three pp ovulatory cycles the peak level of P4 is reported to increase in the
blood from one cycle to the next and in the early weeks of lactation the rate
of this P4 increase is reduced or moderated by NEB. Cows with the most
negative energy status during the first 9 days postpartum still had
decreased plasma P4 levels during their third oestrus cycles corresponding
to the start of the breeding period. Plasma P4 concentrations in cows select-
ed for high milk yield were 25 to 50% lower during the second and third
404
luteal phase than in control line cows.
16.2.8 Post-ovulatory rise of progesterone in inseminated cows.
The ability to produce and maintain optimum P4 concentrations is impor-

tant for fertility due to the regulatory effect of this hormone on endometrial
function by 4-7 days after AI the plasma P4 was found to be higher in cows
that became pregnant than in non-pregnant cows.
Early (by day 5 after fertilization) P4 stimulation alters endometrial
secretions and advances conceptus development. Conversely, a slower rate
of post-ovulatory P4 rise through days 4-5 has been associated with lower
fertility and decreased embryo growth by day 16. At this critical time for
maternal recognition of pregnancy an inadequately developed embryo pro-
duces insufficient quantities of interferon-t to inhibit the uterine luteolytic
mechanism, the oxytocin receptor stimulated PGF2alpha release, which in
itself is made stronger by lower circulating P4. In addition, a delayed rise in
luteal P4 around days 4-5 may allow pregnancy recognition, but will cause
early termination of the pregnancy because the embryo has been compro-
mised during development.
The physiological mechanism by which NEB early in the pp period
translates into reduced P4 production two months later has not been estab-
lished. Possibly ovarian follicles are detrimentally affected by exposure to
NEB during their early growth and development and that ovulation of affect-
ed follicles would lead to lower P4 secretion. This hypothesis may explain
the pattern of plasma P4 concentrations in lactating cows, however, the
effects of dietary intake on P4 clearance must also be considered. In sheep,
high dietary energy intake increases metabolic clearance of P4 from blood
by the liver. The baseline liver blood flow in lactating cows was found to be
twice than that in non-lactating cows and was acutely increased 20-30%
with feeding. High liver blood flow resulted in increased steroid metabolism
and 25% lower plasma E2 and P4 concentrations during the oestrous cycle
and this was associated with a higher incidence of degenerating embryos on
day 5. During early lactation total dietary intake in dairy cows increases
two-fold by the beginning of the breeding period (BAUMAN and CURRIE, 1980).
In this situation increases in P4 clearance due to high dietary intake (both
energy and protein) may be combined with the carryover effects of NEB
result in lower plasma P4 concentrations and to reduce fertility. The uter-
ine environment is dependent on P4, but may be rendered sub-optimum by
405
effects of NEGATIVE ENERGY BALANCE and increased metabolic clearance in high

yielding cows.
16.2.9. The quality of oocytes

Early pp NEB may adversely impact oocytes during the 80-100 days
required for follicle development and, thereby, exert another carryover effect
on fertility. Cows having more severe pp NEB due to over-fatness before and
at calving produced oocytes with lower developmental capacity following in
vitro maturation and fertilization procedures as compared with control cows
during the period of 80-120 days of lactation. High genetic merit cows yield-
ed oocytes with lower developmental potential than medium genetic merit
cows during mid-lactation (125-229 days). Differences in milk yield did not
affect embryo development, but blastocyst formation rates were reduced in
cows with low BCS (1.5-2.5). High genetic merit cows had lower BCS than
medium merit cows. Lower conception rates for the high genetic merit cows
were suggested to be the result of impaired oocyte quality rather than the
currently existing level of any of the metabolic parameters measured at the
time of AI studied. Overall, these experiences suggest a generally adverse
effect of reduced energy status during lactation on oocyte development.
CONCLUSION
Reproductive performance in dairy cows has declined over the past several
decades in association with impressive increases in milk yields. The meta-
bolic demands of high milk production result in greater negative energy bal-
ance, during which blood levels of glucose, insulin, IGF-I, leptin and T3 lev-
els are generally reduced, while triacylglycerols content in the liver and
plasma NEFA (and beta-OH-butyrate) concentrations are increased. NEB
and its metabolic consequences are associated with pp follicular develop-
ment and first ovulation, variability of plasma P4 concentrations, and
impair the oocyte development, resulting in decreased fertility. Fertility in
dairy cows reflects the cumulative influence of metabolic, endocrine, and
health components that have been modified and exaggerated by selection
for high milk yield, as well as by certain diseases (bacterial complications of
uterine involution, mastitis). EB seems the most important factor, but the
complex interactions of all these factors must be considered and controlled
for, if we are to improve our understanding and develop new strategies to
improve fertility. The endocrine signals that most likely can inform the
reproductive axis regarding the postpartum negative energy balance are
406
IGF-I and leptin.
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408
Chapter XVII
INTERRELATIONSHIPS OF FEEDING WITH

IMMUNITY, PARASITIC INFECTION AND GENOMICS
17.1 Nutrition and immunodeficiency

17.2. Metabolic consequences of immune response
17.3.1. Energy and protein deficiency
17.3.2. Deficiency of surplus of minerals
17.3.3. Lack of vitamins and essential fatty acids
17.6. Nutrigenomics
17.6.1. Basics
17.6.2. Fats and fatty acids
17.6.3. Starvation
17.6.4. Dietary essential fatty acids
17.6.5. Conjugated linoleic acids (CLA)
17.6.6. Metal
17.6.7. Vitamins
17.6.8. Nutriceuticals
17.6.9. Possible practical-clinical applications
Chapter XVII: IMMUNITIY- PARASITES-NUTRIGENOMICS
T he immune system is one of the defence mechanisms of animals to

keep the integrity of their body. The other types of protective mech-
anisms are the physico-chemical barrier of the skin, epithelium and
mucous membranes. The non-specific immune response consists of many
humoral factors, like the acute phase reaction. During the acute phase
reaction, starting with fever, anorexia and leukocytosis, the concentration
of some proteins (e.g. serum-amyloid-protein, the SAP; C-reactive proteins,
coeruloplasmin) strikingly increases (HEEGAARD et al. 1998 and 2000;
HORADAGODA et al. 1999), the complement system and the pro-inflammatory
cytokines (IL-1, TNFα, IL-6), produced by the mononuclear phagocytes
(MØ), as well as cellular factors like phagocytes, granulocytes and natural
killer cells are activated. The biological role of the acute phase reaction is to
combat intruders, repair tissue damages and stimulate immune response.
Owing to the adenohypo-physis and the stimulated leucocytes, which in
turn can also release ACTH and thyrotropin, the ACTH, corticosteroids and
thyroxine level increases that of the retinol-binding protein and conse-
quently, the vitamin A level, decrease Mobilization of proteins leads to neg-
ative balance. Owing to the effect of lymphocyte endogenous mediator (LEM)
and the apolactoferrin, the plasma zinc, iron and copper concentration dra-
matically drops, to deprive bacteria from essential nutritional factors. The
causes of drastically decreases in serum zinc concentration is on one hand
a redistribution into the liver and lymphocyte metallothioneins under the
influence of LEM; on the other hand, the calprotectin, compound produced
by the PMN (polymorphonuclar) leukocytes, binds part of the plasma zinc.
The enhanced erythrocyte sedimentation rate (ESR) and the elevation of fib-
410
rinogen fraction get plasma more viscous, aggravating the survival and pro-
liferation of infectious agents.
Owing to the specific immune response, animals get immune to the
given antigen. In case of infections (viral, bacterial, fungal or parasitic), the
developed immunity is a desired effect, because it helps in eliminating the
infectious agent with, minimizing its damaging effect. However, if the anti-
gen in question is of dietary origin, the immune response may be harmful
to the organism. The protection against the pathogenic agents differs,
according to the character of intruder, for example bacteria, acting extra-
cellular (e.g Staphylococcus, Streptococcus, E. coli, clostridia), bacteria act-
ing intracellular (e. g. Listeria, Legionella, Candida) or the viruses (FALUS,
1996). However, there is a common mechanism, active participation of nat-
ural killer cells (NK) by producing toxic oxygen metabolites. The process
includes the so-called „respiratory burst”, while the non-mitochondrial oxy-
gen consumption abruptly increases. First, during phagocytosis the mem-
brane receptors of granulocytes activate the intracellular NADPH-oxydase,
which in turn, by reacting with molecular oxygen, produces superoxyde
anions. The latter, under the influence of superoxyde-dysmutase (SOD)
enzyme transforms into hydrogen peroxyde (H2O2). Then, from the H2O2
substrate the myeloperoxydase (MPO) is capable of producing of the
extremely toxic hypochlorite anion, to kill the pathogens.
17.1 Nutrition and immunodeficiency

Immune deficiency may be congenital, genetic and acquired. The latter can
be caused by virus infection (human and cat HIV), ingestion of mycotoxins
(e.g. T-2 toxin), long-lasting stress and corticosteroid treatment. Immune
deficiency not only increases susceptibility against infections, but also is
connected to some types of allergies and to the autoimmune diseases. At the
same time, immune deficiency correlates only indirectly with the nutrition,
namely if the primary cause of disease leads to chronic diarrhoea, malab-
sorption and therefore there are significant faecal blood, protein, mineral
and vitamin losses. During designing the daily ration for these patients, the
replacement of these substances should be taken into consideration.
17.2. Metabolic consequences of immune response

The so-called „immunological stress” (KLASING, 1987; 1991), beginning with
the acute phase reaction, has a variety of physiological and metabolic con-
411
CHAPTER XVII: IMMUNITIY- PARASITES-NUTRIGENOMICS
sequences, which are at many points different from stress states, caused by
other stressors, like heat or crowding. Events of immunological stress are a
special form of homeorrhetic control and have a significant influence of nutri-
ent utilisation and thereby requirements.
Metabolic changes are mediated the cytokines, produced by the acti-
vated macrophages (see below: TNF-α, IL-1 and IL-6). The intensity of
immunological stress response is proportional to the antigen challenge, but
even a vaccination or subclinical infection exert an important effect in nutri-
ent partitioning, directed them to the synthesis of acute phase molecules
and cells, instead of growth. The immune system may influence metabolism
by three ways: using direct neural connections; by the classic endocrine
system and by the released leucocytic cytokines (RABSON et al. 2005). These
hormone-like peptides directly alter metabolic functions and reduce volun-
tary feed intake.
The physiological changes include an elevation of the basal metabol-
ic rate, strikingly enhanced carbohydrate utilisation and to support this,
stimulated glyconeogenesis and glycogenolysis. Generally degradation
ended in lactic acid, which in turn through the Cori-cycle returns to the liver
for re-utilisation. There is a strong peripheral protein catabolism, to supply
amino acids for the acute phase proteins synthesis in the liver and for the
proliferation of new lymphocyte colonies. Moreover, amino acids, after
deamination, may serve also as a source of energy. A high VLDL blood level
is also characteristic. Starch supplementation has a benevolent effect on
intermediary energy metabolism in monogastric animals.

Not only the extremely insufficient feed intake (general energy and protein
deficiency) alters immune function, but also the ingestion of quantitatively
sufficient, but unbalanced rations. However, the protein-energy malnutri-
tion (PEM) is the primary factor in affecting immune response.
Interpretation of the related data is difficult, because generally there is an
accompanying vitamin and/or mineral deficiency, too (see human
Kwasiorkor). Moreover, the commonly occurring secondary infection not
only weakens further the immune system, but also worsens the general
body condition. Most of the knowledge of this field derives from epidemio-
logical studies, because the results of the experiments, investigating the
effect of one nutritional factor on the immunity, are difficult to extrapolate
412
to the complex situations of the real life.

17.3.1. Energy and protein deficiency
Lack of energy and protein hardly damages the humoral immunity: the
number of B-lymphocytes in the blood is rather independent from the
changes in feeding, the serum IgG and IgM level is stable and the IgA con-
centration generally is slightly increased. However, the concentration of
secretory IgA in the produced nasal, conjunctival, pulmonary discharges
and intestinal secretions decreases. In the latter the species differences are
great, and for example the lachrymal IgA level in rat is a very sensitive indi-
cator of the protein deficiency. Undernutrition fundamentally affects the
cell-mediated immune response and the complement production. Under the
influence of juvenile malnutrition the lymphoid organs, like thymus, lymph
nodes, tonsils and spleen become atrophied. Number and activity of circu-
latory T-lymphocytes decrease, which can be demonstrated by the skin
hypersensitivity reaction. Although the number of leukocytes does not
change, their intracellular bacterial and fungal killing activity decreases.
The susceptibility of the individual immune function is different and as
mentioned above, the lachrymal IgA level in rodents is a very sensitive indi-
cator of the protein deficiency, appropriate even for evaluating the degree of
discomfort: chromorhinodacryorrhea being the first sign of serious discom-
fort in rats.
The injury of the immune system depends not only on the degree of
malnutrition, but also on the presence of other metabolic troubles and also
on the age. Thus, the malnutrition is only an important member of the fac-
tors, besides infective agents, stress, surgical intervention, tumours,
endocrine and metabolic troubles debilitating the patient’s immune system.
At the same time, improvement in feeding supports the appropriate immune
response and decrease susceptibility against infections.
17.3.2. Deficiency of surplus of minerals
Harmonious functioning of the immune system requires the presence of
several macro- and microelements. In experimental animals, the lack of cal-
cium, magnesium, iron, zinc, copper, iodine and selenium has been associ-
ated with signs of immunodeficiency. Mortality of mice, experimentally
infected by Listeria monocytogenes strongly depends on their zinc status
CHANDRA, 1990). The excess of minerals may also be harmful to the immune
system; even a slight exposure to heavy metals (lead, mercury and cadmi-
um) alters immunocompetance, although the exact mode of action is not
413
known in each cases.

There is increasing evidence that the concentrations of trace elements
required for healthy animals are often below what is required for animals
experiencing an immunological challenge. In the past few years there has
been a great deal of research published showing the key role of trace min-
erals in maximizing immunological function. Copper level in the diet has
been shown to affect the resistance of sheep to bacterial infections. WARD
et al. (1993) reported that prolonged exposure to molybdenum (10 ppm) and
sulphur (0.2% sulphur) decreased in vivo cell-mediated immune function in
feeder cattle. The lower in vitro viability of lymphocytes collected from steers
receiving the molybdenum and sulphur suggests that these cells are more
fragile. Either a deficiency or an excess of iron can compromise the immune
system. It has been well documented that serum iron falls early in response
to bacterial and viral infections and rebounds quickly with recovery. This
hypoferraemia is believed to be an important protective component of the
acute phase response to infection. Anaemic animals are much more sus-
ceptible to infections than those with adequate iron supply. Once the infec-
tion is established, iron supplementation has been shown to increase the
bactericidal activity of liver and splenic macrophages. For example, chicks
inoculated with S. gallinarum had increased survival when iron (100 ppm of
diet or more) was added to a basal diet containing 200 ppm of iron.
Substantial evidence has been reported that adding zinc above the
supposed requirement enhances disease resistance. However, excessive
zinc (150 ppm) also depressed immunocompetance. Zinc has both specific
and aspecific role in the immune defence mechanism. As aspecific function,
it protects integrity of skin and mucous membranes (mucosal barrier).
Besides copper and manganese, it is indispensable to the functioning of
SOD activity and thereby to the natural killer activity. Zinc enhances the
chemotaxic sensitivity and intensity of phagocytosis of PMN neutrophils,
monocytes and macrophages, moreover, support complement activity. In its
specific role, zinc regulates the maturation and function of immune cells,
among others by protecting developing lymphocytes from apoptosis. Zinc
counterbalances the glucocorticoid-induced thymocyte apoptosis; by means
of the protein kinase contributes to the activation of lymphocytes, included
NK cells. As part of the zinc-finger proteins, may influence DNA transcrip-
tion. The thymus synthesizes a 9-amino acid peptide hormone, the thy-
mulin, which is activated after having bound zinc (FRADSON et al. 2003). The
414
function of the thymulin is the maturation and activation of the T-lympho-

cytes and the stimulation of the IL-2 production. It also functions as a
transmitter between the neuroendocrin and immune systems. Its degree of
saturation with zinc is an excellent indicator of the zinc supply of the organ-
ism.
Selenium is recognized as an immunostimulant in swine, poultry and
ruminants. Often selenium and vitamin E are supplemented together. COL-
NAGO et al. (1984) challenged broiler cockerels with E. tenella oocysts; grad-
ed selenium levels from 0.1 to 1.0 ppm or 100 ppm of vitamin E/kg were
added to the diet from day one of age. Dietary supplementation of at least
0.25 ppm selenium or vitamin E reduced mortality and increased weight
gains. These authors showed that feeding 0.25 ppm selenium or more,
increased leukocyte number in the blood after infection with coccidia, which
may explain the immune enhancement. BOYNE and ARTHUR (1981) reported
that neutrophils from selenium-deficient calves had a decreased ability to
kill ingested Candida albicans, compared to neutrophils from selenium sup-
plemented calves. The authors linked the decreased effectiveness of the
neutrophils to reduced glutathione peroxidase activity resulting from sele-
nium deficiency. When glutathione peroxidase activity is reduced, peroxides
and lipid hydroperoxides tend to accumulate to toxic levels in the neu-
trophils. Rate of clinical mastitis was negatively correlated to plasma sele-
nium concentration in well-managed dairy herds (WEISS et al. 1990).
The main function of the selenium is the neutralisation of the free
radicals. It interferes to the findings that lack of selenium both alone and
combined with vitamin E deficiency affects immunocompetance. Until the
near past, epidemiologists used to explain the effect of insufficient nutrients
and active ingredients upon the resistance against the infections solely by
the debilitation of the host organism. According to this approach, the mal-
nutrition interacts with several physical and chemical barriers and immune
response, getting human and animal more susceptible to pathogenic
agents. The idea, that the host organism may have a direct effect on the
pathogens, too, has not ever emerged. On the contrary, recently it has been
proved this possibility in rodent, infected by enterovirus. The definitive evi-
dence has been provided for this change of paradigm by BECK (1997). Benign
Coxsackievirus B3 has been inoculated into selenium or vitamin E deficient
mice. The virulence of the virus increased, which could be demonstrated
also by the change of its genetic material. In other words, the benign
415
Coxsackievirus B3, by means of mutation, became capable of inducing

heart lesions. Using this knowledge, by peroral selenium application the
incidence of human cardiomyopathy (Keshan-disease) could have been pre-
vented in a selenium-poor territory of China. Consequently, the traditional
model of nutrition-infection interaction should be modified and include the
possibility of the direct influence of the host organism on the virulence of
pathogenic agents.
Selenium has a vitamin E-independent immunostimulant effect in the
marginally supplied (0.03 mg/kg of feed) calves. The protection could be
demonstrated by the increased plasma gluthathion-peroxydase and IgM
level after artificial IBR (infectious bovine rhinotracheitis) infection. There
are evidences to the increase in total IgG concentration, neutrophil and lym-
phocyte function after selenium application and the stimulation of the
agglutination after vitamin E supplementation. Both selenium and vitamin
E are able to stimulate immune response and to decrease mortality caused
by natural and artificial infections.
Added chromium is mostly beneficial in the periods of high-stress.
MONNSIE-SHAGEER and MOWAT (1993) fed 0, 0.2, 0.5, and 1 ppm supplemen-
tal chromium from high-chromium yeast, to 84 Charolais-crossed feeders
stressed due to shipment. Chromium supplementation decreased morbidi-
ty and rectal temperature at day 2 and 5 after arrival. In dairy cows chromi-
um caused increased anti-ovalbumin response but did not affect the
immune response to the red blood cells. These data suggest that chromium
supplementation may enhance resistance to mastitis in dairy cows.
Chromium reduced serum cortisol levels. Glucocorticoids, which include
cortisol, are known to suppress the immune system. Cobalt deficiency ren-
dered sheep more susceptible to bacterial infections. Reductions in candi-
dacidal activity of ovine neutrophils have been reported as a result of cobalt
deficiency. These authors noted that the changes in the health of the neu-
trophils became evident before vitamin B12 status decreased.
17.3.3. Lack of vitamins and essential fatty acids
In the circle of production or companion animals the occurrence of extreme
vitamin deficiency of overdosage is uncommon. Therefore, data concerning
the effect of the individual vitamins on immune function mostly derive from
animal trials. From the group of fat soluble vitamins, the role of vitamin A
and E is essential. Lack of vitamin A frequently accompanies PEM. Vitamin
A deficiency decreases resistance against infections. This is partly due to
416
the injury of skin and mucous membranes, partly to the impairment of

humoral and cell-mediated immune response. Deficient vitamin A supply of
growing chicken induces atrophy (lower organ weight, compared to the con-
trol) of bursa Fabricii, thymus and spleen (WEST et al. 1991). On the con-
trary, vitamin A supply increases two-three times the number of IgG-pro-
ducing cells in the mesenterial lymph nodes and in the spleen of mice with
Trichinella spiralis infection. Combined effect of supplementary vitamin A
and ?-carotene was able to decrease somatic cell count in milk of dairy cow.
The effect of carotenoids on the immune response is described in Chapter
XXII. Active form of vitamin D3 regulates transcription at cell level, acts as
an immunomodulator and promotes phagocytosis. Insufficient vitamin E
provision reduces both humoral and cell-mediated immune response. On
the other hand, extra large dosages of tocopherols are immunostimulant.
Members of vitamin B group (thiamine, riboflavin, pyridoxine, panthotenic
acid and biotin) promote antibody production. Folic acid and vitamin B12
are involved in antibody and nucleic acid production.
Lack of essential fatty acids in the diet of experimental animals
caused atrophy of lymphoid organs and the reduction both the T-cell medi-
ated and the independent immune response. Thereto, ω-3 and ω-6 fatty
acids are essential in the membrane composition of white blood cells and in
the release of prostaglandins and leukotriens. The ratio of triens to tetraens
(three and four double bonds fatty acids) and that of w-3 and w-6 fatty acids
proved to be also important in relation of immune response. Higher dosages
(5%) of omega-3-fatty acids are able to counterbalance harmful effects of
Eimeria tenella infection in poultry.
Among the water-soluble vitamins the insufficient intake of vitamin
B6 (pyridoxine), the panthotenic acid and folic acid may disturb both
humoral and cell-mediated immune response. Concerning the influence of
vitamin B12, see described at the cobalt. Vitamin C increases the phagocy-
tosis activity of macrophages. Huge amount of peroral intake of vitamin C
(2 gram/day) increased the ascorbic acid concentration of leukocytes, gran-
ulocytes and blood plasma in human (EVANS et al. 1982). Higher lymphocyte
proliferation, measured by 3H-thymidine incorporation in DNA of human
lymphocytes over 18 hours after previous incubation for 2 days with con-
canavalin A at varying ascorbic acid concentration, proved the dose-related
immunostimulant effect of vitamin C (GÜNTHER, 1998). Effect of minerals
and vitamins on the immune status is linked in many points. For example,
417
vitamin B6, in form of pyridoxal kinase, carrying magnesium and zinc, helps
the free radical production in the NK cells.
Although, most of the described data came from animal experiments,
the occurrence of diet-induced immunodeficiency is not uncommon in the
practice, either. If newborn calf delays in receiving the first colostrum por-
tion, becomes susceptible to the slight mineral deficiency of the cow.
Subclinical B12 deficiency of ewes (established by the serum B12 and
methyl-malonic acid level), caused by insufficient cobalt intake, has no
effect on the live weight of newborn lambs, but the perinatal mortality of off-
springs increases. The latter can be explained by the impaired immune sys-
tem of dams and the insufficient passive immunity, given to the lambs. If
pregnant sows of average vitamin supply were provided by 5 mg sodium
selenium and/or by 1000 mg DL-tocopherol on day 100 of gestation, the
amount of transmitted antibodies to the offspring increased. Another prac-
tical application of the new immunological finding is the segregated early
weaning (SEW), which serves the protection of young piglets from the anti-
genic challenges from the dam, weaning them in the age of 10 to 14 days
(CHEEKE, 1998). At that time the passive immunity of piglet is on the peak
and lacking the above described immunological stress, their weight gain
and feed conversion significantly improve (for more details see Chapter XX).

Feed intolerance is a reproducible phenomenon, related to an actual chem-
ical component of the diet. The common manifestation is dyspepsia and/or
diarrhoea. In the background may stand the lack of a digestive enzyme (e.g.
lactose intolerance), a special pharmacological effect (e. g. caffeine stimu-
lated peristalsis) or non-immunological histamine release (ingestion of cer-
tain seafoods, mollusc, octopus, squid, scallop, oyster, crab, lobster etc.).
Frequently and unjustified the latter is also named as allergy or sensitivity.
Feed allergy (or sensitivity) is an abnormal immunological reaction,
while the organism tuned to one of the feed proteins, like milk casein, wheat
gluten, soy protein and others. Feed antigens represent a continuous chal-
lenge, giving chance to the development of a hypersensitivity reaction. The
latter may be an acute, IgE-mediated immediate hypersensitivity reaction
within 1 to 2 hours or a T cell-mediated delayed hypersensitivity reaction
within days, resulting in bronchospasmus (see sudden cot death in human
medicine), severe scour or even death. The chronic hypersensitivity, in turn,
418
can be characterized by increased protein secretion, like secretory immune

globulin A, mucus and epithelial cells in the gut lumen, changes in peri-
stalsis and disturbed permeability of intestinal wall, caused by histological
damages (PEAKMAN and VERGANI, 1997). The specific antibodies are present
in the blood, because there is a possibility for antigen uptake when the pro-
tein digestion is incomplete and macromolecules may remain intact.
Another ways of getting antigen in the bloodstream include the non-specif-
ic absorption by the M-cells in the ileum, defects in mucosal barrier (sub-
chronic enteritis), IgA immunodeficiency or very high concentration of anti-
gens in the intestinal lumen. There are other than intestinal ways (through
conjunctiva or lungs) of intact dietary antigen intake, too. The plasmacytic
colitis with diarrhoea and increased histamine release accompanying by
itching wheals on skin are the common signs of manifestation.
Feed aversion is a learned, psychological process, based on condi-
tioned reflex, which cannot be triggered unless the questionable feed is
offered in the same physical form.

The interaction of parasitoses and nutrition can be well demonstrated on
the example of gastro-intestinal worms. The following data derive from
experiments using the following animal models: Chabertia ovina, in sheep
large intestine; Haemonchus contortus in abomasum of sheep and goat;
Obeliscoides cuniculi in the rabbit stomach; Oesophagostomum
columbianum in the large intestine of sheep and goat; Ostertagia circum-
cincta in the abomasum of sheep and goat; Ostertagia ostertagi in the cattle
abomasum; Trichostrongylus axei in the cattle abomasum; Trichostrongylus
columbriformis in the sheep small intestine and the Trichostrongylus vitrinus
in the sheep small intestine.
Presence of each of the enumerated helminths decreased the average
daily gain of growing animals, even in many cases caused loss of weight.
The characteristic localization of the worms showed no correlation with the
extent of the drop in live weight, but the gravity of infection did. Under a cer-
tain number of worms (“threshold value”) the host organism did not show
detectable changes (TITCHEN et al. 1997). The main reason of the harmful
effect is the decreased feed intake, but the feed utilization also worsened.
Owing to the increased heat production part of the metabolizable energy
lost. The state is reversible, even some adaptation could be observed.
419
Parasitic infection changed the body composition: the water content

increased that of protein and fat dropped; the calcium and phosphorus con-
centration of bones decreased. by 15 to 45%. The wool mostly consists of
protein. This is the reason why even a moderate gastro-intestinal helminthi-
asis decreased the length and diameter of wool fibres. Against PEM the
sequence of tissue susceptibility is as follows: wool, fat depots, muscle,
blood, liver, spleen, heart and the more resistant are the nerves and the
brain (HAMMOND, 1942). Protein-energy malnutrition decreased the resist-
ance against the new infections, but had no influence on the already pres-
ent worms, included their egg production. Resistance to internal parasites
is also compromised with copper deficiency. Copper deficient lambs inocu-
lated with Taenia axei and Taenia colubriformis had maximum faecal egg
counts 2 weeks sooner and became more hypoalbuminaemic than lambs
receiving supplemental copper.
Helminths, developing in the animal, may cause serious local lesions:
for example Ostertagia circumcincta is capable of destroying glands of abo-
masal wall, causing haemorrhage. In small intestine the flattening and atro-
phy of villi, in turn, in the large intestine ulcer and haemorrhage are typi-
cal signs of worm damages. After the appropriate anthelmintic treatment
regeneration took up approximately three weeks. Devasteted intestinal cells
are replaced by functionally undifferentiated cells; in the mucous mem-
brane hyperplasia and inflammation develop and the wall becomes thick
The functionally immature cells, covering the secretory and absorptive sur-
face have a higher permeability to the macromolecules (“leak lesion”), which
results in pathological alterations (BLIKSLAGER et al. 1997). In the abomasum
the pH may change; in the small intestine the production of brush-border
enzymes declines.
Anaemia and the change of plasma proteins are frequent concomitant
of the gastro-intestinal parasitosis. Generally the albumin level drops, that
of globulin and total protein increases. In case of serious infection the con-
centration of the latter drops, too. Infection is frequently manifested also in
oedema, diarrhoea, lower blood pepsinogen level and change in activity of
some liver enzymes.
Effects of worms on digestion
Under the influence of intestinal parasites the gut motility modifies:
the peristalsis generally slows down, unless diarrhoea occurs at the same
time. Worms’ toxins stimulate the production of gastrointestinal hormones,
420
like gastrin and cholecystokinin, causing reduction in voluntary feed intake.

The production of hydrochloric acid in the abdomen, except the immediate
vicinity of helmet, decreases (GUY et al. 2000). Digestion and absorption of
nutrients are also reduced and the endogenous nitrogen excretion elevates.
To replace the amount of plasma proteins, excreted into the gut lumen, the
protein synthesis in blood and liver is enhanced, in turn, in the muscles
lowered. Described alterations of the postabsorptive metabolism can be mit-
igated and shortened by benzimidazol treatment.
As a consequence of reduced feed intake, less volatile fatty acid and
ammonia are produced in the rumen. Part of the amino acids in gut lumen
undergoes deamination; the released ammonia will be absorbed, transforms
into urea, in turn, will be excreted by the urine. Consequently the higher
urinary urea and methyl-histidine from the muscle mobilization reflects
increased protein losses. Higher turnover of the urea cycle involves a high-
er need to arginine and vitamin B6. Extra protein synthesis (to replace the
plasma proteins) uses part of the available metabolizable energy. As a sum-
marized result, both energy and protein balance become negative. As a con-
sequence, the body composition changes and the percentage of protein in
the carcass decline. Gastro-enteral helminthiasis alters the utilisation of
calcium, phosphorus and magnesium. Thereby in growing animals the
measure and density of bone (poor mineralization) decrease. Blood level of
the mentioned minerals generally do not change, except phosphorus, which
in case of a serious infection may drop.
Nutrition and the other parasitic infections
The leading symptoms of the sheep gastro-intestinal helminthiasis, viz. the
negative energy and protein balance and the changes in the body composi-
tion has also been found in wild boars, infected with stomach
(Physocephalus sexalatus and Ascarops strongylina), as well as with lung
worm (Metastrongylus spp.). Basically similar results have been received,
while the artificial infection of poultry with intestinal coccidia or pigs with
different helminths (Strongyloides ransomi, Stephanurus dentatus, Ascaris
suum, Oesophagostomum spp. and Trichuris suis). Infection of rabbits with
Eimeria stiedai, causing biliary coccidiosis (or hepatic coccidiosis), signifi-
cantly decreased voluntary feed intake (possibly owing to TNF-release) and
the digestibility of nutrients, especially that of the fats (YVIRE and GUILAUME,
1976). The latter finding is traceable to the hepatomegalia, liver lesions and
chronic jaundice, caused by the coccidia.
421
ZHU et al. (2000) investigated the effect of Eimeria maxima in growing

chickens. The extent of infection and the oocyte excretion showed a strong
negative correlation with the plasma carotenoid level and strong positive
correlation with the blood nitrogen oxide and ?-interferon concentration, as
signs of protection. Coccidiosis may alter carotenoid metabolism in many
points: only an indirect effect at the ingestion (minor change in feed con-
sumption), a direct alteration of the intestinal absorption of carotenoids,
due to the altered permeability;; coccidia effect transport process into the
liver; coccidia also directly modify the tissue storage (included the eggs) and
the faeces elimination of the carotenoids by speeding of transit time. As pos-
sible consequences, the tissues and the egg-yolk looses colour and general-
ly the carotenoid requirement increases. As mentioned above, higher
dosages (5%) of omega-3-fatty acids are able to counterbalance harmful
effects of Eimeria tenella infection.
17.6. Nutrigenomics
Referring to the process of protein synthesis, the gene expression is regu-
lated by the transcription, translation, mRNA processing, mRNA stability
(half-life) and the protein synthesis of mRNA, ribosome and tRNAs (CLARKE,
1993). Nowadays more and more well-defined pathway provides clear evi-
dences about the effect of feed components on gene expression and made
birth of a new science, viz. the nutrigenomics. Genomics contains knowl-
edge related to the study of the functions and interactions of all genes in the
genome, including their interactions with environmental factors
(GUTTMACHER and COLLINS, 2002; FALUS, 2005). The nutritional science
aligned itself with functional genomics and very recently the new area of
“nutritional genomics” or “nutrigenomics” emerged. “Research and discov-
ery in nutritional genomics elucidate the reciprocal interactions among nutri-
ents, metabolic intermediers, and the mammalian genome. Understanding the
interrelationship among animal and human genetic diversity, genome func-
tion, and dietary components will enable precise manipulation of genome
function and stability throughout the life cycle for optimal health and disease
prevention” (STOVER, 2004).
17.6.1. Basics
Nutrigenomics is a new discipline that examines nutrient-gene interactions
on a genome-wide scale. Nutrigenomics is the intersection of four areas:
health, animal performance, diet and genomics. New tools that take advan-
tage of the numerous available whole-genome sequences to study diet-gene
422
interactions include microarrays (transcriptomics), proteomics,

metabolomics and epigenomics. Microarrays can measure the changes in
mRNA expression of every gene in the genome in a dose-dependent manner
for a particular nutrient or drug. Proteomics measures the changes in the
whole proteome, such as post-translational modifications. Metabolomics
measures the changes in the entire metabolome, which is generally defined
as metabolites with molecular weights less than 2000 Daltons, in a dose-
dependent manner for a particular nutrient or drug (LEE et al. 2005):
Epigenomics measures the changes in the epigenome, the histone post-
translational modifications and DNA methylation pattern, in a dose-depend-
ent manner for a particular nutrient or drug. Finally, the intersection of
health and genomics relates to the identification of markers of aging, dis-
ease predisposition and behavioural genomics.
Beyond inherited genetic abnormalities, alterations in gene expres-
sion can also contribute to disease processes. Recently it has been suggest-
ed that environment may alter such genes and thus be a direct influence on
disease. Diet is a potent mechanism for altering the environment of cells in
most organs, particularly the gastrointestinal tract. Influence of nutritional
factors on intestinal gene regulation derives from the major nutrients, vita-
mins and minerals (included insulin, which is not a dietary component but
responds to dietary changes) and butyrate, a short chain fatty acid produced
by normal intestinal flora (SANDRESON and NAIK, 2000). Manipulation of diet
(nutritional treatment) may be a means of treating intestinal disorders. By
changing the nutrient supply, the functioning of genes in intermediary
metabolism may also be influenced.
17.6.2. Fats and fatty acids
DIETARY FAT has profound effects on gene expression, leading to changes in
metabolism, growth and cell differentiation. The effects of dietary fat on gene
expression reflect an adaptive response to changes in the quantity and type
of fat ingested. SPECIFIC FATTY ACID–REGULATED transcription factors have
been identified in bacteria, amphibians, and mammals. In mammals, these
factors include peroxisome proliferator–activated receptors (PPARα, β, and
γ), HNF4κ, NFαB, and SREBP1c). These factors are regulated either by
direct binding of (oxidized) fatty acids, fatty acyl–coenzyme A or oxidized
fatty acid (eicosanoid) regulation of cell surface receptors as well as activa-
tion of signalling cascades or by oxidized fatty acid regulation of intracellu-
lar calcium levels (JUMP, 1999). At the cellular level, the physiological
423
response to fatty acids will depend on the quantity, chemistry and duration
of the fat ingested; the cell-specific fatty acid metabolism (oxidative path-
ways, kinetics and competing reactions); the cellular concentration of spe-
cific nuclear and membrane receptors; and finally involvement of specific
transcription factors in gene expression. These mechanisms are involved in
the control of carbohydrate and lipid metabolism, cell differentiation and
growth and cytokine, adhesion molecule and eicosanoid production.
17.6.3. Starvation
Increased dietary fat or higher NEFA from STARVATION-CAUSED mobilization
may enhance hepatic oxidation and decrease esterification of fatty acid by
reducing fatty acid synthase expression, preventing thereby triglyceride
accumulation in liver (LEONE et al. 1999). Fasting activates glucose produc-
tion and lipolysis; on the contrary, re-feeding after fasting by fat-free, high-
carbohydrates diet activates fat synthesis by strikingly enhancing fatty acid
synthase transcription rate FUKUDA et. al. 1999). While using for re-feeding
lipids, they will decrease the activity of transcriptional enzymes (phospho-
fructokinase, pyruvate-dehydrogenase and acetil-CoA-carboxylase) in liver,
minimizing fat deposition. The key factors in this pathway are the nuclear
hormone receptor subfamily, the peroxisome proliferators-activated recep-
tors (PPAR), identified first in mouse liver tissue (SCHONNJANS et al. 1996).
Elements, responsive to the peroxisome proliferators have been found in the
promoter regions of several genes encoding proteins that are involved in lipid
metabolism. Activation of PPAR by free fatty acid, eicosanoids or xenobiotics
help to bind PPAR to specific areas of target genes, leading to activation or
repression of gene expression. Interestingly enough, dietary fat does not
suppress body lipid mobilization in fresh lactating dairy cows. Anyway, it is
clearly demonstrated that dietary carbohydrates both independently and
through insulin effect, deeply influences transcription of fatty acid synthase
gene. In rodents, immobilization stress, starvation, diabetes and chemical
compounds (e.g. clofibrate) may also stimulate the expression of PPAR and
peroxisomal β-oxidation of fatty acids (SINGH, 1997). Ingestion of high fat
diets downregulates insulin responsive glucose transporter, GLUT4 protein
expression in adipose tissue, thereto increases leptin gene expression
(HOUSEKNETCH et al. 1998). Ingestion of high amounts of OLEIC ACID OR ω-3
FATTY ACIDS downregulates expression of leptin, fatty acid synthase, lipopro-
tein lipase and phosphoenolpyruvate carboxykinasee in retroperitoneal adi-

pose tissue of pig. There is no effect in subcutaneous adipose tissue. HSU
424
and HUANG (2006) proved that reduced fat mass in rats fed a high oleic acid-
rich safflower oil diet is associated with changes in expression of hepatic
PPARα and adipose sterol regulatory element-binding protein-1c-regulated
genes (SREBP-1c). Feeding protein-rich diet, the mRNA shortage for fatty
acid synthase in the adipocytes, but in liver tissue (CLARKE, 1993), moderat-
ing total body fat. The hepatic fat synthesis, in turn, can be inhibited by
offering unsaturated fatty acids in diet.
17.6.4. Dietary essential fatty acids are the precursors for eicosanoids.
Among the eicosanoids derived from arachidonic acid, prostaglandin E2 is
known to possess immunosuppressive actions. Thus, it has been a prevail-
ing hypothesis that the immuno-modulatory roles of dietary fatty acids are
mediated, at least partly, through the alteration of prostaglandin biosyn-
thesis. Emerging evidence suggests that fatty acids can additionally act as
second messengers, regulators of signal transducing molecules or transcrip-
tion factors (HWANG, 2000). Acylation with long-chain fatty acids can occur
on a variety of signalling molecules and can affect their membrane translo-
cation and functions. Dietary fatty acids can alter functional properties of
lipid mediators by changing the composition of acyl part of these molecules.
Evidence accumulated recently indicates that long-chain unsaturated fatty
acids and their metabolites bind and activate peroxisome proliferator–acti-
vated receptors (PPARs). Summarising, it becomes clear now that multiple
steps in various receptor-mediated signalling pathways can be modulated
by dietary fatty acids. In turn, PPAR expression is nutritionally regulated by
high-fat diet, fasting and linoleic acid.
17.6.5. Conjugated linoleic acids (CLA) are potent activators of PPARγ,
which in turn, this function is related to the anticancerogenic effect.
Moreover, CLAs induce apoptosis in adipocytes. Milk is one of the richest
natural sources of CLAs. Namely, milk fat derives from two sources: approx-
imately half of it comes from the uptake of blood fatty acids, whilst the
remaining part is formed by the de novo fatty acid synthesis in the mam-
mary gland. Composition and function of the rumen microflora have great
influence on the fatty acid metabolism: unsaturated fatty acids undergo bio-
hydrogenation through the action of bacterial isomerases and reductases.
For example, under normal conditions, the linoleic acid (cis-9,cis-12 C18:2)
first becomes conjugated linoleic acid, CLA (cis-9,trans 11 CLA or rumenic
acid), which in turn transforms into vaccenic acid (trans-11 C18:1) and
finally into strearic acid (C18:0). There is also a reverse pathway from
425
vaccenic acid under the influence of Δ9-desaturase enzyme activity in the

liver, adipose tissue and mammary gland to form rumenic acid. The latter,
the cis-9, trans-11 CLA or rumenic acid gives more than two third of the milk-
fat. It is also present in beef meat. In the meantime, using rat model, the
anticarcinogenic effect of rumenic acid had been discovered, namely it was
capable of reducing incidence of breast cancer (AIMUTIS, 2004). During cer-
tain circumstances, the rumen environment and bacterial population differ
from the average, for example while ingestion of diets low in effective fibre.
Altered microflora will produce different CLAs than the rumenic acid, name-
ly from the linoleic acid predominantly the trans-10,cis-12 CLA will be
formed. The latter is an efficient inhibitor of the milk fat synthesis in the
mammary gland: “bIOHYDROGENATION THEORY OF THE MILK FAT DEPRESSION”
(BAUMAN and GRIINARI, 2003). This hypothesis has been confirmed both in
cow and sheep and lactating mice (LIN et al. 2004), by applying the men-
tioned CLA isomer. Thus, a natural bacterial product is capable of influ-
encing mammary gene expression and thereby controlling milk fat produc-
tion.
17.6.6. Metals
Bivalent metals strongly influences gene expression, for example both par-
enteral and peroral ZINC and CADMIUM application enhances transcription
rate in metallothionein gene in intestinal tissue (OULETIE et al. 1982).
Cadmium acts also in prolonging the half-time of metallothionein mRNA in
hepatocytes (DE et al. 1991). Zinc acts as part of the “zinc-fingers”, fixing
activator proteins to the active segments of the DNA. Iron influences trans-
ferrin and ferritin concentration through influencing mRNA stability and the
translation rate (BREMNER and BEATTIE, 1990). The dietary supplementation
with zinc oxide increases IGF-I and IGF-I receptor gene expression in the
small intestine of weanling piglets (LI et al. 2006).
17.6.7. Vitamins
VITAMIN A exerts its regulatory function in form of retinol and retinoic acid.
Target tissues have different types and numbers of intracellular retinol and
nuclear retinoic acid receptor proteins. The most important sites are in the
adrenals, testes, cerebellum, kidneys, prostate, cerebral cortex, skin and in
the visceral tissue. After retinoic acid binds to its receptor, it will stimulate
the transcription and translation of vitamin A-responsive genes, like that
are involved in cell differentiation (GH, glycerolphosphat dehydrogenase
(CLARK and ABRAHAM, 1992) and leptin production among others. Deficient
426
vitamin A state negatively influenced hepatic phosphoenolpyruvate car-

boxykinase (PEPCK) gene expression in mice. By perorally application of
retinoic acid the expression could be restored (SCRIBNER et al. 2005).
Similarly to retinoic acid, the actions of ACTIVE VITAMIN D are mediated by
nuclear hormone receptors. The vitamin D receptor directly binds to DNA at
vitamin D response elements as a homodimer or as a heterodimer to acti-
vate gene transcription. Ligand binding to the vitamin D receptor forms a
complex of co-activators that modulates gene expression in different cell
types (BOHNSACK and HIRSCHI, 2004). Role of BIOTIN in gene expression has
been recognized by the significant depression of ornithine transcarboxylase
gene expression in biotin deficiency. The consequent loss of enzyme activi-
ty is the basis for a hyperammonaemia (YUICHI et al. 1996).
17.6.8. Nutriceuticals
In the experiment of SELVARAJ and KLASING (2006) broiler chicken were fed
LUTEIN and EICOSAPENTAENOIC ACID (EPA) enriched feed mixture. Macrophages
collected at the end of this trial, the treatments were repeated in this cell
culture, previously stimulated by lipopolysaccharide. As a conclusion,
lutein and EPA interact to modify inducible nitric oxide synthase and mRNA
levels through the PPARγ and retinoid acid X receptor pathway. TSAI et al.
(2005) demonstrated that GARLIC ORGANOSULPHUR SUBSTANCES upregulate the
expression of the π-class of glutathione S-transferase in rat primary hepa-
tocytes.
17.6.9. Possible practical-clinical applications

LEAN MEAT PRODUCTION: supplemental CLA in pig’s diet, by activating PPARγ-
responsive genes in adipose tissue, acted as a repartitioning agent, reduc-
ing both voluntary feed intake and whole body fat content (DUGAN et al.
1997). The better knowledge of the metabolism of long-chain fatty acids
helps in designing daily ration for TRANSIENT HIGH-YIELDING DAIRY COW. It
means that instead of supplemental fat for the fresh cow, the provision of
absorbable amino acids and glycogenic precursors, like absorbed glucose,
propionate and isobutyrate should be emphasized (DRACKLEY, 1999). By
application of the inhibitor trans-10, cis-12 CLA, the milk fat production
may be reduced in the critical periods of dairy cow in a controlled manner.
FLUSHING EFFECT (sow, ewe) can partly be explained by the effect of excess
portions (i.e. high-carbohydrates with the concentrate intake), that activate
fat synthesis by strikingly enhancing fatty acid synthase transcription rate.
427
The LYSINE-SPARING EFFECT of sodium bicarbonate (see Chapter XI), or

the beneficial EFFECT OF EPA ON THE CARTILAGE DEVELOPMENT (see Section
21.2.3.2.) can be explained on the level of gene expression.
CLA ISOMERS beyond the described anticancerogenic property have
also proved to have ANTIATHEROGENIC, ANTIOBESITY AND ANTIDIABETIC charac-
teristics both against Type I and Type II diabetes. To prevent obesity and
Type II diabetes, the direct modulation of gene expression by nutrients is
also a potential mechanism. All these pathways converge finally to decrease
expression of protein-glucokinase, key co-activators of PPARα PPARβ, and
PPARγ leading to mitochondrial dysfunction and reduced energy expendi-
ture, all which enhance the risk for obesity and insulin resistance (PATTI and
KAHN, 2004). Genomics can be used also in the detection of latent dog and
cat diabetes (SWANSON, 2004).
FOR FURTHER READING

Colnago, G.L., Jensen, L.S. and Long., P.L. (1984): Effect of selenium and vitamin E on
the development of immunity to coccidiosis in chickens. J. Poult. Sci. 63, 1136-
1148.
Fekete, S.Gy. and Brown, D.L. (2007): Veterinary aspects and perspectives of nutrige
nomic: A critical rewiew. Acta Vet. Hung. 55, 229-239
Fekete, S.Gy. and Kellems, O.R. (2007): Interrelationship of feeding with immunity and
parasitic infection: a rewiew. Veterinari Medicina (Vet. Med- Czech) 52 (4), 131-
143.
Hucker, D.A., and Yong, W.K. (1986): Effects of concurrent copper deficiency and gas
trointestinal nematodiasis on circulating copper and protein levels, liver copper
and bodyweight in sheep. Vet. Parasitol. 19, 67-75
Yvore and Guillaume (1976): The effect in rabbit, of hepatic coccidiosis on digestibility of
fat and energy. Annls. Rech. Vet. 7, 343-348..
Zhu, J.J., Lillehoj, H.S., Allen, P.C., Yun, C.H., Pollock, D., Sadjadi, M. and
Emara, M.G. (2000): Analyse of diseaseresistance-associated parameters in broiler chick-
ens challenged with Eimeria maxima. Poult. Sci. 79, 619-625.
428
Chapter XVIII
REGULATIONS ON FEEDINGSTUFFS
© Josef Leibetseder
18.1 Council Directive 96/25/EC of 29 April 1996
18.2.Council Directive 79/373/EEC of 2 April 1979
18.3.Council Directive 82/471/EEC of 30 June 1982
18.4.Council Directive 93/74/EEC of 13 September 1993
18.5.Council Directive 90/167/EEC of 26 March 1990
18.6. Regulation (EC) No 1831/2003
18.7. Commission Directive 2001/79/EC of 17 September 2001
18.8.Directive 2003/32/EC
18.9.Regulation (EC) No. 882/2004
18.10.Others
18.11.Liability
Chapter XVIII: REGULATION - FOOD SAFETY
I
ntroduction.In the past farmers produced the feed for their livestock
themselves at the farm. Legal regulations on feedingstuffs were,
therefore, not necessary. Later on by-products of different local pro-
ductions like bran, brewer’s yeast or whey were used. Because of the obvi-
ously sufficient information about value and quality of the products due to
the near production and because of every confidence in the producer there
was no need for regulations. About hundred years ago the use of feed-
ingstuffs from afar became more common and feed companies started the
production and the sale of compound (mixed) feed. Consequently, the trad-
ing in feedingstuffs also spread out. Traceability of feed was not possible
anymore. Increasing frequency of problems and subsequent trials due to the
insufficient quality of feedingstuffs and the competition between companies
were the reasons for regulations. In many countries, therefore, national reg-
ulations on feedingstuffs were put into force. Finally the free market of feed
within the European Community led to international regulations and direc-
tives on feedingstuffs. The general intentions of these regulations are the
protection of animal health, the avoidance of risks of the consumers due to
residues of undesirable substances in food of animal origin, and the reduc-
tion of the environmental pollution by livestock. The aim of the regulations
is also to see to the fair competition in order to protect the customer eco-
nomically against incorrect advertising (e.g. by the labelling).
The most elaborate regulations are the U.S. feed law and the regula-
tions and directives of the European Community. The U.S. feed law is avail-
able in Animal Food (Feed) Product regulation (US Code of Federal
Regulations; Federal Food, Drug, and Cosmetic Act). EC Regulations are valid
for all Member States, whereas the Directives are addressed to the States
which have to put them into force by national legislation. The first EC
Directives were published in the Official Journal in the beginning of the sev-
enties of the last century and are in general amended quite often. In regard
to human health the Commission of the European Community adopted the
White Paper on Food Safety in 2000 which is the basis of a new legal frame-
work covering the whole of the food chain, including feed production, estab-
lishing a high level of consumer health protection and clearly attributing
primary responsibility for safe food production to industry, producers and
suppliers. All these Regulations and Directives are available in
430
http://europa.eu.int/eur-lex.
Before the adoption of the EU Regulations and Directives by the
Council of the European Union the competent committees and scientific
panels are asked to put forward their opinion. These committees (Standing
Committee on Feedingstuffs, Standing Committee on the Food Chain and
Animal Health, and the Scientific Committee on Animal Nutrition, SCAN)
belong to the Health and Consumer Protection Directorate General of the
European Commission. With the founding of the European Food Safety
Authority (EFSA) in 2003 SCAN was replaced by some scientific panels
(Scientific Panel on Feed Additives and Products or substances used in ani-
mal feed (FEEDAP), Scientific Panel on Contaminants in the Food Chain
(CONTAM) belonging to EFSA.
In the following a short overview of the most relevant EU regulations
and directives are given (this overview only refers to the most recent by May
2005 amended versions of the regulations and directives):
circulation and the use of feed materials including definitions and descrip-
tions,
18.1. COUNCIL DIRECTIVE 96/25/EC OF 29 APRIL 1996 ON THE CIRCULATION

AND USE OF FEED MATERIALS (Official Journal of the European Communities
(O J) No L 125, 23.5.1996, p.35)
This Directive shall apply to the circulation and use of feed materials
within the Community without prejudice to other Community provisions in
the field of animal nutrition. It defines “feed material”: various products of
vegetable or animal origin, in either natural state, fresh or preserved, and
products derived from the industrial processing thereof, and organic or
inorganic substances, whether or not containing additives, which are
intended for use in oral animal feeding either directly as such, or after pro-
cessing, in the preparation of compound feedingstuffs or as carriers of pre-
mixes and “putting into circulation” or “circulation”: the hooding of any
product intended for animal nutrition for the purpose of sale, including
431
offering for sale, or any other form of transfer, whether free or not, to third
parties, and the sale and other forms of transfer themselves.
Feed materials may be put into circulation in the Community only if
they are sound, genuine and of merchantable quality. Feed materials must
not present any danger to animal or human health or to the environment
and must not be put into circulation in a manner that is liable to mislead.
They have to show on an accompanying document or where appropriate on
the packaging, on the container ore on a label attached thereto the words
“feed material”, the name of the feed material, the net quantity, the name
and address of the producing establishment, the approval number and the
reference number of the batch or any other particulars which ensure that
the fed material can be traced. The names of the feed materials are listed
in Part B of the Annex of the Directive. Other information may be given in the
same way related to objective or quantifiable parameters which can be sub-
stantiated and that it cannot mislead the purchaser. Where, in case of feed
materials from a third country, it has not been possible to provide the nec-
essary guarantees provisional composition data may be allowed.
Feed materials containing a level of undesirable substances or prod-
ucts in excess of that permitted (see Directive 2002/32EC) may be put into
circulation only if they are intended for approved establishments manufac-
turing compound feed and labeled like this.
The indications printed on the accompanying documents, on the
packaging, on the container or on a label shall be written in at least one or
several languages which the country of destination shall determine from
among the national or official languages of the Community. A numerical
coding system for the listed feed materials enabling feed to be identified at
international level may be adopted and a list of materials whose circulation
or use for animal nutrition purpose is restricted or prohibited shall be
drawn up. Member States shall make necessary arrangements for compli-
ance with the requirements of this Directive to be officially monitored, at
least by sampling during circulation. The Directive contains an ANNEX with
three Parts.
Part A gives some general information: Explanatory Notes on the cri-
teria of listing and naming of feed materials in Part B and the division of
Part B into 12 chapters. Provisions regarding botanical and chemical purity:
Provisions regarding designation. Provisions regarding the glossary, which
refers to the main processes used for the preparation of feed (e.g. concen-
tration, decortication, drying, extraction, extrusion, flaking, flour milling,
heating, hydrogenation, hydrolysis, pressing, pelleting, pregelatinisation,
refining, wet-milling, crushing, desugaring). Provisions regarding levels indi-
cated or to be declared as specified in Part B and C: Provisions regarding
denaturing and binding agents:. Provisions regarding minimum tolerance
indicated or to be declared as specified in Part B and C. Provisions concern-
ing the labeling of feed materials comprising protein derived from mammalian
432
Chapter XVIII: REGULATION -FOOD SAFETY
tissue: The labeling must contain the following statement: ”This feed mate-
rial comprises protein derived from mammalian tissue the feeding of which
to ruminants is prohibited.” If a Member State prohibits the use also for
other species of categories of animals, these species or categories must be
mentioned additionally. This does not apply to different products like milk
and milk products, gelatin and hydrolysed proteins with a molecular weight
below 10 000 daltons derived from hides and skins obtained from animals
under certain conditions, dicalcium phosphate derived from defatted bones,
and dried plasma and other blood products. This regulation was introduced
because of the bovine spongiform encephalopathy (BSE).
Additional measures were taken by the Council Decision
2000/766/EC of 4 December 2000 concerning certain protection measures
with regard to transmissible spongiform encephalopathies (TSE) and feed-
ing of animal protein. In this Decision “Processed animal protein” means
meat and bone meal (MBM), meat meal, bone meal, blood meal, dried plas-
ma and other blood products, hydrolysed proteins, hoof meal, horn meal,
poultry offal meal, feather meal, dry greaves, fish meal, dicalcium phos-
phate, gelatine and any other similar products including mixtures, feed-
ingstuffs, feed additives and premixtures, containing these products.
Member States shall prohibit the feeding of processed animal proteins to
farmed animals which are kept, fattened or bred for the production of food.
The prohibition shall not apply to the feeding of: fishmeal to animals other
than ruminants, gelatine of non-ruminants for coating additives, dicalcium
phosphate and hydrolysed proteins obtained in accordance with the condi-
tions to be fixed in accordance with the procedure laid down in Article 17 of
Directive 89/622/EEC, milk and milk products to farmed Animals which
are kept, fattened or bred for the production of food.
With these exceptions Member States shall: prohibit the placing on
the market, the trade, the importation from third countries and the expor-
tation to third countries of processed animal proteins intended for the feed-
ing of farmed animals which are kept, fattened or bred for the production of
food and ensure that all processed animal proteins intended for the feeding
of farmed animals which are kept, fattened or bred for the production of
food are withdrawn from the market, distribution channels and from on-
farm storage. Member States shall ensure that animal waste, as defined by
Directive 90/667/EEC of 27 November 1990 laying down the veterinary
rules for the disposal and processing of animal waste, for its placing on the
market and for prevention of pathogens in feedstuffs of animal or fish ori-
gin, last amended by the 1994 Act of Accession, is collected, transported,
processed, stored or disposed of in accordance with the Directive,
Commission Decision 97/735/EC of 21 October 1997 concerning certain
protection measures with regard to trade in certain types of mammalian
animal waste, amended by Council Decision 1999/534/EC, and Council
Decision 1999/534/EC of 19 July 1999 on measures applying to the pro-
433
cessing of certain animal waste to protect against transmissible spongiform

encephalopathies.
Part B contains a non-exclusive list of 166 main feed materials
arranged in the 12 chapters (Cereal grains, their products and by-products;
Oil seeds, oil fruits, their products and by-products; Legume seeds, their
products and by-products; Tubers, roots, their products and by-products;
Other seeds and fruits, their products and by-products; Forages and
roughages; Other plants, their products and by-products; Milk products,
Land animal products; Fish, other marine animals, their products and by-
products; Minerals; Miscellaneous) including the number, name, descrip-
tion and compulsory declarations (e.g. 1.02; Oat flakes; Product obtained by
steaming and rolling dehusked oats. It may contain a small proportion of
oat husks; Starch).
Part C gives provisions regarding the name and the declaration of cer-
tain constituents of non-listed feed materials. For these feed materials made
of 18 different categories compulsory declarations of certain constituents
must be indicated (e.g. Products and by-products of oil seeds, oil fruits:
Crude protein, if > 10%, Crude fat, if > 5%, Crude fibre).
18.2. COUNCIL DIRECTIVE 79/373/EEC 2 APRIL 1979 ON THE CIRCULATION

OF
OF COMPOUND FEEDINGSTUFFS WITH AMENDMENTS (Official Journal of the
European Communities (O J) No L 86, 6.4.1979, p. 30)
This Directive shall apply to compound feedingstuffs which are mixtures
of feed materials, whether or not containing additives, for oral animal feed-
ing in the form of complete or complementary feedingstuffs. For all com-
pound feedingstuffs except those intended for pets other than dogs and
cats: the feed materials to be declared in this Directive. Compound feed-
ingstuffs may not be marketed unless they show on the label the following
particulars:
The Annex consists of Part A (General provisons) and Part B (Declaration of

analytical constituents in the different types of compound feedingstuffs).
434
18.3. COUNCIL DIRECTIVE 82/471/EEC OF 30 JUNE 1982 CONCERNING CERTAIN

PRODUCTS USED IN ANIMAL NUTRITION, amended by some Directives and
Regulations (Official Journal of the European Communities (O J) No L
213, 21.7.1982, p. 8)
This Directive concerns products which act as direct or indirect protein
sources, are manufactured by certain technical processes and are put into
circulation within the Community as feedingstuffs or in feedingstuffs.
Member States shall prescribe that feedingstuffs belonging to one of the
product groups listed in the Annex or containing such products may be
marketed only if the product in question appears in the Annex and any con-
dition laid down therein are fulfilled. The Directive contains some more
detailed regulations about the duties of the Member States and the amend-
ment of the list in the Annex. In the Annex the following product groups are
listed:
Proteins obtained from the following groups of micro-organisms:
Non-protein nitrogenous compounds:

-Urea and its derivatives
-Ammonium salts
-By-products from the production of amino acids by fermentation
Amino acids and their salts
Methionine, Lysine, Threonine, Tryptophan
Analogues of amino acids
Analogues of methionine
18.4. COUNCIL DIRECTIVE 93/74/EEC OF 13 SEPTEMBER 1993 ON FEED-

INGSTUFFS INTENDED FOR PARTICULAR NUTRITIONAL PURPOSES WITH AMENDMENTS
(OFFICIAL JOURNAL OF THE EUROPEAN COMMUNITIES (O J) No L 237,
22.9.1993, p.23)
‘Feedingstguffs’ intended for particular purposes’ shall mean compound
feedingstuffs which, by virtue of their particular composition or method of
manufacture, can be clearly distinguished from both ordinary feedingstuffs
and medicated feedingstuffs, and are presented as intended to meet specific
nutritional requirements.
‘Particular nutritional purpose’ shall mean the purpose of satisfying the
specific nutritional needs of certain pets or productive livestock whose
process of assimilation, absorption or metabolism could be temporarily
impaired or is temporarily or irreversibly impaired and are therefore able to
derive benefit from ingestion of feedingstuffs appropriate to their condition.
435
18.5. COUNCIL DIRECTIVE 90/167/EEC OF 26 MARCH 1990 LAYING DOWN THE

CONDITIONS GOVERNING THE PREPARATION, PLACING ON THE MARKET AND USE OF
MEDICATED FEEDINGSTUFFS IN THE COMMUNITY (OFFICIAL JOURNAL OF THE
EUROPEAN COMMUNITIES (O J) No L 092, 7.4.1900, p.42)
The following definition shall apply: ‘authorized medicated pre-mix’: any
pre-mix for the manufacture of medicated feedingstuffs which has been
granted an authorization in accordance with this Directive. Member States
shall prescribe that, as regards the medicinal component, medicated feed-
ingstuffs may be manufactured from authorized medicated pre-mixes only.
The Directive also regulates the conditions of manufacturing and of placing
on the market medicated feedingstuffs. Member States shall ensure that
medicated feedingstufs are not supplied to stockfarmers or holders except
on presentation of a prescription from a registered veterinarian on certain
terms.
Where medicated feedingstuffs are administered to animals whose
meat, flesh, offal or products are intended for human consumption, the
stockfarmer or holder of the animals concerned must ensure that treated
animals are not slaughtered in order to be offered for consumption before
the end of the withdrawal period and that products obtained from a treated
animal before the end of such a withdrawal period are not disposed of with
a view to their being offered for human consumption.
Medicated feedingstuffs shall be issued directly to the stockfarmer or
holder of the animals only by the manufacturer or distributor specially
approved. Medicated feedingstuffs for food animals may not be issued
unless they do not exceed the quantities prescribed for the treatment in
accordance with the veterinary prescription and they are not issued in
quantities greater than one month’s requirements.
18.6. REGULATION (EC) NO 1831/2003 OF THE EUROPEAN PARLIAMENT AND THE

COUNCIL OF 22 SEPTEMBER 2003 ON ADDITIVES FOR USE IN ANIMAL NUTRITION
(Official Journal of the European Communities (O J) No L 268,
18.10.2003, p.29)
436
The considerations of the European Parliament for the approval of

this regulation were the following: The free movement of safe and whole-
some food and feed is an essential aspect of the internal market and con-
tributes significantly to the health and well-being of citizens, and to their
social and economic interests. A high level of protection of human life and
health should be assured in the pursuit of Community policies. The basic
principle should be that only those additives approved under the procedure
provided for this Regulation may be placed on the market, used and
processed in animal feeding under conditions set out in the authorization.
In order to protect human health, animal health and the environment,
feed additives should undergo a safety assessment through a Community
procedure before being placed on the market, used or processed within the
Community. Since pet food is not part of the human food chain and has no
environmental impact on arable land, specific provisions for additives in pet
food are appropriate.
It is a principle of the Community law that food and feed imported for
placing on the market within the Community must comply with the relevant
requirements of Community legislation or with conditions recognized by the
Community to be at least equivalent thereto. It is therefore necessary to
subject imports of additives for use in animal nutrition from third countries
to requirements equivalent to those applying to additives produced in the
Community.
Detailed labeling of the product is required since it enables the end-
user to make a choice with full knowledge of facts, creates fewer obstacles
to trade and facilitates fairness of transactions.
The Regulation consists of 26 Articles and four Annexes which are
summarized in the following. The purpose of this Regulation is to estab-
lish a Community procedure for authorizing the placing on the market and
use of feed additives and to lay down rules for the supervision and labeling
of feed additives and premixtures. The Regulation shall not apply to pro-
cessing aids and veterinary medicinal products. It gives definitions for feed
additives, feed materials, compound, complementary and complete feed-
ingstuffs, pre-mixtures, daily ration, processing aids, micro-organism,
antimicrobials, antibiotic, coccidiostats and histomonostats, maximum
residue limit and first placing on the market.
Placing on the market, process or use of feed additives is not allowed
unless the additive is covered by an authorization in accordance with this
Regulation. For experiments for scientific purposes Member States may
authorize the use of substances which are not authorized at Community level,
with the exception of antibiotics, under certain conditions. The mixing of
authorized additives shall not be subject to specific authorization. Any per-
son seeking an authorization for a feed additive or for a new use of a feed
additive shall submit an application. The applicant shall be established in the
Community. No feed additive shall be authorized if it has an adverse effect
437
on animal health, human health or the environment, it is presented in a

manner which may mislead the user or it harms the consumer by impair-
ing the distinctive feature of animal products or misleads the consumer with
regard to the distinctive features of animal products. The feed additive shall
favorably affect the characteristics of feed, of animal products, the color of
ornamental fish and birds, the environmental consequences of animal pro-
duction and animal production, performance or welfare, satisfy the nutri-
tional needs of animals or have a coccidiostatic or histomonostatic effect.
Antibiotics other than coccidiostats or histomonostats shall not be author-
ized as feed additives !
A feed additive shall be allocated to one or more of the following cat-
egories: Technological additives, sensory additives, nutritional additives,
zootechnical additives, coccidiostats and histomonostats.
In ORDER TO AUTHORIZE A FEED ADDITIVE the applicant shall send the fol-
lowing particulars and documents to the Authority: Name and address;
identification of the feed additive, a proposal for its classification by catego-
ry, its specifications, including, where applicable, purity criteria; a descrip-
tion of the method of production, the intended uses, the method of analysis
in feed for the determination of the level of residues, or its metabolites, in
food; a copy of studies and any other material which demonstrate that the
feed additive satisfies the criteria mentioned above; proposed conditions for
placing the feed additive on the market including labeling requirements; a
written statement that three samples have been sent to the Community ref-
erence laboratory; a proposal for post-market monitoring of nutritional,
zootechnical, coccidiostatic and histomonostatic additives and of products
consisting of containing or produced from GMOs. The Authority shall pub-
lish detailed guidance to assist the applicant in the preparation and the
presentation of its application.
The Authority shall give an opinion within six months of receipt of a
valid application. In order to prepare its opinion, the Authority shall verify
that the submitted particulars and documents are in accordance with this
Regulation, and shall verify the report of the Community Reference
Laboratory. In case of favoring the authorization the opinion shall include
the categorization, specification, purity criteria and method of analysis, spe-
cific conditions or restrictions in relating to handling, post-market monitor-
ing requirements and use, including animal species and categories of ani-
mal species, specific additional requirements for labeling, a proposal for the
establishment of Maximum Residues Limits (MRLs) if necessary for the pro-
tection of the consumers. The Authority shall without delay forward its
opinion to the Commission, the Member States and the applicant and shall
make its opinion public deleting any information identified as confidential.
Authorization by the Community shall take place within three months of
receipt of the opinion of the Authority preparing a draft Regulation to grant
authorization or to deny authorization and shall inform the applicant of the
438
Regulation without delay. The authorization shall be valid throughout the

Community for 10 years and shall be renewable in accordance with this
Regulation. The authorized feed additive shall be entered in the Register.
The Regulation lays down the conditions and phasing out for exist-
ing products. With a view to a decision on the phasing out of the use of coc-
cidiostats and histomonostats as feed additives by 31 December 2012,
the Commission shall submit to the European Parliament and the Council
before 1 January 2008 a report on the use of these substances as feed addi-
tives and available alternatives, accompanied, where appropriate, by legisla-
tive proposals. Antibiotics, other than coccidiostats and histomonstats, may
be marketed and used as feed additives only until 31 December 2005; as
from 1 January 2006, those substances shall be deleted from the Register.
After an additive has been authorized any person using or placing on
the market that substance shall ensure that conditions or restrictions are
respected. Where monitoring is required, the holder of the authorization
shall ensure that monitoring is carried out and shall submit reports to the
Commission. On its own initiative or following a request from a Member
State or from the Commission, the Authority shall issue an opinion on
whether an authorization still meets the conditions set out by this
Regulation. The Commission shall examine the opinion of the Authority
without delay. A decision on the modification, suspension or revocation
of an authorization shall be taken in accordance with this Regulation. The
Commission shall without delay inform the applicant of the decision taken.
The Register shall be amended where appropriate.
Authorizations under this Regulation shall be renewable for 10-year
periods. An application for renewal shall be sent to the Commission at least
one year before the expiry date of the authorization containing a copy of
authorization, a report on the results of post-market monitoring, if such
monitoring requirements are included in the authorization, any other new
information with regard to the evaluation of the safety in use of the feed
additive and the risks of the feed additive to animals, humans or the envi-
ronment and where appropriate, a proposal for amending or supplementing
the conditions of the original authorization. In specific cases where urgent
authorization is needed to ensure the protection of animal welfare, the
Commission may provisionally authorize the use of an additive for a maxi-
mum period of five years.
The kind of labeling and packaging of feed additives and premix-
tures is laid down in this Regulation. No person shall place on the market
a feed additive or a premixture of additives unless its packaging or contain-
er is labeled in a conspicuous, clearly legible and indelible manner, in at
least the national language or languages of the Member State in which it is
marketed. The label shall include the specific name of the additive, the
name and address of the responsible person, the net weight or, in case of
liquid additives, the net volume or net weight, where appropriate, the
439
approval number or the registration number assigned to the establishment,

directions for use, and any safety recommendations, the identification num-
ber, the batch reference number and the date of manufacture. Additives and
premixtures shall be marketed only in closed packages or closed containers
which must be closed in such a way that the fastener is damaged on open-
ing and cannot be re-used.
The Commission shall establish and keep up to date a Community
Register of Feed Additives. The Register shall be made available to the
public. The applicant may indicate which information submitted he wishes
to be treated as confidential. Verifiable reasons must be given in such
cases. The Commission shall determine which information should be kept
confidential and shall inform the applicant of its decision. Administrative
review: any decision taken under, or failure to exercise, the powers vested
in the Authority by this Regulation may be reviewed by the Commission on
its own initiative or in response to a request from a Member State or from
any person directly and individually concerned. Data protection: the sci-
entific data and other information in the application dossier may not be
used for the benefit of another applicant for a period of 10 years from the
date of authorization, unless the other applicant has agreed with the previ-
ous applicant that such data and information may be used. The applicant
and the previous applicant shall take all necessary steps to reach agreement
on sharing the use of information, in order not to repeat toxicological tests
on vertebrates. On the expiry of the 10-year period, the findings of all or part
of the evaluation contained in the application dossier may be used by the
Authority for the benefit of another applicant. The duties and tasks of the
Community Reference Laboratory are laid down in the Annex II of this
Regulation. Member States shall lay down the rules on penalties applicable
to infringements of this Regulation and shall take all measures necessary to
ensure that they are implemented. The penalties provided for must be effec-
tive, proportionate and dissuasive. Member States shall notify those rules
and measures to the Commission.
The Regulation contains four Annexes as follows:

Annex I: Additive Groups. In the different category the following function-
al groups are included: Technological additives: preservatives, antioxidants,
emulsifiers, stabilizers, thickeners, gelling agents, binders, substances for
control of radionucleide contamination, anticaking agents, acidity regula-
tors, silage additives, denaturants. Sensory additives: colorants, flavoring
compounds. Nutritional additives: vitamins, pro-vitamins and chemically
well-defined substances having similar effects, compounds of trace ele-
ments, amino acids, their salts and analogues, urea and its derivatives.
Zootechnical additives: digestible enhancers, gut flora stabilizers, sub-
stances which favorably affect the environment and other zootechnical addi-
tives.
440
Annex II: Duties and tasks of the Community Reference

Laboratory . The Community Reference Laboratory is the Joint Research
Centre of the Commission (JRC). The JRC may be assisted by a consortium
of national reference laboratories.The JRC shall be notably responsible for
the reception, preparation, storage and maintenance of the reference sam-
ples;
the testing and evaluation or validation of the method for detection; evalu-
ating the data provided by the applicant for authorization to place the feed
additive on the market, for the purpose of testing and evaluation or valida-
tion of the method for detection; submitting full evaluation reports to the
Authority.
The Community reference laboratory shall play a role in dispute set-
tlements between Member States concerning the results of the tasks out-
lined in this Annex.
Annex III: Specific labeling requirements for certain feed addi-
tives and for premixtures Zootechnical additives, coccidiostats and his-
tomonostats: the expiry date of the guarantee or the storage life from the
date of manufacture, the direction for use,and the concentration. Enzymes,
in addition to the above mentioned indications: the specific name of the
active component(s) in accordance with their enzyme activities, in conform-
ity with the authorization given, the International Union of Biochemistry
identification number, and instead of concentration: units of activity per
gram or per milliliter. Micro-organisms: the expiry date of the guarantee or
the storage life from the date of manufacture, the direction for use, the
strain identification number, and the number of colony-forming units per
gram. Nutritional additives: the active substance level, and the expiry date
of the guarantee or the storage life from the date of manufacture.
Technological and sensory additives with the exception of flavoring com-
pounds: the active substance level. Flavoring compounds: the incorporation
rate in premixtures
Annex IV: General conditions of use. The quantity of additives that
also exists in the natural state in certain feed materials shall be calculated
so that the total of the elements added and the elements present naturally
does not exceed the maximum level provided for in the authorization
Regulation. Mixing of additives shall be permitted only in premixtures and
feedingstufs where there is physico-chemical and biological compatibility
between the components of the mixture in relation to the effects desired..
Supplementary feedingstuffs, diluted as specified, may not contain levels of
the additives which exceed those fixed for complete feedingstuffs. In the
case of premixtures containing silage additives the words “of silage addi-
tives” must clearly be added on the label after “Premixture”.
In comparison with the US feed law the use of hormones for improving
the performance of animals was never allowed in the European Community
and also the use of antibiotics as growth promoters will be forbidden from 1
441
January 2006.
18.7. COMMISSION DIRECTIVE 2001/79/EC OF 17 SEPTEMBER 2001 AMENDING

COUNCIL DIRECTIVE 87/153/EEC FIXING GUIDELINES FOR THE ASSESSMENT OF
ADDITIVES IN ANIMAL NUTRITION. (Official Journal of the European
Communities (O J) No L 267, 6.10.2001, p.1)
This Directive is short consisting of four Articles only, but includes a
comprehensive Annex. Part I of the Annex deals with additives other
than micro-organisms and enzymes. Part II lays down the assessment of
micro-organisms and enzymes. Part II is adopted by SCAN, but not yet
by the Commission. Nevertheless it is used by FEEDAP for the assess-
ment of these categories of feed additives. The Directive is the guidance
for applicants how to prepare the dossier for the authorization of a feed
additive and also a general instruction for the evaluating committees or
442
panels.
induction of cross-resistance to relevant antibiotics, the selection of
resistant bacterial strains under field conditions and shedding or excre-
tion of relevant micro-organism; metabolism and residue studies and
pharmacokinetics, study of residues: identification of those residues
which represent more than 10% of the total residues, kinetic study of the
residues in the tissues, depletion studies, prescription of a withdrawal
period. Studies on laboratory animals should comprise studies on acute
toxicity, genotoxicity including mutagenicity, subchronic oral toxicity,
chronic oral toxicity, reproduction toxicity including teratogenicity, stud-
ies on metabolism and disposition, determination of the bioavailability of
residues, the determination of a no observed effect level (NOEL).
Part I of the Annex is subdivided into eight Sections as follows.

Safety evaluation for the human consumer: an acceptable daily intake (ADI)
as well as the maximum residue limits (MRLs) should, where appropriate be
proposed. The withdrawal period (period after cessation of the administra-
tion of the additive which is necessary to enable the residue levels to fall
below the MRLs with a 95% confidence limit) will be set. An assessment of
risk to workers should be included. Workers can be exposed mainly by
inhalation or topical exposure while manufacturing or handling or using the
additive. Effects on the respiratory system, on eyes and skin should be
studied. The exposure should be assessed. If the risk is unacceptable, then
precautionary measures should be taken to control or eliminate exposure.
Use of personal protection devices should only be regarded as a measure of
last resort.
Consideration of the environmental impact of feed additives is impor-
tant, the risk assessment is therefore necessary. A stepwise approach along
a decision tree should be followed, where in the first phase, additives which
do not need further testing can be clearly identified. This is the case when
the chemical nature and the biological effect of the additive and its use indi-
cate that impact will be negligible (physiological/natural substances, addi-
tives intended for companion animals) or the worst case predicted environ-
443
mental concentration (PEC) is too low to be of concern. In all other cases an

environmental risk assessment should be carried out determining the expo-
sure and the predicted no effect concentration (PNEC).
Part II of the Annex
The general structure of Part II is similar to that of Part I. The biological ori-
gin of each declared enzyme activity or micro-organism should be given. For
enzymes produced using recombinant DNA technology, the recipient organ-
ism (the production strain) and the donor organism should be described in
detail and the nature and purpose of the genetic modification should be
specified. The conditions for the placing on the market of the additive,
including specific conditions of use and handling and a proposal for label-
ing and packaging which should comprise at least the requirements laid
down in Annex IV of Directive 2001/18/EC. Strains of micro-organism
belonging to a taxonomic group which includes members known to be capa-
ble of the production of toxins or other virulence factors should be sub-
ject to appropriate tests to demonstrate at a molecular and cellular level the
absence of any concern.
Enzyme preparations should be free of antibiotic activities relevant
to the use of antibiotics in humans or animals. Micro-organisms intended
as active agents should not be capable of the production of antimicrobial
substances relevant to the use of antibiotics in humans or animals. Strains
of bacteria intended for use as an additive should not contribute further to
the reservoir of antibiotic resistance genes already present in the gut flora
of animals and the environment. Consequently, all strains of bacteria should
be tested for resistance to at least one representative of each of the antibiot-
ic families in use in human and veterinary medicine. Where resistance is
detected, the genetic basis of resistance and the likelihood of transfer of
resistance to other gut-inhabiting organisms should be established.
The introduction of a system similar in concept and purpose to the
GRAS (Generally Recognized As Safe) definition used in the USA is under dis-
cussion in the EU. This system is called Qualified Presumption of Safety
(QPS) (FIGURE XVIII-1). It should not compromise on safety bur should ide-
ally improve, extend, clarify and make more consistent the approval proce-
dure for micro-organisms and, where possible, allow a more generic
approach to be taken in place of a full case-by-case assessment.
444
445
18.8. DIRECTIVE 2003/32/EC OF THE EUROPEAN PARLIAMENT AND OF THE

COUNCIL OF 7 MAY 2002 ON UNDESIRABLE SUBSTANCES IN ANIMAL FEED. (Official
Journal of the European Communities (O J) No L 140, 30.5.2002, p.10)
The Commission intends to review this Directive. The exercise should be
based on updated scientific risk assessments and should take into account
the prohibition of any dilution of contaminated non-complying material
intended for animal nutrition. This Directive shall apply without prejudiced
to the provisions in Regulation (EC) No 1831/2003 of the European
Parliament and the Council of 22 September 2003 on additives for use in
animal nutrition, Council Directive 2002/2/EC on the marketing of com-
pound feedingstuffs, Commission Directive 2000/57/EC and 2202/23/EC
relating to fixing of maximum levels for pesticide residues in and on fruits
and vegetables and on cereals, respectively, Commission Directive
2002/23/EC relating to fixing of maximum levels for pesticide residues in
and on foodstuffs of animal origin and Council Directive 2002/23/EC on
the fixing of maximum levels for pesticide residues in and on certain prod-
ucts of plant origin, including fruit and vegetables, where these residues are
not listed in Annex I to this Directive, Community legislation concerning
veterinary matters relating to public health and animal health, Council
Directive 1999/20/EC concerning certain products used in animal nutri-
tion and the Council Directive 1999/29/EC on feedingstuffs intended for
particular nutritional purposes.
Undesirable substances listed in Annex I may be tolerated in feed only
subject to the conditions laid down therein. In order to reduce or eliminate
sources of undesirable substances of products intended for feed Member
Sates shall carry out investigations to identify the sources of undesirable
substances and shall transmit to the Commission and the other Member
States all relevant information and findings of the source and the measures
taken to reduce the level or elimination of the undesirable substances.
Products intended for feed containing levels of an undesirable substance
that exceed the maximum level fixed in Annex I may not be mixed for dilu-
tion purposes with the same, or other, products intended for feed.
Complementary feedingstuffs may not, taking into account the proportion
prescribed for their use in a daily ration, contain levels of the undesirable
substances listed in Annex I that exceed those fixed for complete feed-
ingstuffs.
With good reasons a Member State may provisionally reduce the exist-
ing maximum level, fix a maximum level or prohibit the presence of that unde-
sirable substance. It shall inform immediately the other Member States and
the Commission thereof, stating the grounds for its decision. An immediate
decision shall be taken as to whether Annex I and II should be amended.
The Commission may define acceptability criteria for detoxification process-
es as a complement to the criteria provided for products intended for feed
446
which have undergone such processes. Member States shall ensure that
measures are taken to guarantee the correct application of any acceptable
process es and the conformity of the detoxified product with the provision
of Annex 1. The Member States shall apply at least the provision of this
Directive to products intended for feed produced in the Community to be
exported to third countries.
Annex I consists of a list of undesirable substances and their maxi-
mum content in products intended for feed. The list is subdivided in three
groups: a) Ions or elements: arsenic, lead, fluoride, mercury, nitrites and
cadmium. b) Products: aflatoxin B1, hydrocyanic acid, free gossypol, theo-
bromine, volatile mustard oil, vinyl thiooxazolidone, rye ergot (Claviceps pur-
purea), weed seeds and unground and uncrushed fruits containing alka-
loids, glucosides or other toxic substances separately or in combination,
castor oil plant (Ricinus communis), crotalaria, aldrin, dieldrin, camphechlor
(toxaphene), chlordane, DDT, endosulfan, endrin, heptachlor, hexa-
chlorobenzene (HCB), hexachlorocyclohexane (HCH) and dioxins. c)
Botanical impurities: apricots (Prunus armaniaca), bitter almond (Prunus

dulcis var. amara seu Prunus amygdalus var. amara), unhusked beech mast
(Fagus silvatica), vamelina (Camelina sativa), mourah, bassia, madhuca,
purghera (Jatropha curcas), croton (Croton tiglium), Indian mustard
(Brassica juncea spp integrifolia), Sareptian mustard (Brassica juncea spp.
Juncea), Chinese mustard (Brassica juncea spp. Juncea var. lutea), black
mustard (Brassica nigra), Ethiopian mustard (Brassica carinata)
In the review process apart from the substances already considered in
the legislation other mycotoxins could be identified in feedingstuffs which
may pose a sufficient risk for animals or humans to require regulation. The
following were considered for a possible full risk assessment before listing
as undesirable substances: ochratoxin A, zearalenon, deoxynivalenol,
T–2 toxin, fumonisins, moniliformin, mycophenolic acid and cyclopia-
zonic acid.
Because of recent problems with dioxin in some European countries
the Annex I to Directive 2002/32/EC was amended as follows: dioxin is
defined as the sum of polychlorinated dibenzo-para-dioxins (PCDDs) and
polychlorinated dibenzofurans (PCDFs) expressed in WHO toxic equivalents,
using the WHO TEFs (toxic equicalency factors, 1977). The maximum con-
447
tents in ng/kg feedingstuff with a moisture content of 12% are for:

18.9. REGULATION (EC) NO. 882/2004 OF THE EUROPEAN PARLIAMENT AND THE
COUNCIL OF 29 APRIL 2004 ON OFFICIAL CONTROLS PERFORMED TO ENSURE THE
VERIFICATION OF COMPLIANCE WITH FEED AND FOOD LAW, ANIMAL HEALTH AND ANI-
MAL WELFARE RULES (Official Journal of the European Communities (O J)
No L 165, 30.4.2004, p.1)
This voluminous regulation (141 pages) laid down the methods of sam-
pling and analysis for the official control of feedingstuffs. It defines the
requirements for reference laboratories of the Member States as well as of
the Commission. Member States have to report the results of their investi-
gations regularly. The Regulation authorizes the Commission to inspect the
control measures of the Member States.
18.10. OTHERS. Besides these most important Regulations and Directives a

large number of other Directives are in force in the European Community
for example
on the use of genetically modified organisms as or in feedingstuffs
(Regulation (EC) No 1829/2003 of the European Parliament and of the
Council of 22 September 2003 on genetically modified food and feed, Official
Journal of the European Communities (OJ) No L 268 18.10.2003, p. 24;
Commission Regulation (EC) No 641/2004 of 6 April 2004 on detailed rules
for the implementation of the Regulation mentioned above as regards the
application for the authorization of new genetically modified food and feed,
the notification of existing products and adventitious or technically unavoid-
able presence of genetically modified material which has benefited from a
favorable risk evaluation, Official Journal of the European Communities (O
J) No L 102, 30.4.2004, p.62),
Many feed regulations are related or based on regulations for food, for
instance the Regulation (EC) No 178/2002 of the European Parliament and
448
the Council of 28 January 2002 laying down the general principles and
requirements of food law, establishing the European Food Safety Authority
and laying down procedures in matters of food safety (Official Journal of the
European Communities (OJ) No L 31, 11.2.2002, p. 1) and the Commission
Regulation (EC) No 466/2001 of 8 March 2001 setting maximum levels for
certain contaminants in foodstuffs (Official Journal of the European
Communities (OJ) No L 77, 16.3.2001, p. 1).
Although the general intentions of feedstuff regulations are interna-
tionally similar, namely the protection of animal health, the avoidance of
risks of the consumers due to residues of undesirable substances in food of
animal origin, and the reduction of the environmental pollution by livestock,
SOME DIFFERENCES EXIST BETWEEN THE EU REGULATIONS AND THOSE
OF THIRD COUNTRIES FOR INSTANCE USA (Federal Food, Drug, and
Cosmetic Act, U.S. Code: Title 21 – Food and Drugs, several Parts; The
Arkansas Feed Law, Act 726 of 1997). As already mentioned HORMONES
WERE EVER FORBIDDEN FOR IMPROVEMENT OF PRODUCTION AND
ANTIBIOTICS WILL NOT BE ALLOWED AS GROWTH PROMOTERS FROM 1
JANUARY 2006. The EU regulations have considerable consequences for the
food production in countries which intended to export food to the Member
States of the European Community because FOOD IMPORTED INTO THE
COMMUNITY FOR PLACING ON THE MARKET WITHIN THE COMMUNITY
SHALL COMPLY WITH THE RELEVANT REQUIREMENTS OF FOOD LAW or
conditions recognized by the Community to be at least equivalent thereto or,
where a specific agreement exists between the Community and the export-
ing country, with requirements contained in the Regulation (EC) No
178/2002.
18.11. LIABILITY
Insufficient quality of feed is often believed to be responsible for impaired
performance of livestock (high morbidity and mortality, insufficient body
weight gain or milk yield, bad performance of layers) or reduced quality of
animal products. In accordance with the White Paper on Food Safety every
single link of the food chain has its own responsibility for the safety and qual-
ity of food including all aspects of animal husbandry. A prerequisite for this
is the traceability, in case of animal nutrition the traceability of feed and
every single component of compound feed. Besides the regulation on the
quality of feedingstuffs by specific regulations and directives like the
Directive on undesirable substances in animal feed feed materials may be
put into circulation in the Community only if they are sound, genuine and
of merchantable quality according to Council Directive 96/25/EC of 29
April 1996 on the circulation and use of feed materials.
In many cases the farmers have to sue for damages. The assessment
of damage as well as the assessment of the causal connection between feed
and damage by expert is necessary. The causal connection can only be
449
assessed by analyzing feed samples correctly taken and analyzed by officials

or experts. Under certain conditions it is wise to take samples at delivery of
feed and to keep the samples until a few weeks after using up the quantity
delivered if this procedure is accepted by the supplier. Most of the feed com-
panies take out indemnity insurance.
FOR FURTHER READING
Regulations and Directives mentioned in the text

White Paper on Food Safety, Commission of the European Communities,
Brussels, 12 January 2000
Animal Food (Feed) Product regulation (US Code of Federal Regulations;
Federal Food, Drug, and Cosmetic Act)
National Feed Laws of the Member States of the European Community
450
Chapter XIX
EVOLUTION OF DIGESTION
19.1. Development of the mammalian digestive system
19.2. Comparison of the digestibility of major nutrients
19.3. Utilization of fibre
19.4. Phenomenon of trophallaxis

Chapter XIX: EVOLUTION OF DIGESTION
n this chapter the evolution of digestion in mammals summed up in
I order to take a right view of the characteristics of digestion in the domes-

tic animals. In course of the phylogenesis, carnivores, omnivores and
herbivores were developed from the small (20-30 g) insectivore-carnivore
primitive representative of the Therapsidae reptiles of the Triassic age (FIG-
URE XIX-1).
FIGURE XIX-1: Digestive system of today’s mole (© KATONA-ZSOMBOR, É. 2007))
19.1. Development of the mammalian digestive system.

About 65 million years ago dinosaurs died out, giving an impulse to mam-
mals which were forced to nocturnal lifestyle up to that time. If the envi-
ronment was rich in prey animals that promoted the development of the
carnivorous-predatory way of life. In carnivores the stomach became big,
the intestinal tract shortened, the caecum is rudimentary (dog, cat) or com-
pletely vanished (mink)
.The “mixed” supply (roots rich in starch, plants, varmints full of pro-
teins, bird eggs, aquatic animals, small rodents etc.) resulted in the devel-
opment of the omnivorous digestion. The increased ingestion of the more
452
fibrous plant material increased more and more the measure and impor-
tance of digestion in the hind gut. At first two autotypes were separated i.e.
the colon and the caecum fermenter groups
In case of “colon fermenters” (e.g. equidae) the main site of the microbial
digestion is the colon. Of course, there is an important bacterial digestion
in the caecum, too). The decomposition of the feed particles is very effective,
and – according to certain authors some of bacterial protein produced can
also be utilized. The species belonging to this group can survive on feeds
with less energy- and high fibre content feedstuffs without developing any
type of coprophagy. FIGURE XIX-2: Vizualizes the development of different
digestive types.
FIGURE XIX-2: Development of carni-, omni- and herbivores (© FEKETE S.GY.)
The “caecum fermenters” (e.g. rabbit, FIGURE XIX-3e) developed a

selective mechanism, which “leaves” and re-transport only the fine, floating
feed particles in the caecum, and excretes the rough, fibrous parts in form
of hard faeces. Animals belonging to this group have smaller body size, and
they can utilize the low energy, fibrous bulk feed without overburdening the
colon. The coprophagy of the soft faeces (caecotrophy) has developed in
order to utilize the synthesized microbial proteins FEKETE and BOKORI,
1984).The anatomy of the hind gut reflects the function (FIGURE XIX-3).
453
FIGURE XIX-3: Comparative anatomy of horse (a), cattle (b), pig (c), dog (d) and rabbit (e)
hind-gut.
Ruminants separated from the first group about 40 million years

ago, since them for many generations only bulky, fibrous mass feed was
available, which was relatively poor in protein. The pre-stomach digestion,
by the help of the utilization of cellulose and non-protein nitrogen (NPN)
compounds - made the survival, and even the spreading of these animals
possible FIGURE XIX-4).
FIGUREXIX-4: Comparison of the stomac of Non-Human Primate (a), horse (b), pig (c),
dog (d) and cattle (e)
454
However, it is very interesting that young, suckling foals ingest the faeces
of their mother even under normal circumstances and coprophagy can be
triggered by the drastic reduction of the protein level in the ration of adult
horses, too.
19.2. Comparison of the digestibility of major nutrients
From the comparison of digestion of the herbivore mammals (Table XIX-1.)
it emerges that horses digest crude fibre less than ruminants, and do it
much better than other monogastric domestic animals.
Table XIX-1. Comparison of digestibility of major nutrients at horses, ruminants and rab-
bits
Digestion of ruminants and monogastric animals consuming fibrous

feedstuff can be compared properly with the aid of the RELATIVE EFFICIENCY.
This index number expresses the transformation percentage of the ingested
digestible energy into net energy. Ruminants are less effective in case of
feeding a low fibre content concentrated feeds, since the fermentation loss-
es (e.g. by eating sugars) in the rumen are significant. Mixing hay (fibre
source) to the concentrated feed improves the feed conversion. (The
described model presumes that the main energy source of animals is car-
bohydrate, not fat!) In case of certain species (elephant, panda bear) this
model is not valid, because they ingest a lot of fibre and it stays in the intes-
tinal canal only for a short period of time.
19.3. Utilization of fibre
The key issue is the utilization of the fibre components. The rumen and gut
flora can provide the cellulase which is not produced by the host organism.
Certain species (termites, e.g. Mastotermes darwiniensis, Coptotermes axi-
naciformis) are able to decompose even the lignin by the help of the aerobe
fungi living in their gut.
In the course of the fibre decomposition volatile fatty acids are pro-
duced and are utilized later by being absorbed. Its measure, quantity and
significance are highly different in cases of omnivores, monogastric herbi-
vores and ruminants. The energy is being released in ruminants (0.2-0.3
MJ/W0.75) in the pre-stomach, in omnivores (swine: 0.01, humans, rats:
0.04 MJ/W0.75) in the caecum and colon, and in rabbits (0.03-0.05
455
MJ/W0.75) exclusively in the caecum. There is no available data about hors-

es, but the figure may be estimated at 0.12-0.18 MJ/W0.75) in the caecum
and colon.
So for example in ruminants the energy released in the pre-stomach
can cover 60-70% of the total requirement, while in rabbits the caecal ener-
gy is enough only for the 15% of the maintenance requirement. The wide
range of the above values largely depends on the feed composition, which is
well demonstrated in swine that can get more fermentation energy from
feeds containing a lot of sugar, pectin and hemicellulose. (see: BFS=bak-
teriell fermentierbares Substanz in the German energy evaluation system).
It is all well illustrated in FIGURE XIX-8 which compares the quantity of
volatile fatty acids being produced in the stomach and the gut by fibre
decomposition, in cases of ruminant cattle, herbivore horses and the omni-
vore swine.
FIGURE XIX-5:
The production
of volatile fatty
acids from fibre
(after BERGMAN,
1990 modified)
19.4. Phenomenon of trophallaxis

The other determining factor in the survival of the species is the way of
selective eaters (coypu, some antelopes, and giraffes), also eating dry grass
(horse), or ingesting all kinds of fibre sources (camel). In the course of evo-
lution, different types of reutilizing the ingested organic matter (recycling,
trophallaxis) have developed besides coprophagy. It holds high importance
to distinguish the physiological trophallaxis and the pathological allotrio-
phagy. Definition of the TROPHALLAXIS is the following: unilateral transmis-
sion or mutual change of bodily fluids, materials having nutritional value
among species companions (Table XIX-2.).
456
Table XIX-2. The main types of trophallaxis
FOR FURTHER READING
Bergman, E.N. (1990). Energy contributions of volatile fatty acids from the gastrointesti
nal tract in various species. Physiol. Rev. 70, 567-590.
Björnhag, G. (1989): Sufficient fermentation and rapid passage of digesta. A problem of
adaptation in the hindgut. Acta Vet. Scand. Suppl. 86, 204-211.
Holecheck, J.L., Pieper, RD. and Herbel, C.H. (2004): Range management: principles and
practices. (5th ed.) Pearson-Prentice Hall. Upper Saddle River. New Jersey
Kirchgessner, M. (1991). Digestive physiology of the hindgut. Verlag Paul Parey. Hamburg-
Berlin
Stevens, C.E. (1988): Comparative physiology of the vertebrate digestive system.
Cambridge Univ. Press. Cambridge-New York-Sydney
457
Chapter XX
PIG NUTRITION AND FEEDING
20.1. Characteristics of pig digestive physiology

20.1.1. Consequences of domestication
20.1.2. Voluntary feed intake,mastication and saliva production
20.1.3.Digestion in stomach and intestine
20.1.4.Digestion in large intestine
20.2. Feeding of suckling piglets; weaning
20.2.1. Basics of digestive physiology
20.2.2. Practical feeding around weaning
20.3. Theoretical basics of pig growth („fattening”)
20.3.1. Energy requirements of growing-finishing pig
20.3.2. Importance of energy: protein ratio
20.3.3. Protein and amino acid needs of growing-finishing pig
20.3.4. Fat requirements of growing-finishing pigs
20.3.5. Ballast (indigestible organic matter) need of growing-
finishing pig
20.4. Applied nutrition of fattening (growing-finishing) pig
20.4.1.Definition of „fattening”; economic considerations
20.4.2.Basic notions of fattening
20.4.3. Nutrient requirements
20.4.4.Feed intake of growing-finishing pigs
20.4.5. Applied feeding systems
20.4.6.Feedstuffs of growing-finishing pigs
20.4.7.Feed additives
20.4.9. Feeding growing-finishing pig and environmental
protection
20.4.10. Feeding growing-finishing pig for quality
20.4.11.Consequences of nutritional failures
20.5. Feeding and nutrition of breeding sow and boar
20.5.1. Feeding of growing–replacement gilts and boars
20.5.2. Nutrient requirement of gestation and lactation
20.5.3. Practical feeding of breeding sows
20.5.4. Diet formulation for sows
20.5.5. Feeding of breeding boars
20.5.6. Possible non-infectious causes of sows’ abortion
20.6. Feed related diseases, herd health aspects
CHAPTER XX: PIG NUTRITION AND FEEDING
n this chapter after an overview of digestive physiology of swine, the feed-
I ing and nutrition of suckling piglet, weaning, rearing of future breeding

gilts and boars, feeding and nutrition of pregnant and lactating sow, the-
oretical basis and practical implementation of fattening pig nutrition, and
last but not least, the most important pig deficiency syndromes and meta-
bolic troubles.
20.1. Characteristics of pig digestive physiology

20.1.1. Consequences of domestication

In most of the animals living in wilderness a capacity developed to be able
to accumulate fat during good nutrient supply. In periods of famine fat
depots are mobilised for survival. This is especially true in case of migrato-
ry birds and winter-sleeping animals, but it is true for wild pig, having lim-
ited feed access in wintertime. This feature did not disappear with domesti-
cation, what is more, through selection for fat production, it became most
intense. Today’s domestic pig is prone to overeating (hyperphagia) and get-
ting fat, which is difficult to avoid using ad libitum feeding. Domesticated pig
is heavier, moves little, reaches puberty earlier than wild pig and instead of
seasonal, has a regular oestrus cycle. Meat producing capacity of modern
swine hybrids significantly improved but the tendency to fat accumulation
did not disappear totally. Pigs are omnivore animal with an excellent capac-
ity for starch utilisation. Thus today’s modern pig requires concentrated
feed of high protein quality.
20.1.2. Voluntary feed intake, mastication and saliva production

Voluntary feed intake of pigs depends to a great extent on the smell of feed.
Olfactory organ of pig is proverbial – let us think of truffle searching sows.
Pigs are highly responsive to changes of their feeds. This is why pigs neglect
rancid feed and overheated green meal and by means of feed flavouring its
appetite can be stimulated. Sweet is the preferred taste. The feed, treated by
organic acid causes transitional decrease in feed intake. DON or T–2 toxin
content will produce feed refusal. Change of composition, even change of
the origin of a constituent may decrease appetite. Dry matter intake may be
460
stimulated by mixing of molasses, milk powder, whey powder and fat.

There are significant differences among corals concerning preference:
barley and wheat are preferred to corn or rye. Bitter, mouldy, musty feed is
refused. The same is true for stale feeds and those high of microbial count.
Physical characteristics of feed are also important: medium particle size
(meal but not flour, particle size above 0.2 mm, low hygroscopic character
and low stickiness are prerequisites of appropriate feed intake.
Pigs chew intensively; incisors serve for grabbing and selecting feed,
molars are used for crushing and only partly for grinding. Animals are not
capable of thoroughly chewing and therefore utilising high fibre feedstuffs
(e. g. corn cob mix, silage and aged grass). Too fine-ground feeds (particle
size smaller than 1 mm) induce less mastication and may lead to the devel-
opment of gastric ulcer on and respiratory diseases. Power production may
be decreased by pelleting or by adding 2-4% fat.
By means of its characteristic olfactory organ, the sensible snout and
mobile tongue pigs are able to take in feed selectively even from uneven sur-
face. For example, mangalitza were able to ingest acorn (“mast”), worms,
insect larvae, snails and slugs while kept in forest or on pasture. With meal,
pellet or hog-wash eating this ability have lost its significance, but in case
of silage, wet sugar beet pulp and pasture regained its significance.
Sensible, mobile tongue with taste buds serves to taste, mix, pass and swal-
low ground feed. The ingested feed in the mouth (mouthful or bite) is lubri-
cated by the product of parotis and sublingual salivary gland. Saliva com-
prises amylase and glycoproteids (mostly N-glycosyl-neuramic acid). For
sufficient grinding and lubricating of dry feed more time is required than in
case of hog-wash eating.
Rummaging or searching is an inherent feature of pigs. By placing
solid feed on a flat plat, transition to solid feed intake of suckling piglets can
be accelerated. Having only small pasture surface, it can be used only as a
walking field; otherwise pigs ruin the grass. Pigs’ need for mastication is
limited, although in case of fibre deficiency breeding sows may eat straw
voluntarily. Wild pigs spend most of their time rummaging and masticating,
which may be replaced by bedding or special feeders in order to assure ani-
mal welfare.
Pig salivary glands are relatively small and saliva produced is suffi-
cient only for a good, concentrated feed. If too much dry feed is given (e. g.
lactating sow) the quantity of saliva is not enough limiting voluntary feed
intake. Crumbling of feed and/or adding of 2-5% of fat may improve
appetite.
20.1.3. Digestion in stomach and intestine

Capacity of pig STOMACH, in comparison of other monogastric animals, medi-
um is, while rabbits have large and horses small stomac.
The gastric parietal cells actively secrete hydrogen and chloride ions.
461
Hydration of CO2 in the parietal cell is responsible for the continuous pro-
duction of H and HCO3 ions. This reaction is catalysed by the Zn-contain-
ing carbonic anhydrase enzyme present in high concentrations in these
cells. The H ions are transported into the lumen by an active pump mech-
anism. Chloride is also secreted into the lumen; metabolic alkalosis that
occurs during active acid secretion following a meal is known as the alka-
line tide.
Under natural circumstances pepsin-hydrochloric acid complex
quickly infiltrate gastric content and the low pH suitable for protein diges-
tion is achieved. If pig receives in a moment high amount of feed of high
alkaline value in a short time or it is under stress conditions, then the pro-
duction of hydrochloric acid decreases and the physiological fall in pH does
not occur. As a consequence, bactericide effect is diminished, availability of
minerals decreases and passage of digesta into small intestine slows down.
In addition to the above mentioned factors, the lenth of time the feed spends
in the stomach is determined by the solubility and fibre content of its com-
ponents. Feeds rich in fibre may spend 7-9 hours in the stomach, while liq-
uid feed, will leave the stomach in a few minutes.
Although fibres stimulate production of gastric and intestinal juice,
roughage cannot be considered as a primary swine feed. Staying for a long
time in the stomach, fibres are mixed with the ingested feed and gastric
overloading occurs. Too much fibre decreases voluntary dry matter intake
and decrease digestibility of nutrients. Under the influence of fibres intes-
tinal passage and ileocaecal flow accelerates, water content, pH and buffer
capacity of CHYME increase. Fibres decrease incidence of caecal dyspepsia
and water absorption increases. Potassium and sodium concentrations in
the faeces and the number of defecations increase. Incidence of constipa-
tion will be minimized. These events are due to the dietetic effect of fibre (for
details see in “Fattening pig” unit). Optimum crude fibre level for suckling
diet is 3%, that for starter-finisher pig and lactating sow is 4.5-5% and for
pregnant sows 8-10%.
In the gastro-intestinal tract of an adult pig all the necessary enzymes
of digestive processes are present. Thus in the stomach pepsinogen A and
D and their activator, hydrochloric acid are found. Intestinal mucosa is pro-
ducing saccharase, maltase, lactase, trehalase, enterokinase and dipepti-
dase. In the large amount of pancreatic juice α-amylase, trypsine, chy-
motrypsine, elastase, carboxy-peptidase, triacyl-glycerine-lipase, choles-
terase and phospholipase A2 are present. Bile contains hyo-deoxy-, hyo and
cheno-deoxy-cholic acids.
Digestion of intact starch (e. g. in raw potatoes) is incomplete. Pectins,
ß-glucans, hemicellulose and cellulose may build a tertiary structure alter-
ing protease activity (“cage effect”). Heat damaged and some intact proteins,
as well as long chained lipids of high melting points are difficult to digest-
ed. Some sugars and polymer carbohydrates are also indigestible in small
462
intestine. Indigestible feed constituents disproportionately decrease ileal

digestibility, thus it is enough to make up feed mixture of 70-75% of prae-
caecal digestibility (see Chapter 7.2.).
20.1.4. Digestion in large intestine

In case of fibre rich feeding the chyme arriving in the caecum has peculiar
composition, compared to low fibre diet feeding: 4-5 times higher fibre con-
centration, low dry matter (13-15%) content and higher pH value. There is
an enzymatic after-fermentation in caecum. Part of arriving organic matter
degrades and in form of volatile fatty acids and ammonia will be aborbed.
(Latter may cause liver damage.) In comparison, pig caecum is smaller than
that of equidae and larger than in humans. (FIGURE XX-1:Vizualizes the
physiological pH- circumstances in the swine digestive tract.)
FIGURE XX-1:Physiologycal pH- ranges in the different section of the pig’s digestive tract.
Five to twenty percent of metabolisable energy of a feed derives from

caecal absorbed volatile fatty acids. In some unique cases (e.g. by feeding
raw potatoes) this proportion may achieve 70-80%. Caecum adapts to gas-
tro-intestinal digestion and to the amount of ingested feed. Large amounts
of fibre will increase caecal content. Slowly degradable fibres slow fermen-
tation intensity and stabilise buffer capacity. The latter may upset microbial
balance in caecum, as well as large amount of readily available substrates
(e. g. starch), infections and stress situation.
Pathological caecal fermentation may lead to watery diarrhoea. Main
factors of prevention are the timing of regular digesta influx and optimised
buffer capacity. These can be assured by setting praecaecal digestibility and
satisfying minimum fibre requirements. Ad libitum feeding, by continuous-
ly “supplying” chyme for caecum is beneficial.
463
20.2. Feeding of suckling piglets; weaning

20.2.1. Basics of digestive physiology

Newborn pig. The average body weight of newborns falls between 1.2 and
1.4 kg and that of a 3-week-old suckling between 4 and 5 kg, and that of 5-
week-old between 8 and 9 kg (in the meantime weaning may happen: wean-
ing pig). Growing is till week 9-10 (20-30 kg) and finisher approximately
from month 3 (30-35 kg). Growing plus finisher may be mentioned together
as fattener. Newborn live weight is rather independent of the nutrient sup-
ply of the pregnant sow, but it negatively correlates with litter size. Newborn
of a modern genotype in a litter of 10 has an average weight of 1250 (0.5%
of dam’s weight). Each piglet over 10 will decrease average birth weight by
100 gram. Litter size of older sows and gilts is smaller.
Although birth weight has a long-lasting effect on weight gain, the
determining factor of average weight gain is the postnatal nutrient supply.
Although newborn piglets physiologically are mature, it did not reach the
state of chemical maturity concerning its body composition. Fat concentra-
tion is very low (1.6%), consequently heat losses are great by radiation.
Piglets have some glycogen for mobilisation as energy source but it is only
enough for 1-2 hours. Thus, it is essential for piglets to receive colostrums
as early as possible. Colostrum provides not only energy (lactose, fat), but
also immunoglobulins and factors of gastro-intestinal development. The
intestinal tract doubles its weight within a few days and undergoes differ-
entiation. This maturation is stimulated by colostral substances. After 46
hours theintestinal wall loses its capacity to take up proteins intact (“clo-
sure”). Differentiation of gut wall tissue is helped and controlled by colostral
peptides, insulin, epidermal growth factor (EGF), insulin like growth factor
I (IGF-I) and biogenic amines (spermin, spermidine).
Sow’s colostrum is rich in protein and vitamins and poorer in fat and
minerals than mature sow milk (Table XX-1).
Table XX-1: Chemical composition of sow colostrums and mature milk
* upper limit
464
During the last month of gestation and first weeks of lactation addi-
tion of fat to sow’s diet (3%) increases the amount of milk fat produced.
During the first 3 weeks of lactation sows produce 7-10 litres milk daily, i.e.
0.7-0.9 litres per piglet per day. In the piglet’s stomach owing to the action
of rennin the milk coagulates and is gradually passed to further gut seg-
ments. If artificial milk replacer contains plant proteins (e. g. soybean), lack
of clotting leaves the solution to arrive in the small intestine earlier. This
may prove dangerous, because of the failure of lactic acid fermentation and
bactericide effect in the stomach. Digestive enzyme activity in the intestine
is time dependent
Newborn piglet has a modest set of digestive enzyme: activity of lac-
tase is important, that of lipase medium and of amylase is very low.
Production of protein splitting enzymes (trypsine, chymotrypsine) is devel-
oping only slowly. Abrupt weaning may decrease concentration of digestive
enzymes for a while.
Depending on environmental temperature and relative humidity
suckling pigs have a different water requirements. Diarrhoea drastically
increases drinking water needs, up to the 5-10% of the body weight. (In case
of water shortage intake of dung water may occur.)
In body composition of young piglet proportion of fat (1.3%), macro-
and microelement (especially iron) are very low and that of water is very high
(82%). In ten days fat concentration sharply increases (ten times more),
whereas water decreases. Later on this tendency continues and the body of
a 35-day-old (8-9 kg LW) piglet consists of 63.3% water, 16% fat, 18% pro-
tein and 2.7% ash and achieves chemical maturity. Glycogen storage capac-
ity of the liver and muscles is limited. At 35 days dry matter accretion cor-
responds to 1.5 kg fat and to 1.2 kg protein, which are supplied from the
mother’s milk.
NUTRITIONAL (IRON DEFICIENCY) ANAEMIA is characteristic for the fast
growing suckling piglets. At birth, in the body of newborn piglets the total
iron content is 42 mg, which provides a suitable iron supply and may
achieve 180 mg till day 21. This means that during the first 3 weeks an iron
requirement of 100 mg appears. Iron depots of newborn piglet cannot be
filled up. (There are some publications about possible transfer of chelated
iron through the placenta, thinking of iron toxicosis, this cannot be pro-
posed for practice. Haemoglobin value of blood may be decreased by a phys-
iological haemodilatation to 5 mmol/l. Sow milk owing to evolutionary
development contains small amount of iron (colostrum: 2-3 mg/l), mature
milk: 1 mg/l), which cannot be influenced by feeding. Besides sow milk,
important solid feed and liquid intake may be expected only at day 20. No
more than 50-70% of the iron from sow milk can be utilised by piglets
because of the relatively high pH and lack of hydrochloric acid (physiologi-
cal achlorhydria) which do not help the production of more effective iron II
ions.
465
For peroral iron supplementation organic iron salts (e. g. iron-fumar-

ic) are applicable and preferable in paste form. The most effective is the par-
enteral treatment by iron-dextran complex within the first 3 days of life. The
two type of treatment can be combined. After iron injection iron toxicosis
and death of piglets may occur, owing to immunodepression. By providing
pregnant sows with high amount of vitamin E, this problem can be pre-
vented.
20.2.2. Practical feeding around weaning

Changes of digestive enzyme production of piglet as a function of age is
shown in Table XX-2. In summary it can be stated that with the advance-
ment of age production of enzymes for degrading milk components decreas-
es and that of solid feed digestion increases.
Table XX-2: Development of young piglet’s digestive enzyme set
** Though it is not an enzyme, for an overview, it is included in table.

Legend: - no detectable; + detectable but practically negligible; ++ medium level of produc-
tion; +++ activity of adult animal
Feeding of suckling piglets should be adjusted to system of rearing-

fattening. In practice there is a variety of systems, but basically they can be
classified into two types: late weaning after relatively long suckling (at least
6 weeks) and early weaning at the age of 4 weeks. The general rule is that
besides dam’s milk piglets should receive supplementary feed from their age
of 2 weeks.
In case of classical weaning (6-8 weeks) 2-week-old piglet receive a
“starter” feed, which is adjusted in its composition to the set of digestive
466
enzymes of young animals and together with dam’s milk meets the nutrient
requirement of piglets till week 5 of life. The starter is followed by “piglet
feed” of lower protein and energy level, which is offered after weaning too,
until changing over to “fattening feed I” (FIGURE XX-2).
FIGURE XX-2: Piglet feeding in case of weaning at week 6 (© FEKETE, L. 1995)
In case of early weaning (week 4) 2-week-old piglets receive

”prestarter”, which, together with mother’s milk, in itself should meet
piglet’s needs until weaning and the week after. At the end of week 5 ani-
mals will receive “piglet feed I and II” (FIGURE XX-4). From of 7 to 20 kg live
weight the average daily weight gain is 400-550 g, the daily feed intake 0.6-
0.8 kg with a feed conversion efficiency of 1.4-1.6.
FIGURE XX-3: Piglet feeding in case of weaning at the end of week (© FEKETE, L.1995)
467
In the US literature the name of piglet is used only till live weight of 20, and
each feed of this period (prestarter, starter, piglet I) is uniformly are called
“starter”. Between 20 and 50 kg is the growing and from 50 to 100-115 kg
of live weight (slaughtering) is the finisher phase.
Between chemical composition of starter and prestarter there is no
significant difference (maybe protein level in prestarter is higher), but
digestibility of prestarter is improved by special processing (rolling or flak-
ing, steam flaking, popping, extrusion, micronization, pelleting etc.). Starter
and prestarter may be called “suckling piglet” diet. From “piglet feed” may
be prepared a higher (“piglet II”, between live weight 10 and 20) and a lower
(“piglet II”, between 20 and 35 kg of live weight) protein and energy concen-
tration form. Table XX-3, using the above nomenclature, gives minimum
nutrient, mineral and vitamin levels of young piglet feed mixtures. (Protein
is given as crude protein, because an excellent biological value is a prereq-
uisite of their use.)
Composition of creep feed depends upon the age of piglets. For 2-3-
week-old-piglets milk protein and lactose containing high energy-high pro-
tein diet (prestarter, starter) should be prepared. Piglets prefer pellets of 0.3-
0.5×0.5-1.0 cm. Nutrient concentrations of creep feeds should be similar to
that of sow milk. Later, at week 2 after weaning it is possible to give the
piglets feed based on vegetable components. It is favourable to mix flavour-
ing into the pregnant sow’s and piglets creep feed, which are excreted by
milk. In this way, the voluntary solid feed intake can be stimulated by
means of imprinting.
In practice it is of high importance to avoid accumulation and over-
lapping of stressors (change of feed, place, loss of dam). At weaning it is the
sow, that should be removed from the pen. The most critical period is the
week after weaning, both in classical and in early weaning. Feed of this peri-
od should be given as creep feed at least one week before weaning (danger
of chronic scour problem). After weaning a feed restriction is proposed in
order to prevent incidence of oedema disease. The starting amount of creep
feed is 100-200 grams per piglet, which, by gradual increase will achieve the
typical average quantity on day 6-8 after weaning. Daily feed intake of
young piglets should not exceed 3-4% of their live weight.
Using the modern genotypes, the optimum weight at the age of 3
month should not be higher than 35 kg. This corresponds to 380-400 grams
of average daily gain in month 2 and 3 of life. Table XX-2 gives the compo-
sition of a typical “starter” and “piglet” diet.
In practice, there are successive series of feeds for piglets, and the
change of feed is linked to the amount eaten from the feed, rather than to
the age of animals. (Attention: the above described two weaning systems are
the most typical, but in practice there is a variety of weaning systems, char-
acterised by different schedules, nutrient composition, natural ingredients
and processing techniques. Nomenclature is often arbitrary, use of codes
468
and numbers are general, thus adhering to the attached technology is

essential.)
Artificial piglet rearing. For experimental purposes or orphan piglets,
after the colostrum suckled, may be raised artificially, too. One of the pos-
sible methods is feeding of a “baby milk replacer”, followed by a “baby
starter” feed in dry form, with ad libitum drinking. Milk replacer should be
made of ingredients with high nutritive and dietetic value (milk powder,
well emulsified animal fat, synthetic amino acids, minerals, antioxidant,
antibiotics) and its energy and protein concentration should be high (14-17
MJ DE, 22-26% crude protein). At the age of 2-3 weeks is the transition to
the above described prestarter. There are some initiatives of using
processed cow colostrum and mixed mature cow milk in liquid form for arti-
ficial piglet rearing.
Weaning system in North America. There is variety of weaning time
and applied feeds. The conventional weaning is at 6 weeks or older, using a
single starter of 18-20% crude protein and 1.0% lysine level. Medium large
producers apply the weaning at 3 to 4 weeks, using three starters, the first
one to approximately 6-7 kg of LW (22% crude protein and 1.4% lysine), the
second to the achieveing of 10-12 kg of LW with a crude protein level of 20%
and lysine concentration of 1.2% and the third with 18% crude protein and
1.0% lysine concentration up to the LW of 18 to23 kg. The first and second
are generally called prestarter and the third as starter diet. From LW of 18-
23 the grower diet (14-16% CP) and from approximately 54 kg of LW up to
slaughtering the finisher diet with 12-14% of crude protein are used.
A very special application of the recent immunological results is the
segregated early weaning (SEW), practised by really great pig operations.
The theoretical basis is that piglets should be removed from the dam hav-
ing their passive immunity on top, to avoid antigen exposure from the sow.
Thus, piglets weaned no later than 1.5 to 2 weeks of age, are removed and
raised in a remote clean facilities. By avoiding – minimizing their immuno-
logical stress, the feed utilisation and growth rate dramatically improve (see
Chapter XVII). SEW piglets have higher amino acid needs than conventially
reared pigs, they require a diet of high in lysine (1.6%) and of protein of ani-
mal origin, like spray dried blood plasma, blood meal, dried blood cells, fish
meal, skimmed milk, whey protein concentrate and processed soy protein
isolates. Soybean should be avoided because of its antigenicity. After eating
7-10 days this special feed mixture, pigs may be switche to the three phase
starter system: “phase feeding (WILLIAMS et al. 1997).
Acidification of feed and drinking water after weaning.

One of the post-weaning problems is the high pH value of feeds and the lack
of sufficient hydrochloric acid in weaned piglet stomach. Feed has high
buffering capacity and a pH of 6-6.5, while desirable stomach pH would be
at least 3-4. Buffering capacity of the feed can be measured by “acid-bind-
469
ing capacity”, which expresses the amount of hydrochloric acid in ml used

to set the pH of 100 gram of feed to pH 4.
A weaned piglet of 28 day and with a live weight of 6-8 kg produces
approximately 20 ml hydrochloric acid during a day, which is “enough” only
to acidify half of the daily feed portion. Since stomach pH has a role in pre-
venting bacteria from the environment from getting into the upper small
intestine and in activating pepsinogens, lack of hydrochloric acid may lead
to dyspepsia, diarrhoea and death.
To select feedstuffs of low buffer capacity cannot be the solution,
because the most valuable ingredients (milk and whey powder, fish meal,
extracted soybean etc.), as well as mineral supplements have high buffer
capacity. Reduction of protein level, use of synthetic amino acids and calci-
um formiate or monocalcium phosphate instead of limestone, may only
partly solve problem. Therefore, in addition, acidification of feed or drinking
water is proposed by means of hydrochloric and organic acids, betain
hydrochloride or lactic acid bacteria. Hydrochloric acid and betain mean a
strong acid loading of the acid base balance, thus their use should be lim-
ited to 5 days before and 5 days after weaning.
Table XX-3: Minimum nutrient allowences for suckling and growing piglets
* Baby pigs require extra iron supplementation !
470
Table XX-4: Natural ingredients and nutritive value of feed mixtures for suckling and grow
ing pigs*
* In most of the cases with synthetic amino acid and antibiotics supplementation
471
20.3. Theoretical basics of pig growth („fattening”)

© Sandor Gy. Fekete)
The starting point of „fattening” (i. e. growing-finishing operation) is a

„weaned” piglet of 3 months age and 30-05 kg of live weight, independently
from the length of real suckling time. Until now all piglets are kept togeth-
er, because too intensive raising is not desirable either for future breeding
animals or even for fattening piglets (danger of oedema disease). Typical
slaughter weight in Europe is 100-110 kg of live weight, when animals are
170-200-day-old. In practice, the old term „fattening” is frequently used,
although this is normal growth. In fact, efforts are made to avoid fat depo-
sition and to enhance muscle (lean mass) accretion.
The metabolisable energy content of the ingested feed is distributed to
heat, bone, muscle and fat production and this repartition with the
advancement of age changes (FIGURE XX-4). Feed composition should be
adjusted to this process.
FIGURE XX-4: Progress of bone, muscle and fat to their mature weight in the body, rela
tive to the progress of the total body to its mature weight (after BERG and BUTTERFIELD, 1985)
20.3.1. Energy requirements of growing-finishing pig

The most common way of expressing it is the digestible energy (DE) in mega-
joule (MJ). For the sake of comparison, there may be a need for converting
it into metabolisable energy (ME). In case of feed mixture the simple equa-
tion (ME=0.96×DE) can be used. In case of raw materials and feedstuffs of
variable protein level the following equation is useful:
ME = (0.997-0.000189 CP) × DE, where CP = crude protein, g/kg.
473
Maintenance energy needs for pigs are 0.477 MJ DE × W0.75, the

accretion of 1 kg of protein requires 45.7, deposition of 1 kg of fat 55.7 MJ
DE. Environmental temperature modifies energy needs: minus 5oC devia-
tion from the lower critical temperature requires approximately 1.1 MJ DE
extra energy. Lower critical temperature for piglets of 30-35 kg of live weight
is 17.6oC, for a finishing pig of 100 kg of live weight 13oC. For more details
see „Effect of environment on nutrient requirements of domestic animals
(NRC, 1981).
20.3.2. Importance of energy: protein ratio

Among the feeding factors basically the energy to protein ratio determines
repartition of energy. Energy is the common expression of the nutritive
value of all major chemical components, which are, within physiological lim-
its, interchangeable and protein, which cannot be replaced by other nutri-
ent. Thus, energy and protein supply cannot be treated separately.
For reflecting energy to protein ratio of the feeds, the E: P ratio used
to be applied when the amount of energy of a unit protein was given.
Nowadays the „protein to energy ratio” (P/E) is mostly used, which gives the
amount of protein (gram) per energy unit (MJ). The higher the P/E value,
the richer the feedstuffs or feed mixture is in protein. Requirements of ani-
mals can also be given in form of P/E, for example piglets of 0 to 3 weeks of
age require a feed mixture of P/E=14 and at the age of 3 to 8 weeks 12. It
means that for each MJ digestible energy 14 or 12 gram crude protein
should go. Amino acid needs can also be given in a similar form, e. g. piglets
of 0 to 3 weeks require 1.12, those of 3 to 8 weeks 0.98 gram lysine per each
MJ DE.
Concerning energy to protein ratio during fattening, there are three
basic situations. The ideal is when energy is just sufficient for maintenance
and protein accretion. In this case there are neither energy nor protein loss-
es. If there is a relative protein deficit, fat surplus forms fat („luxury con-
sumption”). Nevertheless, when the biological value of the protein is not
excellent, extra energy may save some protein (“protein sparing effect”). If
energy is not enough for covering the costs of protein accretion, the remain-
ing protein will be used as energy source for heat or fat („double luxury con-
sumption”).
Therefore, supposing an optimal N supply, protein deposition in grow-
ing pigs responds linearly to increasing energy intake until a maximum is
attained at which the response plateaus. This linear-plateau relationship
assumes that maximal lean tissue growth is determined by intrinsic factors,
which has since been demonstrated for pigs approximately above 50 kg of
live weight (FIGURE XX-5). Where the energy intake of an animal is suffi-
cient to maximize protein accretion (reach the plateau) any extra energy
supplied is deposited as fat, resulting in an overall decline in the rate and
efficiency of growth and marked increase in body fat content. These
474
responses are commonly observed in pigs above 50 kg.
FIGURE XX-5: Relationship between energy intake and N balance in pig weighing 75 kg
(after DUNKIN et al. 1986)
When the diet is improved by increasing the protein content in rela-

tion to the energy it will reduce fatness, as shown in FIGURE XX-6. If the
protein supply is inadequate with respect to the needs of maintenance and
protein accretion, dietary energy which is not used for lean tissue growth
will be diverted to deposite fat.
FIGURE XX-6: Growth

responses of lean and
fat to increasing diet
protein level (after
COLIN and WHITTEMORE,
1986).
475
20.3.3. Protein and amino acid needs of growing-finishing pig

Protein requirements of growing-finishing pig can be evaluated by factorial
calculation and measuring biological answer (growth, feed conversion effi-
ciency, body composition) while feeding feed mixtures of different protein
concentrations. Pigs have qualitative and quantitative demand for dietary
protein. Quality includes digestibility, biological value and the possible
presence/lack of antinutritive substances. Using a corn-soybean diet, the
first limiting amino acid for growing-finishing pig is lysine, then the sul-
phur-containing methionine and cystine, after the tryptophan and threo-
nine. In practice it would be too complicated to calculate quantitatively with
all of the essential amino acids. Instead, the requirement of the first limit-
ing amino acid, the lysine is given quantitatively (g/kg feed, g/MJ DE, % of
CP) and the other essential amino acids according to ideal protein propor-
tions are given as a percentage of lysine, which is regarded as 100%. Table
X-1 (see Chapter X) shows lysine needs of growing-finishing pig and the
appropriate proportions of the remaining amino acids.
POSSIBILITIES OF MEAT AND BONE MEAL REPLACEMENT IN PIG NUTRITION

After the occurrence of BSE use of animal meals has been prohibited. Raw
material which (partly) are able to replace them, mostly derive from places
outside the European markets. EU farmers produce annually 12-14 million
metric tons oilseed, two third of them being rapeseed, and approximately
one fifth sunflower seed and soybean. It means that rapeseed needs are just
met, but the production of sunflower is only enough for 30 and that of soy-
bean for 4-5%. As a consequence, soybean importation and the production
of other European protein sources (pea, bean and sweet lupine) should be
increased. Although the gross amino acid content is relatively high (Table
XX-6), but owing to antinutritive factors, their biological value is lower (30-
50%) than that of proteins of animal origin.
Table XX-6: Crude protein (CP) and the most important amino acids of some vegetable
feedstuffs comparing to pig requirement
* Canola, without eruic acid and glycosinolates
476
There are further concerns, because most of the imported soybean

may be GMO=genetically modified organism, which may cause deep revul-
sion in part of the European population. At the same time, within the EU
the sowing area of oilseed is maximized by the Blair House. In addition, the
feed conversion efficiency may decrease without using animal proteins.
Costs of the described factors difficult to assess.
Applying of ideal protein conception

Protein synthesis in tissues is the most effective if all necessary essential
amino acids are present at the same time and in appropriate proportion.
This situation can be realised by using only vegetable and synthetic raw
materials. BAKER (1993) published ideal amino acid composition for growing
pig (Table XX-7) and the possible performance parameters (Table XX-8). A
comparison of the production traits of pigs fed on natural feed ingredients
or on casein plus synthetic amino acids mixture is given in Table XX-9. It
can be stated that gain per unit of crude protein intake (PER) was vastly
superior in the ideal protein diet compared with the corn-soybean meal pos-
itive-control diet of higher crude protein levels. Nevertheless, it should be
emphasized that natural control was free of animal protein.
Table XX-7. Ideal pattern of indispensable amino acids for pigs in three separate weight
categories
*53% L-Phenylalanine+47% L-tyrosine **50% DL-methionine+50% L-cystine
477
Table XX-8: Efficacy of Illinois Ideal Protein (IIP) when indispensable amino acids are set
at close to requirement levels for 10-kg pigs†
†Means of six individually fed (ad libitum) pigs; 14-day assay: average initial weight 10.6
kg;*53% L-phenylalanine +47% L-tyrosine; **50% DL-methionine + 50% L-cystine.
Table XX-9: Comparison of an ideal protein diet with a standard corn-soybean meal diet
of finishing pigs†
†Means of six individually (ad libitum) pigs during a 14-day assay period (three barrows
and three gilts per diet); average initial weight was 77 kg.
1Contained 79.65% corn, 15.39% solvent-extracted dehulled soybean meal, 2.00% corn
oil and 2.3 g/kg L-lysine-HCl; vitamins and minerals were added to meet or exceed NRC
(1988) requirements.
2Contained 5.58% casein, 5.21% crystalline amino acids, 42.14% corn starch, 30%
sucrose, 5.0% solka floc, 5.0% corn oil and 1.12% NaHCO3; vitamins and minerals were
478
added to meet or exceed NRC (1988) requirements. This diet was formulated to have the
exact same indispensable amino acid profile as the proposed IIP pattern for finishing (50-
100 kg) pigs (Table XX-7)
* Ideal protein diet (casein + amino acids) different (P<0.05) from corn-soybean meal diet
** PER= Protein Efficiency Ratio i. e. body weight gain produced from 1 kg ingested
dietary crude protein
Possible ways of replacing animal proteins include selection of plants

for higher and more valuable protein content, as well as the production of
Single Cell Protein (SCP) by bacteria and yeast on mineral oil, methanol or
other substrate. Unfortunately, preference of SCP for animals is bad, except
furbearer carnivores. Use of ideal amino acid concept in ration formulation
– especially the one based on ileal digestible amino acids (see Chapter VII) –
in experimental circumstances proved to be successful. Nevertheless, feed-
ing growing-finishing pigs on animal protein free diet in practice may lead
to a decrease in performance (less daily gain, worse feed conversion effi-
ciency and higher fat ratio in carcass).
20.3.4. Fat requirements of growing-finishing pigs

There is a minimum fat requirement (approximately 3%) which assures
absorption of fat soluble vitamins and essential amino acids. Added fats
ands oils increase energy density of diet and may improve feed utilisation,
without affecting daily gain. In hot environment extra fat help in replacing
lacking energy due to a decreased feed intake, Too much fat supplementa-
tion leads to the formation of calcium and magnesium sops, inhibiting
absorption of these macro elements.
Needs of growing-finishing pigs for essential fatty acids are deter-
mined by their role in protein accretion, by preventing dermatitis and by
setting trien (linolenic acid) to tetraen (arachidonic acid) concentration of fat
tissues. Piglet with 10 to 30 kg of LW requires 1.5% linoleic (C18:2) acid,
while those with 30 to 90 kg 0.70% linoleic acid in feed dry matter. The
other essential fatty acid can be synthesised from linoleic acid. Under the
circumstances of high corn feeding there is practically no essential fatty
acid deficiency, because approximately 40% of corn oil is linoleic acid. The
situation is similar while feeding oats, but wheat, rye and barley are low in
oils and essential fatty acids.
20.3.5. Ballast (indigestible organic matter) need of growing-finishing

pig
Growing-finishing pig partly utilise fibre as an energy source by hind gut
fermentation, but this is not essential for the animal. Fibre is indispensable
for growing-finishing pig because of its dietetic effects. Optimum fibre con-
centration should be determined on the basis of its effect on digestibility of
other nutrients, on feed utilisation and produced body gain.
479
With regard to digestibility (especially that of protein) there is no fibre

optimum: starting from zero, each additional percent in fibre decreases
nutrient digestibility. This relationship can be expressed by the equation of
Y = 93-1.65 X, where Y= digestibility of organic matter, % and X= crude fibre
concentration, %. It means that the digestibility of a fibre free diet is gener-
ally 93% and each additional percent in fibre content decreases digestibili-
ty of organic matter by 1.65 percent unit. In contrast, by increasing fibre
concentration the feed utilisation increases, then achieving a maximum
point („zenith”) slowly decrease (FIGURE XX-7).
FIGURE XX-7: Effect of dietary fibre on digestibility and utilisation of feed (FEKETE, L. 1979)
The explanation of the described phenomenon is the dietetic effects of

fibre fractions: toxin adsorption, production of antibacterial volatile fatty
acids and pacemaker activity on peristalsis.
FIGURE XX-9: Dietetic effects of fibre fractions
This overall ballast (bulk) effect – to a certain limit – exceeds and com-
pensates losses of decreased digestibility. As a result, until reaching opti-
mum fibre concentration, daily gain steeply increases parallel to increasing
fibre level. By adding more fibre than the optimum, daily gain slowly
decreases (FIGURE XX-9). Crude fibre optimum for piglets is 3% and for
growing-finishing pig 4-5% in air-dry feed mixture.
480
CHAPTER XX: FEEDING OF GROWING-FINISHING PIG
20.4. Applied nutrition of fattening (growing-finishing) pig

© Janos Gundel
peaking about pig fattening does not really mean „making get fat”, but
S rather a way of pig raising with the purpose of producing slaughter-

ing animal. The generally accepted form is to start at 25-35 kg of LW
and to slaughter at 100-110 kg of LW. There are exceptions, for example in
Denmark slaughtering weight is lower, whereas in Hungary (salami), in Italy
and Spain (ham production) may be higher.
20.4.1. Definition of „fattening”; economic considerations

Till the early 90’s the goal of fattening used to be simple: to achieve slaugh-
ter weight economically, as quickly as possible and get a good lean meat
(EUROP) qualification. Nowadays the ain became more complex. Besides the
aims previously described, the environmental concerns should also be con-
sidered, and the body composition of pig should be a healthy food raw mate-
rial for human consumption. The latter includes – among others – meeting
food safety expectation and increasing intramuscular fat against subcuta-
neous one as well as alpha-linoleic and vitamin E concentrations in pork
meat. In addition, depending on the actual farm special goals may be set
like using some local feedstuffs or by products (potatoes, skimmed milk,
whey) or to produce a product of a certain trade mark (Parmian gammon).
To accomplish all these targets genetics and feeding are both essential. Pig
fattening basically is guided by economic considerations and farmer has to
make his decision about the fattening goal.
20.4.2. Basic notions of fattening

although there are different ways of fattening, in the following the most
common system will be described. Available genotypes can be classified into
two groups: the high lean-meat, modern genotypes (this is the majority) and
the traditional, generally predominantly fat-producing ones. Pure breeds
and crossed hybrids are available in both categories. Feeding allowances
differentiate only among the different production types, without taking into
account the differences between the particular breeds and hybrids
There are data and recommendations available concerning the differ-
ent requirements of males, females and castrated animals, which are rarely
applied in every day practice.
Although requirement of growing-finishing („fattening”) pigs – similar-
ly to other species – are known per day and per live weight, in practice an
optimal nutrient concentration of air dry matter is given, and nutrient sup-
ply is controlled through the daily feed intake (daily ration). Ad libitum feed-
ing, especially above 50 kg of LW cannot be proposed because pigs of high
appetite will eat more than their real needs which, in turn, increases fat
481
accretion and reduce feed conversion efficiency. In case of combined feed-

ing (using feedstuffs of different water content, e. g. cereals and whey), the
daily ration should be calculated air-dry matter basis.
In practice, fattening should be divided at least into two parts.
Theoretical basis of this can be seen on the idealised plot of protein and fat
accretion vs. increasing bodyweight (FIGURE XX-10).
FIGURE XX-9: Description of protein and fat accretion in meat animals (after BERGEN,
1974, modified)
During early life, protein and fat accretion occur simultaneously,

whereas in later life the rate of protein accretion becomes negligible and fat
accretion continues. As animals approach mature size fat content of the
body exceeds protein content. Because of the greater energy value of adi-
pose tissue than that of muscle gain (1.0 kg protein and 4 kg water = 23 MJ,
whereas 23 MJ = 0.6 kg fat.), delaying of the „fattening point” increases the
efficiency of body weight gain and lean meat production. Thus, above 50 kg
of live weight, instead of ad libitum feeding, the restricted feeding (giving
daily portions) is proposed. Table XX-11 compares data of the recent
Hungarian Feed Codex and the today’s NRC recommendations. From data
the trends of development can be drawn: aspects for giving nutrient require-
ment are not only the average daily gain, feed conversion efficiency and car-
cass quality, but also the reduction of nitrogen and phosphorous emission
(environmental protection). Vitamin A and D concentrations became lower
and P and niacin availability is also considered.
482
Table XX-11: Fattening pig I. (30-60 kg LW) and Fattening pig II. (60-110 kg LW):
HFC=Hungarian Feed Codex, 1990. Growing (20-50 kg LW), Finishing I. (50-80 kg LW) and
Finishing II (80-120 kg LW): (NRC, 1998).
————————————————————————————————————————————
Items 30-60 (HFC) 20-50 (NRC) 60-110 (HFC) 50-80 (NRC) 80-120 (NRC)
————————————————————————————————————————————
Dry matter,% 86 90 86 90 90
DE, MJ/kg 13.5 14.2 13.5 14.2 14.2
Crude protein, % 16.8 18.0 15.0 15.5 13.2
Lysine, g/kg 8.9 9.5 7.8 7.5 6.0
Met+Cys, g/kg 5.3 5.4 4.7 4.4 3.5
Tryptophan, g/kg — 17 — 14 11
Threonine, g/kg — 61 — 51 41
Crude fat, % 4.0 — 3.5 — —
Crude fibre, % 4.5 — 4.5 — —
Calcium, g/kg 8.6 6.0 7.0 5.0 4.5
Posphorus, g/kg 6.0 5.0 5.0 4.5 4.0
aP* — 2.3 — 1.9 1.5
Mg, g/kg 0.4 0.4 0.4 0.4 0.4
Na, g/kg 1.5 1.0 1.5 1.0 0.1
Vitamin A, IU/kg 2000 1300 2000 1300 1300
Vitamin D3, IU/kg 300 150 200 150 150
Vitamin E, mg/kg 10 11 10 11 11
Vitamin B12, μg/kg 10 10 10 5 5
Niacin, mg/kg 15 10** 15 7** 7**
Zn, mg/kg 50 60 50 50 50
Fe, mg/kg 40 60 40 50 40
Mn, mg/kg 25 2 25 2 2
Cu, mg/kg 5 4 5 3.5 3
I, mg/kg 0.2 0.14 0.2 0.14 0.14
Se,mg/kg 0.15 0.15 0.15 0.15 0.15
Co, mg/kg 0.1 —- 0.1 —- —-
————————————————————————————————————————————
* aP=available P= ** available niacin

During fattening, the pigs are biologically developing organism which are
slaughtered before achieving maturity, thus maintenance and growing
needs are given together. The real requirements of an animal comprise
needs of maintenance and production (weight gain in an expected composi-
tion) of a given genotype. In practice, genetic potential cannot be totally
used, except some experimental initiatives (e.g. use of recombinant Porcine
Somatotropin, rPST).
Genotype is independent, but the level of production significantly
depends on environmental factors (nutrition, size of production unit, tech-
nology, ownership, temperature and humidity etc.) In the modern nutrient
recommendations two series of data are listed: requirement values in abso-
lut numbers and the nutrient concentration of feed mixture potentially able
to meet actual needs. Data are scarce about the influence of environment
on pig nutrient requirements. Prediction of expected voluntary dry matter
483
intake is extremely important, because stomach capacity of pig is relatively

limited, although their nutrient requirements are high. Thus, to avoid the
negative feed back of physical fullness to appetite, concentrated feed mix-
ture with high dry matter content should be given (Table XX-12, XX-13, XX-
14).
Energy requirement of pig feeds and feedstuffs are generally given in
digestible energy, DE (majority of European countries), in metabolisable
energy, ME (German speaking countries, USA), and with experimental char-
acter in net energy (Denmark, France). Data of other nutrients (e. g. amino
acids) can be given in air dry matter, in dry matter and in relation to the
energy.
According to a modern concept, instead of protein supply the amino
acids are taken into consideration. However, for practical use, data of crude
protein (CP) requirement are also listed in allowances. Previously amino
acid requirements were given in total (gross) amino acid, fixed lysine to
crude protein ratio (5.0-7.0%). Today’s amino acid needs are based on the
„ideal protein” concept, introduced van LOEN in 1966. Ideal protein means
that the required amino acid composition is given as a percentage of lysine,
appropriate for a certain production. Nowadays not only faecal, but also
ileal digestible amino acid composition of ideal protein is known making it
possible to calculate the real requirement more accurately (Table XX-15, XX-
16, XX-17 and XX-18).
The majority of nitrogen-free extracts (N-f.e.: starch, sugars etc.) are
important energy source, fibre has dietetic effects. Fats and oils (ether
extract, EE) provide energy, transport store fat soluble compounds and by
their poly unsaturated acid components have specific physiological func-
tions. Thus, in the modern allowances data for linoleic acids (1-2%) are also
available.
As far as calcium and phosphorus requirements are concerned, not
only their quantity, but also the ratio is essential (generally 1.2-1.4 to 1).
Besides, there is a need for sodium, chloride, iron, zinc, copper, manganese,
iodine, selenium, vitamin A, D, E and members of group B supplementa-
tion. Except some mutant genotypes, pigs are capable of synthesising vita-
min C. The summary of recommended nutrient concentrations for three
weight classes are shown in Table-XX-19.
One of the most important tasks is to ensure adequate drinking water
supply (Table XX-20). Generally water is provided ad libitum from self-
drinkers and the expected water consumption is 2.5-3.0 litres per kg dry
matter ingested. Due to the lack of sufficient water first the production
drops and later health problems occur. Too much water causes economic
losses (e. g. production of liquid manure in a large quantity). By using liq-
uid feeding systems, besides self drinkers, dry matter to water ratio should
be below 2.5, otherwise the stomach quickly becomes full and inhibits feed
intake.
484
As mentioned before, in an ideal case, fully meeting the nutrient

requirements helps to realise the full genetic potential. In practice, owing to
the different health status and environmental factors, only 60-80% of the
possible production level can be achieved. Nevertheless, there is an oppor-
tunity to alter composition and quality of swine carcass and meat through
different feeding intensity and composition. By changing amino acid to ener-
gy ratio the amount and distribution of fat accretion can be modified. By
feeding different fats and oils unsaturated fatty acid composition of lard and
intramuscular fat may increase and extra vitamin E (and selenium) supple-
mentation improves oxidative stability of meats etc. As a new trend, the pro-
duction of meat with special (dietetic and pharmaceutic) characteristics for
human consumption (so-called functional food) also deserves attention.
It should be emphasised that tabulated data generally relate to medi-
um capacity genotypes kept in ordinary environmental conditions. However,
production level primarily depends upon the actually ingested feed and its
composition, therefore the issue of voluntary feed intake should be dis-
cussed in more detail.
485
Table XX-12: Expected feed, dry matter, energy and nutrient intake of growing-finishing
pigs
486
Table XX-13: Recommendation for energy, protein and amino acid concentration in grow-
ingfinishing pig diet
DE= digestible energy; CP= crude protein; LYS= lysine; MET+CYS= methionine+cystine;
THR= threonine; TRP= tryptophan
Table XX-14: Recommendation for fibre, fat and macro elements concentration of growing
finishing pigs
CF= crude fibre; EE= ether extract
487
Table XX-15: Recommendation for optimum amino acid concentration according to

„ideal protein” principle, %
Table XX-16: Apparent and true ileal digestion coefficient of the most important amino
acids (%
Table XX-17: Recommended apparent ileal digestible amino acids for growing-finishing pigs
488
Table XX-18: Recommendation for true ileal digestible amino acid concentration in growing
finishing pig diet
489
Table XX-19: Recommended nutrient concentration in fattening pig diet
High lean-meat genotypes; ** Medium lean-meat genotypes

CP= crude protein; EE= ether extract; CF= crude fibre
490
Table XX-20: Average daily drinking water consumption (litre/kg dry matter)
20.4.4. Feed intake of growing-finishing pigs

Among the factors influencing the success of fattening one of the most sig-
nificant one is real feed consumption. Regulation of voluntary feed intake is
a very complex physiological process (for more details see Chapter VI).
Summarising for practice, pigs generally eat according to their energy need
but after reaching the limits of stomach capacity, they stop eating, even
when their energy needs are not fully met. Feed intake is affected by many
genetic (endogenous) and environmental (exogenous) factors. Among the
genetic factors the genotype, gender, constitution and intake capacity
(„greed”) are the most important. Nowadays the genetic intake capacity (an
average of 600-800 g/day) could be increased by 20-40% simply by selec-
tion and crossing as well as feed conversion efficiency by 10-30%.
External factors are strongly related to the given feed, management
system and possibly to economic considerations.
► Factors of feed: composition, quality of raw material, used supple-
mentations, energy density, protein quantity and quality (ideal protein prin-
ciple, ileal digestibility of amino acids), digestibility of dry matter, particle
size and palatability (preference of animal).
► Elements of feeding technology: predicted daily feed intake, fre-
quency of feed distribution, method and form of distribution, change of feed
composition and natural ingredients and drinking water supply.
► Factors of management system: slaughtering weight and age, stall
equipments, dimension and arrangement of pen, feeding regime and quali-
ty of feedstuffs, place for resting and defecation, herd size, settling density
(m2/animal), stall climate, general hygiene and prophylactic measures.
► Economic considerations: rent ability and effectiveness, production
goal and interest and organisation of labour.
491
Table XX-21: Expected daily feed intake and gain
In practice one of the greatest problems of growing-finishing operation

is the insufficient feed intake. Data of Table XX-21 show the average expect-
ed daily feed intake. As a rule of thumb a piglet of 20-30 kg of LW consumes
5%, a pig of 100-120 kg of LW 3% of its body weight from an air-dry feed
mixture daily.
Modern, fully automated feed distributing technologies use softwares
to predict actual live weight and on that basis they correct not only the daily
amount of feed, but also its composition. This method help to avoid over-
feeding, minimize environmental pollution by dung (nitrogen and phospho-
rous) and provide information for economic calculation by pens.
Daily control is a key factor in optimizing feed intake. Maybe, one of
the greatest advantages of the automatic system is that it allows sufficient
time for watching the animals during eating.
20.4.5. Applied feeding systems

Today there is a vast choice of different feeding techniques and technologies;
each of them can be applied correctly but can also be misused. Each of
them requires local adaptation and professional service. The personal
observation of the eating animals cannot be disregarded and should be used
not only to check real feed intake, but also to get information about the pigs’
health state, which leads to a more professional feeding regime („The
farmer’s eye make the pig grow faster!”).
Modes of feeding can be classified by:
492
Feeding of air-dry meal or pellets is common. To avoid dust formation,

it is recommended to moisten them preferably in the trough. There are sys-
tems, where animals can moisten their own feed. The addition of 1-2% veg-
etable oil (e. g. soybean oil) may also be a solution. Viscous, moistened and
liquid feed are considered to be more natural for pigs. Advantage of dry feed
is that it allows higher feed intake; while feed with higher water content is
easy to distribute mechanically, it has less friction resistance and dosage
may be more accurate. In viscous feed dry matter to water ratio is 1:1-1.5,
in liquid feed 1:3 (frequently 1:4-5). For liquid, besides water skimmed milk,
whey and others may be used, if their clearness and temperature, as well as
preference and hygienic status makes possible it. The best mixing ratio is 1
to 2.5-3, with a continuous access to drinking water.
Feeding intensity can be controlled by the daily amount of distributed
feed (i.e. energy). Rationing is a form of restricted feeding, when animals
receive a previously determined quantity independently of their actual
appetite. This is applied for special purposes, for example bacon production.
More frequently used is the semi-ad libitum portioning, which is basically a
feeding according to appetite. Actually, rationing is practised, but the
amount of daily ration is adjusted according to the appetite of to herd every
3-4 days. In ad libitum feeding systems pigs have free access to feed all day
long and they can eat according to their appetite. To avoid overfeeding („lux-
ury consumption”) and consecutive fat accumulation, fibre level should also
be elevated. Advantage of this system is that requires less labour, but ani-
mals may lose strict surveillance. In time-restricted ad libitum technology
feeders are open only for a given time.
The feed can be distributed on the floor, although feed losses are
great, and in case of meal form dust production is health threatening.
Nowadays it is mostly used only for feeding roughages (e.g. silage, green
alfalfa chops etc.). Mangers and self-feeder constructions should prevent
feed spoilage, should be easy to clean and should be resistant against cor-
rosion. Length of trough is significant: in case of rationing all member of
herd should be able to eat at the same time; inner wall should have a
smooth surface to facilitate the spreading of feed mixture. There is also a
combination of group keeping and individual feeding, as in case of breeding
sow. A new development is the self-drinker which is placed above the self-
feeder, with the advantage that dropping water fells into feed and pig can
eat and drink in the same place.
20.4.6. Feedstuffs of growing-finishing pigs

Feed for fattening pigs should meet the animals needs, cannot contain
harmful compounds, have to be balanced and homogenous, should satisfy
food safety requirements and should comply with the relevant regulations.
The range of possibly feedstuffs is determined on the basis of biological and
493
legislative considerations. The pig, being omnivorous animal is able to use

a very wide range of feedstuffs. All of the cereals, agricultural and industri-
al by-products are equally suitable for them to eat. Raw materials are clas-
sified according to their nutrient concentration.
▪ Concentrated feedstuffs are the air-dry grains and seeds (cereals,
corn, oilseeds, leguminous seeds and their extracted meals) and dried ani-
mal products (fish meal, milk powder, etc.). Feedstuffs of this group – phys-
iologically – can be fed without limitation. Soybean and their products
should undergo heat treatment in order to inactivate trypsin inhibitor. Heat
treatment of cereals – by starch processing and sterilisation – may improve
nutritive value, but except for piglet feeds, it is not economic. Genetically
modified organisms (GMO) can be used as feedstuffs in accordance with the
actual regulations.
▪ A series of bulky feedstuffs (potatoes, sugar beet pulp), roughages
and by-products are suitable for pig, but their use in practice is rare. The
reasons for this include their low nutrient density, difficulties of distribution
and seasonal availability. In addition, their transport, storage and pre-treat-
ment require special technology and care. Mostly they are used in close
proximity of their production.
▪ The third group of raw materials are the high moisture grains.
They are conserved by fermentation (silage making). Most of them are based
on corn, especially corn grain silage and the CCM (corn-cob mix: besides
grains containing the upper third of cob too). The dry matter content of both
are 28-32% and the most important difference is in their fibre levels: 2-3%
in grain and 5-7% in CCM, which cause also a decrease in energy and pro-
tein content of the latter. Seldom used is the CSM (corn-soybean mix),
which is a mixture of high moisture crushed corn grain and fullfat soybean
seed fermented together. After grinding and mineral-vitamin supplementa-
tion it would meet the nutrient requirements. Fermentation partly elimi-
nates soybean’s trypsine inhibitor. In some European countries the use of
whole cereal grain (solely or mixed) and the sorghum CCM (silage of whole
canicle are in use.
20.4.7. Feed additives

Without using any feed additives, there is no intensive pig fattening. They
can be technical ingredients (helping for example pelleting process), feed
flavours, pigments, energy sources, growth promoters, coccidiostats and
histomonostats. Most of them have no energy value, but they complete the
feed mixture composition (e. g. synthetic amino acids), improve feed taste or
quality of meat. The use of growth promoter used to be common in feeding
of growing-finishing pig, but for food safety reasons, their application is con-
tinuously decreasing or there is a shift from antibiotics to nutriceuticals. In
the followings most of the ever used growth enhancer are described, refer-
ring to the present legal status too (only generic name is given disregarding
494
trademarks).
COPPER SULPHATE (CuSO4×5H2O): in order to avoid environmental pol-
lution by faeces, only 35 mg/kg feed is allowed, and only in the growing
phase of the animals.
ANTIBIOTICS and ANTIMICROBIAL SUBSTANCES are not allowed in the
European Union. In the United States 15 different antibiotics are in appli-
cation, with an obligatory withdrawal period of 5-70 days before slaughter-
ing. Actual regulations are published in the annual Feed Additives
Compendium.
Natural PLANT PRODUCTS like milling products and extracts are widely
used. Extract of Yucca plant is able to bind ammonia in the gut; other is
anticoccidial, antimicrobial (oregano) and immunostimulant (garlic).
FLAVOUR and AROMA SUBSTANCES: anise, cumin, clove, cinnamon, fennel
oil, sucrose, sodium salt of saccharin (exclusively for piglets till the age of 4
month, maximum in 35 mg/kg feed concentration), ethyl-vanillin, fructose,
mannitol, sorbitol, xylitol, cyclamates, aspartame. Addition of sugar replac-
er dulcin is prohibited. It is worth mentioning that they cannot be applied
to cover taste and smell failure of deteriorated or mouldy feeds. It is a gen-
eral principle that all additives, allowed to be used in human food (see
Codex Alimentarius) can be added to pig feed too.
ORGANIC ACIDS: lactic, fumaric-, citric, formic and propionic acid. They
stimulate gastric acid production, stabilise intestinal flora, and in addition,
propionic acid – through methyl-malonil-CoA pathway – inhibits fat synthe-
sis.
ENZYME SUPPLEMENTATION: cellulase, pectinase, beta-glucanases, pen-
tosanase, alkali protease, pepsin with hydrochloric acid, alpha-amylase,
trypsine, lipase and lysosime.
PROBIOTICS: bacterium or yeast cultures, e. g. Streptococcus faecium
Lactobacullus acidophylus, L. casei, L. lactis, Bacillus toyoi and
Saccharomyces (Pediococcus) cerevisiae. These microbes help in maintaining
optimal balance in gut flora and prevent development of dysbiosis.
REPARTITIONING AGENTS. Under their influence there is a shift from fat to
protein synthesis decreasing body fat and increasing lean meat ratio. They
only act as long as there is an excellent protein and amino acid supply. In
European Union application of growth hormone (GH) injection, immunisa-
tion against somatostatin and supplementation of feed by beta-2-agonists
(clenbuterol, ractopamine) is prohibited.
NUTRICEUTICALS, like L-carnitine, betaine and trivalent organic chromi-
um (chromium-picolinate, chromium nicotinate, and chromium enriched
yeast) also support protein accretion and decrease fat deposition.
Conjugated linoleic acids (CLA) biochemically inhibit fat synthesis, but their
application is only in experimental phase.
For evaluation of effectiveness in growing-finishing operation the
fat-free weight gain is the best parameter. At the beginning (2-4 month of
495
age) the fat-free lean carcass should be calculated (NRC, 1998):

CFFL, kg=0.95 × (0.92 × W – 3.65)
where CFFL= carcass fat-free lean and W= live weight, kg. For exam-
ple in case of a piglet of 35 kg of live weight this value is equal to 27.12 kg.
At slaughtering the fat-free lean index (FFLI), i.e. the fat-free portion
of carcass should be calculated:
FFLI, %= 50.767 + (0.077 × HCW) – (3.55 × BF)
where HCW=hot carcass weight, kg, BF=last rib midline backfat on
hot carcass, cm. If hot carcass weight of a slaughtered pig of 110 kg was 66
kg, then fat-free lean index is 48.75% and the fat-free gain 66-27.12=38.88
kg.
As a matter of fact,are also feed supplement, but they have special
regulations concerning registration, mixing, file keeping, withdrawal time
etc. They cannot be used for prevention and as growth promoters, only for
treatment, under veterinary control.
20.4.8. Balancing the ration: optimizing

Complete feed mixture of growing-finishing pig is generally produced in feed
mill and carried to the farm. Their advantage is that it contains all the nutri-
ents, but makes it impossible to use local feedstuffs and by-products. In
case of on farm mixing only supplements (energy or protein concentrate,
mineral-vitamin premix etc.) are bought and combined with local feedstuffs.
This later practice makes it possible to adjust feed composition to the given
herd. Guidelines for feed composition can be found in national and inter-
national recommendations (ARC, DLG, Hungarian Feed Codex, INRA, NRC
etc.) as well as on the web.
Computing feed formula is made by means of a two-phase simplex

algorithm, where the objective function is the fulfilment of minimum-maxi-
mum conditions with the least costs possible. Calculation can be carried
out for daily ration or for unit of feed (100%). In practice, this process is
called optimization, although this is actually a prize (and not costs!) min-
imisation.
Some typical growing-finishing feed mixture is shown in Table XX-22.
496
Table XX-12: Natural ingredients (%) and nutrient concentration of some typical growing
finishing pig diet
————————————————————————————————————————————
Ingredients Diet A Diet B Diet C Diet D Diet E*
————————————————————————————————————————————
Corn 76.0 45.0 - - 74.44
Wheat - 40.0 - 40.0 —-
Barley - - 83.0 39.0 ——
Soybean
(47% CP) 20.5 11.5 13.5 15.0 22.40
Fat/Oil - - - 2.5 ——
Premix 3.5 3.5 3.5 3.5 2.16
- - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - —- — - - - - - - - - - - - - - - -
DE, MJ/kg 14.4 13.5 12.9 13.9 14.5
Crude protein. % 15.9 15.7 15.0 16.7 16.1
Lysine, g/kg 8.1 7.8 7.4 7.8 8.2
Met+Cys, g/kg 6.1 6.2 5.1 5.9 5.9
Tryptophan, g/kg 1.7 1.0 2.2 2.3 2.3
Calcium, g/kg 9.0 9.0 9.0 9.0 9.0
Phosphorus, g/kg 7.5 7.5 7.5 7.5 7.5
————————————————————————————————————————————
* NRC (1998) „fortified diet”
20.4.9. Feeding growing-finishing pig and environmental protection

Recently in developed countries environmental concerns of animal produc-
tion receive more and more attention. Animal husbandry is an operation in
which is dung and urine excretion inevitable. After theoretical calculation
for the living space of one pig 10.5 kg nitrogen and approximately 2 kg phos-
phorus are excreted into the environment yearly. Therefore, feeding should
be designed to be economic, but causing the minimal environmental load-
ing. possible approaches for that are the improvement of digestibility (by
processing or germination), decrease of endogenous losses (by biotechno-
logical methods), improvement of feed utilisation (by adding synthetic amino
acids and/or enzymes, introduction of phase feeding and last, but not least,
using an accurate feed evaluation system.
Decreasing the amount of ingested crude protein causes decreased
excretion. Thus, by using less, but high quality protein one of the possible
solutions (see ideal protein principle). Table XX-23 shows examples for per-
formance of growing-finishing pigs fed on corn or wheat with different syn-
thetic amino acid (and mineral-vitamin) supplementation.
497
Table XX-23: Effect of cereal and synthetic amino acid feeding on growing pig’s perform-
ance
Phosphorus level of feed mixture may be reduced by calculating the avail-

able P-content, or by adding microbial phytase (Table XX-24).
Table XX-24: Effect of reduced P and phytase supplementation on fattening performance
20.4.10. Feeding growing-finishing pig for quality

The whole growing-finishing operation should be directed to the production
of QYALITY MEAT. Thus, the pacemaker of production is the marked, guided
by quality and food safety. Notion of quality is not clearly defined. Nowadays
the lean meat ratio is in the centre, together with the food safety and its
quality control systems (HACCP). It should underline that food safety does
not include quality.
Quality parameters of pigment are technological (water binding/hold-
ing capacity, pH. state of protein (denatured or not), characteristics of fat
content (saturated and unsaturated fatty acids and their ratio), antioxidants
ands vitamin E concentration, cutting quality, hygienic status (presence of
pathogenic microbes or other harmful materials), antibiotic and drug
residues, heavy metal and pesticide contaminations, ethical (bio or organic
keeping, other considerations), nutritive value for humans (protein quanti-
ty and quality, fat content and composition, minerals, vitamins, digestibili-
ty) and overall acceptability or eating quality (appearance, presence of blood
splash and speckle, colour, tenderness, juiciness, marbling, taste, odour).
By definition of WOOD (1995): “The most important aspect of meat
quality is eating quality, that is, the level of overall eating satisfac-
498
tion, which is a function of the combined effect of tenderness, juici-

ness, and flavour. Less crucial but still important aspects of meat
quality are the appearance of meat when purchased and its handling
characteristics as perceived by consumers and butchers.”
It is worth mentioning that only the quantity of protein can be influ-
enced by feeding, its composition never. In case of fat, both its quantity and
chemical composition can be influenced by feeding. Table XX-25 and XX-26
highlight that the effect of the same feed mixture may differ according to the
pig genotype, too.
Table XX-25: Chemical composition of ham, chop and backfat, %
Remark: intensive feeding according to the Hungarian Feed Codex (2004);

Traditional „Mangalitza” feeding
HLW×HL: Hungarian Large White×Hungarian Large, M: Mangalitza
Nevertheless, data of Table XX-26 clearly shows that fatty acid intake
has a significantly stronger influence on fatty acid composition a the ham,
chop and backfat than genotype. It means that by changing the feed com-
position, the composition/quality of the end product can really be improved.
Table XX-26: Fatty acid composition of ham, chop and backfat, %
Remark: ether extract/linoleic acid/linolenic, %

Intensive feeding: 5.3/3.0/0.3; Traditional feeding: 2.2/1.0/–
As a trend in the near future, the production of high, declared vita-

min E containing pig meat; lard and intramuscular fat of optimized fatty
acid composition can be expected on the functional food market.
20.4.11. Consequences of nutritional failures

First of all, distinction should be made, whether the loss of appetite is a sign
of a disease, or “simply” a feeding failure. Causes of feeding failures are:
Inaccurate amount of daily ration (human error or technical imperfection).
quality of feed (rancidity, deterioration, mouldy etc.) or technical (labourer,
technological.
499
The most frequent “symptoms” are as follows.

Feed refusal is generally caused by high germ count or presence of
mycotoxin. Other typical clinical signs are diarrhoea, erythema, eczema and
possibly death. Average daily gain (by 50-100%) and feed conversion effi-
ciency (by 20-80%) may drastically decrease.
Concerning mineral and vitamin supply, both deficiency and over-
dosage (especially in case of vitamin A and D) may be harmful, decreasing
health status and performance.
Insufficient drinking water will decrease dry matter intake, and may
lead to salt poisoning. Too much water (liquid ingestion) decrease voluntary
feed intake.
The occurrence of oesophageal gastric ulcer may be facilitated by too
small a particle size and stress conditions.
Assuming a common management system and the feeding of isoener-
getic diet, a decrease in protein level by approximately by 2 percent units
(e.g. from 16.8 to 14.8 or 15.0 to13.0%) , and/or decrease of lysine concen-
tration by approximately 0.2 percent units (e. g. from 0.89 to 0.69 or 0.78 to
0.58%) will cause a :
3–15% reduction in feed intake,
10–15% reduction in average daily gain,
10–15% decrease in feed utilisation and
15–30% drop in meat quality grading.
The cited data draw attention to the biological and economic significance of
a balanced and high quality feed formulation.
500
20.5. Feeding and nutrition of breeding sow and boar

Profitability of breeding sow can be characterised by values of reproductive

traits (litter size and weight, weaned number and weight) and the longevity
(few early culling). Table XX-27 summarises statistical guide numbers of
successful sow keeping.
Table XX-27 Statistical guide numbers of sow performance

————————————————————————————————————————————
Parameter Optimum Tolerance level
————————————————————————————————————————————
Farrowing/sow/year
- Day 28 weaning 2.4 2.2
- Day 42 weaning 2.2 2.0
Litter size (total) 11 10
Stillbirth, % 5 10
Slaughtered pig/sow/year
- Day 28 weaning 22 20
- Day 42 weaning 20 18
Interval between weaning and
First insemination, day 6-9 12
Conceived first, % 80-90 80
Barren*, % 2 3
Feed consumption, t) sow/year 1 1.1
————————————————————————————————————————————
*She has not conceived two times
The number of so-called „open” sow days commonly 40-50 day, and
the yearly culling may achieve 100%. This is uneconomical, because the
piglet number weaned per breeding sows decrease owing to the smaller lit-
ter size and milk production of gilts. To achieve optimum numbers, both
genetic and nutritional changes are necessary, because the heritability
value (h2) of the most important economic parameters is low, e.g. live litter
size at birth: 0.05, litter size on day 21: 0.10, litter weight on day 21 (N21):
0.20. The latter is used as a selection index for improving sows productivi-
ty.
20.5.1. Feeding of growing–replacement gilts and boars

Goals of raising the future breeding animals is a physiologically based early
mating or insemination, avoidance of early culling and achieving longevity.
In this way cost of raising is shared by more slaughtered pig. Young breed-
ing animals should be raised with a medium intense feeding and strength-
ening keeping, because breeding gilts, coming from the growing-fattening
herd have lower conception rate and longer weaning to re-conception inter-
501
CHAPTER XX: BREEDING SWINE FEEDING & NUTRITION
val. In overfed pregnant gilt embryonic mortality increases from the average
20-22% to 28-30%. Fat boars are culled earlier as normally owing to feet
problems.
20.5.1.1. Growing–developing breeding gilts

The onset of puberty (the first fertile ovulation) generally depends upon
three factors: genotype, management and nutrition. The beginning of puber-
ty basically is controlled by body weight gain of animals. Generally the first
oestrus occurs on day 200 (within 135 and 250 days). The use of different
stimulations (mixing of different gilt groups, presence of adult boar in the
stall, the use of so-called boar spray) may accelerate sexual maturation by
30-40 days, but this does not mean that animals are really suitable for preg-
nancy (they did not achieve breeding maturity). It is advisable to use for
mating/insemination only the third oestrus, because during the first and
second oestrus the number of released eggs is lower. Today’s modern geno-
types have a high growth rate but a relatively late sexual maturity. Thus,
they reach breeding maturity (when they are ready for insemination) at the
age of 8-9 months with a live weight of approximately 120 kg. From this
time, the earliest time for them to be fertilised is at their third oestrus.
FIGURE XX-10: Effect of nutrient supply on the development of bone, muscle and fat tis-
sue
Keeping and feeding should serve a medium intensive growth. As FIG-

URE XX-10 illustrates in case of ad libitum feeding fat deposition begins ear-
lier. In contrast, poor feeding – especially between 20.60 kg of LW - delays
sexual maturation. An ideal solution is a medium intensive raising, com-
bined with grazing, or if not possible, with walking course/field. In the
absence of these facilities (there is only a runway), animals cannot get rid of
their extra weight and condition control by feeding should receive extra
attention. Principles of this are as follows:
502
From weaning until achieving the live weight of 30-35 kg (approxi-

mately 90 days of age), future breeding animal are kept together with the
growing-finishing pigs, and the receive only a medium intensive feeding.
The remaining 85-90 kg weight gain is characterised by medium
intensity, in order to make the use of flushing possible before insemination.
Although there is a general agreement in scientific literature concerning
principles, the picture is rather varied concerning factual data of recom-
mendation. Recent data of DLG (1987) justified by practice are given (con-
verted into digestible energy). This recommendation divides the raising peri-
od both for gilts and boars into three periods: the first has moderate, the
second medium intensive and the third a restrictive feeding (Table XX-28).
Table XX-28: Raising periods of replacement gilts and boars

————————————————————————————————————————————
Parameter L i v e w e i g h t c a t e g o r i e s, k g
30—60 60—90 90-120
————————————————————————————————————————————
Average daily gain, g 600/700 700/850 500/750
DE needs, MJ/day 20.4/22.5 28.9/29.0
32.2/33.3
CP need, g/day, 280/320 330/420 280/430
Lysine, % of CP 5/5.5 5/5.5 5/5.5
Duration, day 50/43 43/35 60/40
————————————————————————————————————————————
Remark: numbers after fraction stroke relate to growing breeding boars.
The first and the second phase serve medium intensive growth, the
third one assure the success of the ensuing flushing. In case of gilts of (very)
lean genotypes – to make depot building for longevity possible – ad libitum
feeding is recommended during the whole raising period.
For the sake of logical unity, in the followings the preparation of gilts
and sows to mating (or insemination) is treated together. Essence of FLUSH-
ING OR STIMULATING FEEDING IS TO IMPROVE THE CONDITION OF THE ANIMAL. The orig-
inal meaning of the word reflects that the organism is „flushed” by energy.
Flushing of intact gilts means doubling the daily ration (from 1.8 to 3.6 kg)
2 weeks before the expected oestrus and mating. This measure increases
ovulation rate and embryo survival. Weaned sows generally have their first
heat 6-9 days after weaning, therefore there is only approximately one week
for flushing. Thus, the extent of the positive effect (higher litter size) is mod-
erate. Opposite to some previous practice, the elevated portion of flushing
should be decreased on day 2-3 after insemination (for details see sow
nutrition).
20.5.1.2. Raising of growing breeding boars

should also be of medium intensive and natural. Owing to the higher growth
rate of males factual gain data are greater: target body weights are at the
age of 7 months (placing, selling): 120 kg of LW and at the age of 9 months
503
(first service) 150 kg of LW. In Table XX-28 already shown data of growing
boars (after fraction stroke) differ from those of gilts being 700, 850 and 750
average daily gains in the three raising periods, respectively.
Table XX-29 compares recommendation for future breeding animals
and for the corresponding growing-fattening pigs expressing values as a
percentage of the latter.
Table XX-29 Relative nutrient requirements of growing gilt compared to the growing-fin-
ishing pigs value (latter taken as 100%)
————————————————————————————————————————————
Parameter Live weight cate gories, kg
——————————————————-
30—60 60—120
————————————————————————————————————————————
DE, rel. % 96 93
Crude protein, rel. % 98 100
Lysine, rel. % 98 96
Crude fibre, rel. % 125 125
Calcium, rel. % 106 110
Phosphorus, rel. % 108 110
————————————————————————————————————————————
From Table XX-29 the fundamental differences can be seen. Feed mix-
tures for future breeding gilt should contain less energy and more fibre than
those of growing-finishing pigs. The fibre level can go up to 6% by mixing
oats, whole corn plant, hay and alfalfa meal or dried sugar beet pulp.
Supposing an average protein supply (13-16% CP of at least medium
digestibility and biological value), it has no influence on sexual maturation
and ovulation rate. To assure a solid skeletal system, future breeding ani-
mals should receive higher calcium and phosphorus supplementation than
the growing-finishing pigs.
In practice, besides complete feed mixtures it is advisable to feed
grass (by grazing), green alfalfa, gardening by-products, fodder beet, carrot
and high quality silage. Mouldy feed should be avoided, because their myco-
toxin effects (especially F–2 and T–2 toxins) in reproduction and immune
function may last even after finishing feeding of the damaged lot.
Growing breeding boars can be fed on gilt’s diet, using higher daily
portions. Before mating/insemination for both sexes the use of lactation
sow diet is widespread.
20.5.2. Nutrient requirement of gestation and lactation

20.5.2.1. Energy needs of pregnant sow
Increased flushing portion of inseminated sow should be maintained only
till the end of oestrus. Previously it was proposed to keep this level for 2
weeks; nowadays it is known that this too high nutrient supply may affect
uterine fluid composition and thereby negatively influence nidation (implan-
tation) of fertilised eggs. Thus, from the day 2 of gestation till day 84-90
504
requirements of pregnancy is very low and the distributed daily feed amount
is the same continuously. The required daily energy is determined accord-
ing the following consideration:
a) too low energy intake (below 1.7 kg of feed mixture) decreases
subsequent litter size and birth weight of newborn piglets;
b) energy overfeeding negatively affect newborn piglet viability,
farrowing will be long-lasting and more piglets die owing dam’s squashing.
Incidence of MMA (metritis mastitis agalactia) syndrome increases and
sow’s appetite and consequently its milk production are depressed.
The principle is that both emaciation and overfeeding of sows
should be avoided. NRC (1998) recommendations for maintenance is
0.477 MJ DE × W0.75 and 5.4 MJ DE/day for pregnancy needs (from that
foetal growth represents 0.79 MJ DE). Values are valid in thermo neutral
zone. Generally each oC below lower critical temperature (19oC) increases
maintenance energy requirements by 4% (~20 kJ DE/W0.75 =1.3-1-5 g of
feed). For more accurate calculation it should be taken into consideration
that lower critical temperature in individual keeping is 19oC, but in group
keeping only 14oC and the average heat equivalent of -1 oC is 0.6 and 0.3
MJ DE.
In the last 3-4 weeks of gestation pregnancy needs are important
(FIGURE XX-11); therefore energy supply should be increased by 10-20%.
If daily ration remained unchanged the litter size and weight will not be
affected, but the accumulation of body reserves fails to take place and the
subsequent lactation performance and litter weight at weaning decrease.
The same phenomenon could be seen while feeding exclusively pregnant
sows on alfalfa pellets.
FIGURE XX-11: Development of uterus, foetuses and foetal membranes in pregnant sow
505
Inseminated gilt of 120 kg of LW reaches 170-200 kg of LW at the end

of gestation. From the total gain of 40-50 kg is the sow’s own body weight
gain which means further energy needs.
20.5.2.2. Protein requirements of pregnant sows

Deficient protein supply leads to low viability of newborn piglets, their mor-
tality increases till weaning and more sows should be culled. In case of preg-
nant gilt development of own body tissue may also be affected.
The average daily crude protein requirements are 228 grams (NRC,
1998). Adjusted more accurately to different stages with some security mar-
gin (ARC, 1981) 250 grams for the first 77 days of pregnancy; between 84-
98 days 325 grams and from 98 to 114 days 400 gram crude protein per
sow per day are recommended. Lysine requirement is 4.5% of crude protein,
but at least 8.2-8.6 g daily. NRC (1998) proposes similar lysine supply, but
the amino acid pattern of two recommendations is different. If the amount
of lysine is regarded as 100%, the ideal amount of methionine+cystine 67
vs. 53%, from tryptophan 16 vs. 21% and from threonine 84 vs. 70% (ARC
and NRC data, respectively. Arginine is not considered as indispensable for
pregnant sows, because they are able to synthesise the necessary quantity.
The relatively small energy and protein requirement needs more
explanation, because development of gestational products is important
(FIGURE XX-11): weight of foetal membranes at birth is 2.5-3 kg; amniotic
fluid is 6-8 litres and the uterus multiplied 4-6 fold. In addition, the dam’s
own body gain is 20-50 kg. The sow’s gain comprises 25% fat and 15% pro-
tein and their main sites of accumulation are the liver, skin, mammary
gland parenchyma and muscles („pregnancy super retention”). In the back-
ground of this pregnancy anabolism is a changed hormonal balance.
According the recent data protein accretion is limited to the uterus and in
the whole body the water content increases while that of fat falls.
20.5.2.3. Mineral and vitamin requirements of pregnant sows

Phenomenon of super retention can be found in case of minerals too.
Deficient CALCIUM and PHOSPHORUS supply rather decrease the amount of
bone ash and causing lameness than reproductive failures. Piglets’ birth
weight decreases if the sows were fed on a diet deficient in salt (less than
0.4% NaCl). Zinc deficiency of pregnant sows decrease not only birth weight
but also litter size. Signs are very similar in case of MANGANESE deficiency,
because Mn is indispensable in cholesterol synthesis, which, in turn, essen-
tial for sexual steroids building. If IODINE supply is not sufficient during ges-
tation (or there is high amount of thyreostatics in feed) congenital goitre
may appear on newborn piglets. Even a slight iodine deficiency (or presence
of antagonist nitrate, arsenic or rubidium in feed or drinking water) elon-
gates gestation time and leads to the farrowing of weak and hairless piglets.
The addition of 0.17 mg/kg feed iodine (in form of potassium chloride) helps
506
to prevent the condition. Intrauterine selenium and vitamin E supply of foe-

tuses influence newborn piglet’s immune state. There is also a daily require-
ment for linoleic acid (1.9 g/sow).
VITAMIN A (carotene) deficient pregnant sows bore microphtalmic, dead
piglets. Slight carotene deficiency reflects low carotene and vitamin A con-
centration in sow milk. Despite the fact that the main site of
caroteneÞvitamin transformation is the gut wall; there are data for the inde-
pendent role of carotene in sow reproduction. The exact vitamin D require-
ments of pregnant sows are not known; as minimum supply NRC (1998)
recommended 4000 IU/kg air-dry feed.
Intake of higher amounts of choline, biotin and folic acid increase lit-
ter size at birth and prevent feet lesions. In contrast, doubling of common-
ly used micro mineral addition has no similar positive effect.
20.5.2.4. Energy and protein requirements of lactating sows

a) Energy. Maintenance requirements of lactating sows equals to .477 MJ
DE × W0.75. For producing sow milk of average composition (7-9% fat, 5-6%
protein, 5-6% lactose and 5.0-5.4 MJ/kg) animals need 8.4 MJ DE.
Efficiency of ME→NE transformation is 65-75%. The average daily milk pro-
duction varies from 5 to 8 kg reaching its peak on week 3-4. Mobilisation of
body tissue contributes to milk production; 1 kg weight loss is equivalent to
300 grams fat and 130 grams protein and may meet the needs of of 3-3.1
kg sow milk produced. The average daily energy requirements of lactating
sow is 74 MJ DE, being 2.8-3.0 times higher than those of gestation.
During the first 28 days there is a linear correlation between milk
(energy) intake and growth. To assure 46 g weight gain, piglet should suck
the amount of milk containing 1 MJ energy, i. e. 1 gram weight gain requires
22 kJ milk energy. Utilisation of feed for milk production varies according
to authors from 65 to 75%. Therefore, energy requirements of milk produc-
tion can be given by the following equation:
ME, MJ= Nr.×ΔW×0.022/0.65 or 0.75,
where Nr.= number of suckling piglets; ΔW= average daily gain of
piglet, gram and 0.65 or 0.75, according to different author the partial effi-
ciency coefficient. This is the total energy needs of litter, which can be
reduced by the ingested creep feed.
b) Protein. net protein requirements of maintenance is 0.45 g/kg LW,
being the transformation efficiency 70%, the daily requirements is 0.64
gram digestible crude protein per kg live weight. It is worth mentioning that
contrary to energy, protein requirement is proportional to live weight and
not to metabolic body size. For the production of one litre average sow milk
(5.7% protein) animals need 81.4 grams digestible crude protein.
The average daily milk requirements – varying according to live weight
and milk production – amounts to 605-858 grams crude protein. Protein
digestibility is supposed to be 80%. Body mobilisation also contributes to
507
milk production, but to a lesser extent than in case of energy. During the
whole lactation period a 15%-decrease in live weight may be considered as
physiological. Five percent of crude protein should be lysine. Propositions of
the other most indispensable amino acids, compared to lysine as 100% are
as follows: methionine+cystine 55-60%, tryptophan: 19-20% and threonine
70-72%. Arginine became essential for lactating sows.
Assuming an average composition (57 g protein and 5.2 MJ energy) of
sow milk, on the basis of piglets weight gain and their energy intake by milk
(1 gram of weight gain is equivalent to 22 kJ, i. e. to 4 ml of sow milk) the
protein output of the lactating sow can be calculated. It means that
67/5.2×0.022=0.24 gram protein (=4.2 ml sow milk) is equivalent to 1 gram
piglet weight gain. Total protein requirement can be reduced by creep feed
consumed by piglets.
c) Mineral and vitamin requirements of lactating sows
Calcium and phosphorous balance of lactation is negative, because
their excretion by milk exceeds the ingested quantity. Heavy calcium defi-
ciency (0.6 g/day) may affect milk production. In case of phosphorous sup-
plementation availability should be considered, because P in cereals and
oilseeds has a lower availability than in animal or inorganic sources.
Sodium level in sow milk averages 0-03-0.04%, which can be assured by
formulating lactation feed with 0.5% salt level. Potassium concentration in
sow milk amounts to 1 g/kg; knowing the average availability of this ele-
ment (45%), the actual needs can be calculated. There are no sufficient data
concerning MAGNESIUM requirements. SULPHUR cannot be utilised only as part
of organic compounds, like methionine and cystine. Exact ZINC needs of lac-
tation sow are unknown; 33 mg/kg feed proved to be insufficient. Calcium
overfeeding inhibits zinc absorption.
Six mg/kg feed COPPER need has been evaluated. Iodine requirements
are not exactly known, although 20-45% of the ingested iodine excretes by
milk. The maximum recommendable IRON level in lactating sow diet is 0.4
mg/kg, because iron alters P utilisation. In the lack of manganese the milk
production drops, the recommended concentration is 25 mg/kg feed. For
optimum milk production and immune function requires SELENIUM, concen-
tration of 0.1-0.3 mg/kg should be assured. Part of feed selenium is excret-
ed by milk, improving supply of piglets. Fluorine is essential for sows, but
exact recommendations are not available.
Owing to a high storage capacity, the real vitamin A requirement is
difficult to evaluate. Dosage of 7000 IU vitamin A per day and per sow
helped in maintaining blood and liver concentrations, although the more
sensitive parameters (pressure of cerebrospinal fluid and “Relative dose
response” test) were not measured. NRC (1998) recommended minimum
requirement is 2000 IU/kg feed. High nitrite or nitrate concentration in
drinking water or in feed increase vitamin A requirement. There is a low
effective carotenecvitamin A transformation, while 11 molecules of ß-
508
carotene equivalent to 1 molecule retinol. VITAMIN D supplementation –

through milk – can improve piglets supply. In contrast to the daily vitamin
E supplementation for pregnant sows suckling sows requires at least 5.3
grams per day per animal. Deficiency of vitamin E (and/or selenium) makes
animals susceptible to MMA-syndrome. Over dosage of vitamin A affect vita-
min E absorption, but in turn, presence of vitamin E is essential for retinol
utilisation. If feed mixture of lactating sows contains unsaturated (possibly
rancid) fats, vitamin E requirement may double. by vitamin E supplemen-
tation of feed milk tocopherol level may be elevated. Exact data of sow ESSEN-
TIAL FATTY ACID requirement fail, but role of these compounds in
prostaglandin synthesis suggests that an appropriate supply is indispensa-
ble. As minimum requirement (NRC, 1998) recommends 0.1% linoleic acid
in air-dry feed. Optimum recommendation is equivalent to that of growing
pig, i.e. 3% of DE should be given by linoleic acid (this is approximately 15
mg/kg dry matter).
Based on the activity of FAD-dependent gluthation-reductase activity
of red blood cells (FAD= flavin adenin-dinucleotid) daily RIBOFLAVIN require-
ment of lactating sows is 16 mg. The exact THIAMINE need is not known, but
may be similar to that of growing. The influence of extra pyridoxine addition
on sow’s milk production is disputable, but 15 µg/kg feed B12 improves it.
In turn, cyanocobalamine being bound to plasma proteins of sow milk, may
supply suckling piglets too. Abundant BIOTIN supply improves the state of
skin, hoof firmness, milk production and stimulate onset of oestrus after
weaning. PANTHOTENIC ACID requirement is estimated to be 10 mg/kg dry
matter. Folic acid is present in sow milk, bound to plasma proteins, but the
exact need is not known. NIACIN needs of pregnant and lactating sows is not
known. Although 1 g/sow daily ascorbic acid supplementation increased
21-day litter weight, the vitamin C addition is recommended in hot envi-
ronment. In hot environment sows voluntary feed intake decreases, there-
fore, generally, mineral and vitamin levels in dry matter should be elevated
(“fortified diet”).
20.5.3. Practical feeding of breeding sows

Practical realisation of breeding sows’ nutrition is illustrated in FIGURE XX-
12.
Recommended nutrients concentrations are given according to the
Hungarian (1990) and US allowances (Table XX-30).
509
FIGURE XX-12: Scheme of practical sow nutrition
Table XX-30 Minimum nutrients requirement in pregnant and gestant sows’ feed mixtures
(HFC=Hungarian Feed Codex, 1990; NRC=National Research Council, 1998)
———————————————————————————————————————————
For pregnant sows For lactating sows
—————————————————————————————
Items HFC-I. NRC-I HFC-II.NRC-II HFC NRC
————————————————————————————————————————————
Dry matter, % 86.0 90 86.0 90 86.0 90
DE, MJ/kg 12.6 14.2 12.4 14.2 13.5 14.2
Crude protein, % 14.3 12.9 12.4 12.4 16.1 16.3
Lysine, g/kg 6.7 5.8 5.5 5.4 8.1 8.2
Met+Cys, g/kg 4.4 3.7 3.6 3.7 5.3 4.0
Tryptophan, g/kg — 1.1 1.1 1.5
Threonine, g/kg — 4.4 4.5 5.4
Ether extract, % 3.00 3.00 3.00
Crude fibre, % 8.00 9.00 5.00
Calcium, g/kg 10.2 0.75 9.4 7.5 10.9 7.5
Phosphorus, g/kg 7.0 0.60 6.5 6.0 7.5 6.0
aP*, g/kg 0.35 3.5 3.5
Mg, g/kg 0.4 0.4 0.4 0.4 0.4 0.4
Na, g/kg 2.0 1.5 2.0 1.5 0.25 2.0
Vitamin A, IU/kg 6000 4000 6000 4000 5000 2000
Vitamin D3, IU/kg 600 200 600 200 500 200
Vitamin E, mg/kg 20 44 20 44 20 44
Vitamin B12, μg/kg 15 15 15 15 15 15
Niacin, mg/kg 15 10* 15 10* 15 10*
Fe, mg/kg 60 80 60 80 60 80
Zn, mg/kg 70 50 50 50 50 50
Mn, mg/kg 30 20 30 20 30 20
Cu, mg/kg 10 5 10 5 10 5
510
Table XX-30: cont.

———————-————————————-————————————-————————————
I, mg/kg 0.4 0.14 0.4 0.14 0.4 0.14
Co, mg/kg 0.1 — 0.1 — 0.10 —
Se, mg/kg 0.15 0.15 0.15 0.15 0.15 0.15
————————————————————————————————————————————
* available
Remark: HFV- I. and NRC-I stand for pregnant gilts of 125 kg of live weight, HFC-II and
NRC-II stand for pregnant sows of 200 kg of live weight. Predicted lactational weight loss-
es amount 25 kg.
WEANING
In case of weaning on day 42 (or more) over a few days daily ration should
be decreased gradually to maintenance level; on the day of weaning total
fasting, even drinking water deprivation follows. The latter may be replaced
by offering lukewarm water. In case of weaning on day 28 or later 2 days
before weaning the daily ration should be decreased significantly.
PREPARATION FOR MATING AND THE FIRST TWO DAYS OF GESTATION

The actual amount of daily ration depends upon the sow’s condition. In case
of important lactation weight losses, in order to assure flushing effect,
maintenance requirements may be doubled, or feeding ad libitum a feed
mixture of P/E ratio of 12 can be applied. In practice use of lactation feed
mixture (13.5 MJ DE, 16% crude protein) is given in an amount of 3-4 kg
or ad libitum. In ideal cases 6-9 days after weaning sows show fertile
oestrus. If lactational weight losses were minimal (e.g. at sows nourishing
small number of piglets), increase of daily portion will not result in the
described positive reproductive effect.
FROM DAY 2-3 TILL DAY 84-90 OF GESTATION

On day 1-2 after mating the increased flushing daily amount should be
decreased to 2.0-2.2 kg of feed mixture, which is equivalent to a slightly
increased maintenance requirement (+15%). In practice, gestational feed
mixture (12.5 MJ DE/kg feed, 12.14% crude protein) is distributed. A rela-
tively small amount is sufficient, because the growth of foetus and other
gestational products is small and the sow’s feed utilisation is improved.
Described daily amounts are only applicable for sows of good breed-
ing condition, in case of any deviation, corrections are necessary. Condition
scoring may be carried out by the visual and ultrasound Cornell system
(FIGURE XX-13; Table XX-31).
511
FIGURE XX-13: Visual condition scoring of breeding sows
Table XX-31: Breeding sows condition scoring by ultrasound measured of backfat tissu
at the rib 10
————————————————————————————————————————————
Category Condition- Backfat - Change in daily
point (BCS) thickness, amount,
cm kg/day
————————————————————————————————————————————
Emaciated 1 < 1.27 + 0.60-0.75
Thin 2 1.91 + 0.20-0.30
Good (=breeding) 3 2.54* ——-
Fat 4 3.30 - 0.20-0.30
Obese (“whale“) 5 3.81 - 0.60-0.75
————————————————————————————————————————————
*According to the recent data no more than 2.0 cn
Recommended time for condition scoring is the weaning (to get infor-
mation about possible flushing), day 55-60 of gestation, when correction of
daily ration is not too late.
Ad libitum feeding of pregnant sows can be excluded, because they are
prone to becoming fat. Overweight sows are susceptible to MMA-syndrome,
their farrowing, may be delayed and owing to a subsequent decreased
appetite, their lactational performance decreases. Occurrence of MMA-syn-
drome basically depends upon pregnancy feeding; lactational measures
cannot correct sow’s state. From this respect appropriate fibre supply of
pregnant sows has a key role to play. Fibre supplementation can be assured
by mixing green meals (alfalfa, grass, and whole corn plan), dried sugar beet
pulp and possibly straw meal. Control of daily feed intake can be accom-
plished by individual feeding, by the limitation of feed access (while in group
keeping sows of good condition are allowed to through later), by increasing
ballast levels in feed mixture and possibly by using chemical feed refusal
agent.
LAST MONTH OF GESTATION

Foetal and gestational products growth being important, energy require-
512
ment is increased by 15-20%. In practice, in the first half of this period ges-
tational and in the second one lactational feed mixture is given, in an
amount of 2.4-2.6 kg/day. During the last week before the expected far-
rowing daily feed portion should be gradually decreased and on the day of
parturition a small amount of diluted (1:5 with water) feed mixture or wheat
bran can be offered. This procedure facilitates gut emptying, which assures
an easy farrowing by a good presentation of piglets into the uterus and vagi-
na.
LACTATION
It can be calculated from data of the theoretical unit that daily need of a lac-
tating sow would be coved by 6-7 kg feed per day. This amount exceeds feed
intake capacity of sow, consequently weight losses of 15% can be consid-
ered as physiological. In practice, above maintenance of 400 grams lacta-
tional feed mixture is recommended for a sow, but the total mustn’t be more
than 5 kg. The highest quantity should be achieved gradually, by following
the lactational curve. It means that in case of early weaning (day 28-35) this
transition period lasts for 1 week, in case of late weaning (day 42 or more)
for 2 weeks. In the meantime, sow’s condition should be continuously mon-
itored and if the need arise, the amount of daily ration adjusted. Another
practical approach is 3 kg of basic portion for each lactating sow and ¼ kg
feed mixture per suckling piglets.
There are technologies proposing ad libitum feeding during the whole
lactation period, but its only advantage is an economy in labour. In a US
investigation body weight changes of lactating sows have been compared.
Individuals with a daily ration of 1 pound (454 gram) feed mixture per suck-
led piglets showed a physiological 15% of weight loss until weaning; ad libi-
tum sows conserved their farrowing weight, getting irresponsive to flushing
feeding.
It is worth highlighting that sows of very lean genotypes may have a
need for ad libitum feeding during lactation. This is to avoid excessive (more
than 15%) weight losses because poor (thin or emaciated, BCS of 2 and 1)
cannot make fertile oestrus after weaning possible.
OTHER INITIATIVES TO IMPROVE REPRODUCTIVE PERFORMANCE OF SOWS

Growth hormone (GH) treatment of suckling sows may improve milk pro-
duction, but subsequent reproductive performance generally becomes poor-
er. In Europe this practice is prohibited.
Fat supplementation of late gestational and lactational feed mixture
increases litter size and survival rate of newborn piglets. In a Hungarian
field study (FEKETE, S., MAKAI and ZIMMER, 1989) 2.5% mixed animal fat (on
expanded corn carrier) has been mixed 3 weeks before farrowing and dur-
ing the whole 4-week lactation period into sows feed mixture. The treatment
increased litter size at birth by 1.12 and by 0.42 at weaning, besides an
513
unchanged litter weight, mortality and sows feed intake. In sows blood total
lipid concentration elevated by 18%. Supplementation of pregnant and ges-
tant sows diets with probiotics (Saccharomyces cerevisiae, Aspergillus
oryzae) decreased suckling piglets mortality, but only in bad environmental
and hygienic circumstances.
20.5.4. Diet formulation for sows

Table XX-32 summarises some typical feed composition for pregnant and
lactating sow. Corn can be replaced by sorghum or milo, but in those cases
extra linoleic acid supplementation is required (e.g. poultry fat).
Table XX-32 Natural ingredients and nutrient concentrations of pregnant and lactating
sows
————————————————————————————————————————————
Ingredients, % /Nutrient Pregnant Lactating
I. II. I. II.
————————————————————————————————————————————
Corn 47.5 - 44.5 -
Barley - 64.0 - 53.4
Wheat 30.0 22.0 30.0 22.0
Wheat bran 5.0 - 5.0 -
Alfalfa meal 10.0 - 10.0 -
Soybean meal (47% CP) 3.5 10.0 4.5 14.6
Fat/Oil (50% CP)/ - - - 6.0
Supplements 4.0 4.0* 4.0 4.0*
------------------------------------------------—-------—-
DE, MJ/kg 12.1 13.4 12.1 14.3
Crude protein, % 12.9 14.8 14.2 16.1
Lysine, g/kg 4.8 5.9 5.3 6.8
Met+Cys, g/kg 4.2 5.0 4.6 .5.2
Calcium, g/kg 9.0 7.85 9.2 7.85
Phosphorus, g/kg 6.0 .6.0 6.0 .5.9
————————————————————————————————————————————
* 0.5% NaCl, 1.5% DCP (dicalcium-phosphate, 1% limestone and 1% mineral-vitamin pre-
mix
To avoid periparturient constipation the addition of 0.7% MgSO4 or

KCl may be used. Mixing ammonium chloride into the feed in the last three
weeks of gestation, caused the pH and total germ count of sows urine to
decrease. These procedures are effective in preventing the occurrence of
MMA-syndrome, but they do not influence litter size and weight at weaning.
Nevertheless, the key factor of prevention remains the optimal fibre supply.
According to NRC (1998) the future trends in case of breeding sows
include the use of ileal digestible amino acids and the consideration of phos-
phorus and niacin availability in course of practical diet formulation.
Recommended nutrient levels for pregnant sows in feed mixture are 14.2
MJ DE/kg, 5.0 g/kg true ileal digestible lysine, 3.3 g/kg methionine+cys-
tine, 3.5 g/kg available phosphorus and 10 mg/kg available niacin. The cor-
responding data for lactating sows are 14.2; 7.1; 3.5; 3.5 and 10.
514
20.5.5. Feeding of breeding boars
20.5.5.1. Development of breeding boars

In this special field of pig nutrition there is a lack of data, thus generally fig-
ures for female or castrated animals are used. This practice is not scientif-
ically founded, because boars have higher growth rate and their body com-
position is also different (more protein, less water) than that of gilts of sim-
ilar age and weight. Concerning details of development, herby only reference
is made to the unit about raising breeding animals. Nevertheless, it should
be emphasised that the body of growing boars contains more bone than gilts
do, consequently their calcium and phosphorus requirements are higher
(Table XX-33.).
20.5.5.2. Nutrient requirement of adult boar

Energy requirement of resting boar are higher by 15% than that of mainte-
nance. During mating period this may increase up to 30%. In the literature
the significance of appropriate protein supply is also emphasised, because
surplus protein stimulates whole metabolism included testes. In authors
opinion this may be due to the specific dynamic action of absorbed amino
acids. At the same time, protein overfeeding may drastically increase blood
urea level. In the field of protein supply efforts should be made to ensure
quality (taste, rancidity, free of heavy metals and antinutritive substances
etc.) and that a certain part of the protein should derive from animal
sources.
The above described can be illustrated by a trial when feeding a sow
diet (13.08 MJ DE, 14.5% crude protein, 6.8 g/kg lysine and 4.4 g/kg
methionine+cystine) was sufficient for boars in intensive mating (3-4 serv-
ices per week). Sperm quantity and quality have not been improved by
increasing protein intake. Addition of vitamin C and E (with or without sele-
nium) may improve sperm preservation.
According the DLG (1987) recommendation boar of 7-mont-old and
120 kg of body weight can be used for mating, but until reaching 180 kg of
live weight are extra nutrients required for the development of own body tis-
sues. Optimum live weight of adult breeding boars does not exceed 220 kg
of live weight.
Net energy and protein needs of sperm production are not known,
thus recommendations of Table XX-33 are based on empirical data. In case
of intensive use in order to ensure optimum sperm production, daily lysine
and sulphur-containing amino acid intake should be increased by 50%.
Sperm production has an outstanding zinc requirement. This is the reason
why allowances for boars are higher than those for castrated animals with
sows being in between.
515
20.5.5.3. Practical nutrition of breeding boars

Table XX-33 summarized recommended nutrients concentrations of boar
feed mixture (NRC, 1998).
Table XX-33: Recommended minimum nutrients concentration for developing replacement

boars and for active breeding boars of 150-300 kg of live weight
———————————————————————————————————————————
Items 50-80 kg LW 80-120 kg LW Adult, working
————————————————————————————————————————————
Dry matter, % 90 90 90
DE, MJ 14.2 14.2 14.2
Crude protein, % 14.2 12.2 13.0
Lysine, g/kg 6.7 5.3 6.0
Met+Cys, g/kg 3.8 3.1 4.2
Tryptophan, g/kg 12 10 12
Threonine, g/kg 4.4 3.6 5.0
Calcium, g/kg 6.0 6.0 7.5
Phosphorous, g/kg 5.0 5.0 6.0
available P, g/kg 2.3 2.3 3.5
Na, g/kg 1.0 1.0 1.5
Vitamin A, IU/kg 1300 1300 4000
Vitamin D3, IU/kg 150 150 200
Vitamin E, mg/kg 11 11 44
Vitamin B12, μg/kg 10 10 15
*Niacin, mg/kg 10 10 10
Zn, mg/kg 60 60 50
Fe, mg/kg 60 60 80
Mn, mg/kg 2 2 20
Cu, mg/kg 4 4 5
I, mg/kg 0.14 0.14 0.14
Se, mg/kg 0.15 0.15 0.15
————————————————————————————————————————————
* available
Average daily portion is 1.0 kg/100 kg live weight. Where there is no

way to apply a separate boar feed mixture lactating sow feed may be given
possibly with green feed or good quality silage supplementation. This prac-
tice is not accepted by several specialists, because in this case protein,
amino acids and vitamin E needs of boar remain uncovered. The actual
amount of daily ration should be adjusted to the body condition of boars,
because both poor and over condition negatively affects sexual performance.
Accordingly, in practical boar feeding rationing should be followed.
20.5.6. Possible non-infectious causes of sows’ abortion

The direct effect of abortus is an abrupt increase in blood prostaglandin E1-
concentration and contractions of uteral musculature which lead to the pre-
mature rejection of the foetus. Although early embryonic mortality may also
have nutritional origin, hereby only abortion and early parturitions are dealt
with. If any teratogenic or embryotoxic agent kill the foetus, generally abor-
tion also occurs. Though modern feeding techniques normally ensure the
516
meeting all nutrient requirements including macro and micro elements and
vitamins, some possibility of failure always remains (e.g. ingredients×tech-
nology interaction) which may cause deficiency of one or other nutrients.
A possible approach of studying sow abortion may be the extrapola-
tion from other (included human) species, epidemiological studies, search-
ing for correlation between environmental and nutritional factors and abor-
tions and finally following physiological process and regulation of abortion,
trying to find possible causative factors. Latest methods have put prosta-
glandin metabolism in the centre.
20.5.6.1. Habitual abortions

Psychic abortion in humans, although its occurrence is not epidemic, may
be considered as a pathological model, whose components may show resem-
blance to the pathophysiology pf sow abortions. Already HIPPOCRATES noted
that certain women abort as a result of psychic anxiety. In swine herds devi-
ations in the environmental hygiene (e.g. NH3-loading) and social stress (see
classical works of CALHOUN with rats) can maintain a permanently high
ACTH level in blood. Transmitters of social stress are different pheromones;
and besides rodents swine is highly sensitive to their effect.
20.5.6.2. Nutrients, macro and microelements, vitamins

Protein-energy overfeeding rather cause nidation troubles and embryonic
death, especially in gilts after insemination (WEKERELE and SZOLLOSI, 1992).
Energy deficiency generally is compensated by dam’s organism, but in case
of goats it may cause habitual abortion (see Chapter XXVII). Primary or sec-
ondary (caused by high nitrite or nitrate in drinking water, thyreostatic
compounds in cruciferous) iodine deficiency may lead to abortion and myx-
oedema of the foetus. Both overfeeding and deficiency of vitamin A is ter-
atogenic, consequently may lead to abortion. Teratogenic effect of folic acid
(human) and biotin (mice) is also known.
20.5.6.3. Environmental influences

In humans it was observed that in the vicinity of a dump site where numer-
ous chemical wastes were deposited (carbon tetrachloride, trichloroethyl-
ene, benzene) women had a high rate of spontaneous abortions and it was
suspected that exposure to the wastes was involved. Female anaesthetic
doctors also have high abortion rates, which indicate the role of pollutants.
In pig production mostly the causative role of heavy metals (Pb, Cd, Hg),
mycotoxins, polychlorinated biphenyls (PBCs) and other pesticide are sus-
pected.
20.5.6.4. Immunological factors

Immunological processes related to pregnancy are extremely complex: the
foetus acting as graft in the uterus requires the presence of maternal block-
517
ing antibodies, which along with the anti-inflammatory effect of proges-

terone, prevents the rejection of foetal tissues which posses antigens that
are quite different from those of maternal tissues. This antibody is an
immunoglobulin which coats the foreign foetal antigens on the placenta.
How this antibody is produced and which conditions may lead to its defi-
ciency in sows is the task of future studies. A similar immunological factor
is the histocompatibility locus antigen (HLA) which is a critical factor. The
difference in maternal and foetal HLAs may account for the production of
blocking antibodies. It is not known how these immune responses are mod-
ulated or compromised by feed or water-borne pollutants including phyto-
toxins and mycotoxins.
The existence of an early pregnancy factor (EPF) with immuno-sup-
pressive properties appears in the serum of women 48 hours following fer-
tilization and returns to normal levels during the third trimester of preg-
nancy. The pregnancy factor is controlled by the ovary and the pituitary of
the mother and the foetus. Thus, any interference at the ovarial-pituitary
level may lead to abortion, because reduced EPF and immunosuppression
contributes to abortions. Blood group incompatibility between mother and
foetus may also lead to abortion or stillbirth at horse and swine.
Intensive farm performance demand and the associated high level of
protein and carbohydrate intake make the swine an ideal candidate for
allergic diseases. Allergic conditions are associated with the ingestion of
alpha-amylase inhibitors in barley and rye which are related to the symp-
toms of bakers asthma, which is triggered by the inhalation of the meal
dust. A similar allergenic protein was found in rice, too. These inhibitors
have IgE binding capacity in vitro. A second category of allergenic sub-
stances are lectins, which in elevated doses pose danger not only for those
allergic to them. The elevated level of estrogenic substances (PCBs, phytoe-
strogens, mycotoxins) may lead to altered bronchial functions. Estrogenic
substances increase β-2 receptor sensitivity to endogenous catecholamines.
These compounds also compete for binding sites on the Cortisol Binding
Protein (CBP) which may result in elevated plasma cortisol levels. At the
same time progesterone molecules prevent the contraction of smooth mus-
cle fibres by inhibiting the formation of gap junctions.
During the course of a nutritive imbalance when interference with
prostaglandin metabolism, high plasma cortisol level and relative plasma
progesterone level insufficiency lead to bronchospasm and smooth muscle
relaxation failure the endogenous conditions for late term abortion are
favourable. The compound leading to nutritive imbalance may be mycotox-
ins, phytotoxins, phytoestrogens, PCBs and bad ratio of between ω-3 ω-6
fatty acids.
The relationship between progesterone and prostaglandin metabolism
suggest that arachidonic acid metabolism may be involved in nutritional
diseases. It is known that in autoimmune chronic arthritic conditions low
518
arachidonic acid diet alleviates disease symptoms. It is also known that not
only relative progesterone insufficiency induced by estrogenic substances,
but allergic reactions are also involved in arachidonic metabolism through
the cyclooxygenase (COX) system. Since salicylic acid inhibits the COX sys-
tem, the involvement of arachidonic acid metabolism in pathological events
leading to abortion in sows seems to be likely and opportunity of using non-
steroid anti-inflammatory drugs (NSAID) to prevent abortion.
20.5.6.5. Plant substances

There are several plants, especially clovers and legumes, which contain high
levels of plant estrogens, which interfere with reproduction. These effects
may remain below a threshold level on their own, but are able to precipitate
in a clinical condition should other factors exert a synergistic effect. These
so-called borderline effects can be persistently present in the swine indus-
try, causing troubles from time to time.
Sows fed on locoweed (Astragalus spp.) showed reproductive failures
in the offspring. In rats the feeding of locoweeds alters the behaviour of the
offspring. Piperidine containing plant (Conium maculatum, Nicotiana glauca
and Lupinus formosus) induce foetal abnormalities (cleft palate, limb, spine,
neck contractures) which resemble the symptoms of folic acid deficiency.
Folic acid is also involved in prostaglandin metabolism. Thus, possible role
of arachidonic acid, which occurs in feed of animal origin and in certain
plants (flax, algae, wheat germ oil, and garlic) in the therapy or prevention
of sow abortions, remains to be elucidated.
In sow diets raw soybean cannot be used in significant amounts
because they contain biologically active anti-nutritive factors that can
adversely affect health. In addition soybean meal contains residual oil with
estrogenic substances. Another group of compounds the Non-Starch-
Polysaccharides (NSP) in feeds may also contribute to the development of
chronic enteric conditions affecting both nutrition and hormone metabo-
lism. The major detrimental effect of NSP is associated with its viscous
nature and its interaction with gut microflora. The slower transit time of the
digesta may cause a shift in intestinal flora favouring the proliferation of
toxin producing organism. All these processes may contribute for the main-
tenance of a chronic stress syndrome (leading to adrenocortical insufficien-
cy, pituitary ACTH depletion) which turns into an epidemic disease if one
additional factor comes into play.
20.5.6.6. Legionella syndromeand the algae

The air handling and watering systems of swine premises can harbour a
waste range of fungi, algae, protozoa and bacteria. Legionella-like microbes
are natural inhabitants of water and their multiplication depends on water
temperatures, 25-40oC being the most favourable range. In the swine stalls
adiabatic cooling and watering tanks, pipes and valves may provide
519
favourable habitat for these type of micro organism. This means that the
swine stall, where adiabatic cooling may be necessary in the summer peri-
od, can be turned into a breeding ground for unknown microbes, which
cause infections or rather massive allergic reactions. In the summer time
water tanks and tower can be turned into incubators. Legionella-type
microbes, such as cyanobacteria show amplified propagation in the pres-
ence of single cell organism. Other bacteria and algae can provide nutrients
for Legionella spp. organisms and cyanobacteria. Studies indicate that the
chemical environment of the plumbing systems with low level of iron, zinc
and potassium contamination also enhance the proliferation of such
species. Hence metal components and corrosion products of plumbing sys-
tem may play a role in promoting the growth of these bacteria. There is good
reason to believe that cyanobacteria play a still unknown role in porcine dis-
ease syndromes including neuromuscular disorders and reproductive prob-
lems. One of the sources is cyanobacteria and especially their toxins in the
fish meal. Algal blooms results in the accumulation of toxins in fish. It was
observed that small amount of microcystine was produced by the isolates,
but their anticholinesterase activity was as high as 2300-3300 microgram
anatoxin-a(s) g/l neurotoxins, hepatotoxins and skin irritant substances of
cyanobacteria accumulate in fish.
The problem of cyanobacterial toxicosis in swine units may also be
complicated by the observation that during water chlorination practices
sodium hypochlorite decompose microcystin, but during the process many
reaction products are formed. However, pre-treatment of water may cause
toxin release from algae and production of trihalomethanes during water
treatment.
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JAV_20_5_VND_FSGy:MINTAÚJ.qxd 2008.09.03. 15:26 Page 521
CHAPTER XX: PIG DIETETICS
20.6. Feed related diseases, herd health aspects

ACID-BASE BALANCE RELATED PROBLEMS

Feeding a feed mixture of low alkaline value (too acidic) results in decreased
feed intake, depressed average daily growth and worsened feed conversion
in growing pigs (see undetermined dietary anion, Chapter XI). In peripar-
turient breeding sow acidifying of urine will decrease its bacterial count and
will decrease incidence of MMA syndrome.
ANAEMIA) (IRON DEFICIENCY OF SUCKLING PIGLET)

Occurrence of syndrome is just obligatory in intensive keeping system.
Piglet anaemia is due to the low iron and copper content of milk; the small
neo-natal iron stores of piglets; the commercial drive for rapid weight gains
in young pigs; modern pig-housing, which restrict the ability of piglets to
“root” in soil; and cold, damp pig houses. Anaemic piglets are pale and their
immune system is impaired. They are susceptible to infectious agents; aver-
age daily gain and feed conversion decrease. Typically, the 10-20-day-old
piglet are pale, with white mucous membranes (porcelain piglet), breathing
is heavy, pulse rate is high, haemoglobin concentration of blood may fall
below 3.73 mmol/l (6 g/100 ml). Fortunately, the simple deficiency is easi-
ly dealt with. For treatment and prevention peroral or intramuscular or
combined iron supplementation is necessary. Overdosage of parenteral iron
may cause sudden death (iron toxicosis).
BIOTIN DEFICIENCY OF BREEDING SOWS AND BOARS

Hair loss and dry, scabby skin, deep transverse cracks and subsequent
bleeding of the sole and top of each hoof, increasing in severity as margin-
al supply lasts can be seen. Lameness resulting predominantly from lesions
at the junction of heel and toe, frequently complicated by secondary infec-
tions causing swelling and suppuration. Many pigs seem to develop a
hunched stance in which they stand with their hind legs forward of the nor-
mal position.(FIGURE XII-6) During walking, the placing of the feet is very
precautious („sneaker sow”). A flaky dermatitis over the back described as
similar to “greasy cornflakes”. A general softening of the horn of the hoof
and claws so that it can be depressed by the fingernail. Petechial haemor-
rhages under the horn of the hoof, and on the leg above the hoof. In some
cases these spread up the hams to the back of the pig. Of all these condi-
tions, the lesions in the heel region are the most debilitating. Foot lesions
have been attributed to other causes, including selenium toxicosis, physi-
cal and chemical abrasion by concrete, thus there is a need for differential
diagnosis.
In addition to the described physical conditions, there are suggestions
that breeding performance is also affected. Authors of the relating research-
521
es record cases where extended weaning to re-mating intervals, poor con-

ception rates and reduced litter sizes have all been improved by supplying
additional biotin (“inner flushing”). As an opposite, in sows of suboptimal
vitamin E supply infertility, reproduction and lactation problems may devel-
op.
DERMATITIS
caused by riboflavin deficiency: loss of hair, rough, dry skin, gener-
alised dermatitis. Pyridoxine deficiency: generalised exudative dermatitis
and conjunctivitis. Niacin deficiency: rough, dishevelled coat, desquamative
localised dermatitis on back of ears and back. Pantothenic acid deficiency:
locomotor trouble („goose stepping” or „Paradenmarsch”) caused by the
degeneration of the peripheral nerves
GOITRE IODINE DEFICIENCY

Goitre is the non-inflammatory, non-cancerous enlargement of thyroid
gland. The most common form is the iodine deficiency syndrome of newborn
piglets. It is typical that farrowing delays and big (2-2.4 kg) piglets are born.
Piglets have none or less hair and the subcutaneous connective tissue is
infiltrated with water (myxoedema). Thyroid glands of piglets are enlarged 2-
8 times. Within 1-2 days they usually die. Cause of the ailment is primary
or secondary (antinutritive substances like nitrite, nitrate, thyreostatic
plant compounds like glycosinolates) iodine deficiency. In endemic territo-
ries feed mixture of pregnant sows should be supplemented 10-30 mg/kg
potassium iodide, or 1.2 ml/sow/day Lugol solution (tincture of iodine with
5% iodine and 4% of potassium iodide) should be added to drinking water.
ILEITIS (PORCINE PROLIFERATIVE ENTEROPATHY)

Though it is a bacterial infectious disease caused by Lawsonia intracellu-
laris. At the background of this ailment stands an enterocyte overgrowth in
ileum, but the fibre deficiency may predispose animals. The typical (chron-
ic) clinical picture shown by growing, 3-6-month-old pigs are: growth
depression, bad feed conversion and excretion of soft faeces. Besides anti-
bacterial treatment (tylosin per os) the fibre and starch level of feed mixture
should be corrected.
MASTITIS-METRITIS –AGALACTIAE (MMA) SYNDROME

Main clinical sign of syndrome is a partial or totally disturbed lactation at
farrowing and in the following days, followed by the inflammation of mam-
mary gland and uterus. Lack of milk can be seen by the crying suckling
piglets too. In the background, disturbed endocrine function (so-called
“adaptation syndrome”) and intoxication (mostly E. coli) from constipation
can be found. Plant estrogens of soybean, hot and humid environment,
transport and failure in drinking water supply are predisposing factors. In
522
such herds fibre level in pregnant sow diet should be increase (8-9% crude
fibre) in order to prevent constipation; in severe cases addition of Glauber’s
salt (sodium sulphate decahydrate) may be the solution. Acidifying of peri-
parturient sows will help to avoid bacterial infection (for more details see:
Cation Anion Balance, Chapter XI).
MYCOTOXIKCOSIS
Aflatoxin. Sign of acute aflatoxicosis in swine include reduced body weight
gain, depression, haemoconcentration and liver damage. There is icterus,
with a yellow swollen liver, mesenteric oedema and serous atrophy of fat.
The median lethal dose (LD50) for aflatoxin B1 for the pig is 0.6 to 0.8
mg/kg. Chronic toxicosis from low-level ingestion of aflatoxin is much more
common and results in serious economic loss from decreased animal per-
formance, Feeding of high selenium level (2.5 mg/kg of feed) may protect
against aflatoxin toxicity. Haemorrhaging with prolonged clotting time is a
prevalent symptom of aflatoxicosis. Administration of vitamin K is an effec-
tive counter measure. Aflatoxins are highly carcinogenic and cause liver
tumours in swine. Methods of detoxification have been developed, although
the use of decontamination methods for grains and seeds in commercial
channel is regulated. Chemical methods that have been effective include
solvent extraction and formaldehyde, methylamine, sodium hydroxide, cal-
cium hydroxide, hydrogen peroxide, ozone, perchloric acid, methanol or
ammonia addition.
Zearalenone OR F–2 TOXIN and related compounds have estrogenic
activity that interferes with reproduction. Mouldy feeds may produce signs
of oestrus (swollen, reddened vulva) in immature gilts and interfere with the
oestrus cycle in cycling gilts. Zearalenone is different than other mycotox-
ins in that it exhibits estrogenic activity but is not acutely toxic. Its intake
may lead to clinical heat without fertile ovulation in gilts and sows. The
vulva and uterus become swollen and oedematous and vaginal and/or anal
prolapse may occur. The ovaries may partially atrophy. If pregnant sows
receive a feed mixture high in F–2 toxin and low in essential amino acids,
newborn piglet may have oestrogen syndrome and/or spay leg (spreddleg,
Speizbein, Grätschen). In the background of muscle hypoplasia a degener-
ation of spinal column (SZALAY et al. 2001). Abortion does not appear to be
caused by zearalenone alone, but it can result from other toxin present.
Zearalone affects sexual development of young boars. Male reproductive
tracts tended to be smaller when pigs were fed or dosed with high levels (500
to 600 ppm or 5 to 15mg/kg LW) of zearalenone. Prolonged feeding of low
levels (0 to 9 ppm) of zearalenone did not affect libido, but it tended to
reduce semen volume, total motile sperm and percent sperm motility.
Trichothecenes. The trichothecenes, especially deoxynivalenol (DON or
vomitoxin) are primarily responsible for the extraordinarily strong feed
refusal response and occasional emetic response by pigs to grain infected
523
with Fusarium spp. DON contaminated diets did not appear to have delete-
rious effects on gilts or their progeny, or on sexual development in boars or
gilts. T–2 toxin, diacetoxyscirpenol (DAS) and fusarenon-X are nearly (feed
refusal, immunosuppression) an order of magnitude more toxic than DON.
T—2 toxin does not cause abortion when consumed, but it can lead to infer-
tility. Immunosuppression, haematological and histological changes are
also reported. DAS has been diagnosed as causing haemorrhatic bowel syn-
drome in pigs.
Ochratoxin. Ochratoxin A and ochratoxin B are metabolites of several
Aspergillus spp. They are nephrotoxins and has been reported to cause feed
refusal also.
Fumonisin (emphysema of lungs). The presently known 6 fumonisins
are produced by Fusarium moniliforme and proliferatum and Alternaria
alternans. The most toxic is the fumonisin B1 =FB1). Oedema pulmorum,
hydrothorax. Lesion in liver, kidneys and pancreas (necrosis). 10-40 mg/kg
feed fed for 4 weeks caused interlobular and perilobular oedema pulmorum
(ZOMBORSZKYNE-KOVACS et al. 1997).Long-lasting intake of low toxin amount
(1-10 mg/kg feed) causes, after a pulmonar oedema, a fibrosis of lungs from
weeks 6-8 of the treatment. FB1 pass through placenta and in the foetus
also oedema pulmorum develop. Lactating sows excrete the toxin by milk.)
PARAKERATOSIS
It may occur on the whole body surface primary of secondary (calcium over-
feeding) zinc deficiency. Chronic syndrome of pig, characterized by abnor-
mal keratinisation of the skin and tongue epidermis. Enlarged skin is inflex-
ible, rigid and roughly folded with the hair falling out. Basically it is caused
by primary or secondary (calcium, phytic acid overdosage) zinc deficiency.
There is a possibility for bacterial over infection and development of exsuda-
tive, suppurative dermatitis. During differential diagnosis, the vitamin B6
deficiency and the exsudative dermatitis or greasy pig disease (caused by
Staphylococcus hyicus) should be taken into consideration. Treatment:
besides correction of zinc supply an addition of omega-3 fatty acid to diet
may be beneficial.
RICKETS AND OSTEOPOROSIS

Troubles of calcium, phosphorus and vitamin D supply causes incomplete
mineralization of chondrocytes. In growing pig among the symptoms are
stiff, painful gait, immobility, painless hard swellings on limb joints and ribs
(“ricketry rosary”: swelling on the ribs, a characteristic symptom of rickets),
bone distorsion in limbs, backbone, increased bone fragility and soft pliable
bones. In breeding gilts, kept on cereals, without calcium and vitamin D
supplementation, development of pelvic bones is retarded and fracture may
occur in pelvic bones and head of femur.
524
SWINE OESOPHAGEAL GASTRIC ULCER

The direct cause is disputable. Under intensive keeping conditions, espe-
cially if stress feeding failures are present, first themucous membrane of the
pars oesophagica in pig stomach get degenerated after which erosion may
develop. Occurrence of ailment is related to lack of fibre and too fine parti-
cle size in feed mixture. Feeding of large quantities of skimmed milk or whey
makes pig susceptible to this ailment. Fungi may also play a role in patho-
mechanism, especially if dieat is high in sugar. As a typical terminal lesion
is found in the gastric mucosa near the oesophageal opening in a rectan-
gular area of white, glistening, nonglandular, squamous epithelium. It is
common to find a crater encompassing the oesophagus. Crater appears as
a creamy or grey area and may contain blood clots. The healed ulcer
appears as a stellate scar. Anorexia, weakness and black tarry faeces are
typical clinical signs. Increasing the crude fibre content of the diet to 7%
and feeding coarse meal rather than pellets may be of some advantage.
Cimetidine (150 mg/pig/12 hours is useful in treating acute cases. In the
recovery process the positive role of thiamine (vitamin B1) and vitamin U (S-
methyl-methionine) is proved, but they are not effective in prevention (TAMAS
et al. 1975-82).
VITAMIN A DEFICIENCY
Incoordination, emaciation, blindness and posterior paralysis were seen.
Other signs include reduced growth, ataxia, night blindness, elevated cere-
brospinal fluid pressure, abnormal skeletal remodelling, squamous meta-
plasia of epithelial tissue, reduced spermatogenesis, developmental abnor-
malities including microphtalmia and anophtalmia, increased embryonic
mortality, and weakness in newborn pigs. The describe malformation in
newborn piglet may be due not only to deficiency but also to overdosage of
vitamin A in the diet of pregnant sow.
VITAMIN SELEN DEFFICIENCY SYNDROME (VESD)

comprises mulberry heart disease or microangiopathia, myodegeneration in
heart and muscles (Zenker-type), ulcus ventriculi and hepatosis dietetica.
The syndrome is associated with icterus, generalised oedema and death.
Necropsy lesions include pale, swollen, haemorrhagic livers (dietary liver
necrosis) and gastric epithelial ulceration. The most susceptible are the
muscles of the hind limbs (m. quadriceps, m. gracilis, m. adductor, m. psoas
and m. longissimus dorsi). Muscles are pale, soft and exsudative (in vivo!),
clinically like fish meat (“White muscle disease”). Typical is a weakness of
hind legs, locomotor troubles. Syndrome usually affects fast growing pigs of
a live weight of 30 to 60 kg. In the background of the clinical and patholog-
ical picture a severe lesion of cell membranes caused by free radicals can be
detected. Another manifestation of the same syndrome is the sudden car-
diac failure, dietetic microangiopathy and acute circulatory failure. In this
525
condition lesions of the cardiac muscles are prominent though other organs
may also be affected, for example necrotic livers. Before sudden death
depression and inappetance may be observed for several hours. Apathy,
dyspnoea and blue-reddish maculae of irregular shape and size may devel-
op on the skin, especially on the ears (focal cyanosis). Post mortem findings
include mottling and haemorrhage of myocardium, lung oedema, and
venous engorgement, oedema of the gastrointestinal tract and presence of
easily clotting transsudates in the body cavities. For prevention and treat-
ment vitamin E and selenium supply should be corrected. Clinically sick
individuals may receive 0.06 mg/kg LW selenium (in form of sodium selen-
ite or chelate). Five-ten times of recommended selenium concentration in
feed proved to be toxic. Characteristic are lesions on coronet, which being
similar to those of foot and mouth disease, makes differential diagnosis
essential.
Apart from the described syndromes, other pathological states should
also be mentioned, among which vitamin E responsive anaemia in piglets
and iron sensitivity in parenteral iron treated vitamin E deficient piglets are
the most common ones. The later is called “iron toxicosis” when death
results from an iron induced lipid peroxidation in tissues.
FOR FURTHER READING
AFRC Technical Committee on Responses to nutrients (1990): Report number 4, Nutrient

requirements of sows and boars. Agricultural and Food Research Council, Central
Office, Wilts
Cole, D.J.A., Wiseman, J. and Varley, M.A. (1994): principles of pig science. Nottingham
University Press
Fekete, L. (1995): Swine nutrition (Sertestakarmanyozas, in Hungarian). Mezogazda
Kiado. Budapest
Fekete, S. (1999): Non-Infectious Factors in Sow’s Abortion. Reprod. Dom. Anim. 34, 177-
184.
DLG (1987): Ausschuss für Bedarfsnormen der Gesellschaft für Ernhrungsphysiologie.
Energie- und Nhrstoffbedarf landwirtschaftlicher Nutztiere. Nr. 4. Schweine. DLG-
Verlag. Frankfurt am Main
Hollis, G.R. (ed.) (1993): Growth of the pig. CAB INTERNATIONAL. Wallingford, Oxon
Lewis, A.J. and Souther, L.L. (eds). (2001): Swine nutrition. Second edition. CRC Press.
Boca Raton, London, New York, Washington, D.C
Miller, E.R., Ullrey, D.E. and Lewis, A.J. (1991): Swine nutrition. Butterworth-
Heinemann. Boston – London – Singapore – Sydney – Toronto – Wellington
Morgan Jones, S. D. (ed) (1995): Quality and grading of carcasses of meat animals. CRC
Press. Boca Raton - New York – London – Tokyo
National Research Councel (1998): Nutrient requirements of swine. Tenth rev. ed. nation-
al Academy Press. Washington, D.C
Pluske, J. R., Le Dividich, J. and Verstegen, M.W.A. (2003): Weaning the pig. Concepts
and consequences. Wageningen Academic Publishers.
Rafai, P., Bata, A., Vanyi, A., Papp, Z., Brydl, E., Jakab, L., Tuboly, S., Tury, E. (1995):
Effect of various levels of T—2 toin on the clinical status, performance and metabo-
lism of growing pigs. Vet. Tec. 136, 485-489.
Rafai, P., Tuboly, S., Bata, Á., Tilly, P., Vanyi, A., Papp, Z., Jakab, L. and Tury, E. (1995):
526
Effect of various levels of T—2 toxin on the immune system of growing pigs. Vet.
Rec. 136, 511-514.
Szalay, F. Zsarnovszky, A., Fekete, S., Hullar, I., Jancsik, Veronika, Hajos, F. (2001):
Retarded myelination in the lumbar spinal cord of piglets born with spread-leg syn-
drome. Anatomy and Embryology, 203, 53-59.
Ulbrich, M., Hoffmann, M. and Drochner, W. (2004): Fütterung und Tiergesundheit.
Verlag Eugen Ulmer Stuttgart
Williams, N.H., Stahhly, T.S. and Zimmerman, D.R. (1997): Effect of level of chronic
immune system activation on the growth and dietary lysine needs of pigs from 6 to
112 kg. J. Anim. Sci. 75, 2481-2496.
527
CHAPTER XXI: HEALTHY DOG AND CAT
Chapter XXI
FEEDING AND CLINICAL DIETETICS

OF DOG AND CAT
© Sandor Gy. Fekete (if otherwise not indicated)
21.1. Feeding and nutrition of the healthy dog and cat

21.1.1. Anatomical and physiological characteristicof digestive
system
21.1.2. Feed intake, peristalsis, faeces quality
21.1.3. Digestion and absorption of nutrients
21.1.4. Interaction between the individual nutrients
21.1.5. Comparison of some eating behavioural traits of dog and cat
21.1.6. Feed preference
21.1.7. Recommendation about energy, nutrients, mineral and vita-
min requirments
21.1.8. Characteristics of the cat metabolism

21.2.1. Consequneces of inappropriate feeding
21.2.1.1. Excess energy intake (owerweight, obesity)
21.2.1.2. Deficient energy and protein supply
21.2.1.3. Deficient mineral and vitamin supply
21.2.1.4. Hypervitaminoses
21.2.1.5. Unbalanced diet
21.2.1.6. Degenerative diseases of dietary origin
21.2.2. Dietary treatment of pathological conditions
21.2.2.1. Principles of feeding sick, febril animal
21.2.2.2. Diarrhoea and constipation
21.2.2.3. Gluttony, anorexia
21.2.2.4. Gastric dilatation and volvulus
21.2.2.5. Congestive heart failure
21.2.2.6. Feline Liver Diseases
21.2.2.7. Exocrine pancreas insufficienty (EPI)
21.2.2.8. Diabetes mellitus
21.2.2.9. Skin diseases
21.2.2.10. Feed intolerance and allergy
21.2.2.11. Critical care
21.2.2.12. Nutrition and tumours
21.2.2.13. Inherited metabolic troubles in dogs
21.2.2.14. Inherited metabolic troubles in cat

21.2.2.15. Chronic Kidney Disease (renal failure)
21.2.2.16. Nutritional therapy of stones in the urinary
bladder of cats and dogs
21.2.3. Physiological conditions with special dietary needs
21.2.3.1. Reproduction
21.2.3.2. Nutrition and the skeletal system in dogs
21.2.3.3. Feeding of senior dog and cat
21.2.3.4. Raising of orphan puppies and kittens
21.2.3.5. Working dogs’ nutrition and feeding
21.3. Practical aspects of dog and cat nutrition

21.3.1. Types of dog and cat foods
21.3.2. Dietetic and medicinal feeds
21.3.3. Prediction of voluntary feed intake
21.3.4. Prediction of the nutritive value
21.3.5. Feline body mass index
21.3.6. Control of the protein supply and main protein sources of
high BV
21.3.7. Urolith diagnostic and treatment in the practice
21.3.8. Home-made diets
21.3.9. Feeding frequency, feeding practice
he nutrition of the companion or breeding dog and cat is more and
T more based on commercial complete feeds. The evolution of carnivo-

rous animals produced a wide range of digestive types, from the strict
carnivorous cat to the just omnivorous racoon. The dog is situated in the
middle, a carnivorous animal with some omnivorous characteristics. As an
all over consequence is that the digestion of lipids is excellent both in dogs
and cats; that of the proteins only of medium and the carbohydrate utilisa-
tion of dogs is medium to low and very low in cats. The explanation for the
medium protein digestion is that in the nature both species eat animal pro-
teins of high biological value. Dogs, compared to the real omnivorous ani-
mals, like pig, require much more protein and fat; however, he needs car-
bohydrates, too and is capable of utilising them. The difference between dog
and cat is expressed also in the length of gastrointestinal tract, compared
to the body length, averaging 1 to 4 in cats and 1 to 5 in dogs. To under-
stand the real nutrient requirements of dog and cat, it should be borne in
mind that wild carnivores eat not only the meat (muscles) of the prey ani-
mals, but first of all they consume the viscera (liver, kidneys, intestines),
providing in this way themselves by carbohydrates, plant fibres, minerals
and a series of vitamins.
21.1. Feeding and nutrition of the healthy dog and cat
21.1.1. Anatomical and physiological characteristic

of digestive system
The oral cavity of dog, owing the expandable buccal cheek is large; is
enclosed by the smaller lower and larger higher lips (labium). The wide
531
opening of the mouth allows dog to take up huge pieces of feed. Tactile hairs
(vibrissae) are found on the face, mostly on the upper lip (“whiskers”) and
near to eyes and also on the carpus of the cat and the tuft of whiskers on
the dog’s cheek. In the mucous membrane of the oral cavity groups of
mucous glands are situated. From front and from side the buccal cavity is
surrounded by the dental arch: each jaw forms a dental arch, thus there are
four dental arches in total. There are no teeth in pups during the first three
weeks of life. Approximately to the age of six month in dogs develop 42 and
in cat 30 teeth. The milk (or deciduous) dentition are present in the jaw at
birth and erupt as the animal grows. The eruption time for puppies’ incisors
is 3-4 weeks, for the canines 5 weeks and for the premolars 4 to 8 weeks;
there are no temporary molars, giving the dental formulae of I3/3, C1/1,
PM3/3 Î 2 = 28 teeth. In kittens the entire deciduous dentition starts to
erupt at 2 weeks and complete by 4 weeks, giving the dental formulae of
I3/3, C1/1, PM3/2 Î = 26 teeth. The adult teeth or permanent dentition last
for lifetime. The eruption time of dog’s incisors is 3.5 to 4 month, for the
canines 5 to 6 months, for the premolars 4 to 7 months and for the molars
5 to 7 months, giving the dental formulae of I3/3, C1/1, PM4/4, M2/3 Î 2
= 42 teeth. Cat’s incisors push through the gum at the age of 3 months, the
remaining teeth have variable eruption time, but the full permanent denti-
tion is present by 6 months, giving the dental formulae of I3/3, C1/1,
PM3/2, M1/1 Î 2 = 30 teeth. The carnassial teeth pertain to the last upper
premolar and the first lower molar; they are specialized for cutting and
shearing, being very powerful teeth sited close to the angle of the lip; car-
nassials can be found only in carnivores. The roots of the upper carnassial
teeth are prone to infection and easily molar abscess may develop. Generally
the abscesses burst through the bone and the skin to discharge pus below
the eye. Teeth in cats mostly serve for grabbing, tearing and swallowing the
bite.
The well-muscled tongue is spindle-shaped; by means of a dorsal
groove it can be formed into a spoon-form. It is covered in mucous mem-
brane; on the dorsal surface fine, thread-like, rough papillae are found,
which may sometime cause strangulation of the tongue. The tongue carries
the taste buds for gestation, it is important in grooming, thermoregulation
and vocalisation. By stretching out and turned aside, dog and cat is able to
take up (spoon) liquids. The four (parotid, mandibular, sublingual and zygo-
matic or suborbital) salivary glands produce daily on an average 30 ml sali-
532
va/kg LW. The function of saliva is the lubrication of the feed, thermoregu-
lation and antiviral and antibacterial activity by its lysozyme and IgA con-
tent. Oesophagus, especially in the dog, is extendable; its stratified squa-
mous epithelium lining is arranged in longitudinal folds. There is enough
muscle tone around the dog’s lips to retain liquids while they are in the
mouth.
Dog’s stomach is very large and extendable; if it is full, it can reach
the lower abdominal wall. Stomach is closed both at the entrance (cardia)
and at the outgoing (pylorus) by strong muscular ring. Cat stomach has an
elongated, spindle-shaped stomach. Within the surface of the gastric
mucous membrane there are three different zones: the narrow cardia with
glands, producing water-like mucus, the fundus with glands, producing the
real gastric juices and hydrochloric acid and the pyloric zone, with mucus-
producing glands. In the gastric pit three cell types are producing the
secretion: goblet cells (mucus), chief cells (pepsinogen) and parietal cells
(hydrochloric acid). The small intestine is relatively short; its mucous mem-
brane is covered by 0.5 to1.0 mm high leaf-shaped folds (villi) and its last
section, the ileum open in the caecum with a tap-like process (ileocaecal
junction). The small intestinal surface area is enlarged by the tiny microvilli,
extending from each epithelial cell border, forming the “brush border”.
Intestinal wall is well-muscled and there are digestive glands (Brunner’s
glands in the duodenum and Lieberkuhn crypts in the jejunum and ileum)
and in the dog even lymphoid patches in the wall. The pancreatic juice and
the bile are secreted into duodenum separately. The caecum of dog is small,
that of the cat just virtual (see Chapter XIX); short colon ends in the anus,
which is closed by a well-developed anal sphincter.
Around the anus orifices, lying between the internal and external anal
rings, the peas to grape large glands open (anal sacs). These are modified
sebaceous glands, producing greyish, mucous secretion, which serves to
coating the faeces and for territorial scent marking. The impaction of the
anal sac may cause painful inflammation, which in turn, manifest in con-
stipation. The large liver goes beyond right arch of the ribs; its relative
weight is 3 to 4% of LW. As a practical conclusion of the anatomy, the dog
and cat will require high density low fibre diet for the normal digestion.
21.1.2. Feed intake, peristalsis, faeces quality

Dogs (and generally Canidae) are such type of carnivorous, which are capa-
533
ble of ingesting huge amount of feed for once. This characteristic has been
developed owing to the haphazard, periodic feeding, the bad predictability of
the next prey and the competition with the herd mates present.
Consequently, the dog is prone to greedily eating and superficially chewing.
On the other hand, to help herd mates, that are unable to hunt (lactating
bitches, weaning pups), a special form of trophallaxis developed. It means
that hunters eat huge amount from the killed prey, but they vomit it for the
others. Sometimes it occurs also by jealousy or enviously in kennels.
Felidae, except lions, eat many times smaller portions in a day.
The mode of dog’s feed intake primarily depends upon the consisten-
cy of its feed. Small bites are taken up by the incisors and swallowed with-
out thorough chewing. Larger pieces are fixed by its legs and with the head
turned aside, shred it with his carnassial teeth. Secretion of some digestive
juice began already during the feed intake. Excellent sight and smell of cat
facilitate the catching of the prey. The maintenance requirement of an adult
cat is approximately 1.8 mice/kg of LW (indoors) and 2.1 mice/kg LW (out-
door) of 27 g of LW, containing 32% DM, 18% crude protein, 9.5% ether
extract, 3.4% ash and 0.1% N-free extract (FEKETE et al. 1996), the calcu-
lated ME-value is 0.16 MJ/mouse (“mouse unite™”, FEKETE, 2003). this
explains the feeding habit: many times, smaller portions, having the daily
needs of 7 to 10 mice. The calculation based on the requirement data of
NRC (2006), taking 0.293 MJ ME/kg of LW for indoors and 0.335 MJ
ME/kg of LW for outdoors adult cat of optimum BCS.
Well-developed papillae on the tongue help in tearing off the meat
from the bones, but also the process of grooming (washing himself).
Although the strong molars suffer injuries with the age, it has small influ-
ence on the efficacy of the digestion. Drinking is carried out by the tongue,
forming in a spoon-shape. Swallowing, even gulping of carnivorous is easy
and there is no danger of false-deglutition into the larynx. Saliva, by lubri-
cating (moistening) feed, helps the process. The bite readily goes through
the expendable oesophagus and enters the stomach.
Notwithstanding that the gastro-intestinal tract of the dog (FIGURE
XXI-1) and cat is short and simple; the stomach is relatively large (see also
Chapter XIX) and chiefly in the dog, is appropriate for storing of huge
amount of feed. In the fundus part starts the protein digestion of the accu-
mulated feed.
534
FIGURE XXI-1: Schematic presentation of the dog’s digestive system
The material is soaked by digestive juice, and then moderate contrac-

tions push it slowly toward the pylorus. Emptying of the stomach is regu-
lated by the composition of gastric juice (mucus, pepsinogen, hydrochloric
acid, pepsin), by the osmotic pressure of the content, by the particle size,
the viscosity, acidity (pH), the presence of soluble fibres (e.g. pectin, glu-
cans, arabinoxylans) as well as the fat and fatty acid content of the duode-
nal fluid. Gastrin, produced by the stomach mucosa, stimulates, entero-
gastrin of the gut wall on the contrary, inhibits gastric motility. There is also
a neural control of the emptying and the mentioned soluble fibre fractions
have delaying effect. The fullness and composition of the small intestine
have also important influence. The more firm the feed particles are, the
longer they stay in the stomach. For example liver pieces of 50 grams stay
for approximately four hours in the stomach. Owing to the regulatory mech-
anisms, high-fat diet remains more time in the stomach than the carbohy-
drate-rich feeds. Generally, no later than 15 to 20 hours the stomach con-
tent reaches the small intestine. Occasionally, owing to an antiperistaltic
movement of the duodenum, gut content may return in the stomach
through the opened pylorus, initiating some fat digestion there, too.
As mentioned above, the dog and cat vomit easily. The most common
causes are the viral or bacterial infections, long-term inhibition of the gas-
tric emptying, gastric irritation, travel sickness, the presence of hardly or
indigestible objects (e.g. foreign bodies, bone, grass), endoparasites, abdom-
inal neoplasm and metabolic diseases (!). Vomiting is also common in the
lactating bitches in the third-fourth weeks of suckling. The passive vomit-
535
ing (regurgitation) is caused by an overflow of oesophageal content. The con-

genital overgrowth of the pyloric sphincter causes a pyloric stenosis, which
in turn a projectile vomiting in newborn puppies and kittens.
In physiological circumstances, the frequent gastric movements (in
every 15 to 20 second) push the whole stomach content into the duodenum
during some hours. The speed of the gut content movement is approxi-
mately 4 cm/min; in the meantime, nutrients are partly digested and
absorbed. After a physiological small intestinal digestion only a few indi-
gestible fractions (hair and horny materials, plant fibres, ash, occasionally
plastic) get into the large intestine. Total lack of indigestible particles, sim-
ilarly to human, results in colon dilatation, obstipation, liver damage and
possibly in secondary diarrhoea (EASTWOOD and KRITCHEVSKY, 2005). This is
the reason, why both dog and cat require some (1-3%) indigestible fibre in
their feed mixture. In the caecum and colon microbes of flora ferment indi-
gested compounds. Part of the released fatty acids, small amount of amino
acids, organic acids, ammonia, water and minerals are absorbed. At the
same time, there is also an excretion of minerals into the lumen by the large
intestinal glands. There are generally some volatile fatty acids in the faeces,
too. Increased VFA concentration is an indicator of the intensity of the bac-
terial fermentation in the large gut. In case of small intestinal resection
there is some functional compensation of the large intestine in helping
digestive functions.
Composition of faeces is variable; basically it consists of the undi-
gested feed particles, secretions of digestive glands, sloughing off/desqua-
mating enterocytes and microbes of intestinal flora. While feeding an aver-
age, common diet, and the first defecation occurs approximately 12 hours
after ingestion, it achieves the peak in 24 to 30 hours after feeding and
ceased in hours 40 to 60. The average daily defecation occasion in dogs is
1.7, which may increase to four in case of eating hardly digestible feed;
while eating solely meat or viscera, there is defecation only on every 2 to 4
days. Water content of the faeces is 55 to 75%. The Waltham Centre for Pet
Nutrition developed a faeces consistency scoring system, where score 1
being the very dry and 5 the watery diarrhoea. (“Grade 1: “Hard, dry and
crumbly, ‘bullet-like’, Grade 1.5: Hard and dry, Grade 2: Well-formed, does
not leave a mark when picked up, ‘pickable’, Grade 2.5: Well-formed, with
a slightly moist surface, with leaves a mark when picked up, ‘almost sticky
to touch’, Grade 3: Moist, beginning to lose form, leaving a definite mark
536
when picked up, Grade 3.5: Very moist, but still has some definite form,
Grade 4: The majority, if not all the form is lost; poor consistency, viscous,
Grade 4.5: Diarrhoea, with some areas of consistency and Grade 5: Watery
diarrhoea”). The precise definition of diarrhoea is frequent evacuation of
watery faeces, caused by maldigestion, malabsorption, increased peristal-
sis, gastrointestinal irritation, dietary mismanagement and infectious
agents. Colour of the faeces, according to the ingested feed, may be tan or
tawny, like the clay, light or dark brown. After eating much bone, it gets
light grey, that of milk or dairy products yellowish. Blood makes faeces
colour dark brown or black; after feeding of much vegetables, faeces may be
olive green. Table XXI-1 compares the chemical composition of faeces after
ingestion of different feeds.
Table XXI-1: Average faeces composition after ingestion of different feeds

(after BOKORI, 1993).
21.1.3. Digestion and absorption of nutrients

The main features of digestion in dog and cat are similar to the other mono-
gastrics (see Chapter XX at the pig); therefore these processes are described
only shortly. The three most important physical factors, influencing the car-
nivorous digestion are the pH, the water content of digesta and the intes-
tinal flora. The pH affects both the activity of digestive enzymes and the
composition of the microbial flora. The pH-value of the gastric juice before
feeding is approximately six, which will drop postprandial, under the influ-
ence of produced hydrochloric acid up to pH 1.5-3. In the duodenum the
acidic content will quickly be neutralized (up to pH 6. to 6.5) by the pan-
creatic juice. In the following sections of the gut the pH remains within the
range of 6.0 to 7.0. The quantity of liquid, present in the stomach, primari-
ly depends on the ingested feed, by having mixed with the saliva and gas-
tric juice, even in case of dry feed eating, will achieve the 30-40% of the total
content. Owing to its absorption, the water concentration in the colon con-
537
tent is lower (25 to 35%) and that of the faeces even lower, unless there is
diarrhoea. Gut content is never sterile; the intestinal flora is relatively poor,
the average germ count in the duodenum is only 106-7/g content, which
rearward continuously increases, attaining the 1010-11/g colon content.
Although the host organism is decisive (by the IgA production) in determin-
ing the composition of the intestinal flora, in turn, the chemical composition
of the ingested feed has also a great influence. Namely, the carbohydrate-
rich diet favour the proliferation of Lactobacillus acidophilus,
Bifidobacterium bifidum, Streptococcus spp., Enterococcus spp. and
Clostridium perfringens, in turn, the protein-rich diet facilitate the colonisa-
tion of Clostridium sporogenes, Cl. putrefaciens, Cl. novy, Bacterium mesen-
tericus, Proteus spp., staphylococci, Pseudomonas fluorescens and
Pseudomonas aerugenes. In case of mixed diet Escherichia coli appear, too.
Unbalanced feeding, predominant offering of a single feedstuff or abrupt
changes of feeding regime significantly influences the flora composition.
While feeding high-lactose or high-starch diet (e. g. milk, bread, potato,
bean and rice), the number of sugar-degrading bacteria (e. g. Bacterium aci-
dophilus) increases, parallel to the repression of protein-degrading bacteria
and the coliforms. Protein-rich diet elevates concentration of Clostridium
perfringens. Dietary lipids have small influence on bacterial flora; however,
too much fat ingestion drops the number of streptococci and coliforms.
Feeding of feed mixture, high in vegetable oil may result in diarrhoea.
Digestion realizes through the action of the products of salivary
glands, stomach mucosa, pancreas, liver, brush border enzymes and the
intestinal flora. Composition of the saliva depends upon the feeds charac-
teristics: feeding raw meat, the saliva is mucous, while eating meat-meal
(meat-and-bone meal, MBM); the diluted saliva of the parotis dominates.
The sodium and bicarbonate content of the saliva has buffer capacity.
Saliva of dog and cat does not contain digestive enzymes, it serves chiefly to
soak the bite, but its biologically active compounds, like IgA and lysozyme
are essential in defence mechanism against pathogens.
Gastric juice is produced by the fundus and pyloric glands.
Hydrochloric acid, amounting approximately 0.5% of the gastric juice is
responsible for the appropriate pH of the gastric content. Mucous mem-
brane of stomach is protected by the on site synthesized mucus.
Pepsin(ogen) is synthesised in the fundus and pyloric zones has important
role in protein digestion. Its pH optimum is within the range of 1.5 to 3.5.
538
The daily gastric juice production, according to the species, breed and feed
quality, accounts 20 to 50 ml/kg LW; the daily amount depends also on
linked conditional reflexes and the emptiness-fullness of the stomach. There
is basically only protein digestion in the stomach, pepsin degrades proteins,
splitting mainly bonds at the aromatic amino acids (phenylalanine, tyro-
sine); up to water soluble olygopeptides; practically no free amino acids are
released. In the first phase of the digestion, owing to the activity of
Lactobacilli, there is also lactic acid fermentation in the stomach, producing
lactic acid. In abnormal feeding conditions (high-carbohydrate diet or inges-
tion of yeast etc.) the produced large amount of lactic acids and gases may
cause gastric dilatation and many times volvulus, too.
Pancreatic secretion is a clear, water-like liquid; its pH is within the
range of 7.0 to 8.0. Its sodium bicarbonate content contributes to the neu-
tralisation of the very acidic stomach content. However, there is also mucus
in the pancreatic fluid. The pH optimum of the produced digestive enzymes
is at about 7.0. Protein-degrading enzymes of the pancreas are excreted in
inactive form and become activated in the gut lumen into trypsine, chy-
motrypsine, elastase and carboxypeptidase. In pathological circumstances,
in case of some pancreas ailments the enzymes get active in the site of the
production, causing autolysis of the glandular tissue. Amount and activity
of lipase is strikingly strong in both species. Part of the released free fatty
acids reacting with the sodium content of the medium, forms fat soaps.
Among the carbohydrate-degrading enzymes the amylase is the most impor-
tant. This enzyme occurs already in pancreas as active form. Production of
pancreatic fluid is continuous but during feed intake its amount temporar-
ily increases. Trypsine degradation produces mostly olygopeptides, besides
a small amount of free amino acids. Breakdown of olygopeptides are ended
by the aminopeptidase, carboxy-peptidase and dipeptidase. Free amino
acids, di-, tri and tetrapeptides enter the enterocytes (epithelial cells); but
only free amino acids get into the blood of vena portae.
Enzymes of the mucous membrane are the intestinal lipase,
aminopeptidase, dipeptidase, nucleotidase, and enterokinase. The latter
releases trypsine from the pancreatic trypsinogen. Besides trypsine, pan-
creas-originated digestive enzymes are the chymotrypsin, carboxypeptidase,
nuclease, pancreatic lipase, alpha-amylase and cholesterol-esterase.
Trypsine breaks down olygopeptides by splitting peptide bonds at the car-
boxyl group of basic amino acids (lysine, arginine, histidine). Enzymes of the
539
brush borders (maltase, lactase and saccharase) contribute to the more

effective digestion in the small intestine. The most intensive period of the
lactase production is the first three weeks of age and then there is a con-
tinuous decline in the available quantity, especially in cat. The lactose-
degrading capacity of the adult cat is not higher than 1-2 gram lactose/kg
LW daily (i.e. 20-40 ml cow milk/kg of LW (PIBOT, 2005). Maltase and sac-
charase activity slowly develop from the birth. Disaccharase activity even in
the adult dogs and cats is low; digestion of saccharase (sucrose) is limited
both in dog and cat, above the ingestion of 6 to 8 gram/kg LW it may cause
diarrhoea. Generally, for dogs and especially for cats, starch and other com-
plex carbohydrates should be given only after heat treatment. During
absorption from the small intestine both active and passive transport mech-
anisms occur. Metabolism of enterocytes is extremely fast, the renewal time
is 2 to 7 days, compared to the 2 to 3 months of the skeletal muscles.
Bile is produced by the liver; its original dry matter content is 3.5%,
which get four-six fold more concentrated in the gall bladder. Bile contains
conjugated (primary) bile acids (cholic and chenodeoxycholic acid, which
are secreted in conjugated form with glycine and taurin (dog) or only with
taurin (cat). The synthesized daily amount of bile acids is high, 100 mg/kg
LW, but the majority of that is reabsorbed from the small intestine, forming
an enterohepatic circulation. Continuous bile production is stimulated by
the protein and lipid components of the feed and endocrine influences, too.
Hepatic bile secretion and mucus production in the Brunner-glands of the
mucous membrane are stimulated by the compounds of chyme (e. g. lipids,
as well as peptide hormones (secretin and cholecystokinin). From one hand,
bile keeps fats and oils in emulsion, on the other hand, activates lipase. As
long as dog able to conjugate bile acid both with glycine and taurin, the cat
is capable of binding only by the taurin (taurocholic acid), for details see
later.
As mentioned, caecum is small in both species, in the cat just rudi-
mentary; either in fibre degradation or in vitamin synthesis it cannot be
taken into account. Bacterial enzymes, like cellulase and pectinase help in
splitting part of the fibre in the large intestine. End-products of bacterial
carbohydrate fermentation include organic acids (e.g. butyric acid) and
gases; that of the proteins ammonia and biogenic amides. Bacterial
enzymes contribute also to the degradation of the mucus content of the
digesta, excreted into the lumen of the small intestine. Through the coloni-
540
cytes of the hind gut there is only passive diffusion of compounds towards
bloodstream. Colonic microbial fermentation produces volatile fatty acids
(acetic, propionic and butyric), which after having absorbed provide energy,
the acetate in the tissues and the propionate in the liver. Moreover, butyrate
regulates metabolism and proliferation of colonicytes. Digestion of dog and
cat is influenced by the stress during the feed intake. The accelerated peri-
stalsis negatively influences feed utilisation. High feed intake also decreas-
es digestibility of nutrients. The preparation of feed has especially a huge
influence on the utilisation of carbohydrates and in less extent of proteins.
In cat trial, HULLAR et al. (1998) has found a significant improvement of the
digestibility of raw soybean-based diet after extrusion (organic matter from
58 to 76%, crude protein from 58 to 76%, ether extract from 88 to 91% and
N-free extract from 34 to 72%.
21.1.4. Interaction between the individual nutrients.
There are interactions between the digestibility and absorption of feed com-
ponents. Although protein overfeeding has little influence on the digestibil-
ity of other major nutrients, its deficit decreases the digestibility of the oth-
ers. Diets, high in fat cause a prolonged staying of feed in the stomach, pro-
moting in this way the gastric protein degradation. Crude fibre decreases
the digestibility of organic matter, especially that of the protein, 1% extra
crude fibre (above 3%) may cause a decline of 2-3 percent units. The ratio
of different fibre fractions (NDF, ADF and lignin) dramatically modifies the
effect of dietary fibre on voluntary feed intake and the digestibility of major
nutrients (FEKETE et al. 2004). High ash content may neutralize gastric juice,
inhibiting in this way the activity of pepsin.
21.1.5. Comparison of some eating behavioural traits of dog and cat.
Dogs, but especially cats during grooming, swallow hair, which may form
hairball. To trigger vomiting, both species graze. Consequently, for the
indoor cat “cat grass” (commonly germinated cereals) should be provided.
Lack of available, appropriate “cat grass”, cats are inclining to gnaw orna-
mental plants. There are many plants and household products that are
toxic to cats, for example Aloe vera, Dieffembachia (dumbcane), onions,
tylelol. It has been proven that many times cats will chew on the house-
plants out of sheer boredom. Vomiting helps also to get rid of other indi-
gestible materials, like bones, plastic etc.). Differences of the postabsorptive
metabolism of dog and cat see later.
541
21.1.6. Feed preference

Voluntary dry matter intake depends upon a series of external factors. The
general paradigm (“eating on energy”) is less decisive for the carnivores,
especially for cats. Cats tend to it approximately the same amount (per kg
LW) of feeds having different energy density (MORRIS et al. 2006). Adding cel-
lulose to the feed mixtures, dogs significantly increased their dry matter
intake; on the contrary, cats ate practically the same amount of dry matter
(PROLA et al. 2006). From the endogenous factors the gastric emptiness, the
sight, smell, touch and taste, previous experiences and the composition of
blood plasma (the level of individual nutrients, hormones, peptide, the ratio
of amino acids etc.) should be mentioned. Important exogenous factors are
the access to the feed, feeding frequency, amount of portion, physical form,
temperature. colour and tastiness. Feed preference basically depends on the
lipid content, (it is stimulated also by the presence of sweet amino acids
(glycine, alanin, lysine, histidine, cystine and prolin), nucleotide, the uami-
taste and the level of common salt. Dog and cat, being carnivorous, prefer
animal protein better, than the plant. They feel high-fat and high-protein
diet as more tasty. “Digest” is a product of partial protease fermentation of
slaughter house by-products (viscera, chiefly intestine and liver) and it is
applied to improve the preference of dog and cat feeds. Sweet sugar taste
(from glucose, sucrose, fructose and lactose, but not from maltose, as well
as from sodium cyclamate) is a positive stimulus for dog, but neutral for the
cats, having no sweet receptors (LI et al. 2006). Cats are also insensitive to
salt (BRADSHAW, 2006), rather prefer some amino acids and nucleotides;
dogs have an aversion towards sodium saccharin.
Both species, but especially cat is prone to develop an exclusive pref-
erence to the taste of the feed (addiction). Cats remember to the diet- linked
negative experiences (e.g. LiCl stimulated vomiting), dog, on the contrary,
makes the same feed choice failure repeatedly. The excellent taste and pref-
erence of a commercial diet does not automatically means an outstanding
nutritional value! At the same time, amino acid imbalance reduces volun-
tary feed intake. The final preference of a feed is the outcome of the physi-
cal form, texture, water content, and dustiness, measure of bites, taste,
microbiological status, mouldiness and rancidity. In case of a very taste and
concentrated feed, sometimes the rationing is necessary to prevent develop-
ment of obesity.
542
21.1.7. Recommendation about energy, nutrients, mineral and vitamin

requirements. In the every day practice the deficient nutrient supply is not
common and it is more characteristic for home-made diet feeding.
Nevertheless, using commercial diet, overfeeding or unbalanced mineral-
vitamin intake may occur. During covering the nutrient requirements, is
essential to take into account not only the absolute quantities, but also the
appropriate ratio between them, considering the existence of several inter-
actions. Antagonism occurs during the absorption of minerals (e.g. calcium
vs. phosphorus, calcium vs. zinc, zinc vs. copper etc.). The practical signif-
icance of the antagonism between amino acids of similar side-chain (e.g.
lysine vs. arginine, valin vs. leucin vs. isoleucin) is smaller. Synergism can
be observed between fat content and the utilisation of the fat-soluble vita-
mins. Based on the similar biological effects, complementary effects can be
found between selenium and vitamin E, methionine and cystine, among the
members of the methyldonator group (methionine, choline, folic acid, and
vitamin B12), phenylalanine and tyrosine, tryptophan and niacin.
Nutrient requirement can be given for live weight and for 24 hours or
as a concentration of the feed dry matter content. “MINIMUM REQUIREMENT
(MR)” is necessary to prevent manifestation of clinical deficiency signs. In
the practice is more useful to add some safety margin, to compensate occa-
sional differences in the biological availability of nutrients in the individual
raw materials, to enhance or support higher resistance of animals against
stress influences or pathogenic agents. The widespread accepted and used
RA-NRC (National Research Council) recommendations (2006), contrary to
the previous editions, gives not only the MR, but propose an ADEQUATE
INTAKE (AI), RECOMMENDED ALLOWANCE (RA) and SAFE UPPER LIMIT (SUL). The
AAFCO (Association of American Feed Control Officials) in turn, applied
mostly by the feed industry, proposes the „OPTIMUM REQUIREMENT”. The NRC
RA is similar to the AAFCO minimum and the NRC SUL value to the AAFCO
maximum recommendation (SWANSON and FAHEY, 2005).
Dogs’ maintenance energy requirement (MJ ME) according to the NRC
(2006) is 0.544 × W0.75, but the breed-, age-, housing- and activity-related
differences are big, ranging from 0.393 to 1.046 MJ ME per W0.75 . Cat: the
energy requirement data of NRC (1986) are 0.293 MJ ME/kg of LW for
indoors and 0.335 MJ ME/kg of LW for outdoors adult cat of optimum
BCS. The AAFCO (2003) recommendation (Tables XXI-2) are appropriate for
practical use.
543
Table XXI-2: Nutrient, mineral and vitamin requirement of dogs and cats (AAFCO, 2003)
1Methionine+Cystine; 2Phenylalanine+Tyrosine; 3No recommendation (NR)
544
21.1.8. Characteristics of the cat metabolism

Cats, owing to the less evolutionary pressure, evolved to strict, obligate car-
nivorous animals. Compared to dogs, concerning the metabolism, nutrients
needs, feeding behaviour and intermediary biochemical pathways, cats
show the following species characteristics. The protein requirements are
higher by at least 30% than in dogs. The ileal endogenous amino acid loss-
es are 2-3 times higher in cats than in the omnivorous mammals. The car-
bohydrate needs are low; moreover cat can base its energy supply on fats
and by means of glyconeogenesis, on proteins. Consequently, the intensity
of urea-cycle (ornitine-citrullin circle) is higher, with the concomitant high-
er needs to arginine and biotin (see biotin effect on the gene expression of
ornithine carboxylase, Chapter XVII). The reason for the high arginine
requirement is, that cat is able to produce ornithine exclusively from argi-
nine in the liver and not also from proline and glutamate in the gut wall, like
dog (MORRIS and ROGERS, 1978). Specific feature of glucose and energy
metabolism, namely their starch and NSC utilisation is very low, but on the
contrary, they can survive without carbohydrates. Cats, like several mem-
bers of the Felidae family, are excreting in the urine a sulphur-containing
amino acid, the FELININE. Its production is high in the entire mature tom cat,
lesser and lesser in the castrated males, entire females and the lowest is in
the spayed females. (HENDRICKS et al. 1995). Felinine excretion is stimulated
by the testosterone and it is a powerful pheromone (MIYAZAKI et al. 2006).
ISOVALTHINE is also a sulphur-containing amino acid in the cats’ urine. It
canot be found in the urine of healthy omnivores, but it is present in the
urine of humans with hyper-cholesterolemia (OOMORI and MIZUHARA, 1960).
Although there is some taurin synthesis in the cat’s liver, it is not
enough for the conjugation of bile acids and for the other biological function
(retina, ovaries, immune system), it is an essential compound for the cat,
required in the offered feed. Moreover, Maillard reaction during heat treat-
ment of commercial feeds (see Chapter IV) the availability of taurin dramat-
ically drops. Long-chain fatty acids with double bound at ω-6 and ω-3 posi-
tions cannot be synthesized endogenously by the carnivore species, there-
fore, they should be provided in the diet. Dog is able to produce from linole-
ic acid (18:2ω6) and a-linoleic acid (18:3ω3), by a series of desaturases and
elongases arachidonic acid, ARA (20:4ω6), as well as eicosapentaenoic acid,
EPA (20:5ω3) and docosahexaenoic acid, DHA (22:6ω3), respectively. On the
contrary, cats having low Δ5 and no Δ7 desaturase activity in their liver,
545
require exogenous arachidonic acid, too (RIVERS et al. 1975). In human, an

increased blood ratio of eicosatrienoic acid (20:3ω9) to arachidonic acid
(20:4ω6), i.e. if triene to tetraene ratio is higher than 0.2, it is a diagnostic
sign of ω6 fatty acid deficiency (HEIRD and LAPILLONE, 2005). Characteristic
symptoms of deficiency include growth depression, and scaly skin lesions.
Lack of ω-3 fatty acids results in neurological abnormalities, disturbed visu-
al and cognitive functions.
Lack of carotenase enzyme, the transformation of carotene to vitamin
A cannot be accomplished; therefore carotene supply cannot replace the
vitamin A intake. The relative exogenous vitamin D needs of cats are high-
er than in dog, because of less amount of the precursor 7-dehydrocholes-
terol in the skin. Generally eating more unsaturated fatty acids, the vitamin
E requirements of cat is high; even the harmful effect of taurin deficiency
(the incidence of dilated cardiomyopathy) is reinforced having marginal
tocopherol supply (FOX et al. 1993). Their niacin needs are high, because
tissue niacin synthesis from tryprophane in the liver is insufficient. Low
manganese intake causes infertility in queens; maybe, this phenomenon is
related to the low absorption rate in the short and less acidic intestinal
tract.
They are more susceptible to the harmful effects of antinutritive sub-
stances, like trypsine inhibitor or iron absorption inhibitor in soybean. In
the other animal species the ruminal and intestinal flora contributes more
efficiently to the neutralisation of plant antinutritives; moreover, the poten-
tially harmful compounds can be conjugated both with glycine and taurin
and excreted by the urine. Intestinal flora of the cat is poor and there is no
conjugation with the glucuronic acid. Sulphur containing compound of
onion may cause haemolytic anaemia, characteristic for the presence of
Heinz-bodies. Dogs are much less susceptible to these compounds of the
onion. Differences in the feeding behaviour (frequency, way of ingestion,
addition), stress susceptibility during eating are also strikingly different in
cats, compared to dogs. Application of diazepam in cat, but not in dog, is
capable of improving appetite.
546
FOR FURTHER READING
AAFCO (2003) Association of American Feed Control Officials Inc, Official Publication.
Atlanta, GA, USA
Bradshaw, J.W.S. (2006): The evolutionary basis for feeding behaviour of domestic dogs
(Canis familiaris) and cats (Felis catus). J. Nutr. 136, 1927S-1931S.
Eastwood, M. and Kritchevsky, D. (2005): Dietary fiber: how did we get where we are?
Annu. Rev. Nutr. 25, 1-8.
Fekete, S. G., Hullar, I, Andrasofszky, E. and Kelemen, F. (2004): Effect of different fibre
types on the digestibility of nutrients in cats. J. Anim. Physiol.a. Anim. Nutr. 88,
138-142.
Fekete, S., Szakall, I., Andrasofszky, E., Kosa, E and Hullar, I. (1996): Body composition of
mice of different condition score and sex. Acta Vet. Hung. 44, 399-410.
Fox, P.R., Trautwein, E.A., Hayes, K.C., Bond, B.R., Sisson, D.D. and N.S. Moise (1993):
Am. J. Vet. Res. 54, 563-569.
Heird, W.C. and Lapillone, A. (2005): The role of essential fatty acids in development. Annu.
Rev. Nutr. 25, 549-571.
Hendricks, W.H., Moughan, P.J., Tarttelin, M.F. and Woolhouse, A.D. (1995): Felinine: a
urinary amino acid of Felidae. Comp. Biochem. Physiol. B, Biochem Mol. Biol. 12,
581-588.
Hullár, I, Fekete, S.. and Szőcs, Z. (1998): Effect of extrusion on the quality of soybean-
based cat food. J. Anim. Physiol.a. Anim. Nutr. 80, 201-206.
Li, X., Li, Weihua, Wang, H., Bayley, D.L., Cao, J., Reed, D.R., Bachmanov, A.A., Huang,
L., Legrand-Defretin, V., Beauchamp, G.K. and Brand, G. (2006): Cats lack a sweet
taste receptor. J. Nutr. 136, 1932S-1934S.
Miyazaki, M., Yamashita, T., Suzuki, T., Saito, Y., Soeta, S., Taira, H. and Suzuki, A.
(2006): A major urinary protein of the domestic cat regulates the production of feli-
nine, a putative pheromone precursor. Chem. Biol. 10, 1071-1079.
Morris, J.G. and Q.R. Rogers (1978): Ammonia intoxication in the near adult cat as a result
of a dietary deficiency of arginine. Science 199, 431-432.
Morris, P.J., Calvert, E.L., Holmes, K.L., Hackett, R.M. and Rawlings, J.M. (2006): Energy
intake in cats affected by alteration in diet energy density. J. Nutr. 136, 2072S-
2074S
National Research Council (2006): Nutrient requirements of dogs and cats. The National
Academy Press. Washington, D.C.
Oomori, S. and Mizuhara, S. (1960): Structure of a new amino acid isolated from the urine
of hypercholesterolemic patient. Biochem Biophys. Res. Com. 3, 343-345.
Prola, L., Dobenecker, B. and Kienzle, E. (2006): Interaction between dietary cellulose con
tent and food intake in cats. J. Nutr. 136, 1988S-1990S.
Rivers, J.P.W., Sinclair, A.J. and Crawford, M.A. (1975): Inability of the cat to desaturate
essential fatty acids. Nature 258, 171-173.:
Swanson, K.S. and Fahey, G.C.Jr. (2005): An update on the energy and nutrient require-
ments of companion animals. In: Garnsworthy, P.C. and Wiseman, J. (eds): Recent
advances in animal nutrition -2004. Nottingham University Press. Nottingham, p.
49-66.
547
CHAPTER XXI: DOG AND CAT DIETICS
21.2. 1. Consequneces of inappropriate feeding
21.2.1.1. Excess energy intake Þowerweight, obesity (obesitas)

SUMMARY. The nutrient requirements of dog and cat may vary within
extreme ranges. During urban way of living the balance of the three main-
taining factors of the normal live weight, namely the intensity of basal
metabolism, physical activity and energy intake gets upset, and the animal
grows fat. Degree of overweight, beyond weighing can be evaluated by con-
dition scoring and if possible, by the evaluation of the total body composi-
tion, using body mass index or DXA-measurement. There are scoring sys-
tems of 1 to 9 in use too, in these cases the target, optimum state is that
of body condition score (BCS) of 5. The generally accepted design of the
slimming diet is of cyclic-periodic one, which may leads to the achievement
of the target weigh during some months. The used feed should have a high-
er protein, amino acid, vitamin and mineral level (“fortified diet”) amount
The time available for feeding can be shortened and the physical activity
should be increased. The extra, uncontrolled feed intake must be prevent-
ed. The indigestible (ballast) ratio of the daily ration may be increased by
adding cooked vegetables, like carrot, sugarbeet pulp, eventually alfalfa
meal. The adjuvant role of high-fibre level is discussed and instead, the use
of hardly digestible carbohydrates (e.g. sorghum or barley starch of low
glycemic index) is applied. Anyway, the fat level should be reduced approx-
imately up to 10%. Moderate vitamin A suplementation, through the stim-
ulation of the expression of glucagon-like peptid-1 gene, support the weight
loss. The use of fat burner substances, like L-carnitine and organic chromi-
um, may also prove effective and liver protective at the same time. Author
shortly describe the available therapeutic opportunities, too.
INTRODUCTION. Today the obesity is a widespread diseases among dog and cat
population. According to a survey in 2006 34% of the adult dogs is obese or
overweight in the United States (LUND et al. 2006). The main reason for that
is that the dog is living together with man for 100 thousand and the cat for
8 thousand years, sharing more and more the accomodation, food and cer-
tain habits. It is just considered as natural that the eats not only if he is
hungry but also according to the dictate of customary law. Eating became a
549
social event. Human already eats not simply to cover its nutrient require-
ments. For him the food is at the same time the source of pleasure too and
the common meal is a collective-social event reinforcing the solidarity.
Parallel to the descibed citizens of the developed contries are less and
less forced to make physical and manual work and from „self-diligence” they
are less doing sport. Alarming and continually growing numbers of the
human obesity are known. Although with a time lag but the „medical histo-
ry” of the dog and cat shows a very similar picture. Really, with the domes-
tication these phenomena appear parallel to the domestication, let’s think
on the dog fed from the family table or on snacks. Similarly to its owner
(master) dogs and cats are not only eating more than necessary but they
also are moving less than in the nature. The third factor of the obesity devel-
opment is the basal metabolism and the consequent heat production are
decisively inheritable. This is the same in case of the companion animals,
since the level, combination and activity of background enzymes and hor-
mones differ among individuals and even amon breeds, too. This is the rea-
son that there are breeds inclined to obesity, for example the Labrador
Retriever, Dachshund, Beagle, Cocker Spaniel, Dalmatian. The more and
more common castration, aging and the concomitant diseases (joint,
endocrin and cardiovascular) could intensify the susceptibility to get obese.
Our pets are fed on better and better commercial feeds which in many
cases is topped by table scraps, in the best possible case by industrial
treats. To make matters worse mot of the owner cannot accurately evaluate
his animal’s condition. Newertheless, one third of the companion animals is
overweight of obese in the developed countries. And which is more remark-
able and more important that in the majority of cases, especially elderly
people, the owners cannot be called slim. This will explain that for the suc-
cessful treatment of dog and cat obesity a change in living habit of the
owner is required, too.
ETIOLOGY AND PATHOGENESIS OF THE OBESITY

Obesity is the prolonged retention of more than necessary fat in the organ-
ism. Both criteria are important because a threshold amount of fat in the
females is necessary for the regular sexual cycle and conception (see
Chapter XVIII). Still the fat accumulation is explicitly necessary and useful
and serves as „fuel” for the migratory birds and as „feed replacer” for ani-
mals in winter sleep, like hedge hog or bear (WIDMAIER, 1998). Namely the
550
obesity develop if functioning of the homeorrhetic control assuring the con-

stant adult live weight and healthy body composition (icluded fat) is dis-
turbed. For each warm-blooded animal is characteristic a certain percent-
age of body fat content that is inherited and individual like the constant
body temperature. The three main members of supporting regulation (the
so-called „triad”) are the feed intake, the basal metabolism with the heat
production and the physical activity, the movement. Knowing that is under-
standable why drawing geese, migratory storks or swallows get never obese.
The most important physiological factors (only as a list) are as foloows: the
uncoupling proteins (UPC-1, UPC-2, UCP-3 enzyme proteins) producing
heat from the ingested feed instead of fat, the thyroid hormones controling
the intensity of metabolism, the GH, insulin and corticosteroids regulating
the carbohydrate, protein and fat metabolism. Concentrating on the mem-
ber of triad the most susceptible to influence, the feeding, it can be stated
that healthy animals basically eat according to their energy needs. This is
assured by the indicator porotein, the leptin, which is produced by the fat
cells (adipocytes) and which acts on the hypothalamic hunger-satiety cen-
tre.
The organism realizes its constant fat content basically by the regu-
lation of the feed intake. The discovery of the leptin was a landmark of
knowledge of the above mechanism (ZHANG et al. 1994). This adipocyte-pro-
duced protein hormone inhibits the hypothalamic hunger centre. The effect
realizes through decreasing the production of neuropeptide Y (NPY). Excess
energy intake increases the dimensions of adipocytes, in turn, the leptin
production is proportional to the size of fat cell. Through the melanocortin
4 (MC-4) high blood leptin level decreases appetite, increases basal metab-
olism and heat production, furthermore favours reproduction and the orig-
inal body composition will be restituted. During fasting opposite processes
can be observed. Shrinking fat mass is producing less leptin, consequently
the NPY secretion, the hunger and the feed intake increase, the basal
metabolism and the reproduction decrease. If sufficient feed is available, the
original body fat content will be re-established (FIGURE VI-1) Summarizing,
large fat tissue of the obese animal produces a high amount of leptin, which
in turn, influencing brain stops voluntary feed intake. If inherited causes or
bad habit prevent regular functioning of that mechanism, the individual
may get obese even besides a low feed intake.
Individual body composition is regulated by several genes; for exem-
551
ple leptin production of fat is controled by the OB (obes) gene. During evo-
lution after longer hunger periods the frequency of less leptin-producing
OB/ob, as well as the ob/ob genotype concluding to a total lack of leptin
increased in human and companion animal population. Partial or total lep-
tin deficiency signals for the hypothalamus if there were no fat tissue: the
hunger and the feed intake continually increase and the individual get
obese.
The descibed basic mechanism is refined by the discoveries of the last
decade: the gastric wall produced grhelin stimulates the appetite and the
hypothalamic centers are able to perceive the ratio of leptin to ghrelin in the
blood (EPELBAUM, 2007). Besides leptin adipocytes are producing a series of
hormon-like substances, like TNFα (tumour necrosis factor alfa), IL-6 (inter-
leukin-6), adiponectin (complement-dependent adipocyta-protein or Acrp-
30), sexual steroids, glucocorticoids, blood clotting and complement factors,
angiotensiogen and resistin. The latter being the antagonist of insulin may
have a particular role in development of secondary diabetes. According the
localization and type (α, β/γ, δ) of the PPAR (peroxisoma-proliferator akti-
vated receptor) the metabolic role of hepatic, brown fat tissue, visceral and
subcutaneous fat is different (FEKETE and BROWN, 2007). This is the visceral
fat mass which predisposes to metabolic troulbles (LAFONTAN and BERLAN,
2003). In addition to that several compounds are synthetized in the hypo-
thalamus regulating voluntary feed intake. It is useful to classify these com-
pounds according to their effect, i.e. appetite-stimulating (orexigen) and
appetite-inhibiting (anorexigen) (BESSESEN, 2004). The corticotropin releas-
ing hormone (CRH), the pro-opiomelanocortin (POMC), e alpha-melanocyte
stimulating hormone (α-MSH), the cocain-amphetamine-related transcrip-
tion factor (CART) and the serotonin decrease, the melanocortins (MC4,
agouti peptide, agouti-like peptide), the melanin concentrating hormone
(MCH), the above mentioned neropeptide Y (NPY), the galanin and orexin A
and orexin B increase feed intake (SCHWARTZ et al. 2000). The gastrointesti-
nal peptides from the gastric and intestinal wall have also an important part
in regulating feed intake (NEARY et al. 2004). The above mentioned ghrelin,
the YY peptide (PYY), the glucagon-like peptide-1 (GLP-1) and the cholecys-
tokinin (CCK) are involved in the control of appetite, too. The CCK causes
satiety only for some hours; PYY at the same time maintains satiety feeling
for half day. The GLP-1 is produced in the enterocytes from the pro-
glucagon. Beyond its anorexigen effect it stimulates the insulin and inhibits
552
the glucagon secretion in the pancreas.

Let’s ake a simple calculation to evaluate how many genotypes may
exist if only the following factors are involved: body composition, factors of
voluntary feed intake, the orexigen and anorexigen compounds, UCP-1,
UCP-2, UCP-3, T3, T4 and T3-dejodinase, all together approximately 25 to
30 elements. If the localization of genes is disregarded the number of possi-
ble combinations equals to 2n-1=230-1=229 =536’870912 (more than five
hundred million) in the dog and cat populations. This is why the direct con-
nection of feeding, lifestyle and obesity cannot be recognized in a given indi-
vidual and why the dietary treatment does not prove efficient in some cases.
These facts suggest the involment of veterinarian in the treatment of obese
animals, who is trained to prescribe the personalized diet and is able to
decide about the possible medical treatment. FRAYLING et al. (2007) have
found the FTO gene predisposing to obesity. The name came from the leg
deformity of mouse (fused toe), localized on the chromosome in a homolo-
gous position. It is not clear that the FTO gene is indicator or regulator one.
RECOGNITION AND EVALUATION OF OBESITY

Despite the general view that this is obvious, it is more complicated in the
praxis. Namely, the overweight owners are inclined to excuse more to their
animals, too; moreover several non-infectious diseases may lead to similar
appearance. The golden rule is the live weight should be related to the breed
standard. In the light of the above described it is evident that after the
occurrence of body weight changes (getting fat, possible loss of weight) the
dietary medical history may be very helpful in finding the triggering causes.
In all cases is useful the answering of the questions of dietary anamnesis
(Table XXI-3) in finding the causes and deciding about the necessary meas-
ures. How can be the overweight or obesity quantified? Certainly, the most
simplest is the weighing and comparison of the result with the breed stan-
dard. It is an overweight if the live weight exceeds the breed standard by
15%; above that it is called obesity. The body condition scoring (on a scale
of 1 to 5), the feline body mass index and the dual X-ray absorptimetry
(DXA) measuremt give more accurate result. In the human medicine or in
the experimental veterinary medicine impedance analysis, isotope dilution
techniques, CT and (N)MR measurements are in use, too.
Both for dogs (LAFLAMME, 1997a) and for cats (LAFLAMME, 1997b) a
body condition scoring systems on a scale of 1 to 9 are developed. For the
553
classification of animals the palpability of ribs, lumbar vertebrae and hip

(pelvic) bones, thickness of the subcutaneous fat layer and the filling of tail-
head with fat tissue, the abdominal line from side and contour (outline) of
the waist (from above) should be evaluated. Emaciated, cachexic individual
receives the body condition score 1 (BCS 1), while there is not only a loss of
the fat, but also that of the muscle mass. The BCS 2 is the very thin, the
BCS 3 the thin, the BCS 4 is the slight loss of weight, the BCS 5 the ideal,
the BCS 6 the slight overweight, the BCS 7 the overweight, the BCS 8 the
obese the the BCS 9 the heavily obese animal.There is also a more simple
system, using only five catagories (Table XXI-4) Although there are predic-
tion equations using zoometric data both for dogs (BURKHOLDER, 1994) and
for cats (STANTON et al. 1994), in the practice only the more simple but still
reliable feline body mass index (FBMI) (BUTTERWICK, 2000) came into use.
The Body Fat, % =(Rib Cage/0.7067-LIM)/0.9156-LIM, where the Rib
Cage=girth length at rib 9 in cm, the LIM=leg index measurement, i.e. the
distance between the patella tuber calcanae in cm. For example if the
parameters for a cat of 3.45 kg LW, and of 5.1 kg LW the measure Rib Cage
is 36.5 and 44 cm, the LIM in turn, 13.9 and 14.8 cm, then the calculated
total body fat content is 27 and 37%, respectively. Cats having more than
30% of total body fat are considered as obese.
Recently GERMAN et al (2006) developed reliable method for dog and
cat body condition evaluation. This SHAPE-method is simple enough to be
familiarized with the owners, too. The acronym „shape” came from the
words „size”, „health”, „and”, „physical” and „evaluation”. Basically the
SHAPE-system uses the same physical and visual examinations than the
previous systems (see www.petslimmers.com/shape. htm), but by means of
exact instruction and yes/no decidable questions leads the investigator to
determine properly the thickness of the fat layer covering the ribs, spines of
vertebrae, subcutaneous, abdominal and waist. The algorithm asks for the
possible locomotor troubles (lameness) and health impairment, too. The
elaborated 7 condition categories are marked by capital letters. The „A”
stands for the very thin, the „B” for the thin, the „C” for the slightly thin, the
„D” is the ideal, the „E” means the slightly overweight, the „F” the obese and
the „G” is the heavily obese. To the individual categories belong dietetic
advices and the recommended time of the veterinarian’s supervision (imme-
diately or during the next regular visit etc.)
554
DIFFERENCIAL DIAGNOSIS. Given the fact that several other syndrome, meta-
bolic and endocrine troubles may cause obesity, the following diseases and
physiological states should be excluded by clinical investigation and analy-
ses during the differencial diagnosis: Cushing’s syndrome (hyper-adreno-
corticism). constipation, pregnancy, (stationary) corpus luteum persistent,
hypothyroidism, diabetes, acromegaly, abdominal tumours, ascites (e.g.
due to congestive heart failure, lack of protein, pyridoxin, L-carnitine and
the taurin in cats) The long-term applied, weight gain stimulating drugs
should also be taken in account. The most important among them accord-
ing BRAY (2004) are as follows: neuroleptics (tioridazin-HCl, olanzepin, que-
tiapin fumarate, risperidon, clozapin), antidepressives like the tricyclic
drugs (amitriptilin-HCl), the mono-amino-oxidase inhibitors (nortriptilin-
HCl, imipramin) and the inhibitors of selective serotonin re-uptake (mir-
tazapin, paroxetin-HCl), the anticonvulsive medicines (valproat, carba-
mazepin, gabapentin), the antidiabetics (insulin, sulphonilurea, gitason),
the serotonin antagonists (pizotifen), the antihistamines (cyproheptadin-
HCl), beta-adrenerg blockers (propranolol-HCl), alfa-adrenerg blockers (ter-
azosin-HCl) and the steroid hormones, (progestins, glucocorticoids.
In the manifestation of obesity the genotypeÎenvironment interaction
is fundamental. In this case the major components of the environment are
the feeding and the physical activity. The obesity is a chronic, recurrent
metabolic trouble that facilitates at the same time the development of sev-
eral other ailments like that of the cardyovascular system, joints, dys- and
hyperlipidaemia, atherosclerosis in human and cats, the disposition of the
parenchymatic organs for tumour augments, there is an increased suscep-
tibility for Type II diabetes, even the impairment of immune system is com-
mon and the life span shortens. The insulin sensitivity of adipocytes is
inversely proportional to their size. The consecutively or concomitant occur-
ring hyperinsulinaemia, diabetes, hyperlipidaemia, high blood pressure,
gout and a high blood concentration of plasminogen activator inhibitor is
called metabolic syndromes.
DIETARY TREATMENT. After all, obesity is not only an aesthetic question, but it
damage the quality of life and shortens the life span of of our companion
animals, therefore it should be treated. The successful slimming diet and
the subsequent maintenance of live weight are based on the accurate feed-
ing technique, diet composition, included the the functional feeds, not for-
get of course to the regular exercise, either. Perhaps the most important
555
prerequisite is the compliance of family members to avoid the so-called

„begging dog syndrome”. There are several appropriate feeding techniques:
a slow, gradual decrease of the daily ration or the low energy supply (dog:
60%, cat 75%) or the rotation of just sufficient (maintenance) portions with
the low energy supply every 2-3 weeks. Essential is that the weight reduc-
tion should not be drastic because, particularly in case of obese cats, may
lead to a fatty infiltration of the liver, which is life threatening! In optimum
cases weight loss in dogs is equivalent to maximum 2% LW and 1 to 3 % in
cats weekly.
Among the sliming diets the cyclic periodic schedule proved suitable.
First a target body weigth is determined which corresponds to the breed
standard. After that the corresponding maintenancs energy requirement is
calculated and 60 % (dog) or 75% (cat) of that is given for 2 to 3 weeks. This
reduced energy requirement can be calculated directly by using the fol-
lowing equations: 0.544×0.6×W0.75 MJ ME for dogs and 0.293×0.75×LW=
0.22×LW MJ ME for indoors and 0.335×0.75×LW= 0.25X×LW MJ ME for
outdoors cats, where W0.75 is the metabolic body size and the LW is the live
weight in kg (data from NRC, 1986 and 2006). Daily rations containing the
reduced energy contant is given for 2 weeks, followed by the daily rations of
100% maintenance energy requirement in order to help liver in recovery
from the fatty infiltration.
The feed of the slimming diet should be low in energy and especially
in lipids (no more than 10% of the dry matter) and rich in protein, amino
acids, complex carbohydrates, minerals and vitamins („fortified diet”).
Increased level of protein, minerals and vitamins is necessary to assure the
satisfactory supply even on the low dry matter intake. Furthermore a high-
er than usually fibre concentration (10 to 13% crude fibre) and/or the use
of hardly digestible starch of low glycaemic index may decrease the diets
energy concentration. The lowest postprandrial glucose peak were detected
after feeding sorghum, corn and barley grain. The lowest insulinaenia was
caused by feeding barley, corn and sorghum; the wheat starch moderately
and the rice strongly raised the blood sugar and insulin concentration
(BOUCHARD and SUNVOLD, 1999; 2000). In the dog foods rather barley and
sorghum and in the cat food preferably corn and sorghum is recommended.
Differences in the physiological effect of starch from the various sources are
due the differences in starch crystal structure. Owing to the low fat content
the preference of slimming foods are generally bad, therefor a flavouring is
556
frequently necessary.
Functional feed additives (nutriceuticals). Similarly to the human „fat
burner” preparations, the supplementation of dog and cat food by vitamin
A, L-carnitine or organic chromium efficiently help the slimming process.
A slight overdosage of the vitamin A actually stimulates the expression of
beta-adrenerg agonist UCP-1 gene: a a consequence, the heat production
and energy expenditure will increase. L-carnitine brings the long-chained
fatty acids into the mitochondria, the site of oxidation. Trivalent chromi-
um (chromium nicotinate, picolinate or in form of yeast) as part of the glu-
cose tolerance factor (GTF) by means of the phosphorilation of the receptors
increases the insulin sensitivity of tissues. As it is obvious from the above
described, the home-made diet with its unknown and unpredictable com-
position cannot be recommended for the purposes of slimming diet.
Otherwise the whole process should be realized exclusively under the con-
trol of a veterinarian.
After a successful, 3 to 12-month-long slimming diet there is a threat
of the so-called „yo-yo” effect: after having accustomed to the poor energy
supply the organism utilizes nutrient more efficiently and even fed on a nor-
mal diet, will put on weight again. For the prevention of the „yo-yo” effect,
light, maintenance and weight control diets were developed with a moderate
energy concentration (ranging between the normal and the slimming diet).
During this period the regular walk, exercise, outdoor activity is essential.
According to established opinions in „post obes” conditions the amount of
physical activity should be increased by 50% to assure the maintenance of
the ideal body weight. The keeping of the calculated daily ration is extreme-
ly important, because the quite small but regularly ingested amounts of
surplus energy are added. Since 1 kg adult weight gain corresponds to
approximately 32 MJ ME (NEWMAN, 2004) or 2 kg of dry food, the regular
daily intake of 10 gram will result in 200 days in a overweight of 1 kg! If the
social relation of owner and pet should be in any way supported by snacks
or treats, their amount should be included into the daily ration. If no con-
tra-indication like some cases of urolithiasis, the moist diets are preferable
to the dry ones.
Supplementation of the diet by fermentable fibre or fructose
oligoszacharides (FOS) sources like sugar beet pulp, carrot, Jerusalem arti-
choke improves the sugar metabolism by stimulating the proglucagon pro-
duction in the colonicytes through the released bacterial short-chained ??
557
fatty acids in the large gut. The proglucagon, in turn, transforms into
glucagon in the epithelium cells and will increase the insulin secretion and
decrease appetite (MASSIMINO et al. 1998).
CAT – NEW APPROACH. Cats, compared to the less strict carnivore dog, can be
considered as an absolute carnivore. It is showed, among others that except
the pregnancy, practically cat does not require carbohydrates, because the
necessary blood sugar can be produced by gluconeogenesis. In an average
adult cat food one-third of the energy is given by protein, one-third by fat
and one-third by carbohydrates. Under such circumstances the energy
needs are covered in the first place by the carbohydrates and secondly by
the fats; proteins are not or hardly used as a source of energy. While
decreasing the proportion of carbohydrates in the diet (up to the 10 to 15%
of the total energy) and increasing the percentage of proteins and fats (up to
40 to 45% of the total energy), cats will use fat as the primary energy source
and the excess protein as the secondary energy source. Nevertheless, this
method slightly and within the physiological ranges increased the blood
urea level (from 6.8 to 7.8 mmol/l), a weekly approximately 1% weight loss
could be achieved (LAFLAMME and HANNAH, 2005). This approach is similar to
the human Atkins- diet that consequently and undoubtedly proved efficient,
but considering that humans are omnivore, in a long-term it is strongly dis-
cussed in view of the possible side effects (ARBOR, 2003).
MEDICAL TREATMENT. In the human medicine, beside the diet and the appro-
priat exercise, the medical treatment is permitted if the body mass index
(BMI=body weight in kg/height2 in m) is higher than 27. The amphetamine-
like in structure phenteramine (Fastin, Apipex-P) enhances the effect of
epinephrin and norepinephrin the the brain leading to a decrease of
appetite. The sibutramine (Meridia) is an effective inhibitor of the serotonin,
dopamin and norepinephrin re-uptake and in a peroral dosage of 5 to 15 mg
it is a good drug of the human obesity. The sibutramine is not exempt from
side effects: it may cause increased blood pressure and rapid pulse, there-
fore it is not recommended for hypertensic patients. The phenfluramine
(Pondimin) and DEX-PHENFLURAMINE (Redux, Isolipan capsule) have a similar
mode of action. Their authorization were withdrawn owing to the side effect
on the cardiac valves and the danger of primary pulmonary hypertension
(PPH). The combination of phenfluramine and phenteramine, the „phen-
phen” was substituted for the combination of „ma huang” plant ephedrine
558
alkaloid and the St. John’s wort (Hypericum perforatum) („herbal fen-phen”).
The orlistate (Xenical) is a gastrointestinal lipase inhibítor, altering fat
absorption. Taken in before or simultanously the meals in a dosage of 120
mg, it decrease fat absorption by approximately one-third. In case of a high-
fat diet it may cause flatulance and a loose faeces consistence.
Although high doses (60 mg/kg LW) of dehydro-epiandrosterone
(DHEA) lower body fat, but owing to the limited knowledge about the side
effects it cannot yet be used. The situation is the same concerning the
course of injections using recombinant human leptin (DIEZ and NGUYEN,
2006). Promising studies are carrying out by using the PYY-analogue (PYY
3-36), the GLP-1 receptor antagonist exendin-4 and the inhibitor of the
dipeptidil peptidase-4 (DPP4). By means of the latter the biological half-life
of GLP-1 can be prolonged (NEARY et al. 2004). The fat-reducing effect of the
conjugated linoleic acids (CLA) is still proved only on pig (FEKETE and BROWN,
2007). There are promising experiences in the human medicine about the
lipogenesis inhibition of alpha-hydroxi-citric acid from Garcinia cambod-
gia. (WESTERTERP–PLANTEGA and KOVACS, 2002). Phytanic acid from the par-
tial degradation of the plant chlorophyll acts against fat accumulation
through the peroxysome-proliferator activated receptors (PPAR). According
to the experiments this branched-chain fatty acid and its metabolit, the
pristanic acid are promising both in the tratment and prevention of dog
and cat diabetes (DEKEYSER et al. 2003).
The mitratapid (Yarvitan®) placed recently on the market inhibits the
functioning of microsomal tryglyceride transferprotein (MTP) in the entero-
cytes. As a consequence the lipid absorption will be inhibited and it will
remain in the enterocytes cytoplasma. Desquamation of the gut surface
these cells will eliminate the encapsulated fat, too. The feeling of fullness
increases. Treatment should be realized according to the cyclic-periodic
schedule, but exclusively at dogs. As side effects, vomiting, diarrhoea and
appearance of soft faeces; in this cases the application of medicine should
always be done after meals. Contra-indications include liver injury, preg-
nancy, lactation and young age (below 18 months). Certainly, it cannot be
used if the obesity is caused by endocrine disorder like Cushing’s syndrome,
hypothyreosis etc. The effect of dirlotapid (Slentrol® has a similar mode of
action as the mitratapid. The compound acts on the energy metabolism in
the enterocytes. Since the side effects are not uncommon, mitratapid and
dirlotapid should be used only by the veterinarian.
559
Table-XXI-3: Dietetic questionaire
560
Table-XXI-3: Dietetic questionaire (cont)
561
Table XXI-4 Body condition scoring of dogs (adapted from Ohio State Univ,
www.nssvet.org/food/bcs.html)
———————————————————————————————————————————
1. Emaciation
Enfeeblement, general weakness.
Striking signs of emaciation: ribs, processus hamatus of scapulae, lumbar verte-
brae, pelvis and hip bones and all prominences evident from a distance.
Lack of palpable subcutaneous fat. Obvious loss of muscle mass.
2. Thin
Hip bones are not prominent
Thurl and pelvis (hook or hip bone and thurl) is rounded; easily seen and felt.
Only some fatty tissue felt under the skin.
3. Optimum (moderate)
Neck and shoulder are well separated from the rump, but the transition shows a
smooth line.
Obvious waist and abdominal tuck when viewed from above and from aside.
Ribs, spines of lumbar vertebrae, ??lapockatövis, pelvic bones are not visible, but
easily palpable.
Slight fat layer is palpable under the skin of chest and abdomen.
4. Overweight (stout)
Ribs are invisible, but palpable with difficulty.
Pins visible with difficulty, but palpable, covered by slignt fat deposit.
General fleshy appearance; waist and abdominal tuck may be absent.
Noticeable fat layer on ribs, lumbar spine and on tailhead
5. Obese
No part of ribs, lumbar spine and pelvis can be felt even with firm pressure
Large fat layers over chest, spine, neck and limbs.
Tuck and abdominal wait absent; oval shape is visible from above
Large fat deposit under the skin of chest and abdomen. At the loin area folds of
fatty tissues are over short ribs and the bone structure cannot be felt
Tailhead is buried in fat tissue, skin is distended.
———————————————————————————————————————————
21.2.1.2. Deficient energy and protein supply

The insufficient energy and protein supply (PEM=protein energy malnutri-
tion) leads to emaciation (inanitio). It is common in animals of advanced
chronic kidney disease and different tumours; for details see in those units.
21.2.1.3. Deficient mineral and vitamin supply

The most common mineral deficiency, especially of growing puppies and
kittens, the lack of calcium, alone or combined with vitamin D deficiency.
In young animals rickets (rachitis), in adults osteomalatia develop. The
growing plate of the long, tubular bones shows incomplete mineralization;
bone density decreases. Limb bones become bent, twisted, the epiphyseal
cartilage overgrows and in serious cases „ricketsy rosary” form at the car-
tilageal border of rib bones. The most importan part of the therapy is the
provision of sufficient dietary calcium in form of limestone (calcium car-
562
bonate), calcium hypophosphoricum or calcium lacticum. For more details

see „Nutrition and skeletal system” subchapter.
Lack of sodium may occur, if the home-made diet is not supple-
mented with common salt. The main consequence of this situation is a
reduced water intake and the danger of urilith formation. The vitamin E
(and/or selenium) deficiency is also more common in cats, than in dogs.
In most of the cases the fat or oil supplementation of feed is not accompa-
nied by the necessary elevation of vitamin E. From the poly-unsaturated
fatty acids free radicals emerge and affect, among others, the fat cell. A gen-
eral infalmmation (pansteatitis), later necrosis of adipocytes develop. As
secondary, degenerative process, yellowish ceroid pigment is deposited into
the fat tissue, giving the syndrome the „yellow fat disease” denomination.
During the clinical examination, cats show pain while palpating; diagnosis
can be reinforced by biopsy of the subcutaneous fat. Fever and lethargy are
common; the prognosis is bad. Long-lasting vitamin E and selenium treat-
ment may cure animals.
The lack of thiamine (vitamin B1) is not common in dogs; the most
susceptible are the growing kittens. In the nintieth years, the extreme heat
treatment of caned feed caused in the UK high incidence of ailment in kit-
tens. The first symptoms are atypical (anorexia, loss of weight), then after
1 to 2 days ataxia and short seizures characterize the syndrome.
Afterwards opistothonus develops, varying from time-to-time by falling
aside. In this phase there are brain damages like small petechia in the cere-
bellum and midbrain, but no cerebrocortical necrosis as in case of polioen-
cephalo-malacia of calves and lambs. Sinus bradycardia may also develop,
the prostreation and shortly will end in death of the animals. Treatment in
the middle phase is yet successful, by giving 50 mg thiamine per cat bid,
either iv, im and/or perorally. Vitamin B12 (cobalamin) deficiency may
occur in in cats fed on feedstuffs of plant origin. The poor intestinal flora of
the cat cannot provide host animal and if the serum cobalamin concentra-
tion drops below 100 ng/l, deficiency sighns, like anaemia develop (GROSS
et al. 2000).
In dogs, receiving little feed of animal origin, deficiency of L-carni-
tine may develop, manifesting in dilatation cardyomyopathy. Brachio-
cephal breeds, like boxer are especially susceptible to this ailment. Cats,
fed on vegetable feedstuffs develop taurin deficiency. Taurine deficiency
may cause central retina degeneration (CRD), dilated cardyomyopathy,
563
reduced litter size and weight and impairement of the immune system. If
the taurine concentration in blood is less than 200 nmol/ml, there is a
marginal supply and clinical signs will be manifested if the plasma taurine
concentration drops below 40 nmol/ml (KIRK et al. 2000). Cats living on
vegetarian diet or on a feed mixture, consisting of predominantly vegetable
component, have ahigh risk of taurine and vitamin B12 deficiency.
However, deficiency syndromes are less and less frequent, because com-
mercial diets are supplemented by synthetic taurin and cobalamin
(WAKEFIELD et al. 2006). In the nature, the highest taurine concentration
can be found in the see-food („frutti del mare”), like oyster, molluscs, clams,
salt-water fish, but the fresh-water fishes and the viscera of warmblooded
animals contain large amounts of taurine, too. In feedstuffs of plant origin
virtually there is no taurine. In vegetarian families the occurrance of cat
taurin deficiency is more common.
21.2.1.4. HYPERVITAMINOSES
Dangerously high vitamin intake may occur owing to the failure of produc-
tion, when the commercialized diet contains extremely large amounts of
fat-soluble vitamins, but in the practice it is rather the vitamin A poison-
ing problem pertains to cats, ingesting huge amount of liver for a longer
period of time. For the prevention, it is a thumb rule, thath the daily vita-
min intake should not exceed 100 to 300 IU/kg LW in case of growing kit-
tens and 100 IU/kg at the adults. The index of nutritional quality (INQ, see
Chapter I) of the liver moves within the range of 100 to 120, which means,
that eating exclusively liver, the vitamin A ingestion is 100-200 fold of the
requirement. Owing to the developmental troubles of bone and periosteum
exostoses grow on the limbs and vertebrae. The latter may compress and
affect afferent and efferent nerves of the cervical spinal cord. Sick cats are
anorectic, limp and because of the painful movement of the neck, do not
„wash”; as a consequence, their hair coat is dishevelled and frequently turn
into exsudative eczema. Besides shuting down the vitamin A intake, the
regeneration of liver can be supported by lipotropic substances (methion-
ine, choline), but the prognosis generally is unfavourable.
Vitamin D oversupply will cause the calcification of the soft tissues,
primarily of vessels and kidneys. It is not uncommon, that the ailment is
iatrogenic, if the veterinarian, giving vitamin D injection, do not prescribe
the peroral calcium supplementation. The daily calcium and phosphorus
564
requirement for growing puppies 175 to 460 mg/kg LW calcium and 120 to
290 mg/kg LW phosphorus, for adult dogs for maintenance 100 mg/kg LW
calcium and 85 mg/kg LW phosphorus. Since the raw materials of feed
mixture (meat, cereals) have enough phosphorus, generally only calcium
supplementation should be given.The daily vitamin supply should not
exceed the 20 mg/kg LW for growing and 10 mg/kg LW for adults dogs.
21.2.1.5. UNBALANCED DIET

Surpassing the sucrose (saccharose) tolerance limit (5-6 g/kg LW), for
example by giving cakes and chocolate („snacking”) results in osmotic diar-
rhoea. With the advancing of age, especially in cats, lactose intolerance
can be observed. The intake of dairy products should be limited, in order
to have no more than 1 to 2 g/kg LW lactose daily (practically 20 to 40 ml
cow milk/kg LW). Milk derivates, having no lactose content, like cotton
cheese, chees, yogurt naturally can be fed. Eating high amount of bone
(more than 10 g/kg LW), especially accompanied by poor physical activity,
will lead to grave obstipation.
Exclusive, or predominant feeding of boneless (lean) meat results in
calcium and iodine deficiency („all-meat syndrome”). Calculating with the
above numbers, the daily calcium requirement for a growing puppy of 10
kg LW is 1.75 to 4.6 g Ca; whereas 100 gram of lean beef contains 0.02,
100 gram of liver 0.01 g Ca (N.B. 100 g of cow milk has 0.74 g Ca!).
Continuous calcium deficiency stimulate parathyroid gland to overproduc-
tion (secondary alimentary hyperparathyroidism). High blood parathor-
mone (PTH) concentration cause bone mobilization in the whole body.
Decreased bone density frequently leeads to cracking of vertebrae or/and
long, tubular bones. Diagnosis can be reinforced by X-ray or CT-picture.
Dietary treatment means a transient calcium overdosage: besides assuring
the absolute phosphorus needs, the calcium to phosphorus rati should be
set to 2 to 1, by using limestone (calcium carbonate) or calcium lactate.
(Application of calcium phosphoricum is not advisable, bacause of its phos-
phorus content.) After recuperation (some weeks), the Ca:P ratio should be
adjust to the optimal 1.2-1.4 to 1. Iodine deficiency can be alleviate by
adding potassium iodine, iodinized salt, iodin-starch complex, or iodine-
EDTA.
565
21.2.1.6. DEGENERATIVE DISEASES OF DIETARY ORIGIN

High fat diets. Although. the dog is used as a model animal during human
atherosclerosis research, the occurrance of hyperlipidemia and atheroscle-
rosis in the practice is uncommon, it can be rather found as an accompa-
nying disease of hypothyreosis. The cat’s lipid metabolism is more similar
to that of human, consequently the incidence of aetherosclerosis is higher,
too. High phosphorus level of feed, especially in old dogs and cats with
chronid kidney disease, may result in osteodystrophy and nephrocalci-
nosis.
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21.2.2. Dietary treatment of pathological conditions
21.2.2.1. Principles of feeding sick, febril animal

Dogs and cats with a higher body temperature commonly are anorectic.
Therefor, the first taks in their treatment is to stimulate voluntary feed
intake. The fundamental rule is, that the fortified diet should be given, i.e.
high in energy, vitamins and minerals, having high digestibility and excel-
lent biological value, good tast and low indigestible fibre content. In the
practice the use of junior feed (designed for puppies and kittens) may satis-
fy these requirements. Feed should be given at body temperature (38-39oC),
because tastes are more intensive in this state. Fat or oil addition is
favourable, because not only increase energy density, but also improve pref-
erence. Feed shoul be offered frequently. Feeding more dogs at the same
time also imcrease feed intake („jealousy appetite”). Residues should be
removed after 15 minutes of distribution. In final case the enteral tube feed-
ing (using naso-oesophageal, pharyngostomy, gastronomy or jejunostomy
tubes) should be applied (Table XXI-8, see later).
21.2.2.2. Diarrhoea and constipation

Ingestion of too much bones (in case of indoor-kept cat: hide and feather)
results in obstipation. The animal excretes less frequently and the faeces
consitency is dry (according to the scoring system, 0.5 to 1.5, see
Subchapter 2.1.). The natural treatment is the giving of cooking or paraffin
oil by spoon, perorally, or in serious cases the use of purgative medicines,
like sodium or magnesium sulphate. In very refractory cases parenteral
application of gastrointestinal prokinetic drugs (e.g. ranitidine, nizatidine)
may help. For dogs and cats, especially in old age, if they are prone to the
constipation, the daily ration should contain more fibre (whole bread, fruits,
vegetable, wheat bran) and organic acids (cottage cheese, yogurt, buttermilk
etc.).
Diarrhoea may be caused by infectious agents, like bacteria and
viruses, but it can be created as a consequence of unbalanced diet alone.
Ingestion of too much readily fermentable carbohydrate results in fermen-
tative (sucrsose diarrhoea), too much protein in putrefying diarrhoea. Too
low indigestible fibre (ballst) level (2 to 3% in dogs and 1.5 to 2% in cats), as
well as the abrupt change of feed or feeding regime predispose to dysbiosis
and to a secondary diarrhoea. Water content of the excreted faeces increas-
569
CHAPTER XXI: DOG AND CAT DIETETICS
es up to 90% (the normal being 20 to 30%). For the exact description of the
faeces consistence the use of faeces consistency score (Waltham Centre for
Pet Nutrition) is appropriate, which takes into account the outside, the
hardness, the water content and the possibility how to pick up from the
ground (with or without leaving mark). Feaces of scores 3.5 to 5 signify mild
to watery diarrhoea.
Non-infective diarrhoea can be treated by a 2 to 3 days of fasting,
besided ad libitum drinking water supply. If the excretion of watery faeces is
accompanied by vomiting, instead of water, black tea, or equal mixture of
black and camomile to should be offered. Additional therapy may includes
mucosal protectans and absbants (kaolin-pectin, activated charcoal, bis-
muth subsalicylate). In serious cases tannic acid-containing preparation
(e.g. Pulvis quercus adsorbens FoNoVet; Farmatan plus® from Tanin
Sevnica d.d. and Matischa Ltd.) can be added to the tea. As fluid therapy,
0.5-0.6% sodium chloride and 0.2-0.3% sodium bicarbonate solution or tea
can be given. Solutions of human preparation, like Normolyt powder (con-
taining glucose, sodium chloride, sodium citrate and potassium chloride)
can be used with success. Use of motility modifying drugs (diphenoxylate,
loperamide: but in case of infectious diarrhoea they are contraindicated)
may help in attenuating the clinical signs (symptomatic treatment).
21.2.2.3. Gluttony, anorexia
Dogs’ gluttony is basicly a conditioned reflex, built up by the master; how-
ever, some breeds are more predisposed, like beagle. Decrease of the ener-
gy density of feed by mixing fibreous feeds (vegetables,wheat bran etc.) and
grabing of the dog’s attention using bones, artificial bones or treats may
prove helpful.
The independently occuring anorexia the most common at male dogs,
when in the neighbourhood there is a (rutting) bitch in heat. In these cases
3 to 4 days of fasting can be observed. It can be useful to apply 10 -15 min-
utes before the planned feeding preparation, stimulating gastric secretory
function (e.g. Tinctura stomachica FoNoVet or Tinctura amara FoNo, or
pressed orqange peel), but in persistent cases the parenteral stimulation
(0.02-0.05 mg/kg of LW diazepam im. or iv.) cannot be excluded.
In cats the development of feed aversion is easy, if around the feeding
they receive medicines, causing unpleasent feelings (e.g. tetracyclins, eryth-
romycin, metotrexa, doxorubycin). One of the most important side effect of
the hepaticus lipidosis the feed aversion; the tube feeding must be started
570
immediately and for 48 hours all feed (residue) should be removed from the
animal. If changing the feed, the new brand should be offered only in famil-
iar environment. The enzyme-treated animals viscera (digests) are effective
in enhancing the preference of commercial cat feeds.
At the same time, the anorexia is the most common introductory and
accompaniyng signs of most of the cat diseases; having feed refusal, the
appropriate diagnose and the correct treatment of the main disease is
essential. Special attention should be given to the alleviation of pain of any
origin and to the fluid therapy. As a practical measure, pulpy feeds (e.g.
smashed meat, oil fish, corn beef, human baby food) can be smeared on a
paw, because it will be licked off with great chances. Feeder should be shal-
low and wide to avoid the touching the rim of feeder dish by the whiskers.
Only in final cases (e.g. toumor patient) is proposed the medicinal tratment:
diazepam (0.1 to 0.4 mg/kg of LW iv.) to induce the direct feed intake,
oxazepam (twice a day 2 mg/cat per os) or cyproheptadine (once of twice 2
to 4 mg/cat per os daily) for a continuous, long-lasting stimulation of eat-
ing (MARKS, 2002).In case of the troubles of liver functions the application of
diazepam-derivates is contraindicated!
21.2.2.4. Gastric dilatation and volvulus

This ailment is commonly found at the individuals of dog of large or giant
breeds, having deep, broad-chestedbody, for example Great Dane, Borzoi,
Bloodhounds, St. Bernard, Irish Wolfhound, German Shepherd etc.). There
is an air accumulation in the stomach, which has been related to aeropha-
gia. The role of diet type is unknown and the irregular functionning of the
smooth musculature of the stomach is involved into the development of the
ailment. The prevention consists of the offering of the daily ration into 2 to
3 portions; the feeder should be placed high, in order to force dog to eat with
an uprisen head. Calm environment should be assured during eating and
after feed intake at least 1.5 hours rest should be applied. According many
oppinions, commercial feeds, containing high level of vegetable raw materi-
al (soybean, corn gluten, corn starch etc.) should be omitted.
21.2.2.5. Congestive heart failure

Etiology. Circle of causes of congestive heart failure is very varied, more
than 30 different heart insufficiency causes can be mentioned. Among oth-
ers the dilatation cardyomyopathy can occur as deficiency syndrome (L-car-
571
nitine in dogs, taurin in cats, vitamin E/selenium in both). Accordingly, the

primary treatment is also diversified. Anyway, because of congestive heart
failure in most of the cases is accompanied by sodium and water retention,
which can be influenced by dietary treatment, preventing, alleviating or cur-
ing ascites and hydrothorax.
Dietary treatment. The sodium intake should be reduced to a mini-
mum (max. a sza. 0.07% of DM), i.e. if using home-prepared diet, only
boiled noodles, pasta, rice and potato can be given. From the dairy prod-
ucts, the cream and the white of the egg may also be applied to decrease
sodium ingestion. The (common) salt concentration of the daily dry matter
cannot be higher than 0.175%. (To compare: a common feed contains at
least 0.5% salt.) From this point of view it is important to know that chees,
bread or table scraps have high sodium level. Low-sodium veterinary (pre-
scription) diets are also available. Since these animals generally receive also
a diuretic treatment, the excereted B vitamins and potassium should also
be replaced. To avoid the pressure of the stomach on the diaphragm, the
daily ration should be distributed into many small portions. According to
the recent date the additional arginine supply improves the cardiovascular
performance.
572
21.2.2.6. Feline Liver Diseases

© Géraldine Blanchard
METABOLIC FUNCTIONS OF THE LIVER. Due to its central place between the diges-
tive tract and the blood, and because of the variety of roles it plays, the liver
is one of the central metabolic organ in the body (FIGURE XXI-2).
FIGURE XXI-1: Metabolic functions of the liver
After digestion, the organic nutrients are absorbed. Schematically, the

hydrosoluble (HS) nutrients (amino-acids, glucose, hydrosoluble vitamins)
go thru the small intestine to the portal vein, when the lipoluble (LS) ones
(fatty acids, cholesterol, liposoluble vitamins) are transferred to lipoproteins
into the enterocytes, and secreted into the lymphatic system, and join the
blood flow via the thoracic duct, without going immediately thru the liver.
Thus the HS nutrients enter the liver before entering the whole organism,
when the LS nutrients have access to the peripheral cells first. The same
track may be followed by ingested drugs, depending in part on their solu-
bility. The liver is also irrigated by arteria coeliaca (a.c.), coming from the
aorta, and blood ends from the Iiver into the vena cava caudalis (v.c.c.),
573
which is the only vessel to leave from the liver.
The liver is involved in so many parts of the metabolism that it’s total
dysfunction cannot be compensated by any medical treatment and is rap-
idly fatal. Indeed, it is involved in process as divers as:
-digestion, by the secretion and excretion of bile and the entero-hepatic
cycle, intermediate metabolism, acting as a regulated filter of the nutrients
and other substances absorbed and entering the organism via the portal
vein,
- via the synthesis and the catabolism of carbohydrates -glyconeogenesis,
glycogen storage, lipids (incl. fatty acids and cholesterol), amino-acids and
proteins metabolism (Table XXI-5), the activation of vitamins (vitamin D),
but also by the possible storage of different nutrients (vitamin A, copper,
glycogen, cholesterol, triglycerides, etc.),
-detoxification of a variety of endogenous (amino acids catabolism, ammo-
nia and urea cycle, bilirubin) and exogenous compounds, excreted into the
bile or the blood flow.
Table-XXI-5: Major proteins synthesized by the liver (MICHEL, 1995) and their role
Finally, the liver has the ability to regenerate after injury, but its cen-
tral metabolic role in the organism makes its like of functionally impossible
to compensate and rapidly mortal.
The unique position of the liver, between the outside, represented by
the digestive tract, and the inside of the organism, makes it quite vulnera-
ble to pathologic and toxic agents and exposed to a variety of inflammatory
574
conditions. The liver diseases encountered in cats include so inflammatory

states, associated or not with infectious agents, metabolic disturbances due
to an overload like hepatic lipidosis, neoplasia, and unusual infectious con-
ditions (Table XXI-6).
Table XXI-6: Main hepatic and hepatobiliary diseases encountered in cats (CENTER, 1998;
WEISS et al. 1997; GAGNE et al. 1999; BEATTY, 2002)
————————————————————————————————————————————
Inflammatory
Cholangitis/cholangiohepatitis (suppurative or not
Chronic bile obstruction (of intrahepatic or extrahepatic aetiology)
Hepatic vacuolar change (amyloidosis)
Biliary cirrhosis (rare)
Not inflammatory
Hepatic lipidosis (primary or secondary)
Neoplasia (lymphosarcoma, carcinoma, round cell tumor)
Polycystic hepatic disease (Persians, Himalayans)
Porto vascular abnormalities (rare)
Toxic hepatopathy (endotoxins, drugs like diazepam, viral like FIP)
————————————————————————————————————————————
The general lack of hepatic function is, fortunately, unusual in cats. The
main liver diseases encountered in everyday practice are cholangiohepatitis
and hepatic lipidosis. The general nutritional response to a deficient hepat-
ic function will be presented, with a special regards to the nutritional treat-
ment of choloangiohepatitis and finally the management of cats with hepat-
ic lipidosis.
GENERAL NUTRITIONAL RESPONSE TO LIVER FAILURE

It is obvious that the primary cause of the liver dysfunction should be iden-
tified and treated as soon as possible. But as the diagnosis, and the full
recovery, can take days or weeks, the daily priority is to feed the cat, in
order to not add a malnutrition state to the underlying liver disease.
Anorexia is frequent in sick cats, thus, cats presented for a liver disease are
often already anorectic at presentation. At that time, the risk of hepatic lipi-
dosis is real (see further in this subchapter), if not already present, espe-
cially if the cat is or has been obese.
As a general consideration, the components of the diet, in any case of
liver dysfunction, must be of high quality. This goal is to minimize the waste
and the amount of nutrient that arrive to the liver, from the digestive tract,
but also those from the entire body. This goals can be reached by four com-
plementary ways: first by providing a high quality, high digestibility diet
575
either formulated for cats or home-made (Tables XXI-7 and XXI-8) (MARKS et
al.1994); second by providing enough food to cover the essential nutrition-
al requirement; third by feeding small amounts of food at a time; and forth
by insuring an appropriate hydration.
Table XXI-7: Example of balanced home-made diets for a cat with liver dysfunction. Diets
1 to 3 contain 0.8 MJ ME each. They can be prepared for several days, frozen by daily por-
tions or by meals –the daily portion can be distributed in the ice cube vats, and defrost
overnight in the fridge to be distributed the day after, gently heated with hot water.
Ingredients with a * can be replaced by a mineral-vitamin supplement bringing Ca, all vita-
mins and trace elements.
Table XXI-8: Composition of the home-made diets presented in Table XXI-7, expressed
as g or mg per MJ ME
DM= dry matter; EE= ether extract; CP= crude protein; NFE=N-free extract, Arg=arginine
a) Providing a food of high quality

Different petfood companies propose diets for cats with liver disease. The
difficulty is generally to convince the cat that such a diet is good for him! To
improve the chance of success, it is better to keep the form of diet the cat is
used to (dry, canned or home-made); to propose a very slow change by mix-
ing more and more of the new diet with the previous one at each meal; not
to use syringe gavage, but prefer first a tube feeding until the clinical con-
dition ameliorates and then propose a diet the cat appreciate until it’s
requirement is covered and then move slowly to a diet appropriate consid-
576
ering the cat condition.

b) To cover the energy requirements
The maintenance energy requirement of an adult cat ranges from 60 to 40
kcal of metabolizable energy per kg of ideal body weight per day, for an adult
indoor cat. To consider the ideal body weight facilitates the covering of the
proper nutritional requirements, whether the cat is obese or too thin.
Nevertheless, in the last case, the body weight considered should not exceed
120% of the actual body weight, to prevent a risk of overload of the diges-
tive tract. The covering of the energy requirement of the cat implicates the
estimation of its ideal body weight, the calculation of its energy require-
ments and the knowledge of the energy density of the diet (MJ ME/ kg of
fed).
c) To feed small amounts of feed at a time
The distribution of small meals may be recommended to minimize the
amount of nutrients the liver has to manage at a time. To start with 5 to 6
meals per day seems reasonable. When a dry food is distributed, it can stay
available anytime. But even dry food can oxidized, and it’s better to renew
it regularly at least twice a day to keep it fresh and attractive. The amount
of food really ingested has to be controlled daily. If necessary, a tube feed-
ing procedure has to be started. The limit can be 24 hours of anorexia, the
ingestion of less than 70% of the energy requirement 3 consecutive days, or
a weight loss of 10% or more.
d) To insure an appropriate hydration
A good hydration is a warranty for a good perfusion of all the organs, and
ameliorate the elimination of toxics and metabolism byproducts from the
body. If the perfusion of isotonic solutions may be necessary in case of
dehydration, it can not be a issue. The best way to hydrate correctly the cat
is to make it drink or ingest water. To enhance the ingestion of fluids, a wet
food may be preferred, but only with the „agreement” of the cat! If the cat is
not used to it, small amounts of wet food can be proposed very progressive-
ly. During a tube feeding procedure, the tube may also be used to hydrate
the cat. After calculating the daily water requirement, once deducted the
water brought by the food, we input the water in the tube in at least 4 times
a day, at least 2 hours after the previous and half an hour before the next
meal. The volume of water should not exceed the volume of the coming
meal.
577
NUTRITIONAL CONSIDERATIONS ASSOCIATED WITH HEPATOBILIARY DISORDERS

The first step is definitely to feed the cat (see above). Some nutrients have
certainly to be discussed further, but, except for the energy source, very few
publications are available to attest their benefit in cats with a liver disease.
Protein and amino acids. Considering the obligate catabolism of pro-
teins in cats, the minimal protein requirement has to be covered, even in
cats with a liver disease. Except in case of hepatic encephalopathy, a high-
protein 30 to 40% protein in the dry matter (BUNCH, 2003), corresponding to
a protein to energy ratio of 17 to 21 grams protein/MJ ME may be benefi-
cial to cats with hepatic lipidosis, hepatobiliary disease and neoplasia. In
case of hepatic encephalopathy (except when associated with hepatic lipi-
dosis), the protein amount should be close to the minimal i.e. 15 to 17
grams protein/MJ ME, when the total energy requirement is covered. This
basal value can be increased progressively as far as the cat can afford. The
protein source determines the amino-acids provided to the cat. Some
authors (BUNCH, 2003; LAFLAMME, 2000) suggested to lower protein of animal
origin and provide legumes (soybean meal) and dairy (cottage cheese and
whey) protein, especially in cats with hepatic encephalopathy, but no pub-
lication is available to justify this proposition to date and the composition
in terms of amino-acids may be very close to those of lean meat.
Furthermore, the two amino acids which could be considered as lim-
iting for the synthesis of protein in cats with prolonged anorexia are argi-
nine and methionine (BIOURGE et al. 1994b), two essential amino-acids rep-
resented differently upon current feed ingredients (FIGURE XXI-3). As
shown in this FIGURE XXI-3, the most interested ingredients, which can be
consider as protein sources (those with at least 18% protein as fed) and
which are rich in both arginine and methionine are horse meat, whole egg
content and coalfish. The richest in arginine are the legumes, but they are
poor in methionine and their low protein content (7.8% in lentils, 3.6% in
peas) except for soybean meal, their richness in fermentable fibre and their
low palatability in cats make them of low interest except as complementary
protein sources in commercial diets.
578
FIGURE XXI-3: Arginine and methionine content of different feed ingredients, expressed in
g/100g proteins (ingredients containing as fed* between 10 and 20%*** more than 18% of
protein) modified from SOUCI et. al 2000)
Fats and oils. Lipids are the best source of energy for a cat and are
very palatable, too. Their is no known reason to lower fat in the cat’s diet
except in 2 cases, namely the cholestasis associated with fat malabsorption
clinically associated with steatorrhoea and cats with a lowered lipoprotein
lipase activity and hyperlipaemia. Cats with hepatic lipidosis are an excep-
tion to this feature (see below) as hyperlipaemia is current in this disease
but fat still has to be a preferential energy source.
Carbohydrates. Fat and protein are much better tolerated by the cat
than high-carbohydrate diets, even in pancreatitis (SIMPSON and MICHEL,
2000), as well as in hepatic lipidosis (BIOURGE et al. 1994) Thus, rapid sugar
should be avoided, but they are rarely given to cats. Complex sugars are
usable as the third energy source after lipid and protein. Starch sources
have to be cooked to insure a perfect gelatinization of starch granules and
enable its digestion. Dietary fibre is provided enough to stimulate the tran-
579
sit in the digestive tract, not too much to not dilute energy density.
Minerals. Kalaemia has to be followed carefully and potassium pro-
vided adequately with at least 0.6% K in the DM of the diet, and enteral sup-
plementation with potassium gluconate syrup (2 mEq K/day per os) when
necessary (LAFLAMME, 2000). A sodium restriction, less than 0.25% DM in
the diet can be beneficial in case of ascites. Zinc is a nutrient of special
interest in case of anorexia, as zinc deficiency itself may induce anorexia. If
the diet is not rich enough (minimal requirement of 75 mg Zn/kg DM), or in
case of deficiency, an oral supplementation with zinc gluconate (3 mg/kg
LW/day) or zinc sulphate (2 mg/kg LW/day) divided in 3 doses may be con-
sidered (LAFLAMME, 2000). Copper is normally stored in the liver, but in case
of chronic liver disease, the copper store may not be usable anymore and
can itself induce hepatic fibrosis. Copper level in the diet should be reduced,
and copper chelators (10-15 mg/kg LW D-penicillamine or 50-100 mg zinc
acetate separately from food, per os twice a day) may be of interest
(LAFLAMME, 2000).
Vitamins. Vitamin deficiencies are commonly found is patients with
liver disease. As cats are very easily anorectic, may suffer of malnutrition
but can generally not consume or receive enough food within the first few
days to cover their nutritional requirements, a supplementation is certain
vitamins may be beneficial. Vitamin K deficiency is a frequent feature in
cats with liver disease (LISCIANDRO et al.1998; CENTER et al. 2000) and a sup-
plementation shall be consider as beneficial in cats with impaired coagula-
tion. The recommended dosage is 1.0 to 1.5 mg/kg of LW, im. or sc., 1 to 3
doses daily, up to the recovery of a normal coagulation.
The allowance of water soluble vitamins (especially from the B group)
may be doubled compare to the maintenance requirements. Brewers yeast
can advantageously be added to the meals as a source of B group vitamins
(except vitamin B12). It is also highly palatable for some cats. One to 3 tea-
spoons a day (2 to 6 grams) of yeast flakes is suitable for an adult cat.
Considering its role o antioxidant, a Vitamin E supplementation is suggest-
ed (10-100 UI/kg LW/day per os) but with no evaluation of its efficiency
(CENTER, 2005). Commercial diet designed for cats should contain all vita-
mins A, D3 and E. Both vitamins A and D are stored in the liver. A supple-
mentation over the minimal recommendation is not recommended, and
probably to avoid.
None of the supplementations can really compensate a balanced diet
580
with all nutrients provided at the same time, several times a day, in order
to cover energy and other nutritional requirements. The digestive tract suf-
fers much of anorexia, the digestive capacity decreases, atrophy of the
mucosa may happened as well as bacterial translocation. In case of anorex-
ia, a progressive re-feeding plan has to be established (FIGURE XXI-4), in
order to not overload the capacity of the digestive tract and prevent entero-
toxemia. The cat is first proposed its usual diet, or an other one, the clos-
est to its preferred one, especially of the same texture. If there is no contra-
indication, a tube feeding procedure has to be considered after 24 hours of
anorexia especially in hospitalized cats (see below). When enteral nutrition
is not possible, in critically ill patients with gastrointestinal dysfunction,
partial or total parenteral nutrition may be indicated to provide an intra-
venous nutritional support (LIPPERT et al. 1993; CHAN et al. 2002; PYLE et al.
2004).
FIGURE XXII-4: Rhythm of re-feeding the anorectic cat
581
NUTRITIONAL CONSIDERATIONS ASSOCIATED WITH HEPATIC LIPIDOSIS

Hepatic lipidosis (HL), either idiopathic or secondary, is a disease frequent-
ly associated with a period of anorexia of variable duration (FIGURES XXI-
5a, 5b,5c; FIGURES XXI-6a, 6b, 6c).
Even in presence of a primary disease, the treatment of HL has to be
started very quickly otherwise the chance of survival are even reduced.
CENTER et al. (1993) could show very few differences between survival and
non survival groups of 77 cats with HL. One of them was the delay between
the diagnosis of HL and the start of treatment, which appeared longer in
case of secondary HL. The treatment of hepatic lipidosis in cats involves
fluid and electrolytes therapy, and nutritional support via tube feeding.
a) Fluid and electrolytes therapy
Fluid therapy is necessary to correct hydration and compensate the loss
when vomiting is present. Isotonic solutions, mainly isotonic sodium chlo-
ride shall be used. Fluid supplementation with dextrose or lactate are con-
traindicated as cats in HL are glucose intolerant and such a supplementa-
tion leads to an increased glycemia, and impaired lactate metabolism is sus-
pected in some cats with HL. Potassium chloride may be judiciously added
t the fluids according to conventional sliding scale based on serum K (no
more than 0.5 mEq/kg of LW/h).
b) Nutritional support
Energy source. The first diets used for enteral forced feeding of cats
with hepatic lipidosis were those designed for humans, poor in protein and
lipid and rich in carbohydrates, preferably glucose. The work of BIOURGE
(1994) indicates that dietary protein is much more efficient to reduce hepat-
ic lipid accumulation in cats in negative energy balance. Carbohydrates are
usually less tolerated than lipids in cats, therefore diarrhoea and hypergly-
caemia may be associated with their extensive use, as confirmed by SIMPSON
and MICHEL (2000) and our own experience. As a consequence, the diet shall
be chosen rich in protein (30 to 40% of the energy), moderate in lipids
(around 50% of the energy) and relatively poor in carbohydrate. Liquid diets
balanced for cats fitting these criteria are nowadays available.
Other nutrients. A supplementation with other nutrients has been sug-
gested by CENTER (1998 and 2005) and other authors (ROUDEBUSH et al.
2000), but above the coverage of nutritional requirements, their use is still
poorly scientifically documented in the cat. L-carnitine is the one the most
studied and seems to have an interesting effect in fatty acid oxidation in
582
cats in negative energy balance and in the early treatment of hepatic lipi-
dosis (CENTER, 2005; IBRAHIM et al. 2003; BLANCHARD et al. 2002). L-carnitine
may be provided per os at a dose of 250 to 500 mg per day (CENTER, 2005).
Vitamin K may be used as defined in the previous paragraph (General nutri-
tional considerations)
c) Re feeding program
Feeding plan. In case of long term anorexia, the digestive capacity but
also the gastric volume may be reduced, down to 1/10th (FIGURES
7,a,7b,7c,7d). The amount of food by meal should thereafter be adapted
(Table XXI-9).
Table XXI-9: Re-feeding plan for cats with hepatic lipidosis.
* with MER (maintenance energy requirement) of 0.293 MJ MJ/kg optimal live weight (NRC
(2006)
Enteral forced feeding. Oral forced feeding with the finger or a syringe
directly put into the mouth may be dangerous by 3 ways: first by the high
risk of false-route which can induce a broncho-pneumonia and even death,
second by the risk of feed aversion easily inducible in anorectic cats or cats
with hepatic lipidosis; third because in the very few cats which accept to be
fed this way, the amount of food is very small compare to the requirements
of the cat and the delay between the start of the treatment and its success
is delayed.
The method of choice is at first a naso-esophagus tube feeding. Then,
if the cat does not recover its appetite within 7 to 10 days, and depending
on the context, the placement of another kind of tube may be consider
(Table XXI-10). Enteral feeding of cats with HL may be realized via a naso-
esophagus tube, an esophagostomy tube or a gastric tube. The first one is
easy to place, cheap, and does not require any anesthesia of the cat, the two
others may stay longer but requires more technique and certainly to anes-
thetize the cat. The naso-esophagostomy tube is certainly the first to try as
it is much less invasive and a number of cats will recover using only this
method. A esophagostomy or a gastric tube may also be placed in second
583
intention, once the state of the cat has stabilized thank to the naso-
esophagostomy tube feeding.
Naso-oesophageal tube: 6 to 7 CH tube is used, preferably radio-
dense. The lenght of the tube to mark with a peace of plaster or a pen is
taken from the nose to the elbow (4th rib) on the side of the standing cat.
The cat has to be maintained sit in a physiological position. The extremity
of the tube is lubricated with a few topical lidocaine gel, and a local anes-
thetic is gently sprayed at the nasal passage (FIGURE XXI-8a) too much
induces salivation and discomfort-. Then the tube is advanced ventrally and
medially (FIGURE XXI-8b), down to the mark. It can be fixed with a point of
surgical glue on the nose and a single suture to the skin between the ears.
An Elizabethian collar may be useful to prevent self-removal of the tube. The
tube is removed once the cat eat itself enough food to cover almost all its
energy requirement.
Pancreatitis may be associated with HL or prolonged anorexia and so
high risk of HL. It is now admitted that fasting cats with pancreatitis is of
no benefit (MANSFIELD and JONES, 2001). In case of vomiting, the feeding plan
may be adapted, but to treat hepatic lipidosis (i.e. to feed the cat) is a pri-
ority. JUSTIN and HOHENHAUS (1995) reported about hypophosphatemia asso-
ciated with enteral alimentation, which may cause seizures and haemolytic
amaemia. They propose that cats with high alanine aminotransferase activ-
ity, hyperbilirubinaemia and weight loss should be closely monitored for
hypophosphataemia during the first 72 hours of enteral alimentation.
As a conclusion, general rules may be proposed for cats suffering of a
liver disease are as follows: the cat must be fed with a high quality and bal-
anced diet, in several meals and water balance correctly restituted. (hydrat-
ed)
584
Table XXI-10: Naso-esophagus tube feeding: how to do?

———————————————————————————————————————————
-The tube is installed and fixed. Its location is checked with a radiograph.
The diet is chosen: not too rich in carbohydrates, balanced for cats, liquid (canned food
can be blended with a small amount of water but try to not dilute energy too much: a
minimum of 0.9 kcal/g is suitable).
- The maintenance energy requirement (MER) of the cat is calculated based on his opti-
mal body weight (0.20-0.25MJ ME/kg LW/day)
- The amount of energy the first day of re-feeding is calculated (see table 3 MER / N, with
N the number of days of anorexia)
- The number of meals is the first day of re-feeding is chosen: if feeding starts too late in
the day to allow several meals, the amount provided is preferably those of one meal, so
the amount of the all day in just one time!
- A syringe of 5-10 ml of water is prepared; water should be at room temperature –do not
use an isotonic NaCl solution, which can induce emesis.
- A syringe of diet is prepared; lukewarm by putting the food container in a bain-marie:
the liquid diet coming canned has to be kept in the fridge once open. Ten minutes before
each meal, take it out from the fridge, shake the contain, sample in a clean glass the
required amount for a meal, and put the can back in the fridge. The glass containing the
food can be put for a few minutes in a larger but lower recipient previously full with hot
water.
- First the syringe of 5 ml of water is instilled slowly. The first time, only water may be
instilled and a meal prepared for 3 to 4 hours after, depending of the state of the cat. For
instance in case of history of emesis. Otherwise, the syringe of liquid food is instilled
slowly after the water. Then a last syringe of water is instilled to flush the tube. The
extremity of the tube is carefully closed and protected from the cat claws.
- The complications associated with a naso-esophagus tube are: self-removal, nasal and
pharyngeal irritation, tube displacement by retroflexion during emesis with a naso-esoph-
agus tube.
————————————————————————————————————————————
Differences between esophagostomy and gastrostomy tubes
√ Anesthesia is required to insert the tube, and to take it out. A sur

gical procedure is definitely engaged (FLANDERS,1994).
√ The lumen of the feeding tube is larger, i.e. 10-12 CH and 20 CH or
more respectively. This permit the use of a larger variety of diets, even
dry food blended with water or home-made diets, but the require
ments of the cat are the same, the calculation of the amount of food
to provide still has the same importance.
√ During 24 hours after placement and after withdrawal-of the tube,
it is better to not feed, to permit the start of healing around the inser
tion site.
√ The complications due to the surgical procedure are added to the
previous ones.
585
FIGURE XXI-5a FIGURE XXI-5b FIGURE XXI-5c
FIGURE XXI-6a FIGURE XXI-6b FIGURE XXI-6c
FIGURE XXI-7a FIGURE XXI-7b
FIGURE XXI-8a FIGURE XXI-8b
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21-3 - 21-4 közötti.indd 586 2008.09.03. 15:45:57

Legend to FIGURES XXI-5–8
FIGURE XXI-5a: Cats with hepatic lipidosis may still look overweight even after several
weks of anorexia and severe weight loss;
FIGURE XXI-5B: Siallorrhea and amaurosis, generally asociated with hepatic encephalpa-
thy are clinical signs associated with severe hepatic lipidosis;
FIGURE XXI-5c: Hepatomegaly is obvious in cats with hepatic lipidosis
FIGURE-6a: Histology of a healthy cat liver (H-E, 40x
FIGURE-6b: Histological picture of the hepatic lipidosis (HE, 40x)
FIGURE-6c: Histology of a cat liver after 4 month treatment for hepatic lipidosis (HE, 16)
Gastric volume desreased with the long-lasting anorexia:
FIGURE XXI-7a: Stomach volume of a healthy cat (LW=2.5 kg, gastric volume 150 ml)
FIGURE XXI-7b: Stomach of a sick cat (LW=2.5 kg). Gastric volume after 6 weeks of anorex-
ia: 15 ml)
FIGURE XXI-8a and 8b: Placement the naso-oesophageal tube
587
21.2.2.7. Exocrine pancreas insufficienty (EPI)
The incidence of exocrine pancreas insufficiency in dogs is higher than in

cats. Affected animals have frequent defecation, with the excretion of large
amount of faeces, containing undigested feed components, like fat and fat
soluble vitamins. They generally show emaciation, the condition of their
hair coat is bad, however, their appetance and feed intake may remain
unchanged. The fatty faeces is light-coloured, having unpleasant smell, with
light microscopical examination undigested fat particles can be revealed. In
the backgound, the congenital pancreas hypoplasia, inherited degenerativ
pancreasatrophy (an autosomal recessive genetic failure of German
Shepherd breed) and pancreatitits equally may be found. Deficient produc-
tion of pancreatic digestive enzymes cause indigestion, which in turn, a
chronic osmotic diarrhoea.
Medical treatment is palliative, viz. the lacking digestive enzymes
should continuously replaced. From the pulverized preparation 8,000 lipase
units/kg of LW should be given twice a day. Veterinary preparation are
available (e.g. Pancrex-Vet®, Pharmacia & Upjohn), but human products
(e.g. Kreon® or Panpur®) may also applicable. Theoretically the feeding of
raw cattle pancreas is also a possibility, but in the EU, owing to the veteri-
nary rules and regulations, it is practically excluded. The suitable dietary
treatment consists of feeding readily digestible, low-fat, high vitamin diet.
(In case of acute pancreatitis the type of used diet is similar, but is is given
after a preliminary 4-day total fasting.) The mineral and vitamin supply,
with special regard to the fat-soluble vitamins and the vitamin B12, extra
peroral or parenteral provision can be practised.
21.2.2.8. Diabetes mellitus

The chronic disorder of carbohyrate metabolism, the diabetes is caused by
an absolute or relative lack of insulin and/or glucagon abundance (KANEKO
et. al. 1977). Peripheral tissues like skeletal and cardiac muscle and adi-
pose tissue) require insulin for the glucose transport into the cell cytosol.
Main forms of diabetes are the Type I, autoimmune insuline-dependent
diabetes mellitus (IDDM) and the Type II obesity-related, refractory to the
insuline, non-insuline-dependent diabetes mellitus (NIDDM). Dietary treat-
ment of obesity has been described above; hereto the diabetes itself will be
treated. Diabetes can be found not only in dogs and cats, but also in rep-
tiles and cage birds, too.
588
Etiology and Pathogenesis. Owing to the deficient insuline production

of the pancreas β-cells, the blood glucose is constantly high and glucosuria
develops. Amyloidosis damages the pancreatic tissue included the beta
cells, especially in cats. The amyloid derives from the islet-associated
polypeptid (IAPP), which is also a product of β-cells. Accumulated amyloid
and/or IAAP destruct the β-cells and cause insulin resistance. Diabetic
cats has a higher percentage of amyloid accumulation in the islets.
Clinical Findings-Diagnosis. The first striking clinical signs are the
pathological thirstiness (popydipsia) and excretion of large amount of glu-
cose-containing urine (polyuria). The renal threshold for glucose is approx-
imately 10 mmol/l (180 mg%) in dogs and 13 mmol/l (240 mg%) in cats.
The exaggerated lypolysis leads to the accumulation of keton bodies in the
blood and in some cases to hepatic lipidosis in cats. Precipitation of
ketoacidosis is partly due to the action of the hyperglucagonaemia. The
smell of exhaled air is similar to the acetone. In long-lasting, untreated
cases diabethic cataract and polyneuropathy are common. Cataract devel-
ops commonly only in dogs; it is charactarized by lenticular opacities of
stellate shape.
Treatment. The medical cure can be initiated by oral antidiabetic
agents like sulphonylurea family, meglitinid analogues and D-phenylalanin
derivates. In more developped cases insulin preparations (rapid or regular,
intermedier, lente or ultralente) or their combinations shlould be applied.
Dietary Treatment. The aim of the treatment is the regulation of live
weight by controlled feed intake and the synchronisation of the carbohy-
drate ingestion with the insulin dosage. Energy needs of the diabetic dog
and cat should be basically covered by feeding complex, long-realease car-
bohydrates, like starch. While selecting raw materials, the glycaemic index
of feedstuff should be taken into account. The glycaemic index (GI) shows
the capacity of given feedstuff to increase blood glucose 2 hours after eat-
ing, compared to the chemically pure glucose, which has a GI of 100.
(Technically, the blood glucose values of series blood sampling are plotted
against the time, presented in a graph and the territories below curve are
compared.) The glycaemic index of wheat bran is 42, of milk 27 and of the
soybean carbohydrates 18. Heat-treated potato has a relatively high GI of
70 to 80%. The barley, oats, rice and according to the recent data the
sorghum, milo and backwheat have a medium-to-low GI, consequently
they can be applied in diabetic diets. Notwithstanding that dog and cat vir-
589
tually do not digest cellulose and lignine, the fibre level in the feed of dia-
betic patiants (especially in case of dogs) should be increased at least up to
10%. The main role of fibre is to delay the sugar release from the starch in
the gut lumen. Thus, the diet for diabetic animal should contain 30 to 45%
starch, 8 to 17% of crude fibre, medium level of good quality protein (15 to
25% for dogs and 28 to 45% for cat of a true digestibility of at least 85%)
and low amount (generally 10%, but no higher than 20%) lipid. The con-
straint of the latter is the risk of production of keton bodies. The soluble
fraction of total dietary fibres is more efficient is this ailment. Another
approach in case of cat that instead of dietary fibre the level of protein is
increased with low level of carbohydrates. A fixed and constant formula-
tion is preferred to the home-made diet.
Distribution of the daily ration should be scheduled in order to pre-
vent the postprandrial blood glucose peak, but the blood glucose does
should remain above a minimum value. Meals should be timed that the
absorption of nutrients to coincide with the insulin peak in the blood. The
portion of 24 hours should be divided into two unequal parts: ¼ of the daily
ration should be given early in the morning (0. hour). If the animal con-
sumed the whole amount, the subcutaneous insulin injection (generally 0.5
U/kg of LW) follows. (If dog and cat refused to ingest the offered feed, the
insulin injection should be omitted to avoid the abrupt drop in blood glu-
cose.) It is advisable to check every morning the glucose and keton concen-
tration of the first urine using strip kit and adjust the dosage of insulin. If
the glucose level in urine is 0.25%, the dosage of the previous day may be
used; if the urinary glucose concentraion is more than 2%, 2 U insulin
should be added to the dosage, if there is no glucose in the urine, 2 U
insulin should be reduced from the due amount. Generally combined
insulin preparations are applied, whith a combination of 30% of fast-
absorbing amorphous and of 70% of a long-release crystalline insulin form.
In this case, 7 to 8 hours after the first feeding the remaining ¾ part of the
daily ration is given. Using this scheme, blood insulin concentration is long-
lasting high enough to prevent the outstriking blood glucose concentra-
tions. Organic chromium supplementation of feed increases the sensibility
of insulin receptors and thereby may improve glucose metabolism (LAI et al.
2006). The American Diabetic Association (1996) is sceptical in this respect
and stated that „chromium replacement has any beneficial effect on
glycemic control is for people who are chromium deficient”. On the other
590
hand, NAKHOUL et al. (2006) demonstrated in rat that the brewer yeast-
derived GTF (glucose tolerance factor) was able to inhibit the development
of nephropathy that is induced by diabetes.
To sum up the main species differences between dogs and cats are as
follows: the prevelance at dogs is higher in females, at cats, on the contrary,
in males. Constantly high blood progesterone concentration increases the
GH secretion causing hyperglycaemia and insulin resistance only in dogs.
The IAPP and amyloid deposition in the pancreatic islet cells as well as the
hepatic lipidosis occur only in the cats. The development of cataract is typ-
ical for dogs. Dogs response better on the higher dietary fibre in their feed.
The use of oral antidiabetic agents (e.g. glipizide, glimepiride or acarbose, an
oral a-glycosidase inhibitor) is recommended mostly for the cat patients.
Veterinary diets both for diabetic dog and cat are available.
21.2.2.9. Skin diseases

During treatment of dermatological cases of nutritional origin or that of
influenced by diets, the general scheme of skin disease treatment should be
followed. Thus, the whole dietary and clinical history should be made, clin-
ical investigation should be performed with special regard to the skin and
its appendixes, analysing the data of performed laboratory and other sup-
plementary tests, stating the diagnosis, evaluation of prognosis and finally
prescription of medicinal and diatary treatment.
Main causes of itching dermatosises may be ectoparasites (flea,
cheyletiellosis, Sarcoptes itch mite, pediculosis, trombiculidiasis), bacterial
infections (pyoderma, superficialis folliculistis), hypersensitivity (flea saliva,
atopy, feed allergy, contact irritation), otitis externa and anal sacculitis.
Main causes of non-sitching dermatosises are ectoparasites (demodi-
cosis), endocrinopathies (hypothyroidismus, Cushing-syndrome, Sertoli-cell
tumour), dermatophytosis, deep pyoderma, nutrients, mineral and vitamin
deficiencies.
One of the most common cases is the hair fall caused by energy and
protein deficiency (telogen and anagen defluxion), which is frequently
accompanied by a depigmentation of hair and skin. (In some breeds, like
puli it is limited only to the forepart of the body). Gestation, or rather lacta-
tion, chronic kidney diseasse or insufficient protein ingestion are the main
causes. Full recovery can be expected after dietary supplementation. In the
practice, breeder use for these purposes high energy and protein puppy and
591
kitten feed for lactation bitches and queens.

ZINC DEFICIENCY. In case of zinc deficiency the manifestation of clinilac
signs is facilitated by carotenoid deficiency. Type I, so-called zinc-responsive
dermatitis occurs generally in Alaskan Malamut and Siberian Husky. The
main cause if an inheritable trouble (very low rate) in zinc absorption.
Symptomes includews parakeratosis of some predilection anatomical
regions (mouth, paws, perineum, chin, rim of lips, ocula around the eyes,
hock joints. The dietary treatment consits of the daily provision by 10 mg
zinc sulphate/kg LW or 1.7 mg zinc methionate/kg of LW, but the high zinc-
containing yeast preparations are also applicable. Acrodermatosis is the
extreme form of inability to absorb zinc, caused by autosomal genetic defect,
which besides the skin signs will lead to deat at the age of 7 to 8 month of
dog, having insufficient immune functions, too. Its incidence is relatively
high at bull-terriers. (For more details see below). Type II dermatosis is
caused the primary dietary zinc deficiency; it may occur in all dog and cat
breeds. Calcium overfeeding frequently causes secondary zinc deficiency.
LACK OF VITAMIN A AND CAROTENOIDS is not uncommon in spaniels,
which can be easily revealed using therelative dose response (RDS) method.
Owing to the trouble of keratinisation in skin, acne develops.
Supplementation of diet by vitamin A (5000 IU/kg DM), or application of
400 IU vitamin A injection daily, during two weeks favours recovery. While
overdosing vitamin A, fatal poisoning may develop!
DEFICIENCY OF ESSENTIAL FATTY ACIDS, besides dermatitis, manifests in
delayed wound healing and in impairment of immune system. The latter in
cats can be aggraveted by a lack of taurin. The linoleic acid requirement of
dog and cat is 2% of the energy intake; beyond that cats, having no ∆7-
desaturase enzyme activity, at least 0.04% arachidonic acid of the total
energy intake. Appropriate essential fatty acid supply is especially impor-
tant, if the fat&oil level of the feed is high. Developed skin symptoms can be
improved by giving 1.5 ml vegetable oil (maize, oliva, pumpkin seed, grap
seed, sunflower and cottonseed) for a month for replace linoleic acid, plus
for the cat poultry fat as a source of arachidonic acid. To set the optimal w6
to w3 fatty acid ration (5-10 to 1), fish oil supplementation may become nec-
essary. Established cases of atopic dermatitis can also be improved by
adding fish oil (3 mg/kg of LW) and evening primrose oil (13 mg/kg of LW),
in guise of omega-3 fatty acid sources.
Among the water-soluble vitamins, the BIOTIN DEFICIENCY may manifest
592
in dermatitis. Shortage of biotin can be caused not only by an absolute

dietary deficiency, but in rancid feeds peroxides deteriorate biotin, thereto
avidin, presnt in the raw egg white binds biotin in the gut lumen, prevent-
ing in absorption. Besides, biotin in cereals has a low availability. The early
clinical signs appear at the tailhead region, expanding backwards (caudal,
on the tail) and onwards (cranial), symmetrically on the hip and back. Signs
in dogs are commonly depigmentation of hair and dry, scaly dermatitits; in
cats the dermatitis is more serious, and it results in loss of hair on the men-
tioned body parts. Symptoms should be distingueshed from the hairles tail
(„rat-tail”), signs of diabetes or hypothyreosis.
OTHERS. It was found that daily application of 5 mg biotin comple-
mentthe medicinal treatment of the cat’s miliar dermatitis. For the treat-
ment of autoimmune canine skin diseases, besides the 500 mg tetracyclens
bid, the daily ½ to1 gram niacin supplementation proved to be benevolent.
Huge amount of vitamin E dosages may contribute to the cure of discoid
lupus erythematosus.
21.2.2.10. Feed intolerance and allergy

The background information concerning the feed intolerance and allergy is
treated in Chapter XVII. During the evolution of Life the existence of ani-
mals were basically threatened by the wounding and the infections and does
nowadays, too. As a counter manoeuvre the mechanism of wound healing
and the immune system developed. The task of the latter is the recognition,
destruction and elimination of foreign matters (in the first place the protein
components of pathogen viruses and bacteria). There are cases when the
immune system „fulfil” or the defence mechanism glides into a wrong path-
way and pathologic symptoms occur like asthma and food allergy.
Today we have a totally new and astonishing knowledge about the
immune system. White blood cells in and around the gut wall are called
„gut-associated lymphoid tissue” (GALT). Another of the immune mecha-
nism, the microbes of the gut content may have beneficial, neutral or harm-
ful to the health of the host organism. Why proteins of the ingested food do
not cause any injury? Why does not initiate the dietary protein allergic reac-
tions, diarrhoea, dermatitis or asthmatic attack? This is due to the oral tol-
erance which assures the protection by a complex mechanism. First of all,
the well-digested dietary protein is broken up into amino acids that even
after absorption will not stimulate immune system to antibody production.
593
Absorption of big, undegraded molecules is inhibited from the absorption

consequently they are eliminated by the faeces. Tiny amounts of dietary
proteins getting undegraded through the M-cells in the circulation will force
leukocytes to produce the special antibody (IgA), secreted by the intestinal
mucous membrane. If one of these factors had dysfunction, harmful
immunologic processes occur and the food allergy develops. Factors facili-
tating the manifestation of food allergy include
-ingestion of too much protein that cannot be digested by the animal;
-sub acute inflammation of the gut wall caused by parasites, bacteria
or viruses;
-infections paralysing the immune system like cats’ AIDS, ingestion of
certain mould toxins (e.g. T–2 toxin), endocrine disorders and anti-inflam-
matory steroid treatment.
-If the dietary protein avoids the alimentary tract and get into the
blood stream through the conjunctivae or the bronchial surface (milk drops
in case of puppies or kittens).
At such a case the immune system perceives it as a foreign protein
and especially at the second meeting will react powerfully but abnormally
that means dead-threatening anaphylactic reaction shock, broncho-spas-
mus and in chronic, prolonged cases the classic food allergy. Under the
influence of the latter mainly the colon, skin and its appendices get into
inflammation. Manifestation includes plasmacytic colonitis with diarrhoea,
torturing itching of the skin, otitis externa, conjunctivitis, dermatitis of the
axillar and interdigital skin.
The food allergy should be marked off the hypersensitive reactions to
a feed component (e.g. too spicy home-made diet, eating of octopus, squid,
coffee-ground etc.) as well as off the enzyme deficiency diarrhoeas. Many
older dogs and especially cats are not able to digest the milk sugar (lactose)
and the intake of milk will cause prolonged and untreatable diarrhoea. If a
certain feed form or taste is linked in the memory with a bad memory the
intake may provoke vomitus or diarrhoea, too. This is the so-called food
aversion.
In case of the feed allergy, the first task is the clearing up the basic
cause. Vets are able to identify several proteins but the most secure is the
elimination diet. It means that the dog or cat should receive solely a pet food
or home-made diet containing only one protein and one energy source,
namely that is unlikely met the animal before. For example the lamb meat
594
CHAPTER XXI: DOG AND CAT DIETETICSr
and rice, the duck and the tapioca meal, horse, rabbit or turkey meat with
potatoes. This diet should be given for 2 to 4 weeks until the clinical signs
like erythema, itching and diarrhoea cease. After that, but only under strict
veterinarian control, the organism should be challenged. It means that
besides the energy component one of the previous protein sources is given,
changing weekly. If an allergic reaction occurs, the given feedstuff should be
prohibited for a life.
The beef meat, milk, poultry meat and soybean are the most common
cause of the feed allergy, but the situation is dynamic and may change
according to the frequency of actual raw material usage. There is even food
allergy to a certain type of commercial food, owing to the special composi-
tion, conservative and colorant content. The main protein (prolamin) of the
wheat, barley and rye may trigger the so-called gluten allergy. The latter is
characterized by an untreatable diarrhoea and small intestinal inflamma-
tion. It should be mentioning that the stale water by means of its pollution
and algal growth may also cause allergy.
The dietary treatment of the feed allergy is that either as a home-
made diet or as a veterinary diet should be fed which contains exclusively
„unknown” protein for the animal. There are also hypoallergic diets avail-
able that are pre-treated by digestive enzymes to degrade proteins up to the
measure (>16000 Dalton), less to that would be sufficient enough to elicit
antibody response. Treatment of the clinical signs like diarrhoea and skin
alterations should be treated by the veterinarian.

Critical care means the feeding of sick, pulled down, anorectic animals or
patients during preoperative or postoperative periode. It may happen by vol-
untary ingestion of the offered special feed or a force feeding, using tube-
feeding or parenteral route of application. Critical care is justified when
▪ live weight abruptly drops by 10% or there is a chronic state of
malnutrition;
▪ trauma or postoperative period;
▪ anorexia, existing during 3 to 5 days;
▪ diagnosed hepaticus lipidosis in cats,
▪ increased nutrient losses (e.g. vomiting, diarrhoea, malabsorp
tion, intestinal resection);
▪ intensified energy and protein requirements (burns, infections,
595
trauma, fever);
▪ besides application of certain drugs (corticosteroids, immunsu
pressive preparations, antibiotics etc.);
▪ chronic organ diseases, tumours and
▪ peculiar laboratory findings (less than 2% blood albumin concen
tration, decreased albumin to globulin ratio and drop in the
lymphocyte number).
The main types of enteral feeds are as follows: gruel, polymeric diets,
mixed, hydrolysed protein, human polymeric liquid diets, elemental diets
(mixture of amino acids) and protein module (protein-fat concentrate). In
the majority of cases it is sufficient to apply the general rules of feeding of
sick and febrile dog and cat. Even among the fortified diets the most con-
centrated are the „instant diets”, that should be dissolve in hot water. They
have two main types, members of the first are intended basically to correct
electrolyte household, containing salts, sugars and amino acids in mal-
todextrin carrier (e.g. Electrolyte Instant Fluid Canine Pedigree and
Elektrolyte Instant Fluid Whiskas Feline Diets etc.). Besides to giving the
instant diet, a low-fat, high protein feed should be fed. The so-called „recov-
ery” powders (e.g. Prescription diet canine p/d, Hills; Eukabana Nutritional
Recovery Formula for Dogs; Reanimyl, Virbac; Robor CPD, Species;
Concentration Instant Canine Pedigree; Concentration Instant Whiskas
Feline Diets, CliniCare Canine Liquid Diet, PetAg; RenalCare Canine Liquid
Diet, PetAg; Nutrison powder and Nutridrink, EGIS etc.), that basically con-
tain milk and whey powder, supplemented by oils and active ingredients.
Their energy density and protein level is high and they should be given sole-
ly/alone. The higher arginine, glutamine and zinc concentration still merit
mentioning, considering the extra needs of immunological stress (see
Chapter XVII).
Forced feeding is commonly executed by using syringe or naso-
oesophageal tubes, but it is possible to use pharyngostomy, oesophagosto-
my, gastrostomy, and enterostomy tubes, too. Beyond the mentioned pre-
scription diets there are formularies for home-prepared mixtures, like
shown in (Table XXI-11).
596
Table XXI-11: Mixture for tube-feeding

———————————————————————————————————————————
Ingredients Quantity, gram
———————————————————————————————————————————
Low-fat cotton cheese 300
Sunflower oil 130
Glucose 20
egg yolk (1 piece) 18 (approximately)
Water 100
Calcium lactate 15
Orto-phosphoric acid (85%) 6
NaCl 2
Potassium sulphate 2
Micro-elements 0.25
Vitamin premixt 0.07
<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<
Nutrient content in 100 gram in 100 ml
Dry matter 38.0 36.8
Ash 2.04 1.97
Crude protein 6.60 6.39
Ether extract 20.8 20.1
N-free extract 8.56 8.29
Organic matter 36.0 34.8
Calcium 0.48 0.47
Phosphorus 0.37 0.36
——————————————————————————————————————————
21.2.2.12. Nutrition and tumours

In case of tumour patients an essential distinction should be made accord-
ing that the patiant is inoperable or on the contrary, he is just before or after
a surgical intervention. In the first case the aim is the elongation of life
assuring a good life quality by minimizing pain and suffering. In this case
the diet should have a high energy density and protein level; the latter
should be of excellent quality (high digestibility and good biological value) in
order to minimize urea production and loading liver and kidneys. In any
case the elimination of the end product of protein metabolism may be facil-
itate by supplemental arginine provision, which help function of urea cycle.
Additional L-arginine (0.23 g/kg of LW), L-glutamine (0.23 g/kg of LW) and
beta-hydroxy-beta-methylbutyrate (0.05 g/kg of LW) may increase lean
body mass (muscles). Functional state of liver can be supported by adding
L-carnitine.
If the tumour is operable and the patient is still in condition, the aim
of dietary treatment is the minimizing of formation of recurrences. The the-
oretical basis is that tumour cells have the highest energy, proteins, miner-
al and vitamin needs in the organism. Namely, they practice only anaerobic
597
energy yielding, which has en efficacy of 1/7 to 1/8 that of oxidative phos-
phorilation. Their fast multiplication requires continuous amino acid and
active ingrediaint supply. These facts, supported by experimental data too,
suggested that before and after operation a drastic feed restriction should
be applied (approximately giving the half of the maintenance requirement).
The scheme shrunkens the life expectances of tumour cells and decrease
the incidence of recurrences.
21.2.2.13. Inherited metabolic troubles in dogs

Metabolic troubles are characterized by the incapability of digesting,
absorbing, retaining or metabolize a certain compound. The majority of
inherited metabolic troubles is an enzymopathy. Primary enzymopathy
derives from a gene mutation, which in turn cause the lack or deficient
function of an enzyme. As a consequence, the substrate of the given enzyme
accumulates in the organism. Secondary enzymopathy generally develops
as a a result of the damage of cell membranes, which in turn leads to the
release of the until then compartimentised enzymes in the extracellular
space or into the bloodstream. In this case the pathological effects are due
to the lack of the enzyme on its suitable spot.
HYPERLIPIDAEMIA
The postprandrial hyperlipidaemia is a physiological phenomenon, caused
by a transitional increase of chylomicron concentration in blood, during no
more than 6 to 10 hours. On the contrary, the hyperlipidaemia in a fasting
dog is a pathological phenomenon and may act as a risk factor of many dis-
eases. In case of dog hyperlipidaemia the triglycerid concentration in blood
plasma is higher than 1.7 mmol/l and the colesterol level is higher than 7.8
mmol/l. The corresponding values for cats are 0.15 mmol/l for the trigly-
cerids and 5.2 mmol/l for the cholesterol. Chronic hyperlipidaemia mani-
fests in abdominal pain, vomiting, diarrhoea, anorexia, cerebral haemor-
rhage, hepatomegalia, lipidosis of the individual organs, thereto the risk of
the development of acute pancreatitis increases. In dogs lipid deposits may
appear in the cornea. Atherosclerosis, however, is both in dog and in cat
uncommon.
The pathological hyperlipidaemia is more frequently secondary, devel-
oping as side-effect of diabetes, hypothyroidism, pancreatitis, renal failure
and liver diseases. Incidence of primary hyperlipidaemia is lower, generally
598
is inherited, in the Dwarf Schnauzer breed. In affected, clinically apparent-

ly healthy individuals hyperlipidaemia can be detected. The incidence is
higher in the age of older than 4 years and its occurrence is independet from
the gender. In developed cases the symptoms resemble to that of acute pan-
creatitis, thereby it is called alos as pseudopancreatitis. The blood bio-
chemistry is helpful in the differential diagnosis.
Dietary treatment of primary hyperlididaemia includes the decrease of
the energy density and fat content of the diet. The threshold values, when
the feeding of this low-fat diet should be started, if the fasting serum triglyc-
erid attains the 5.6 mmol/l and the cholesterol concentration of 18.1
mmol/l. In three weeks the blood analysis should be repeated and the diet
adjusted. The low-fat (8-12% in the dry matter) facilitate the decrease of
chylomicrons and VLDL in blood. It worth mentioning that affected dog can-
not receive any treat or table scrap, because ingestion extra fat may induce
an acute pancreatitis. In older animals the dietary treatment generally
should be supplemented by medicinal treatment, giving clofibrate, gemfibro-
syl, niacin in pharmacological dosage and 30 to 60 mg/kg of LW fish-oil.
INHERITED TROUBLES IN PURIN METABOLISM OF DALMATIANS

Purins are components of nucleic acids, therefore the organism is capable
of synthesizing them from precursor molecules, but the endogenous and
dietary nucleotids are utilised, too. Excess purin bases first are degraded to
uric acid, then under the influence of the hepatic uricase enzyme they are
transformed into allantoin. Undegraded uric acid is found in tissues in form
of sodium salt. Most of the mammalian species, with the exception of man,
apes and Dalmatian, transform the decisive majority of uric acid into allan-
toin and only very low is the urinary uric acid concentration. On the con-
trary,by the urine of Dalmatians the uric acid excretion is much higher (400
to 600 mg daily) than in the other breeds (10 to 60 mg daily): Accordingly,
the blood uric acid concentration in Dalmatians is 2 to 4-times higher (17.8-
23.8 μmol/l) than in the other dog breeds. The high blood value of
Dalmatian is caused by a reduced hepatic uricase activity; which in turn
will not be reabsorbed in the renal tubuli, causing the high urinary con-
centration.
The troubles of the purin metabolism in Dalmatians is autosomal
recessive genetic failure (SIMKIN, 2005). Because the water solubility of uric
acid is much lower than that of allantoin, it can easily be precipitated in tis-
599
sues. The gout is caused by the precipitated uric acid crystals in the joints
or the visceral gout in the body tissues and cavities will not developped in
the Dalmatians, because of the low efficiency of tubular reabsorption. As a
consequence, the uric acid concentration of blood generally is below the
gout formation threshold; at the same time, the high urinary concentration
predisposes dog to urolithiasis. crystal prescipitation in the urine is only
1/20 of the total urolithiasis cases, but half of that occurs in Dalmatians.
Development of urate urinari calculi is influenced by the pH, ammonia con-
tent, the ingested purin amount and by the presence of microorganism. The
incidence of urate urolithiasis is 1% in the Dalmatian population.
Dietary treatment of the urate urolithiasis means the feeding of feed of
low purin content, having alkalizing character. Considering, that proteins
acidify urine, low protein diet should be fed, using high at leat 80% of bio-
logical value. Daily application of the xanthin oxydase inhibitor allopurinol
is also efficient in preventing the formation of urate stones.
MALABSORPTIONS
Vitamin B12 malabsorption of Giant Schnauzers is an inherited ailment.
The vitamin malabsorption is not accompanied by the defective utilisation
of other nutrients. Perorally applied vitamin B12 is inefficient, because of
the lack of intrinsic factor, namely the cause of the metabolic trouble is
localized at the receptors of the small gut epithelial cells. Consequently, the
parenteral vitamin B12 supplementation (0.5-1 mg in every three months) is
indispensable for the affected individuals.
Zinc malabsorption. The autosomal, recessive genetic failure of bull-
terriers manifests in the lethal acrodermatosis. Practically there is no zinc
absorption, even feeding diets of high zinc concentration lacking the carrier
protein in the gut wall. Afflicted animals have an impaired immune func-
tions too. At the age of 10 weeks the growth of puppies practically stops and
the serious parakeratosis develops primarily on the nose and end of the
legs, giving the name of the ailment (acro= peak). Immundeficiency results
in pyoderma , trivial infections and death till the age of 7 months.
Less serious is the partial zinc absorption incapability of Alaskan
malamut, Siberian husky, less frequently of Great Dane and Doberman
pincher breeds. The inheritary charicter is supposed by the observation that
the autosomal recessive dwarf individuals exhibit generally insufficient zinc
absorption, too. Concerning the clinical signs and the treatment see above
600
(skin diseases).
COPPER STORAGE DISEASE OR INHERITED COPPER ACCUMULATION

In some breeds (Bedlington terrier, West highland white terrier, Dobermann
pincher, Cocker spaniel) the copper may get accumulated in the liver, grad-
ually increasing the physiologically 200 to 400 mg/kg liver DM concentra-
tion. The cause of the metabolic trouble is the failure of a gene, coding the
a copper transport protein, similarly to the human Wilson-disease. This pro-
tein is responsible for the elimination of copper from the liver, mostly bound
to metallothionein molecules. When hepatic copper concentration achieves
the 2000 mg/kg DM value, centrolobular hepatitis develops. The early
symptomes are the Parkinson-like neurological dysfunction, copper deposi-
tion in the eyes (see human Kayser-Fleischer ring) and the coeruloplasmin
level of the blood drops, below a critical value. It is a pathognostic sign, if
after the application of D-penicillinamine the urinary copper concentration
strikingly increases (up to tenfold).
Afflicted animals should be fed on a low-copper diet lifelong (it means
that liver, kidney and other viscera, mushroom and cocoa-containing prod-
ucts cannot be given). The dietary therapy includes perorally provision by
zinc salts (e.g. zinc sulphate, zinc acetate – Galazin, 100 mg, bid, zinc in
lipophyl matrix) or tetrathiomolibdate. Zinc compete with copper in the
enterocytes for carrier proteins, preventing in the absorption. Moreover,
zinc induces the synthesis of metallothionein in epitheial cell, which in turn,
binds both zinc and copper, and with the exfoliated cell will be excreted by
the faeces. Molibdenate form insoluble compound in the gut lumen, inhibit-
ing in this way copper absorption. By the application of chelating agents,
like D-penicillamin (with vitamin B6) or trientine the urinary copper excre-
tion can be enhanced.
21.2.2.14. Inherited metabolic troubles in cat

Among cats, supposedly owing to the later domestication, the number and
incidence of inherited metabolic troubles is lower than in case of the dogs.
Diabetes is more frequent in the Burmese breed. The gangliosidoses are
degenerative diseases of the neural system, caused by enzyme deficiencies.
Affected animals die at the age of 8 to 10 months. Mannosidosis is a lethal
enzimopathy (sign: „man”), resulting in stillbirth or hepatomegalia, tremor
and ataxia. The hyperoxaluria (sign: „ho”) favours in cats of 5 to 9 months
601
old the accumulation of oxalate crystals in the renal tubuli, which in turn
will cause fatal acute renal insufficiency. In case of the hyper-chylomicron-
aemia (sign: „hce”) of kittens the lipidaemia become chronic resulting till the
age of 8-9-month several haematomas all over the body, through that to the
compression of the peripheral nerves, loss of reflexes and finally to death.
The hypothyroidism has an inherited form in cats too; the supportive, pal-
liative therapy is generally successful. In the occurrence of different types of
hurolithiases the familiar predisposition is also suspected. Following from
the charicter of these ailment, the dietary support is limited only to a tem-
porary palliative treatment.
602
21.2.2.15. CHRONIC KIDNEY DISEASE (RENAL FAILURE)
The urinary system has metabolic and excretory function. In this the kid-
ney performs a number of functions that help maintain the physiological
integrity of the extracellular fluid (ECF) volume. These processes are a) con-
servation of water, fixed cations, glucose and amino acids, conservation
being used in the broad sense to imply the return of body fluids, of the
amount of the substance required by the body’s needs, the excess being
excreted into urine, b) elimination of the nitrogenous end products of pro-
tein metabolism, primarily urea, creatinine and ammonia, c) elimination of
the excess hydrogen ions and the maintenance of the physiological pH of the
body fluids, and d) elimination of complex organic compounds both endoge-
nous and exogenous. Two important endocrine substances are secreted by
the kidney, erythropoietin and renin which assume a role in the regulation
of aldosterone secretion by the adrenal cortex.
In chronic kidney disease, loss of functional renal tissue is prolonged,
significant, and usually progressive. It usually occurs in older animals,
although congenital renal disease may cause renal failure in animals
younger than 1 year. There is no sex predilection. Although it has been
described as chronic interstitial nephritis, this term essentially describes
the morphologic appearance of kidneys with chronic, progressive and irre-
versible disease, and does not contribute to the understanding of the under-
lying cause. Identifiable causes include pyelonephritis, amyloidosis, chron-
ic obstructive uropathy, congenital lesions, glomerulonephritis, allergic and
immune-mediated nephritis, and neoplasia.
CLINICAL FINDINGS. Polydipsia, polyuria, and occasional vomiting are
the early signs. Renal failure being progresses over weeks or months to
years, anorexia, weight loss, dehydration, oral ulceration, vomiting, and
diarrhoea are seen. In the terminal stages, severe dehydration, vomiting,
convulsions, and coma lead to death. Mucous membranes will be pale if the
animal is anaemic. Loose teeth, deformable maxilla and mandible („rubber
jaws), or pathologic fractures may be seen with renal secondary osteodys-
trophy. Careful palpation may reveal small, irregular kidneys, in animals
with end-stage renal disease or large kidneys in animals with tumours or
hydronephrosis.
DIAGNOSIS. The blood nitrogen, serum creatinine, and inorganic phos-
phorus levels are increased. A moderate to severe no regenerative anaemia,
603
metabolic acidosis, and hypertension develop as renal function decreases.

Osteoporosis may be seen radiographically. In normal dogs, urine specific
gravity is usually 1.016-1.060; in dogs with renal dysfunctions, it may be
fixed at 1.008-1.012. Controversely, dogs, with primary glomerular disease
may become azotaemic, while retaining some ability to concentrate urine.
Cats with chronic renal failure usually produce urine with a specific gravi-
ty of 1.008-1.034, (normal range for cat 1.020-1.040) probably because of
their normal ability to produce very concentrated urine. The polydipsia and
polyuria of chronic kidney disease must be differentiated from those asso-
ciated with diabetes (insipidus or mellitus), pyometra, pyelonephritis with-
out renal failure, and hyperadrenocorticism. The normal range for urine vol-
ume is 10-20 ml/kg LW for cats and 20-200 ml/kg LW/day for dogs.
Adrenal insufficiency may be confused with primary renal failure because
prerenal azotaemia may be caused by the vomiting, diarrhoea, and polydip-
sia associated with the former.
TREATMENT. With appropriate therapy animals can survive for long
periods with only small fractions of functional renal tissue, perhaps 5-8%,
both in dogs and cats. The recommended treatment varies with the stage of
the disease. The systemic hypertension found in 20% of animals with
chronic kidney disease may be observed at any stage and is not effectively
controlled by the simple feeding a low-salt diet. The usual antihypertensive
medications are a calcium-channel blocker such as amlodipine besylate
(0.1-0.25 mg/kg, perorally, twice a day), or an angiotensin-converting
enzyme (ACE) inhibitor such as enalapril (0.5 mg/kg, once or twice a day).
While these may be administered together, a calcium-channel blocker is
usually recommended as initial therapy in cats and an ACE inhibitor in
dogs.
At the same time to providing a continual supply of fresh drinking
water and encouraging and documenting adequate dietary intake, body
condition scoring should used routinely to assess adequacy of intake.
The aim of the DIETARY TREATMENT is to provide a low–protein diet,
replace the water-soluble vitamins and calcium which have been lost and
prevent accumulation of phosphorus and sodium.
The low-protein diet will help to reduce both the accumulation of
nitrogenous waste and the intake of phosphorus, which is high in meat
(Ca:P in meat equal to 1:20; in cereals 1:9). It has also been suggested that
restriction of dietary protein and/or phosphorus may affect the rate of pro-
604
gression of renal disease. The ideal level of protein for dogs with mild to
moderate chronic renal failure is approximately 2.0-2.2 grams of high qual-
ity protein/kg of live weight per day. This should be considered a starting
point, and adjustments made to suit the individual case to ameliorate clin-
ical and biochemical manifestations of uraemia, whilst avoiding malnutri-
tion. Cats with renal failure: an intake of approximately 3.3-3.5 grams pro-
tein/kg live weight in a day has been recommended for this species. The
concept of „ideal protein” should be applied: dietary protein of high
digestibility and of excellent biological value (BV), i.e. its amino acid profile
absolutely adequate for the needs of tissue synthesis. Using the AAFCO
optimum data, the most important amino acids of their ideal protein for
maintenance are as follows (first the dog, after the cat data): Met+Cys
68/133, Trp 22/23, Tre 74/88, if the lysin needs are taken as 100%. (It is
interesting the strikingly high requirements of cat towards sulphur-con-
taining amino acids.) The absolute lysine requirement for maintenance is
8.0 g/kg air-dry feed at the dog and 8.3 g/kg air-dry feed at the case of cat.
If using home-made diet, egg, cottage cheese, cheeses and last, but not least
the lean chicken, rabbit, mutton and beef meet are applicable.A number of
veterinary diets with restricted protein and phosphorus content designed to
assist in the medical management of chronic kidney disease in dogs and
cats are available to the clinician (e.g. for mild and moderate cases
Prescription Diet Canine k/d and in case of advanced renal failure, with
uraemic encephalopathy: u/d; for cats Prescription Diet Feline g/d and k/d.
Although these diets significantly reduce the phosphorus intake by
renal patients, this can be further enhanced by giving aluminium hydroxide
orally to chelate the phosphorus in the diet thus reducing absorption from
the intestine. The use of aluminium hydroxide should be with caution, espe-
cially in the cat, starting within the range of 30-90 mg/kg/day. Calcium and
calcitriol (with precaution) supplements should only be provided once calci-
um to phosphorus ratio has been restored. When calcium addition is pro-
vided too early, soft tissue calcification (calciphylaxis) may develop. Fat and
carbohydrate should be used to provide all the energy requirements of dog
and cats. This helps to reduce tissue catabolism, weight loss and nitrogen
waste accumulation. Fats and especially oils are particularly valuable
because of its energy density and ability to improve the palatability of low-
protein diets. The last is almost important in the anorexic patient.
605
SUMMARY OF DIETARY REQUIREMENTS OF THE NON-INFECTIOUS RENAL

Chronic Kidney Disease: low protein of high biological value (BV=high
digestibility and ideal amino acid composition), reduced phosphorous,
increases B vitamins; insure adequate fat/oil and carbohydrate supply to
meet energy requirements and spare protein
Glomerular Nephritis: low protein of high BV, reduced sodium (less than
0.3% of dry matter, i.e. maximum 0.75% sodium chloride as a total).
Acute Renal Failure: in oliguria or anuria phase use a very low protein diet
(1.5 g/kg LW for dog, and 2 g/kg LW for cats; apply mannitol or dextrose
solution (10-20%) intravenously to ensure energy intake; in polyuric phase
treat as in chronic kidney disease.
ADDITIONAL MEDICAL TREATMENT.
Administration of H2-receptor antagonist such as famotidine (5 mg/kg, per
os, 2-4 times a day) decreases gastric acidity and vomiting. Anabolic steroids
such as oxymethalone or nandrolone, have been administered to stimulate
red blood cell (RBC) production in anaemic animals, but this is not really
effective. Recombinant erythropoietin is effective in stimulating RBC pro-
duction, but antierythropoietin antibodies develop in approximately 50% of
animals and may result in refractory anaemia, until species-specific prod-
ucts become generally available, erythropoietin administration is now rec-
ommended only for animals showing clinically apparent signs of anaemia
(e.g. weakness, marked lethargy not attributable to other factors), which
generally occurs at a hematocrit less than 20%. Fluid therapy with polyion-
ic solutions, given intravenously or subcutaneously in the hospital or s.c.
by owners at home, is often beneficial in animals with intermittent signs of
uraemia. Although controversial, vitamin D administration requires prior
resolution of hyperphosphataemia and it may induce hypercalcaemia. Some
therapeutic renal diets tend to alkalinize blood pH (e.g. using sodium bicar-
bonate or MgO) as acidosis begins, which may minimize the effect of acido-
sis and help the animal feel more energetic and eat better (for that the
dUA=dietary undetermined anion or the CAB=cation-anion balance should
be positive).
Feeding tubes may help manage chronic anorexia. Euthanasia or
renal replacement therapy (renal transplantation and/or dialysis) should be
carefully considered if therapy does not improve renal function and allevi-
ate signs of uraemia.
606
21.2.2.16. Nutritional Therapy of Stones in the Urinary Bladder

of Cats and Dogs
© Tony C. A. Buffington
Stones occasionally occur in the urinary tract of cats and dogs. These
stones may from and grow from the ionic components of mineral and organ-
ic material commonly present body and in the diet. The common ions found
in stones include calcium (Ca), magnesium (Mg), nitrogen (NH4) oxalate
(Ox), phosphate (PO4) and urate (Ur). Urinary stone disease may result from
variable combinations of 1) an unsatisfactory diet (diet-induced) or 2) some
abnormality in an animal consuming a satisfactory diet (nutrient sensitive).
The distinction may be helpful in recommending therapy; an unsatisfacto-
ry diet may be changed to any diet known to be satisfactory for the patient,
whereas a diet formulated to accommodate the (largely unknown) disease-
related limitations present in individual patients with nutrient sensitive
stone disease may be necessary. In the author’s experience in the US, nutri-
ent sensitive cases appear to be far more common than diet-induced cases.
Although the mechanisms of crystal and stone formation in the urine
are surprisingly complex and poorly understood, some appreciation of the
basic processes of stone formation in the urine is useful for understanding
therapeutic recommendations. The first step in stone development is crys-
tal formation. This occurs when the concentrations of the ionic components
of the crystal become supersaturated, exceeding the amount that can be
held in solution in the urine. The urine is commonly supersaturated with a
variety of ions however, so observation of crystals in a single urine sample
does not mean the patient is at high risk to form a stone. In such cases no
particular treatment is necessary unless a stone is present, or has formed
in the patient sometime in the past.
Urine supersaturation depends on the amount of the ion ingested and
excreted in the volume of urine produced, and can occur for several possi-
ble reasons. Reasons related to diet include nutrient composition of the
food, food and water intake, and effect of diet on urine pH. Animal factors
include inadequate concentrations of inhibitors or excessive concentrations
of promoters of crystallization, alterations in turnover of mineralized tissues
or in intermediary metabolism, abnormalities resulting in altered urine pH
such as renal tubular acidosis, and activation of the stress response sys-
tem. Factors that predispose to bladder stasis also can play an important
607
role in stone formation, because crystals must reside in the urinary tract for
a sufficient period of time for a stone to form.
Supersaturation of urine with crystal-forming ions depends on the
interaction of dozens of ionic species in the urine. Moreover, the probabili-
ty of crystal formation is further complicated by contributions from other
organic molecules in the urine that may inhibit or promote crystallization.
The pH of the urine also affects crystal formation; struvite, calcium carbon-
ate, calcium phosphate, and urate are less soluble in alkaline urine, where-
as cystine and uric acid are less soluble in acid urine. Urine pH per se does
not appear to have a major effect on the solubility of calcium oxalate,
although most calcium oxalate stones occur in cats with acid urine.
Additionally, drugs such as anticholinergic agents that inhibit normal blad-
der function can play a role in stone formation.
When the solubility product of a particular crystal is exceeded, crys-
tals may form, aggregate, and grow into a stone if the urine is saturated for
prolonged periods. These relationships are shown in FIGURE XXI-9
Methods of estimating the risk of crystal formation using in vitro methods
include relative supersaturation index and activity product ratios. A risk
index that could reliably predict the likelihood of stone recurrence in indi-
vidual patients might help clinicians select appropriate preventative thera-
py. Unfortunately however, none of the indices currently available has suf-
ficient predictive power to be useful to the clinician in making individual
treatment decisions, although they do provide general guidelines. The “gold
standard” for therapy remains the prospective randomized double-blind (to
control for the powerful non-specific effects of diet change) trial in patients
with naturally occurring disease. (FIGURE XXI-11)
608
FIGURE XXI-9: Factors involved in the growth of crystals in urine. From: DRACH, G.W.
(1978): Urinary lithiasis. In: Campbell's Urology (4th ed) HARRISON, J.H. et al. (eds) WB
Saunders, Philadelphia p. 793.
Urinary tract stones contain primarily organic or inorganic crystal ions

and a small amount of organic matrix. When 70% or more of the stone is com-
posed of one type of crystal it is named for that crystal. Secondary crystal
ions can comprise up to 30% of the total weight. For this reasons, increas-
ing the urine volume and frequency of urination, and modifying ion con-
centrations as appropriate are important nutritional factors in managing
stone disease in cats and dogs. In 1996, BORGHI et al. reported results of a
study of 101 controls and 199 stone forming patients. After a baseline study
period, patients were randomized and one group was carefully trained to
achieve a urine volume of at least 2 liters per day. The other group did not
receive any treatment, and no dietary advice was provided. During five years
of follow-up, significant differences were identified between the groups;
urine volumes increased, (from ~1 to greater than 2 vs. to 1.0–1.2 liters/24
609
h), the recurrence rate declined (12 of 87 vs. 27 of 73), and the time to first
recurrence (38.7±13.2 vs. 25.1±16.4 months) was longer in patients in the
intervention group.
Background
The prevalence of all forms of bladder stones in cats and dogs seems to be
roughly 5/1000 cases (0.5%). The two most common types of stones are
struvite and calcium oxalate, with other types such as urate and cystine
occurring occasionally.FIGURE XXI-10 and 11 vizualize and the following
paragraphs provide a brief review of formation of these stone types in cats
and dogs.
FIGURE XXI-10: Calcium sulphate, cal-

cium oxalate, struvite and two types of
calcium carbonate crystals (left)
FIGURE XXI-11: Sodium urate, uricacid, hipp-

uric acid and ammonium urate crystals (right)
610
Struvite. The chemical formula of struvite is magnesium ammonium phos-

phate (MgNH4PO4-6H20). Calcium phosphate also commonly occurs in
small amounts (less than 10%) in struvite stones. These stones formerly
were called “triple phosphate” stones because of the three cations - Ca+2,
Mg+2, and NH4+ - detected by early qualitative analytical methods. Struvite
stones are variable in shape and may be present singly or in large numbers
of varying sizes. They most commonly occur in the bladder, although they
may be found at any site in the urinary tract. Struvite formation and disso-
lution is strongly affected by the pH of the urine. The solubility is marked-
ly reduced when the urine pH rises above ~6.7 due to removal of protons
from di- and monobasic phosphate ions.
Struvite stones and urethral obstruction have been induced experi-
mentally by feeding diets containing large amounts of magnesium to healthy
cats, which led to the conclusion that magnesium was a primary cause of
naturally occurring struvite stones in cats. Subsequent studies found that
these struvite stones dissolved when the urine pH was reduced to approxi-
mately 6, suggesting that the effect of magnesium on struvite formation
depended on the urine pH. The research implicating magnesium and pH as
a potential causes of struvite stones led some cat food manufacturers to
reduce the magnesium content of diets, and to add ingredients to promote
more acidic urine in the cats consuming them. The added ingredients
included variable combinations of corn gluten meal, animal digest (a palata-
bility-enhancing product containing phosphoric acid), methionine, and
phosphoric acid. Some manufacturers also replaced alkalinizing anions of
mineral salts such as carbonate with acidifying ones such as chloride and
sulfate.
Unfortunately, these modifications appear to have increased the fre-
quency of calcium oxalate stones. For example, one laboratory reported an
increase in calcium oxalate stones from 2 to 40% between 1984 and 1995,
and subsequent data suggest that the proportion of calcium oxalate and
struvite stones in the United States currently is approximately equal.
Because no detailed epidemiological studies of urinary stones in cats in pri-
mary care practice currently are available, any effects of changes in com-
mercial diet composition on urinary stone prevalence are not known. The
increased proportion of calcium oxalate stones could be related to increased
consumption of urine acidifying, low magnesium diets however; these mod-
ifications seem to increase urine calcium and decrease urine magnesium
611
concentrations, which might increase the likelihood of calcium oxalate

stone formation in susceptible individuals (see later).
In dogs, urinary tract infection by urease-positive bacteria (especially
species of Proteus spp. and Staphylococci) play the primary role in struvite
stone formation. Hydrolysis of urea by these bacteria liberates ammonia
and carbon dioxide, which increases the pH of the urine and the availabili-
ty of ammonium and phosphate ions for struvite formation. Struvite forma-
tion also is more likely in animals with persistently alkaline urine, even in
the absence of urinary tract infection. In such cases, predisposing factors
include a positive family history, a diet based on vegetable proteins, and dis-
tal renal tubular acidosis.
Calcium oxalate. Calcium oxalate, which is the most common (kidney)
stone type in people, has been found with increasing frequency in cats and
dogs since the mid-1980s. These stones are composed of calcium oxalate
monohydrate (Whewellite) or calcium oxalate dihydrate (Weddelite). Oxalate
frequently is not detected by qualitative analysis, making quantitative stone
analysis necessary. Calcium oxalate stones usually are white, very hard,
and often have sharp, jagged edges. One to many stones may be present.
They are found most often in the bladder and urethra, but recently have
been identified in the ureters and kidneys of cats. Oxalate is derived main-
ly from the diet, with small amounts produced from the metabolism of
ascorbic acid (vitamin C) and the amino acid glycine. In human beings,
increased oxalate ingestion, increased absorption of oxalate from the colon
secondary to fat malabsorption, vitamin B6 deficiency, and inherited defects
of oxalate metabolism may predispose to the formation of calcium oxalate
stones. The role of most of these and other diet-related factors in naturally-
occurring calcium oxalate stone formation in cats and dogs has not been
adequately investigated, and so remains poorly understood.
Altered calcium metabolism also may play a role in development of
calcium oxalate stones. Increased urinary calcium concentrations (hyper-
calciuria) can result from increased absorption of calcium from the intes-
tinal tract (absorptive hypercalciuria), from increased loss of calcium via the
kidney (renal leak hypercalciuria), or from increased release of calcium from
bone (resorptive hypercalciuria). In absorptive as compared to renal leak
hypercalciuria, urinary calcium concentrations are higher after feeding than
during fasting. Chronic acidosis also may be associated with increased uri-
nary calcium concentrations due to increased calcium release from bone.
612
Long-term feeding of acidifying diets may also contribute to resorptive

hypercalciuria. In humans, citrate ions form soluble complexes with calci-
um in the urine, which may inhibit calcium oxalate formation. The role of
citrate, if any, in urinary stone formation in cats and dogs however, is not
yet understood. In contrast to struvite, experimental diet manipulations to
induce calcium oxalate stone formation in healthy cats have not been
reported. In fact, the recent increase in calcium oxalate stone incidence
suggests that some cats may be genetically predisposed to form calcium
oxalate stones, and that diet changes have only unmasked an underlying
predisposition. Previous commercial cat diets that did not restrict magne-
sium or reduce urine pH may have obscured this susceptibility by resulting
in formation of urine in which calcium oxalate stones were unlikely to form.
When manufacturers added acid-forming ingredients and reduced the
magnesium content of foods to prevent struvite formation, the diets may no
longer have protected cats prone to calcium oxalate formation. Cats sus-
ceptible to calcium oxalate formation were now consuming a “provocative”
diet. This situation is somewhat analogous to individuals that cannot digest
lactose because of inability to express the lactase enzyme. If no lactose is
ingested they appear normal, whereas consumption of lactose-containing
products promptly and dramatically reveals the absence of the enzyme.
Calcium oxalate stone disease in humans has been suggested to result from
just such a “powder keg” (the susceptible individual) and “tinderbox” (the
provocative environment) scenario. This hypothesis may be even more com-
pelling in cats, because the overall occurrence of calcium oxalate stones in
the United States appears to be no greater in cats than it is in humans,
despite the fact that much of the cat population consumes similarly formu-
lated diets. Moreover, looking back on earlier studies from this perspective,
struvite stones induced to form in the bladder of healthy cats may have led
to the misperception that commercial diets were the cause of naturally-
occurring struvite stone disease. These diets too may only have unmasked
cats intrinsically susceptible to struvite stone formation.
Studies have shown that differences in breed, age, sex, and repro-
ductive status do not seem to contribute to the apparent reciprocal rela-
tionship between occurrences of calcium oxalate and struvite uroliths in
cats. Indoor housing, however, may be an independent risk factor for calci-
um oxalate stone formation. Indoor cats could void less often, which would
allow a longer period for stone forming products to aggregate in the bladder.
613
Activation of the stress response system also might play a role in some of
these patients. Differences in breed specific risk for calcium oxalate forma-
tion have been reported, however. For example, obesity increases the risk,
and cats between seven and ten years of age are reportedly more likely
develop calcium oxalate stones than are cats younger than two years of age.
Male cats may be 1.5 times more likely to develop a calcium oxalate stone
than are females, and neutered cats are at greater risk than are sexually
intact cats.
Altered calcium metabolism may play a role in development and/or
maintenance of calcium oxalate stones in some cats, so serum total calci-
um should be measured to determine if hypercalcemia is associated with
the stone. Correction of hypercalcemia is important to prevent recurrent
stone formation, and early intervention might avoid development of compli-
cations such as kidney failure. The recurrence rate of calcium oxalate
stones may be as high as 50% in cats. Urinary tract infection, when it
occurs, is thought to be a complicating rather than a predisposing factor to
calcium oxalate stone formation.
As in cats, some breeds of dogs also are at increased risk for calcium
oxalate formation, stones occur about twice as often in males than females,
and neutering and obesity appear to increase the risk. In one study,
Miniature Schnauzers had higher urinary calcium concentrations during
fasting than did Beagle dogs, which increased 3-fold after feeding (i.e.
hypercalciuria seemed to be absorptive). Dogs with hypercalcemia due to
primary hyperparathyroidism may develop calcium oxalate (or calcium
phosphate) stones due to parathyroid hormone-mediated mobilization of
calcium from bone (“resorptive” hypercalciuria).
Urate.
Urate stones are composed of uric acid and its monobasic salt ammo-
nium acid urate. The cause(s) of urate urolith formation remain largely
unknown. Calcium oxalate may be a secondary component, and those
found in patients with Porto systemic shunts often also contain struvite.
Urate stones are small, brittle, and spherical with concentric laminations,
usually multiple in number, and light yellow, brown, or green in colour.
They are found most often in the bladder and urethra, with a recurrence
rate as high as 50%. When present, urinary tract infection occurs as a com-
plication of the urate stone, rather than as a predisposing cause. Some 6%
of stones from cats in the US reportedly are composed of uric acid and
614
urate.
In dogs, urate stones are found most commonly in the Dalmatian and
English bulldog breeds, although other breeds also may be affected. Urate
stones may be found in dogs with portosystemic shunts, possibly due to
reduced conversion of ammonia to urea, and of uric acid to allantoin. A
defect in uric acid metabolism predisposes some male Dalmatian dogs to
form urate stones. Although the defect may be a predisposing factor, it does
not appear to be a primary cause of urate stone formation. This is because
Dalmatian dogs that do not develop stones excrete as much urate as those
that do, and because other breeds (e.g. English bulldogs) that do not have
this defect also develop urate stones. Uric acid is a metabolic degradation
product of purines, which is then converted to allantoin in the liver by the
enzyme uricase. Although uricase is present in the liver of Dalmatian dogs
in amounts comparable to those found in other breeds, they have higher
plasma uric acid concentrations and excrete much more uric acid in their
urine than do non-Dalmatian dogs. This suggests that impaired transport
of uric acid into liver cells may reduce the ability of the liver to metabolize
purines in Dalmatians (SIMKIN, 2005).The proximal renal tubules of the kid-
neys of Dalmatian dogs also appear to reabsorb less and secrete more urate
than do those of non-Dalmatian dogs.
Cystine.
Cystine stones vary in prevalence between species and by geographic
location. The prevalence in the United States is ~0.2% in cats, and less than
2% in dogs, whereas the prevalence among dogs in Sweden is reportedly
8%, and as high as 26% in central Spain. Although the cause(s) of cystine
stone formation remain largely unknown, increased urinary concentrations
of cystine (cystinuria) often results from a hereditable defect in renal tubu-
lar reabsorption of cystine and other dibasic amino acids (ornithine, lysine,
and arginine). Since not all animals with cystinuria develop cystine stones,
the presence of crystals may be a predisposing factor rather than a cause.
Cystine stones are most commonly found in middle aged to older domestic
short haired cats. In dogs, nearly all cystine stones occur in middle aged
males. As with other stones, bacterial urinary tract infection usually occurs
as a consequence of cystine stone presence rather than as a cause.
DIAGNOSIS
Patients with bladder stones present with variable combinations of signs
615
referable to the lower urinary tract. Frequent attempts or straining to uri-

nate, blood in the urine (hematuria), or urinations in inappropriate places
may be reported, or the owner may observe no signs at all until the stone
obstructs some portion of the urine outflow tract. Once a stone is identi-
fied, neither urinalysis nor urinary tract imaging can positively identify the
type of stone present. Moreover, both false-positive and false-negative
results can occur in the urinalysis. For example, hematuria found in a sam-
ple collected from the bladder by needle and syringe (cystocentesis) may be
the result of trauma during collection as well as a stone. The presence or
absence of crystals also can be confusing. Healthy cats and dogs can have
crystals in their urine, many patients with bladder stones have no crystals
in their urine, and some 10% (of cats) with a stone reportedly have crystals
of a composition different from that of the stone. Crystals also can appear
in urine after collection if it is cooled between the time the specimen is
obtained and evaluated.
The pH of the urine can also be difficult to interpret. Increased urine
pH, previously thought to be caused only by diet or urinary tract infection,
also can rise above 7.0 as a result of the stress of travel and examination in
cats (this also may occur in dogs, but to my knowledge has not been report-
ed). Urine pH values measured by dipstick and by pH meter also do not
agree well; a dipstick reading of 6.5 can occur with meter readings from 5.8-
7.0. Thus, none of these parameters is diagnostic for stones. A plain abdom-
inal radiograph can be obtained to identify the presence of radiodense (e.g.
struvite or calcium oxalate) stones larger than 3mm in diameter.
Ultrasound can be used to detect the presence of stones regardless of radio-
density if the stone is sufficiently large. A double contrast radiographic
study or abdominal ultrasound may be necessary when stones are not
radiodense (e.g. urate). Neither imaging technique can positively identify the
type(s) of stone(s) present. When macrocrystals or very small stones (<5
mm) are seen in the urinary bladder of cats, surgery may not be necessary
and voiding urohydropropulsion may be used to remove the stones. The
reader is referred to the recommended readings for a description of this
technique. Stones as large as 5 mm sometimes can be removed from female
cats, but the operator should be prepared to take the cat to surgery for
removal of stones that cannot be successfully removed by urohydropropul-
sion.
616
TREATMENT
The frequency of stones does not appear to be greater in cats or dogs than
in human beings, so no modification of the diet is necessary prior to the for-
mation of the first stone, which announces the presence of a susceptible
animal or an unsatisfactory diet. Therapy of patients with a stone includes
acute therapy to remove or dissolve the stone, and chronic therapy to
reduce the risk of stone recurrence. Stone-specific treatment recommenda-
tions should be based on quantitative analysis of spontaneously voided
stones, or stones obtained by catheterization, urohydropropulsion, or sur-
gery whenever these can be obtained for analysis.
General recommendations
Acute treatment. The first steps in management of urinary stone disease are
to relieve any urinary tract obstruction to re-establish urine flow, and to
correct associated fluid, electrolyte, and acid-base imbalances. Since stru-
vite stones in dogs usually are associated with urease-positive urinary tract
infection, alkaline urine, struvite crystalluria, and a radiodense stone, all
dogs with these findings should have their urine cultured. If infection is
present, appropriate antibiotic or sulphonamide therapy and careful follow-
up should be instituted to ensure elimination of the infection.
Surgical removal and/or medical dissolution may be used to elimi-
nate stones, depending on their composition. Consumption of a canned (but
not dry), magnesium-restricted, urine acidifying diet has been shown to dis-
solve naturally occurring struvite stones in cats and dogs. No medical regi-
men has been shown to successfully dissolve calcium oxalate uroliths how-
ever, so surgery is recommended for these patients. When the stone type is
unknown, the choice of medical or surgical therapy may be offered to the
client after the risks and benefits have been explained. At The Ohio State
University Veterinary Teaching Hospital, the expense of surgical and med-
ical treatment is comparable when all costs are considered.
Chronic therapy. All patients that have formed a stone are at
increased risk to form another one; recurrence rates of stone formation of
50% are common. Approaches that decrease the concentration of potential
stone-forming ions in the urine and increase voiding frequency by increas-
ing urine volume provide the foundation of primary therapy to reduce the
risk of formation of another stone. Unfortunately, many stone forming
patients still are fed dry food. In a recent case series, all cats with a urinary
bladder stone had consumed dry food. Of eight cats with struvite stones,
617
three had been fed dry veterinary foods designed to dissolve or prevent for-
mation of struvite stones, and one of seven cats with calcium oxalate stones
had been fed a dry veterinary food recommended for cats with calcium
oxalate stones.
Increased water intake to reduce the urine specific gravity and
increase the frequency of urination is the cornerstone of therapy for stones
in both human and veterinary medicine. When a stone is diagnosed, we
recommend to clients that the urine of their stone-forming pets be made
consistently COCO - Clear, Odourless, Colourless, and Often by encourag-
ing the patient to consume enough water in canned foods or moistened dry
foods and drinking to increase urine volume. The aim is to reduce the urine
specific gravity to less than 1.030 in cats, and less than 1.020 in dogs,
which corresponds to approximately doubling urine output. Urine specific
gravity should be monitored at periodic re-evaluations to ensure adherence
to the recommendations.
Author recommends that patients that have formed a stone never be
fed on dry food again. This is not to say that they cannot be fed dry food,
just that at least one volume of water should be added to each volume of dry
food before feeding so that enough time can pass for the food to completely
absorb the moisture. Patients also may be fed canned formulations, or
water, other liquids, or sodium chloride or other salts may be added to the
diet to induce a diuresis. Patients must be allowed frequent opportunities to
urinate to avoid urination in an inappropriate location. Provision of sodium
chloride should be avoided in patients with chronic kidney disease or at risk
of fluid retention. Dogs with oxalate and cystine stones also should not be
fed large amounts of sodium chloride because increasing urinary sodium
concentrations may increase urinary concentrations of calcium and cystine.
Recommendations to clients for increasing water intake are presented in
Table XXI-12, and for changing diet when necessary in Table XXI-13.
Most cats do not seem to enjoy eating dry food that water has been
added to, and seem to prefer canned foods. If the cat refuses to accept the
new diet after appropriate introduction to it, water intake can be increased
by the use of drinking fountains, flavoured juices (meat, fish) and the addi-
tion of ice cubes to the cat’s water. (Please see Table XXI-12 for further sug-
gestions). Regimens to reduce environmental stressors may be useful
adjuncts to management of patients with urinary stone disease. In gener-
al, providing diet consistency and feeding time predictability seem to reduce
618
stress responses in most captive animals that depend on their owners for
their food supply. Additional manoeuvres to reduce environmental stress for
cats include providing places to hide and opportunities to express natural
predatory behaviour, such as climbing posts and toys that can be chased
and caught. More information on this aspect of management (for cats) is
available at http://www.indoorcat.org.
Stone-specific recommendations
STRUVITE. Until relatively recently, urinary acidifiers played an important
role in management of cats with struvite urolithiasis without a urinary tract
infection. During the past 20 years or so, most commercial cat foods have
been reformulated to result in a urine pH of approximately 6.5. Urine acid-
ifiers should only be given to cats if the urine pH is greater than 6.5 when
measured by meter from urine collected at home under ad libitum feeding
conditions. Since travel to a veterinarian can increase the urine pH into the
alkaline range due to stress-induced hyperventilation, urine should be col-
lected from the cats in their own environment, and the sample brought to
the hospital for assessment of urinary pH and specific gravity whenever pos-
sible.
Ingestion of large quantities of acid can cause anorexia as well as sys-
temic acidosis. Because so many commercial diets for cats have been mod-
ified to restrict the formation of struvite, acidifying agents such as ammo-
nium chloride or DL-methionine should not be routinely prescribed for cats
with struvite stones or crystalluria. These agents only impose an additional
acid load that may contribute to the development of metabolic acidosis.
Acidifying agents are contraindicated in cats with calcium oxalate urolithi-
asis, and have no known value in the treatment of cats suffering from other
lower urinary tract disorders. Nutrient analyses of a number of veterinary
diets to help prevent the recurrence of struvite stones are available here:
http://www.nssvet.org/food/search.html. These diets may be of benefit in
reduction of risk of recurrence, although few have evidence-based outcomes
documenting their effectiveness in prevention of recurrence of struvite
urolithiasis. Feeding patterns also may influence the development of stru-
vite uroliths due to an increase in pH (postprandial alkaline tide) that may
occur after a large meal, depending on the composition of the diet, so ad libi-
tum or multiple small meal feeding of patients with struvite stones seems
prudent.
Feeding a canned form of a diet designed to dissolve stones has been
619
reported to induce dissolution of recurrent struvite stones in female cats,

which usually occurred within one month. These diets should not be fed to
growing kittens, to pregnant or lactating queens, or for prolonged periods
after dissolution of the stone to avoid the adverse effects of chronic acido-
sis. The use of urinary acidifiers to maintain urine pH in the range of 6.0-
6.5 has been suggested in dogs because struvite and hydroxyapatite are
most soluble in acidic urine. In most dogs with struvite urolithiasis, eradi-
cation of urinary tract infection will return urine pH to this range. Use of
urinary acidifiers in the presence of infection by a urease-positive organism
is futile. If the urine pH remains alkaline after elimination of urinary tract
infection, other potential causes (e.g. dietary, familial, metabolic) of alkaline
urine should be investigated.
CALCIUM OXALATE. Although a decreased risk of recurrence of oxalate
stones related to diet change has never been documented in cats, changing
to a diet that is less acidifying and that has not been magnesium-restricted
seems reasonable, as long as the resulting urine specific gravity is <~1.030.
Many pet food manufacturers offer diets designed to reduce the probability
of stone formation. These diets have been tested in healthy cats, and it is
hoped that they will prove effective in cats with naturally occurring stone
disease. Unfortunately, the frequency of recurrence of stone formation in
cats is not known, so the safety, efficacy, and cost-effectiveness of these
diets cannot yet be determined. Research to answer these questions is sore-
ly needed.
Although provision of a diet restricted in calcium and oxalate seems
logical to reduce the risk of another stone, no evidence based studies in cats
or dogs with naturally occurring disease currently are available to support
or refute this recommendation. Moreover, intake of calcium with food
appears to decrease risk for symptomatic kidney stones in humans, where-
as isolated intake of supplemental calcium may increase the risk. Thus, the
timing and association with intake of other nutrients may also influence the
role of calcium and other nutrients on stone formation, and these effects
may be different in animals with naturally occurring disease than in healthy
ones. Reducing only one of these minerals could alter the availability of the
other, further complicating interpretation of effects. Dietary phosphorus
should not be excessively restricted because reduced phosphorus intake
could result in increased activation of vitamin D3 to calcitriol by 1-α-hydrox-
ylase in the kidney and cause increased intestinal absorption of calcium.
620
Also, urinary pyrophosphate derived from the diet may function as an

inhibitor of calcium oxalate formation. Magnesium intake should not be
restricted because it may serve as an inhibitor of calcium oxalate formation.
Current recommendations on salt intake are controversial, but sodium
chloride supplementation has been reported to increase urinary calcium
excretion and increase the risk for oxalate uroliths in people. Avoidance of
vitamin C has been recommended because ascorbic acid is a metabolic pre-
cursor of oxalate, but to my knowledge no stones have been reported to
develop in cats or dogs as a result of vitamin C ingestion.
Administration of citrate as potassium citrate (Urocit-K®) at a dosage
of 100 to 150 mg/kg/day has been recommended since urinary citrate
seems to act as an inhibitor of calcium oxalate formation and its alkaliniz-
ing effect may reduce bone release of calcium, but it is unclear if supple-
mentation of the diet with citrate actually changes the risk of stone recur-
rence in cats or dogs. To date, only the feeding of the Royal Canin S/O diet
has been shown to result in urine that is undersaturated for both struvite
and calcium oxalate in patients with naturally occurring stone disease. To
what extent this approach will reduce the risk of stone recurrence as pre-
dicted remains to be documented.
URATE Attempts to dissolve ammonium urate stones so far have been
unsuccessful and surgery or voiding urohydropropulsion is required to
remove these stones. Dissolution and prevention protocols for dogs include
some combination of urine dilution, low purine diet, alkalinisation of urine,
and allopurinol. Dissolution of an ammonium urate stone in a cat was
reported to occur using a combination of allopurinol and a diet relatively low
in purine precursors. Although allopurinol reduces the formation of uric
acid, xanthine uroliths can occur, and the efficacy and potential toxicity of
allopurinol in cats are not known.
Prevention of urate stone recurrence requires some combination of
urine dilution, reduced purine intake, and allopurinol if the previous efforts
are inadequate. Diets low in organ-derived meats reduce the ingested purine
load. Feeding a low protein, low purine diet reduced urinary excretion of
urate in normal dogs, and a purine-restricted, non-acidifying diet (e.g., Hill’s
u/d) has been recommended for dogs with urate urolithiasis, but no pub-
lished clinical studies confirming its efficacy are available. If the dry form of
the diet is fed, at least one volume of water per volume food should be thor-
oughly mixed with the diet before feeding. The value of the recommendation
621
of urine alkalization to avoid urate stone recurrence is uncertain. Canine

urate calculi nearly always are composed of ammonium acid urate and,
whereas uric acid becomes more soluble in alkaline urine, urate becomes
less soluble. The pKa (point where concentrations of uric acid and urate are
equal) of the reaction uric acid → urate + H+ is approximately 5.4, but the
pKa of the reaction NH4+ → NH3 + H+ is nearly 9.4. As urine pH increases
from 5.4 to 7.4, the amount of urate doubles, but the amount of ammoni-
um decreases by only a negligible amount. Thus, alkalinization of the urine
may be of limited benefit. Hydrogen and ammonium ions are thought to
cause flocculation of ammonium urate in urine. Administration of NaHCO3
or potassium citrate decreases urinary hydrogen and ammonium ion con-
centration. Administration of NaHCO3 reduces renal tubular ammonia pro-
duction, and the sodium load contributes to the induction of polyuria.
Hence, administration of NaHCO3 at a dosage that keeps the urine pH of
7.0-7.5 has been recommended.
Allopurinol is a competitive inhibitor of the enzyme xanthine oxidase,
which converts hypoxanthine to xanthine and xanthine to uric acid during
the course of purine metabolism. One of its own metabolites, oxypurinol,
also is an inhibitor of xanthine oxidase. Allopurinol therapy reduces the
amount of uric acid formed from hypoxanthine. It is recommended for use
in dogs with urate urolithiasis at a dosage of 30 mg/kg/day divided bid. A
dosage of 30 mg/kg/day divided bid for one month, followed by 7-10
mg/kg/day has been recommended for prevention of recurrence. XANTHINE
STONES may develop in some dogs receiving allopurinol at > 30 mg/kg/day.
Ideally, the dosage of allopurinol should be adjusted so that 24-hour uri-

nary excretion of urate is approximately 300 mg. If 24-hour urate excretion
exceeds 300 mg, the allopurinol dosage should be increased; if it falls below
300 mg, the dosage should be decreased. The dosage of allopurinol should
be reduced in the presence of renal failure since the kidneys excrete it.
Allopurinol should be prescribed only after the desired increases in urine
volume and frequency have been achieved to avoid formation of xanthine
crystals or stones.
CYSTINE. In addition to increasing urine volume in patients with cys-
tine uroliths, cystine becomes more soluble as the pH increases. Therefore,
a non-acidifying canned food seems appropriate for these patients. N-(2-
mercaptopropionyl)glycine (2-MPG), which decreases the concentration of
cystine by a thio- disulfide exchange reaction resulting in a more soluble
622
compound, has been reported to be of benefit in preventing recurrent cys-

tine uroliths in a cat. No adverse effects were documented in this one cat.
Potassium citrate can be used to further alkalinize the urine if necessary.
Sodium bicarbonate cannot be recommended because dietary sodium may
enhance cystinuria.
Summary
Diet therapy may benefit patients with urinary tract stone disease. Some
also seem benefit from provision of a single, moist(ened) diet, if such a feed-
ing plan is not too stressful to the patient or the owner. The issues sur-
rounding stress, diet change and disease currently are controversial, and
further investigation of these relationships is needed. Cats with naturally
occurring urolithiasis seem to form stones despite consumption of dry diets
that should inhibit their formation. This observation emphasizes the signif-
icance of that most important nutrient, water, in the treatment of patients
with urinary stone disease. Whether veterinary foods designed to treat
patients with a stone reduce the (unknown) rate of stone recurrence to a
greater extent than the patient’s usual diet when appropriate maneuvers to
ensure adequate dilution of the urine are implemented remains to be deter-
mined. In humans, “simply” diluting the urine by increasing water intake
has been shown to reduce the 5-year-rate of calcium oxalate recurrence by
55%. Patients who have had cystic calculi initially should be monitored reg-
ularly for recurrence. In addition to ongoing encouragement of clients to
adhere to therapy protocols, it is ideal to obtain an occasional imaging study
for a quick “stone check” for silent recurrence.
623
Table XXI-12: Diluting the Urine by Increasing Water Intake
624
Table XXI-13: Tips to Help You Help Your Pet Change Its Diet
625
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Weiss, D.J., Armstrong, P.J. and Gagne, J. (1997): Inflammatory liver disease. Semin Vet
Med Surg (Small Anim). 12(1), 22-27.
Westropp, J., Buffington, C.A.T. and D.J. Chew, D.J. (2005): Feline Lower Urinary Tract
Diseases. In: Ettinger, S.J, and E.C. Feldman (eds): Textbook of Veterinary Internal
Medicine. St. Louis: Elsevier-Saunders, pp 1828-1850.
627
21.2.3. Physiological conditions with special dietary needs
21.2.3.1. Reproduction
Susceptibility of the reproductive functions of bitch and queen is much
higher that it is in case of male dog or tomcat. Nevertheless, the extremely
high zinc needs of sperm production have to be emphasized; thus before
and during the mating season the animal protein, viscera (in the first line )
liver, or zinc supplementation (pill, zinc sulphate powder, ZnO) is essential
for them.
Feeding level during gestation and lactation should be describe
according to the actual body condition. For the pregnancy needs bitches
require significantly higher nutrient provision only in the last three weeks,
the queen, on the contrary, already from the midterm. Pregnant queen has
an appropriate weight gain, if her live gain on the day of parturition equal
to the following equation: ∆W, g = 890 + 107 n (where “n” = litter size). For
example a queen of 4 kg of LW having a kitter size of 6 have to achieve the
5.532 kg to the day of parturition, showing the important nutritional needs.
Gestational needs of pregnant bitch are comparatively lower, in turn, dur-
ing lactation, especially having large litter, may attain the quadruple of the
maintenance requirement. To cover that, a very concentrated feed should be
fed, unless loss of hair (see above) and because of the exhaution of calci-
um storages, puerperal eclampsia will occur. The typical incidence of the
latter falls between the week 2 to 4 of nursing.
In memoriam Vitéz (1996-2004)
629
21.2.3.2. Nutrition and the skeletal system in dogs

©Richard C. Nap
Developmental abnormalities related to the bones and joints are often diag-
nosed in dogs and turn out to be the reason for serious clinical problems,
which may significantly influence not only the dog’s health, but also the
relationship between the dog and the owner. Impaired locomotory function,
disability and pain, significantly impact the so-called human-companion-
animal or human-pet bond. Pups of large and giant breeds are the fastest
growing species and within a few weeks they add incredible size and volume
to their birth measures. They can weigh well over 40 kilos at 6 months and
within one year they grow from 500 grams to 60, 70 kilogram bodyweight.
Rapid longitudinal growth takes the radius from 9 centimeters at the age of
9 weeks, to 20 centimeters at the age of 6 months, a doubling in length in
a period of 4 months. However, this spectacular growth process has been
identified to be at risk for genetic and environmental influences. Many pop-
ular dog breeds have been diagnosed with skeletal disorders and a heredi-
tary background has been confirmed for hip dysplasia (HD), elbow dyspla-
sia (ED), osteochondrosis (OCD) and radius curvus syndrome (RCS).
Breeding programs for a variety of practical reasons have limited success in
reducing the consequences of the conditions.
It has been shown that nutrition can play a significant role in the
manifestation of the developmental skeletal conditions such as HD and ED.
This subchapter will discuss these influences and will give recommenda-
tions for feeding of large and giant breeds during growth. Size and growth
in people is expressed and measured in centimeters (or inches) whereas size
and growth in dogs is often expressed in kilogram bodyweight (pounds or
stones). This may be confusing because by expressing growth as a measure
of weight, the longitudinal (vertical) and bodyweight (horizontal) growth are
mixed up. An overweight immature dog that has normal height but excess
kilo’s did not grow too fast, but became overweight during growth. It is bet-
ter to state that the growth rate in such case is normal, based on the fact
that we consider growth to be the result of longitudinal skeletal growth orig-
inating in the long bone physes in the extremities.
SKELETAL GROWTH
Long bone growth is characterized by transformation of cartilage into bone
in the epiphyseal areas at both ends of the long bone, as well as bone
630
remodeling (FIGURE XXI-12). This process is called endochondral ossifica-

tion. Cartilage in the epiphysis contributes during maturization to the
length of the metaphysic and the diaphysis as well as to the epiphysis itself,
as the subchondral bone of the joint cartilage. Organized in columns the
cartilage cells (chondrocytes) mature towards the hypertrophic and miner-
alization zones of the columns. Disturbed chondrocyte maturation in the
hypertrophic zone is found in clinical conditions of OCD in the joint carti-
lage, and cartilage cones in the growth plates. These have a similar back-
ground as the maturation process ceases and the cartilage layer becomes
thicker in the affected areas, resulting in interference with normal physical
properties and function. Bone remodeling is an essential part of normal
bone growth because the metaphyseal area of the long bone is much wider
than the diaphyseal area, while the medullary canal is absent and needs to
be widened throughout the diaphyseal growth. The same is true for the
spinal canal and the vascular canals that traverse bones. This bone remod-
eling takes place at a very high turnover rate and forms an essential part of
the active calcium metabolism during the growth phase.
FIGURE XXI-12: Longitudinal bone growth.

Legend: Schematic representation of the proximal end of a long bone (A). including epiphysis
(Eph), physis (Ph), metaphysic (Mph) and proximal part of the diaphysis (Dph). The enlarged
section (B) represents the physeal zone (from proximal to distal): the resting zone (RZ), the pro-
liferation zone (PZ), the maturation zone (MZ), and the zone of hypertrophy and provisional
calcification (HZ).
Chondrocytes are oriented in longitudinal columns and move away from the resting cell zone
at the epiphyseal side, to the zone of calcification at the metaphyseal side of the growth plate
during proliferation, maturation and hypertrophy. During endochondral ossification physeal
cartilage is replaced by primary spongiosa, which in turn will be transformed into secondary
spongiosa during metaphyseal (re)modeling. During longitudinal growth ( ↑ ) skeletal remod-
eling takes place and includes peripheral bone resorption ( ─ ) in the proximal part of the
metaphysic, and bone formation ( + ) in the more distal metaphysic and diaphysis.
——————————————————————
631
DEVELOPMENTAL BONE AND JOINT DISEASES IN THE GROWING DOG

Hip dysplasia (HD) is a malformation of the various components of the hip
joint that result in poor congruency and loose fitting of the joint. There can
be a flattening of the acetabulum and the femoral head, and a significant
joint laxity in the clinical exam. The condition is mostly diagnosed in large
and giant breed dogs, but does occur in small breeds and cats. It has a
hereditary origin and in the vast majority of cases both hips are affected. In
case of unilateral HD other diagnoses are most probably involved. Similar
to Elbow Dyslasia (ED), HD being a developmental condition, it is by defini-
tion a disease of the growing animal, although it may have serious conse-
quences for later life due to the development of osteo-arthritis (also called
[osteo-] arthrosis in some countries). The poor and loose fitting of the hip
joint results in instability and may cause serious discomfort or pain,
although this is not the case in all affected dogs. At least it is not recognized
as such. The diagnosis is made based on the history, the clinical signs and
exam, and the radiographs of the hip joints. One of the clinical problems
with HD is the fact that the clinical signs are not always consistent with the
radiographical signs. Nutrition has been proven to have a significant impact
on the development of HD.
ED is a group name for at least 4 independent conditions affecting the
elbow. They all have a hereditary back ground and can occur in the same
elbow, or dog, but have been proven to transfer independently to the next
generation. In many cases the elbows are affected bilaterally. The diagnoses
under the umbrella of ED are: 1. Ununited Anconeal Process (UAP), 2.
Fragmented Coronoid Process (FCP), 3. osteochondritis dissecans (OCD)
and 4. Elbow Incongruency (INC). In all 4 of these conditions disturbed
endochondral ossification is part of the ethiology. For 3 out of 4 (1, 3, and
4) of the conditions nutrition is a significant environmental risk factor in the
development. Radius Curvus Syndrome (RCS) is the result of a-synchro-
nous growth of the radius and ulna. Under normal circumstances the
radius and ulna grow in sink and form the antebrachium and compose
elbow joint. In case of RCS, the distal ulnar growth (over 90% of ulnar
length) is retarded due to the presence of a cartilage cone, which interferes
with normal growth velocity. Trauma to the distal ulnar growth plate can
have similar (often unilateral) effect. As a result, the radius will be relative-
ly long compared to the fellow ulna, resulting in RCS (shortening of ante-
brachium, dorso curvature of the radius, valgus deformity, exorotation of
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the foot, and elbow joint incongrously).

Bone remodeling is an important part of overall bone growth with high
calcium turnover as a result. After ingestion and absorption, calcium is
deposited immediately in the skeleton (accretion) and reabsorbed based on
the needs during the mineralization of newly formed and remodeling bone
processes of the growing skeleton. Calciotropic hormones like parathyroid
hormone (PTH), calcitonin (CT) and vitamin D3 balance the plasma calcium
(Ca) levels within strict levels, in order to allow physiological processes such
as nerve conduction, muscle and cardiac functions, enzyme activity and the
blood coagulation cascade, while securing proper skeletal growth. Vitamin
D3 and PTH contribute to increased plasma Ca levels, whereas CT lowers
Ca levels and increases accretion.
Biosynthesis of PTH in the parathyroid glands and secretion are stim-
ulated primarily by a reduction of plasma Ca. Typically, plasma PTH levels
are higher during the first months of life in giant breed dogs compared to
levels during adulthood. PTH is the principle hormone involved in the
minute to minute fine regulation of plasma Ca. The target organs include
the bone and kidneys. Increased PTH levels result in increase in an plasma
Ca via increased bone resorption and enhanced renal reabsorption of Ca.
PTH has also been found to promote the absorption of Ca from the gastro-
intestnal tract (Table XXI -14). This effect is most likely to be indirect via
PTH induced renal 1,25-dihydro-cholecalciferol synthesis. The D vitamins
(vitamin D; calciferols) are fat soluble substances of plant (vitamin D2; ergo-
calciferol) or animal sources (vitamin D3; cholecalciferol). Vitamin D3 is syn-
thesized in subcutaneous tissues in herbivores and omnivores following an
exposure to sun ultraviolet B. However, dogs do not seem to be able to pho-
tosynthesize their vitamin D3 in sufficient amounts and therefore they need
vitamin D3 from nutritional sources. After intestinal absorption vitamin D3
is bound to plasma globulin and transported to the liver for hydroxylation
to 25(OH)D, the most abundant vitamin D metabolite in the circulation. In
renal tubular cells it is again hydroxylated to either calcitriol [1,25(OH)2D]
or to 24,25-dihydroxyvitamin D [24,25(OH)2D. The rates of the later two
metabolites seem to be reciprocally related and controlled by similar factors,
i.e., high dietary Ca intake results in decreased 1,25 and increased 24,25
formation. The 1,25(OH)2D synthesis is well controlled and regulated by
plasma Ca concentration, by PTH and possibly CT. The 1,25 plasma con-
centration is increased by low plasma Ca and P, and enhanced Ca needs
633
during pregnancy, lactation and growth. Plasma 1,25 levels do not have sig-
nificant daily fluctuations. The 1,25(OH)2D activates osteoid synthesis and
mineralization of osteoblasts. In addition, it works as a “permissive factor”
for PTH effect on osteoblast. Under the influence of 1,25 both the activity
and the number of osteoclasts increase. 1,25 stimulates active intestinal Ca
and P absorption, osteoclasia and renal reabsorption of Ca and P.
Calcitonin (CT) is produced in the C-cells in the thyroid and parathy-
roid glands. Relatively large amounts of CT are stored in the C-cells to act
as an emergency hormone to protect against hypercalcaemia. The plasma
and extra cellular Ca concentration are the main stimulus for CT secretion,
and CT levels rise after Ca intake, under the influence of the gastrin induced
CT secretion. The net effect of CT secretion is to lower the plasma Ca and P
concentrations. Under CT influence osteoclasts decrease their activity and
decrease in number. Similar to PTH, CT decreases renal absorption of P to
cause an increased P excretion via the urine.
Table XXI-14: Source and effects of parathyroid hormone (PTH), 1,25 dihydroxyvitaminD
(1,25(OH)2D)), and calcitonin (CT) on skeleton, kidney and gastro-intestinal (GI) tract.
------------------------------------------------------------------------------------------------------
PTH 1,25(OH)2D CT
Effects on parathyroid liver & kidney thyroid
-------------------------------------------------------------------------------------------------------
1 skeleton osteoblast ▼ ▲ no
osteoclast ▲ ▲ ▼
__________________________________________________________________________________
2 kidney Ca-excretion ▼ ▼ no
P-excretion ▲ ▼ ▲
1,25(OH)2D ▲ ▼ ▲
__________________________________________________________________________________
3 GI tract Ca absorption no ▲ no
P absorption no ▲ no
--------------------------------------------------------------------------------------------------------
▲ = stimulation, ▼ = inhibition, and no = no effect
THE NON-NUTRITIONAL FUNCTIONS OF NUTRITION

Nutrition is mostly regarded from the aspect of providing energy and nutri-
ents to support basic body metabolism or maintenance, body growth, phys-
ical activity, pregnancy, lactation etc. Textbooks, symposia, seminars and
lectures on nutrition deal in great detail with the minimal and optimal
requirements and extensive debate is going on about the optimal and toxic
levels of ingredients, minerals, vitamins and trace elements. Significant
research investments are made to enhance our understanding in term of the
interaction between nutrients and the interaction of nutrients and body
634
function. As an example, this has in recent years significantly increased our

understanding of the importance of fatty acids and has contributed to our
knowledge that not only omega-6 (n-6) but also n-3 fatty acids are essential
for optimal health, both in humans and in animals. A new field of science
is developing called nutrogenomics, and deals with how nutrition influences
gene expression (See Chapter XVII). However, a very important aspect of
nutrition is often overlooked although it has significant impact in our soci-
eties. The food in restaurants is often decorated in such a way, that it looks
more like a piece of art than a meal. By enjoying a meal under these condi-
tions we communicate to those that are important to us.
Over the past 25 to 40 years, pets have come a long way from being
kept outside the house in a barn, moving gradually into the house, and now
in many cases sharing the house, and often the bedroom (and the bed?)
with their owners. Pet owners are called “pet parents”. Pets share the house
with us, and they have become family members. Owners look at them as
children and companions. A recent (2005 American Kennel Club) survey
showed that when asked “is a person who is in a house, with only a dog”,
alone, or not alone?, 79% of pet-owners consider such a person NOT alone,
while this number is still a significant 62% for non-owners. The conclusion
is that for the vast majority of the population (USA) a person with a dog in
the house is no longer alone. This trend is spreading rapidly and will be the
norm in many countries within a few years. Grand parents bring chocolate
bars for grandchildren not because the kids have an energy or chocolate
requirement that needs to be met, but because they want to communicate
that they like them (but have little in common to talk about). Owners want
to communicate to pets that they like them, but have little to talk about. So
they feed them. And it is so nice to be rewarded by the pets behavior when
preparing the meal. Owners often times talk while preparing the food and
pets respond both verbally (especially cats) and non-verbally (behavior).
This is really funny and hard to resist. Before the age of 1, most of us have
been taken out in our stroller to go and feed the ducks. In the Zoo the man-
agers strongly request (it is forbidden) visitors „NOT TO FEED” the animals.
However despite the „DO NOT FEED” signs all over the place, people bring
food and they feed the animals. They cannot resist the temptation although
they know that they should not do it because it is bad for the animals’
health. The emotional desire to communicate via the food is too strong.
Pets, who are the lowest in the hierarchic ranking in the house, like
635
the attention given by a higher (and even the highest) ranked person in the
group, and will repeat the behavior that has led to the attention (food), inde-
pendent of hunger or need for nutrition. Food is available and both the
owner and the pet like the feeding ritual. So it is very likely that this emo-
tionally driven behavior will be repeated. Many of the complete and bal-
anced diets and treats offer high quality, highly digestible nutrition. The
result is that our pets receive too much food compared to their real needs.
On top of that, people like “teddy bears” and baby faces. This preference for
round shapes compared to squares first shows when people choose a puppy
from the litter. Teddy bear type puppies are preferred (New Foundlander,
Chow Chow, Bernese Mountain Dog, e.o.) and the new owners almost
always choose the most “teddy bear like” puppy from any litter. Also judges
who are involved in dog (and cat) shows like teddy bears. As a result, the
breed standard (what the judge wants to see) is an overweight dog (Labrador
Retriever) with too much belly compared to the optimal body condition. The
standard dog that people have in mind as “how the dog should look”, is
overweight, and this includes the growing dog and the young adult.
Overweight and obesity are problems in our society, and our pets also have
become part of that, after becoming our family members.
NUTRITION DURING GROWTH

Supplied by mother milk during the lactation period, puppies start to eat
solid food from week 3 or 4. First in small amounts, but soon it contributes
fairly significantly to the total energy and nutrient intake. Weaning is com-
plete at the age of 8 weeks when the pups are separated from the mother.
The total daily energy requirements increase rapidly during growth, but the
energy requirement per kilogram bodyweight is highest in the pup at the age
of weaning. After that, the requirements decreases on a per kilo bodyweight
basis as they are linked to the metabolic body weight (W0.75). In case of a
complete and balanced diets is provided to the growing pup, the require-
ments for all nutrients need to be balanced to the energy content of the food.
When energy needs are met, food intake stops, and all other nutritional
components need to be presented in optimal amounts and balance. This
puts a significant responsibility on pet food companies that market com-
plete and balanced diets for growing dogs, especially for the fast growing
pups of large and giant breeds.
636
NUTRITIONAL INFLUENCES ON GROWTH, SKELETAL DEVELOPMENT
AND JOINT INTEGRITY
Universities and pet food companies have contributed significantly to the

understanding of nutritional needs of growing dogs. Based on previous work
in Scandinavia (GRONDALEN, KASSTROM, OLSSON), HEDHAMMAR (Uppsala,
Sweden) contributed a mile stone publication in 1974, when he finished his
research on the influence of overfeeding growing Great Dane pups.
Compared to non-overfed pups, his ad libitum fed test group showed signif-
icantly more skeletal problems. This included more HD and more OCD, as
well as more RCS. All other conditions between the groups were kept iden-
tical. The composition of the food was the same but the amount was differ-
ent. This resulted in excess intake of both energy and all other ingredients
because the dogs were fed a “complete and balanced” diet. Starting with the
publication of his thesis in 1986, HAZEWINKEL (Utrecht, Netherlands) and co-
workers (NAP, SCHOENMAKERS, TRYFONIDOU) have contributed to clarifying the
effect and role of different components of the feed for the growing dog.
Extensive research has shown that intake of excess calcium (Ca) is respon-
sible for most of the developmental problems of the growing skeleton. The
reason behind it all is that growing puppies cannot down-regulate Ca intes-
tinal absorption enough to protect themselves from the “toxic effects” of
excess Ca supply. When feeding standard diets containing 1.2% Ca (15 MJ
ME/kg DM) during the growth period in both small and large breed pups
about 45% of Ca is absorbed. If the diet contains smaller or minimal
amounts of Ca the body is able to up-regulate the absorption efficiency to
95% of ingested Ca. However, when excess Ca is provided, the ability for
adequate down regulation is lacking. The reason being that below the min-
imal 40 to 45% the intestinal absorption is passive and not vitamin D3
dependent. The excess Ca that is absorbed under these circumstances is
routed to the bone tissue immediately, and PTH, vitamin D3 and CT man-
age to keep plasma Ca levels within narrow margins. The excess Ca deposi-
tion (accretion) is not balanced by higher Ca resorption during the process
of bone turnover. The excess Ca interferes with the endochondral ossifica-
tion and the bone remodeling resulting in increased frequency and severity
of growth disturbances such as OCD, ED (OCD, UAP, FCP and Inc), RCS
and Enostosis (Panosteitis) and Wobblers’ disease respectively, in high Ca
fed dogs. It has also been shown that excess Ca supply before the age of 8
weeks completely disrupts the normal balance between calciotropic hor-
637
mones and can upset the regulatory system even when the mineral intake
is normalized after weaning. Pups in the first weeks and months of life,
proved to be most sensitive to excess Ca intake.
Phosphorus (P) intake is balanced at 90% of the Ca level. As long as
the ratio between Ca and P does not exceed the 60% to 150% margins, no
problems are foreseen. However, in case of low Ca intake in combination
with relatively high P intake (all meat diet) skeletal problems will occur. The
low Ca level in meat (0.03%) will drain Ca from the skeleton deposits built
up during pregnancy and lactation. Ca resorption from the bone is further
increased by CT-driven P losses via the urine. Ca is bound to the P and lost
via the urine despite the low intake. The result is a rapidly demineralization
and weakening of the skeleton, and pathological “spontaneous” fractures
occur sooner (in large and giant breeds) or later (in small breeds).
Vitamin D3 plays an important role in the Ca homeostasis and is an
essential component of the food. Vitamin D3 increases plasma Ca levels by
Ca absorption from the intestinal tract and reabsorption from the pre-urine
in the kidney. Different to the metabolism in humans and other species, it
has been proven that in the dog, vitamin D3 is not metabolized in the skin
under the influence of ultraviolet light. Nutritional intake is the only vitamin
D3 supply for dogs. The good news is that the vast majority of pet foods pro-
vide more than enough vitamin D3 to meet the requirements, and the over-
supply by “concerned pet-owners is more of a concern than deficiency. The
research in Utrecht has shown that high (10Î×normal) vitamin D3 intake
during growth can result in disturbances in skeletal development similar to
those seen in case of high Ca intake. When excess vitamin D3 intake is
started at the age of 3 to 4 weeks, dogs in the high vitamin D3 test group,
show physeal disturbances, similar to those diagnosed during rickets
(rachitis) which is typical for low vitamin D3 intake during growth but at a
higher age. The change in Ca setpoint is supposed to be reason for these
findings (TRYFONIDOU).
Since nowadays complete and balanced diets provide Ca, P and vita-
min D3 in sufficient amounts and well balanced, the mineral and vitamin
D3 supply for growing pups seems to be a problem of the past except under
specific conditions. The risk for problems increases when
638
In addition to the above listed potential causes of bone and joint prob-
lems during growth, may be the biggest risk of all related to the role of
nutrition in the communication between the owner(s) and the pup (or kit-
ten). The intake of excess feed results in rapid (horizontal) bodyweight
growth while the longitudinal (vertical) growth is similar to dogs receiving
normal or adequate amounts indicated by their real requirements. But not
only do the dogs ingest too much energy, they also consume higher total
daily amounts of minerals. Supported by the results of scientific research
(Hedhammar [Great Danes], and Kealy [Labrador Retrievers]) it is obvious
that owners who do overfeed their pups during growth, significantly
increase the risk for skeletal problems such as OCD, ED and HD. The con-
sequence of the relative overweight during the growth phase is, that the
immature cartilaginous hip joint gets overloaded and changes shape. The
femoral head and the acetabulum flatten and the position of the acetabu-
lum is more vertical (increased inclination angle). The clinical consequence
is a significantly higher grade of HD in overfed overweight dogs. The excess
intake of daily minerals including Ca, results in the above mentioned
increasingly more severe OCD and related diseases in case of overweight.
The veterinarian and her/his staff play an important role in the education
of the client in order to try and prevent such overfeeding. Offering weight
management programs and advising during the growth period for optimal
condition, are a service that people can expect from their veterinarian. Not
639
only is this important for the short term well-being of the dog, but also
these bone and joint conditions seriously impact the long term health and
life expectancy of the pet. Overweight results in the development of more
significant osteo-arthritis (OA) or arthosis, and reduces life expectancy by
more than 3 years. The OA results in pain and disability and can be the
reason for life long medication and treatment, major surgeries and pre-term
euthanasia. Anything within the circle of influence of the veterinary com-
munity that can be done to reduce the risk of such conditions should be
developed and offered to the clients.
NUTRITIONAL CONSIDERATIONS IN THE AGING DOG’S SKELETON

Meeting the nutritional requirements for adult dogs is less critical compared
to growing dogs, especially in small and medium size breeds. The detri-
mental consequences of feeding an all-meat diet to an old small breed dog,
often does not show until during an annual dental (extraction) procedure at
the veterinary hospital, a “pathologic” fracture occurs in the mandible, or
when the dogs fractures a leg when jumping from the lap of the owner. This
can be years after the poor feeding habit was introduced.
Most complete and balanced pet foods supply the adequate levels of
nutrients in a balanced formula. Differences between foods can be based on
quality and digestibility, but are often more difficult for the owner to judge.
Some indication can be obtained from the stool consistency and volume aa
well as from coat and fur conditions, but none of these will provide black-
and-white type indicators. In addition to their own observations, owners will
rely on information by others that they trust and the veterinarian for sure
is among the most trusted source of nutritional advice. Dogs are not carni-
vores and even if they were, they need more than meat (protein). Carnivores
eat their whole (often herbivore) prey and this includes the content of the
abdomen, which is filled with pre-digested vegetables. The mature predators
prefer the belly content over the muscles of the legs, which are left for the
juniors.
The main concern for mature and senior pet nutrition is the amount
of food consumed daily. Owners have a tendency to give more than the pet
needs. In addition, the enthusiasm of family members to walk the dog
decreases with time compared to when the puppy entered the house and
owners tend to compensate attention with additional food. The need for
communication with the pet continues and, as a result, the mature pet most
640
likely will end up being overweight or even obese. Similarly to the situation
in people, it is much easier to prevent obesity than to restore optimal
weight. Once the vicious circle has been launched it spirals down (or may
be “up” is a better description) and is hard to reverse. Older people have
lower basal energy requirements per kg LW and typically are less active in
sports. The same is true for pets, and in most cases the owner and the page
age simultaneously. A complicating factor is the fact that people find it hard
to appreciate how small their pet actually is. Especially long hair pets look
much bigger than they really are, and this may add to the feeding behavior.
Everybody knows how little the grandparents eat. But compared to a 70
years old grand mother with a bodyweight of 60 kilograms, the nutritional
needs of a 10 years old 12 kg pet are incredibly small. Too small for many
owners to satisfy their own(ers) needs to communicate with the pet, and to
care for it. The result is overfeeding, overweight and obesity.
Overweight and obesity add significantly to the pain and discomfort
of animals suffering osteo-arthritis (OA). Most owners of pets that have suc-
cessfully completed weight reduction programs, report that the pets appears
to be years younger. They are more active, show more interest and have a
higher endurance when taken out for a walk. Obviously this sets off a
vicious circle in the opposite direction and will contribute to a healthy
weight management and exercise pattern. It is not unusual for an adult
Labrador Retriever to have up to 8 kg overweight. Just imagine how it would
be like for the owner to loose 25 kg. When owners insist or emotionally need
to add “something” to the complete and balanced diet, it is advised not to
have this exceeding 5% of total daily intake, with the exception of minerals,
trace elements and vitamins. These should not be added at all. Some own-
ers are convinced that fruits or cheese should be added. When the owner
needs it, and dog likes it and the amount is small, there are no problems to
be expected.
NUTRITIONAL MANAGEMENT OF OSTEO-ARTHRITIS (OA)

Nutritional management of OA (also called arthrosis in some countries)
starts with the prevention of joint disease in the (fast) growing puppy.
Reducing the risk for HD and OCD and ED to develop at young age, reduces
the risk for OA to develop at later age. A dog that did not develop HD or ED
during growth will not develop these conditions afterwards, as these are
developmental diseases by definition. However, when HD and ED did devel-
641
op before the age of maturity, the dog will invariable suffer from OA later in
life. Although the degree of clinical problems cannot be predicted, there is
no doubt that some degree of discomfort and pain is present in all OA affect-
ed dogs.
OA is an irreversible, painful and often debilitating condition, and full
cure of affected joints is not an option. The best available option is to slow
the progression once it has started and to try to minimize the discomfort.
The veterinarians’ role is to educate the owner on the nature of the condi-
tion and to advice on the management. Nutrition can play an important role
in the management of OA. First and foremost by avoiding overnutrition and
by managing a healthy bodyweight. Secondly by providing nutrition that is
optimally formulated to meet specific demands of pets with OA. Ingredients
that have been reported to have beneficial effects on OA affected joints, are
1. the so called Disease Modifying Osteoarthritis Agents (DMOAs) also called
neutraceuticals, or “chondroprotectives”, containing chondroitin sulfate
and glucosamin, and 2. omega-3 (n-3) fatty acids, and especially eicosa-
pentanoicacid (EPA).
DMOAs have become common place in the treatment of OA despite
the lack of definitive scientific studies confirming their efficacy. The mixture
of chondroitin sulfate and glucosamine supposedly enhance cartilage health
by providing the necessary precursors to maintain and repair. They are
reported to have a positive effect on cartilage matrix, enhance proteoglycan
production, and inhibit catabolic enzyme production or activity in OA joints.
These agents are marketed as oral nutritional supplements and not “drugs”,
and for that reason the standard drug efficacy data for the products do not
have to be provided by the manufacturer or distributor. Current research
suggests that glucosamine and chondroitin sulphate may be prophylacti-
cally beneficial in patients that are prone to develop OA as a result of HD,
ED, OCD or cranial cruciate ligament rupture. Positive results are available
from in-vitro and in-vivo studies in several species including men, horse and
dog. They argue for a “benefit of the doubt” type approach in prescription of
DMOAs despite the lack of information on dosages and indication. However,
many studies lack the double blind placebo controlled standards, and they
are often subjective or do not exclude bias. Further controlled clinical stud-
ies are welcome to fully support the ambitious claims. Despite positive anec-
dotal reports, there seems to be less scientific support for the use of
hyaluronic acid (sodium hyaluronate, HA) for the prevention or treatment of
642
OA.
N-3 fatty acids are poly unsaturated acids (PUFAs) that serve as sub-
strate for eicosanoids production. Eicosanoids are local chemical mediators
such as leukotreines, thromboxanes, prostaglandins (PGs) and prostacy-
clins. They are typically produced from omega-6 (n-6) PUFA arachidonic
acid. Depending on the nutrient profile, dietary fatty acids can effectively
change eicosanoid production. Inreasing the ratio of n-3 fatty acids com-
pared to n-6 fatty acids, will result in the replacement of arachidonic acids
in the cell membrane with EPA. The eicosanoids derived from Arachidonic
Acid (PGs E2, leukotreine C4 and Thromboxane A2) are pro-imflammatory,
whereas eicosanoids synthesized from n-3 PUFAs such as EPA, DHA and
α–linoleate or ALA (PG E3 and Leukotrein B5) are less inflammatory.
Research in dogs showed that feeding diets relatively high in n-3 PUFAs
increases leukotrein B5 and decreases leukotrein B4 synthesis in the skin.
With regard to this effect on inflammatory mediators, the recommended
ratio n-6 to n-3 is suggested to be 5 to 1.
In addition to the above mentioned n-3 effect with regard to inflam-
mation, recent reports claim a specific effect of EPA with regard to OA.
Research has shown that EPA supplementation abrogates canine articular
cartilage degradation in in-vitro explant culture systems (CATERSON).
Cartilage cells pretreated with EPA show significant dose dependent
decrease in glycosaminoglycan (GAG) release under in-vitro test conditions,
indicating abrogation of cartilage degradation. The reason for this EPA effect
was not completely clear but involves interference with the aggrecanase-
mediated cartilage proteoglycan catabolism. It is suggested that EPA inter-
acts with mRNA for the aggrecanase, and thereby alters the expression of
the genome, and therefore can be characterized as “nutrogenomics” (see
Chapter XVII). The authors involved in these first studies suggest that
dietary supplementation of dog food with EPA may prove to be efficacious in
slowing down the rate of cartilage degradation in canine OA. Again, placebo
controlled double blinded clinicals trials in dogs with OA are needed to pro-
vide conclusive evidence.
643
21.2.3.3. Feeding of senior dog and cat

Process of aging is caused and controlled by a series of factors, like genetic
determination, stepwise, gradual decrease of vital functions, troubles of
some physiological systems (mostly the immune and the neuroendocrine)
and accumulation of harmful substances (e.g. lipofuscin). Theories can be
classified into stochastic (accumulation of transcription failures in protein
synthesis, cross-linkages in extracellular collagen, wearing out: telomeres
and telomerase and the long-term damages of free radicals) and determin-
istic (Hayflick-Moorhead limit, 1961), pacemaker theories in hippocampus
and hypothalamus, genetics, i.e. aging gene, see Cenorhabditis elegans) one.
All of them are related to the nutrition. The general basics of aging is given
by the telomere-hypothesis, according to the cells, having capable of multi-
plicating only a limited times, are programmed to a death (apoptosis). Aging
will clinically be manifested by through the impaired immune competence
and the life-long deleterious effects of free radicals. This is the reason, why
the application of immunostimulant and antioxidant compounds (e. g.
carotenoids, vitamin E and C, rutin, plant flavonoids etc.) may partly
improve the health state of older individuals.
Clinically the aging means the slow, but progressive reduction of vital
functions, for example the physical fitness, the respiratory capacity, cardial
performance, renal insuffiency, cognitive dysfunctions, bone, teeth and
joints problems, decreasing immunocompetence, other (collapses, chronic
cough, vomitus etc.) There are not uncommon the cases of endocardiosis
(atrio-ventricular valves), of chronic interstitial nephritis, diminished num-
ber of chondrocytes and the calcification of back (dorsal) vertebrae. Obesity
is a frequent consequence of getting older, especially in urban, indoor dogs
and cats. The predisposing factors for that are the endocrine changes,
decreasing the basal metabolism, the sexual inactivity (especially in neu-
tralized bitches), the lack of sufficient physical activity and many time also
the high energy intake.
WHEN CAN A DOG OR A CAT CONSIDERED AS OLD ? Although basically of
course the advancement of time determines old age, but the body size
influence it significantly, which in turn depends primarily upon the breed.
According to an internationally accepted classification dogs and cats may
be taken as old, in case of
644
PHYSIOLOGICAL BASICS OF OLD-AGED NUTRITION.

The maintenance energy requirements of the senior dog is approximately
lower than that of young, active individuals. This is not valid for cat, where
animals even generally increase feed intake to counterbalance the declin-
ing fat digestibility, having practically unchanged energy requirement.
High digestible proteins of good biological value should be offered to avoid
loading of liver and kidneys. To prevent constipation, a medium fibre level
(at least 3-5% should be assured; fibre supply is especially important in
case of an diabetes. The continuous, ad libitum access to drinking water is
essential, because the water metabolism of older animals is more labile,
than in the adult, middle-aged dogs and cats. Senior diets generally have
lower sodium and phosphorus level, considering the possible renal insuffi-
ciency. In older cats the incidence of struvite urolithiasis decreases and
that of oxalate increases; consequently their feed does not need acidifica-
tion. Because of the lower utilisation of vitamins, the relative vitamin needs
are higher than in younger individuals.
Commercial and prescription senior diets are formulated by taking
into account the general symptoms of aging. Thus, for dogs the energy den-
sity has been diminished by 20% and to spare kidneys, the protein level
has also been decreased. The latter is not necessary in the light of the
recent data, unless there is a serious chronic kidney disease. Nowadays it
is known that aging basically cause peculiar changes in two fiels, víz. in the
functioning of the induvidual organ systems and in the activity of, mainly
metabolic enzymes.
Efficacy of digestion can exactly be measured and expressed by
means of the digestion coefficients (see Chapter VII). Effectiveness of dog
digestion, unless there are no teeth injuries, practically does not change by
aging. On the contrary, in cats the lipid digestion drops by 10 to 15 per-
645
cent units. Both in older dogs and in cats gradually a glucose intolerance
develop, causeb by the fall of number and sensitivity of insulin receptors.
As a consequence, ingestion even a small amount of sugar cause long-last-
ing postprandrial glucose elevation. Insulin affinity of liver drops, too and
as a consequence, the gluconeogenesis diminishes and the fat synthesis
will be enhanced. There is a change also in the rate of fatty acid metabo-
lism and among others the desaturation decreases, leading to a trouble in
the metabolism of eicosa-derivates. The concentration of prosztaglandin E2
(PGE2) and leukotrien B4 increase, predisposing organism to the inflam-
mative processes and smooth muscle spasms. The latter can be counter-
balance by a high ω-3-fatty acid and vitamin E provision. Old age makes
animals prone to dysbiosis and the composition of intestinal flora will be
shifted and more biogenic amines (scatol, histamin, putrescin, kadaverin
etc.) are produced. In the practice it can be improved by feeding yogurt or
culture of lactobacilli. Supplementing the daily ration by prebiotics
(Fructoz OligoSacharides, FOS), for example dried beet pulp, boiled carrot
or speciel senior diets, containing synthetic FOS). In this way the unpleas-
ant smell of the faeces can be moderated or prevented.
The capacity of immune system decline with the age, too, primarily
the cell-mediated immune response (mitogen stimulation, chemotaxis,
phagocytosis, natural killer cell activity) is inhibited. Before all, the energy
intake should be limited and the amino acid supply should be optimized
(e.g. extra arginine, glutamin etc.). By offering immunostimulant and
immunomodulant substanses, like grapefruit, red beet, garlic extract, toco-
pherols, germinated cereals, carotenoids (steamed carrot, pumpkin), min-
erals (Mg, Cu, Zn, Se), synthetic antioxidants and vitamins (vitamin C-vita-
min, piridoxin), the capability of immun system can be supported.
Summarizing the principles of old-aged nutriton, the energy intake
should be reduced (dog) or maintained (cat); assure sufficient proteins of
high biological value; avoid carbohydrates of high glycaemic index; increase
the proportion of ω-3 fatty acids. Concerning the mineral and vitamins, use
fortified diet. During applying diets for the particular diseases (diabetes,
congestive heart failure, chronic kidney disease etc.) the quality of protein
cannat be overemphasized. Using home-made diet, the raw material of
choice are the egg, cheese, cotton-cheese, lean beef, muton, chicken and
rabbit meat. It is advisable to avoid sugars and long-release, complex cart-
bohydrates of medium-to-low glycaemic index (e.g. boiled rice, whole cere-
646
als, buckwheat, sorhum, milo etc.) should be used, to minimize postpran-

drial glucose and insuline peaks. To favour heart functioning, the sodium
intake is moderated (e.g. using farinaceous products); on the contrary, the
supplemental potassium chloride and magnesium citrate are advanta-
geous. L-carnitine intake improve muscle (especially heart muscle) capaci-
ty (dogs), in turn, the taurine in cats. The commonly occurring obesity may
be moderated by the fat burners (organic chromium, L-carnitine). Nowaday
prescription senior diets are available, considering the above described
data.
21.2.3.4. Raising of orphan puppies and kittens

While nursing the orphan puppies or kittens, the starting poing should be
the Bunge-Abderhalden rule, formulating that the redoubling time of the
birth weight is in close, positive correlation with the composition of dam’s
milk. It means that the composition of the milk replacer should be as simi-
lar as possible to that of the mother’s milk and the frequency and mode of
application should also imitate the natural way. Table XXI-15 helps in com-
parison of mature milk of different species.
Composition of milk of the shown species differs not only in the major
nutrients, but also in the minerals and vitamins, too. The fine physical
characteristics of the components (for example the casein or fat structure
and particle size) may also different. This is why only in case of necessity
can be used the home-prepared milk replacers (Table XXI-16), because the
risk of development of deficiency symptoms cannot be excluded. As a typi-
cal example is the copper deficiency of kittens, nursed basically by cow
milk, developping depigmentation, loss of hair, even in serious cases leg
deformaties.
647
Table XXI-15: Average composition of the milk of bitch, queen, cow and goat
Table XXI-16: Formularies for home-made dog and cat „milk-replacers”
* distributed at least to 6, but rather to 8 occasions
The density of cow milk can be increased by adding cream, meat

meal, cod-liver oil or maize-oil. Commercial products are or used to be also
available, for example Pedigree Canine Milk Substitute Instant Diet,
Whiskas Feline Milk Substitute Instant Diet, Litterlac, Welpi, Supermilk,
Canovel Milk Mix, Sherleys Lactol, Cimicat etc.
.
21.2.3.5. Working dogs’ nutrition and feeding
Strong physical activity primarily increases the energy requirement of dog,
for example military and police dogs, sled god, hunting dogs, greyhound etc.
The daily energy needs may be as high as 2 to 4 fold that of maintenance
requirements. The dry matter intake capacity is limited, therefore this huge
nutrient ingestion can be assured only by high density, high digestibility
648
and tasty feed mixture. Higher amounts of ballast is not desirable, but con-
sidering the dietetic effects of fibre, the absolute fibre-free diet should be
avoided. Table XXI-17 gives an example for a formula satisfaying the above
principles.
Table XXI-17: Recommended feed composition for dog of hard work
Possible ingredients are the fish, meat, viscera, fatsoils and the pre-
treated cerals. Only raw material of excellent (at least 90%) digestibility can
be applied. Extra energy is primarily given by the fat, the higher protein level
serves the gluconeogenesis. To support the haemoglobin synthesis, more
iron should be given and to prevent the fragility of red blood cells, more vita-
min E and selenium should be supplemented. If the forced work induced
some related ailment (e.g. diarrhoea-dehydration syndrome, rhabdomyoly-
sis, gastric dilatation and volvulus, colonic dilatation, haemorrhagic diathe-
sis in colon, stress anaemia and fractures of metacarpal or metatarsal
bones), the recovery should be supported besides the special medical or sur-
gical measures, by appropriate dietary treatment, too.
FEEDING REGIME. The main feeding is the „dinner”, víz. ¼ to ⅓ of the
daily ration should be given early morning, 10% after a short rest at noon
and all the remaining is offered in the evening. Watering is indispensable
many times during the day.
Carbohydrate (starch) loading of the muscles did not prove success-
ful in case of the dog, instead muscles should be adapted, using special
training, to the enhanced fatty acid oxidation (for details see Chapter XXIV).
L-carnitine supplementation support the fatty acid utilisation of muscles;
by feeding creatin-monohydrate (1-4 g/dog/day) the energy economy can be
649
improved.
In human trials the supplemental creatine improved muscle strenth
for short duration and it can also increase lean mass. At the same time,
supplemental arginine may help in endurance and can definitely increase
exercise tolerance in patients with cardiovascular disease, included conges-
tive heart failure. The possible explanation for these positive effect is that
arginine stimulates nitric oxide and GH production.
The described principles are valid for the cat, too, but in this case
extra taurin and arginine should also be added.
FOR FURTHER READING
Hazewinkel, H.A.W.H. (1985): Influences of different calcium intakes on calcium metabo-

lism and skeletal development in young Great Danes. Thesis Utrecht University.
Hedhammer, A. (1974): Overnutrition and skeletal disease. The Cornell Veterinarian. 64,
Suppl. 5.
Meyer, H, and Zentek, J. (2005): [Ernährung des Hundes – Dogs‘ Nutrition – in German].
Parey Buchverlag Berlin 5th Ed.
Nap, R.C. (1993): Nutritional influences on growth and skeletal development in the dog.
Thesis Utrecht University. ISBN 90-393-0256-1
Richardson, D.C., Zentek, J. and Hazewinkel, HAWH, et al. (2000) Developmental orthope-
dic disease of dogs. In: Small Animal Clinical Nutrition. (Eds): Hand, M.S., Thatcher,
C.D., Remillard, R.L., and Roudebush, P. (200): 4th Ed. Mark Morris Institute. pp
505-528.
Davies, M. (1996): Canine and feline geriatrics. Blackwell Science. Oxford-London-
Edinburgh-Cambridge-massachusetts-Carlton Victoria
Walser, K. and Bostedt, H. (Eds.) (1990): Neugeborenen- und Säuglingskunde der Tiere
[Newborn and suckling animal science – in German] Ferdinand Enke Verlag Stuttgart
650
21.3. Practical aspects of dog and cat nutrition

Dietetic feeding of dog and cat earlier encompassed two great topics, name-
ly to replace some deficient nutrient and to correct some imbalance or to
support the medicinal treatment by dietary treatment. Nowadays the dietary
nutrition is not limited to the feeding of sick animals, but occasionally com-
prises also the supply of senior dog and cat or indoor living urban pet and
the nursing of orphans.
21.3.1. Types of dog and cat feeds

Commercial diets can basically be classified into two groups: the complete
diets contain all of the nutrients and active ingredients, which are able to
cover the requirements of the dog or cat of the given age and physiological
state. Besides ad libitum drinking water access they can be fed exclusively,
alone, without any supplementation (e.g. Chappi, Pedigree Complete Menu
etc.). Representatives of the second group either contain the necessary pro-
tein, amino acids, minerals and vitamins in a concentrated form and they
require only energetic supplementation (e.g. in form of boiled rice, potato,
vegetable oils, e.g. the Pal cans), or they are full of pre-treated starch (e.g.
popcorn-like feed) and they need protein supplementation. There is a spe-
cial field of „treats”, when the goal of application is not to meet the nutri-
ent requirements, even, to provide as low energy as possible to the animal.
Commercial diets for healthy dogs and cats, according to their water
contents, are denominated as moist or canned (72-85% water content), dry
(6-7, maximum 12% water level) and semi moist (15-30% water concentra-
tion, chiefly in form of sausage). According to a more accurate classification,
feeds of approximately 17% of water content are the semi-dry (e.g. Frolic
etc.) and the semi-moist (having 25% water, like the Hap feeds). The appear-
ance of dry feeds may be pressed, pelleted or extruded.
The water content of DRY FEEDS is within the range to 12%, their form
may be meal, pelleted, extruded, cooked, cake/biscuit- or kibbles-like. The
main components of the dry feeds are the cereals and the by-products of the
milling industry (wheat, corn, barley, oats grains, wheat germ, bran and
middling), extracted soybean and soy protein isolate, slaughter house by-
products, cattle liver, fish, meat-and-bone meal, dairy products, corn
gluten, corn starch, fats and oils, and the so-called “chicken broth” from
poultry slaughter house, digest, as well as mineral and vitamin mixtures. As
mentioned, dry feeds may be complete (“complete food”) or only supplemen-
651
CHAPTER XXI: PRACTICAL DOG & CAT NUTRITION
tary feeds (“mixers”). The „mixer-type” diets are generally mixture of cooked
cereal grains, with added fats or oils. The level of mineral-vitamin additions
shows their real value. The advantage of dry feeds is that they can be stored
6 to 12 months, but their preference, especially for cats, is not the best and
sometimes they make animals prone to urolithiasis, by decreasing the
amount of excreted urine. Water content of the SEMI MOIST (“semi moisture”,
“soft-moist”) diets is approximately 20-25%; they consist mainly of slaugh-
ter house and dairy by products, as well as soybean derivates. To the
preservation and the setting of the water activity of the products organic
acids, glycerine, carboxyl-methylcellulose, sorbate, propionic acid, as well
as colorants are used. The film-closed package generally contains the
dosage of a feeding. The advantages of semi-moist diets are that they are
preferred by cats, too and if not yet opened; they can be stored in room tem-
perature. Normally they are complete and do not need supplementation.
MOIST OR CANNED FEEDS normally contain 70-80% water. They consist of
slaughter house by-products, fish, soybean derivates and supplements.
While feeding commercial dog and cat feeds, the ad libitum access to
drinking water is essential. Some authors (like the author) propose to soak
dry feeds with water, milk or soup; unless it may cause greed swallowing
and digestion problems in some gluttonous breeds and individuals digestive
problems, like gastric dilatation and volvulus. Anyway, one should be care-
ful, because the dry feeds, diluted by water or other liquids, can easily be
spoiled and in this way may cause diarrhoea. As a new trend, the commer-
cial and breeder diets are more and more specific and adapted to the live
weigh (diets for giant, large, medium and small animals), to the sex (special
diet for spayed or neutered pets), as well as for breeds of peculiar requests
(Persian). There is a very special group of the dog and cat feeds, the mix-
tures of dietetic purposes („veterinary diets”, “prescription diets”).
21.3.2. Dietetic and medicinal feeds

OR „feeds for special dietary goals “, veterinary dietetic feeds). This group of
commercial diets would be applied only after veterinary prescription and

feed them under the control of veterinarian (prescription diets). According to
the EU-comform definition of Hungarian Feed Codex ‘Their trade does not
require veterinary prescription, however, on the label it is advisable to draw
the attention of consumer, that before using, it is recommendable to ask for
the advice of veterinarian, because feeding of these products may irreversible
652
affect life processes of animals’.

Form of package may be canned or pouch (moist) and bag (dry). In the
following the main types will be presented, without mentioning the trade
marks, because from a basic type more trademarks are available. Most of
the diets have closed formula, i.e. only the nutritive value is guaranteed, the
natural ingredients may change from batch to batch. In some special cases
(e.g. antiallergenic diets), the formula is open and the contents of natural
ingredients are given, too (e.g. chicken and rice). While labelling, the ingre-
dients used in feeds must be listed on the package in the order of their pre-
dominance by weight. This means that the first ingredient listed is present
in the largest amount, the second in the next largest amount, etc. It is espe-
cially important in case of substances, giving no actual concentration.
Advantages of using dietetic feeds, compared with the home-made diets are
the continuous assurance of the desired composition and quality. Basic
characteristics and fields of application are given in Table XXI-18.
The presented basic types, complying with the goal of ditary treat-
ment, may appear combined in a commercial prescription diet. In the
European market the following important companies are present (in alpha-
betical order, without claiming to be comprehensive): Acana, Hill’s, Iams-
Eukanuba, Purina, Royal Canin, Waltham. (The two latter merged and the
veterinary prescription diets are put into circulation under the name of
Waltham-Royal Canine.) US-labelling generally does not apply SI-system;
the following conversion factors may help in calculation: 1 kcal=4.184 kJ,
1000 kJ= 1 MJ; 1 oz (ounce) = 2.53 gram; 1 lb (pound) =453.59 gram, or 1
kg = 2.2 lb. Producers generally are using a characteristic logo. To be able
to predict the actual intake from feeds of different water content, the knowl-
edge of voluntary dry matter intake may be helpful.
653
Table XXI-18: General categories of dietetic feeds
21.3.3. Prediction of voluntary feed intake

To be able to rationing the daily amount of feed, especially in case of weight
control or slimming diets, the expected, physiological daily dry matter
intake should be calculated. On an average, dogs ingest approximately 2%
dry matter of their live weight. In case of small or large animals, it is more
accurate to see the value of the given body weight (Table XXI-17). At the
same time, pregnant bitches may eat 3.8% dry matter of their live weight,
lactating do 7.0% of their LW.
654
Table XXI-19: Voluntary dry matter intake in dogs, %
Cats tend to eat according to the stomach fill, having their dry mat-
ter intake within the range of 16.5 to 24.2 g/kg of LW (PROLA et al. 2006).
Adult, non-pregnant cats ingest for maintenance on an average 0.335 MJ
ME/kg LW (approximately 20 to 22.5 g dry matter) (FEKETE et al. 2002).
Although pregnant queens and lactating queens increases their dry matter
intake, the extra needs should be covered by giving more concentrated diet.
21.3.4. Prediction of the nutritive value

1. Lack of actual digestion coefficients, the metabolizable energy (ME) can
be calculated as follows. The approximate (or so-called wendee) analysis of
feedstuff or feed mixture should be done. The accepted average digestion
coefficient of crude protein is 80% (or in case of high-collagen content: 75%),
of ether extract 90%, of N-free extract (N-f.e.) 85% and zero for crude fibre.
After calculating the digestible composition, the major chemical com-
pounds should be multiplied by the ME-factors (or modified Atwater fac-
tors), i.e. each gram digestible crude protein by 0.0184, each gram
digestible ether extract by 0.0394, each gram N-f.e. by 0.0174; summing
the items, the results is given in MJ ME/kg.
E. g. the energy content of poultry meat, if it contains 19% crude
protein, 6.5%; ether extract and 0.4% N-f.e., then
190×0.80×0.0184+65×0.90×0.0394+4.0.85×0.0174 =5.2 MJ ME/kg.
If the given batch may contain higher amount of hydroxyproline (ani-
mal fibres), skin (hide), hair or horny material, the protein digestibility
should be measured by means of an in vitro pepsin-hydrochloric system or
in animal trial.
2. Energy concentration of commercial feeds can be evaluated by
simplified methods, using data of the chemical analysis. The results are
given in MJ/kg, using one decimal number. From the point of view of war-
655
ranty, no more than 15% deviation from the declared value is acceptable.
The method of calculation is as follows. The abbreviations: CP= crude pro-
tein, EE= ether extract, N-f.e. = N-free extract.
Dog and cat feeds (except cat feed of higher than 14% water con-
tent): ME, MJ/kg=0.1464×CP%+0.3222×EE%+0.1464×N-f.e.
For example the Hill’s r/d a veterinary dry diet has a ME-content
(MJ/kg), if the crude protein concentration is 34.2%; the ether extract level
7.5% and the N-free extract content 28.4%,
then 0.146 ×34.2+0.3222×7.5+0.1464×28.4=11.6 MJ/kg. – e.g. the
Eukanuba Veterinary Diet (Restricted Calorie Diet): 94.2% DM, 29.8% CP,
10.1% EE, 1.6% CF and 52.1% NFE, GE=17.76, ME+14.89 MJ/kg DM
(experimentally measures in vivo; NFE is mainly from sorghum carbohy-
drate).
Cat feeds of higher than 14% water content:
ME, MJ/kg=0.1632x×CP%+0.3222×EE%+0.1255×N-f.e.-0.2092
For example the Hill’s r/d moist veterinary diet has a ME content
(MJ/kg), if the crude protein is 8.5%; the ether extract 2.1% and the N-
free extract content 7.5%;, then
0.1632×8.5+0.3222×2.1+0.1255×7.5–0.2092= =2.8 MJ/kg.
21.3.5. Feline body mass index.

To control the way of the weight-reducing process of obese cats, besides the
weighing, the use of the Feline Body Mass Index (FBMI), developed by the
Waltham Centre for Pet Nutrition, is extremely useful. The FBMI gives the
actual body fat concentration. Equation use two zoometric (felidometric)
measures: one, which is related to the fatness and an independent one. For
the validation of this method DEXA measurements were used.
FBMI (fat%)=(RC/0.7067-LIM)/0.9156-LIM
where: RC= ribcage circumference or heart girth (cm, at rib 9), LIM=
limb index measurements or length of the lower leg, tibia (cm, from mid-
patella to heel bone or tuber calcanei). For example a castrated adult
short-haired cat of 3.45 kg LW, and another of 5.1 kg of LW the RC data
are 36.5 cm and 44 cm, the LIM 13.9 cm and 14.8 cm, thus the calculat-
ed total body fat concentration are 27% and 37%.
21.3.6. Control of the protein supply

and main protein sources of high BV
656
In case of marginal protein feeding (glomerulonephritis, chronic kidney dis-

ease), especially if home-made diet is used, the protein status of animal is
advisable to control by blood analysis (Table XXI-20).
Table XXI-20: Physiological value of blood parameters of the protein metabolism (fasting),
after MERCK, 1998
* pups and kittens: 4.5-5.5
PROTEIN SOURCES OF HIGH BIOLOGICAL VALUE

In many cases (e.g. chronic kidney disease) there is a need for the reduc-
tion of protein intake. In these cases, the protein deficiency cannot be pre-
vented unless ideal, i.e. well-digestible, of excellent biological value and
amino acid profile raw materials are fed. Table XXI-21 provides help in
choosing the appropriate ingredients.
Table XXI-21: Protein sources covering the daily protein requirements of dog (cat’s data in
parantheses)
657
21.3.7. Prediction of the urinary pH

DOG. Having a larger stock or kennel, before starting feed a new batch, a
representative sample may be sent to laboratory analysis. Based on the data
of most important minerals, the expected pH-value of urine can be calcu-
lated: pH= 6.5+0.0019×CAR. The CAR (cation-anion ratio, mmol/l) =
(500×Ca+820×Mg+430×Na+260×K)–(650×P+280×Cl), where elements stand
in g/kg (ZENTEK et al. 1994). Sulphur content of diet is supposed to be of
average for more details see Chapter XXI). If the urinary pH is above 6.5, the
formation of struvite urinary calculi is facilitated. Below pH 6.0, the inci-
dence of oxalate stones is higher. For example, a dry cat feed, formulated
according the previous AAFCO (1984) recommendation, it would contain
0.6% Ca; 0.04% Mg; 0.2% Na, 0.6% K; 0.5% P and 0.3% Cl; using the for-
mula, 6.5+0.0019×[(500×6+820×0.4+430×2+260×6)–(650×5+280×3)] = 9.6,
the expected urine pH is high, to avoid the struvite urolith formation, the
pH should be decreased to approximately pH 6 by means of methionine pills
or ammonium chloride but in case of senior cats.
CAT. YAMLA et al. (2006), based on a large set of data, developed equa-
tions to predict the urinary pH of adult cats. To achieve the desirable accu-
racy, separate formula should be used for wet and another for dry cat foods
(elements are given in % of dry matter).
pHurine=7.03+(4.7×Mg)+(0.93×K)+(1.43×Na)+(0.34×Cysteine)-
-(1.16×Cl)-(0.30×S)-(0.92×P)-(0.41×Met) (WET foods) and
pHurine=7.03+(0.89×Ca)+(1.58×Mg)+(1.00×K)+(1.00×Na)-
-(0.93×Cl)-(1.61×S)-(1.04×P) (DRY foods).
21.3.7. Urolith Diagnostic and treatment in the practice

(See also Chapter 21.2.2.16.) The exact type of stone can be determined by
the microscopical investigation of the urine sediment and the chemical
analyses of stones. For example, the struvite stones are coffin-like , the
oxalate crystalls small toothed, star-like ) urinary calculi.(FIGURE XXI-10)
Treatment-Prevention. Formation of struvite crystals can be prevented by
four measures: by decreasing magnesium ingestion, setting the maximum
magnesium level as 0.3% of the feed dry-matter. The general dosage for
mammals is 0.4% of dry matter. Urine pH should be continuously kept
slight acidic (pH»6.5) by supplementing the diet with ammonium chloride,
or methionine. By increasing the salt level (up to 1.02% sodium, i.e. 3.0%
658
NaCl) the drinking can be stimulate and in this way a large amount of dilut-
ed urine will be excreted. (As a side effect, in some individuals, the high salt
level may predispose to haemolysis.) Finally, by feeding well-digestible
feeds, like caned feeds, lean beef meat and viscera, the faecal water losses
can be minimized. Chances of oxalate formation can be decreased by mini-
mizing calcium intake and/or alkalizing the using sodium bicarbonate or
potassium citrate supplementation. If mannan-derivates are added to the
acidifying feed mixture, the increase water intake will keep the so-called
„relative super saturation” (RSS) of urine below the level of the formation
both of struvite and of oxalate stones. Prescription (veterinary) diets are
available for all purposes.
The cystine stone have a shape of benzene-ring, giving a positive uro-
cystin test (BENDE et al. 2001). The continuous dietary alkalizing of urine
using potassium citrate is capable of keeping the urinary pH above 7.5, pre-
venting the precipitation of cystine crystals. High blood and subsequent uri-
nary uric acid concentration favour of the formation of urate stones (ammo-
nium urate). Urate urolithiasis gives only approximately 5% of all the canine
urolith cases, but because of genetic background, half of them can be
observed in Dalmatians.
21.3.8. Home made diets

21.3.8.1. General considerations.
Home-made diets may be useful if the appropriate veterinary diet is not
available or in case of economic necessity. Practically it is impossible to
assure the continuously standard chemical composition and nutritive
value, therefore their use can be recommended only for emergency cases.
The main energy sources of the home-made diets are the boiled rice, pota-
toes and generally the farinaceous products (pasta, noodles, vermicelli,
spaghetti and dumpling), the latter especially, if the sodium intake should
be limited (e.g. congestive heart failure). Being the fact, that both dog and
cat are carnivores, feedstuff of animal origin is obligatory component of the
home made diet, too. As the available supplements (preservatives, flavours
etc.) concern, the general rule is that is should be from the group, that is
authorised for human.
21.3.8.2. Characteristics of potential raw materials for home made diet
MEAT AND SLAUGHTER HOUSE BY-PRODUCTS
Meat consists of muscle tissue, fat, connective tissue and blood vessels.
659
Nutritive value of meat from the different regions of the body depends upon
the ratio of muscle fibres, the intra- and intermuscular fat, because most
cuts of meat consists of muscle combined with some connective and fatty
tissue. Muscle tissue is made up of bundles of muscle fibres. Individual
muscle fibres are made into bundles by a thin network of connective tis-
sue. A sheet of connective tissue surrounds the bundle and forms the ten-
don that attaches a muscle to a bone. Connective tissue contains the pro-
tein collagen and elastin. Tough cuts of meat tend to have more collagen
and elastin than do the tender cuts. As animals are fattened, fat is deposit-
ed in connective tissues, first around certain organs such as the kidneys
and under the skin of animal and later the marbling within the muscles.
Fat-free muscle generally contains 25% protein and 75% water; protein
concentration in the fat-free dry matter is approximately 80%. Protein con-
tent of raw lean meat (having only intramuscular fat) is 20 to 22% and that
of fat 3 to 9% and the water content is 70 to 75%. Fat concentration shows
the largest variation. The so-called white meats (rabbit, poultry and veal),
as well as the fat-free beef have lower (3 to 5%) fat than pork or lamb (6 to
10%). Beef for human consumption generally has 24% fat, the meat of the
legs, of the plate and flank contain approximately 18% protein and 16 to
18% fat. Deep-frozen yearling mutton has 30 to 36% fat.
The pig lard and poultry fat are softer and the melting point lower,
than the beef tallow or mutton tallow, because they are higher in unsatu-
rated fatty acids. Carnivores are capable of utilising all of them, owing to
their excellent fat digestion. Slaughtering by-products (liver, kidneys, lung,
technological cuttings etc.) from different meat animals generally are good
protein sources, but their chemical composition shows great deviations.
Besides protein and fat, there is originally some glycogen in the meat,
but generally it is fermented into lactic acid. All meat and meat products
are high in phosphorus and very low in calcium (the ratio may be Ca to P
equals to 1 to 20), instead of the desirable 1.3 to 1; consequently, the cal-
cium supplementation is indispensable, unless the so-called “all-meat syn-
drome” develops (secondary alimentary hyperparathyroidism). Except liver
and kidneys, meat products are low in fat-soluble vitamin. Liver and kid-
neys may contain very high amounts of vitamin A, making animals, espe-
cially cats prone to vitamin A poisoning. Meats and particularly viscera are
good sources of vitamins B.
Cats generally prefer beef, veal and chicken meat, consumes horse
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meat, but don’t like sheep meat and pork. Otherwise, the raw pig meat may
contain viruses of Aujeszky disease, which is mortal for carnivores. Liver
has high nutritive value and preference for cats (used with preference dur-
ing the re-convalescence period), but there is a danger of addiction.
Ingestion of large amounts of raw liver causes diarrhoea, however, in
cooked form, it may result in constipation.
Chicken (generally poultry) meat is a dietetic feed for the carnivores,
especially for puppies and kittens. Feeding of bone-containing poultry meat
can cause injury of throat, oesophagus and even the anus, because of the
strong, tubular bones (e.g. femur, tibia) may crack into splinters. At the
same time, feeding of soft, boiled bone is preferred and to this practice can-
not be objected.
Based on slaughter house by-products, especially that of poultry,
meat-and-bone meal (MBM) is produced. The nutritive value of these meals
strongly depends on the proportion of employed raw materials (blood, head,
legs, bones, intestines, liver, kidney, confiscated/seized organs and cadav-
ers). The protein content varies within the range of 50 to 60%, that of the
fat within the range of 10-20%, the ash between 20 to 25% and the crude
fibre 0.5 to 1.0%. Since the BSE outbreaks, the regulatory pre-treatment of
raw material happens (under 3 bar pressure, at 136-138 oC, 18 minutes
or 121 oC, 150 minutes; maximal particle size 150 mm). This technology
causes an irreversible denaturation and of proteins, decreasing the biolog-
ical value of the product. Anyway, their use is prohibited only in the feeds
of food-producing animals, many dog and cat feeds may contain poultry
MBN. Development of rancidity is a frequent problem of these products.
MILK AND DAIRY PRODUCTS
The different dairy products (e.g. cream, sour cream, butter, cottage
cheese or curds, cheese, yogurt, sour and sweet whey, buttermilk) contain
the original components of milk (fat, proteins, lactose, minerals, vitamins)
in very different proportions. According to the observations, dogs show
higher preference to the dairy products than cats. However, milk contains
all of the nutrients, needed by dog and cat, except iron and vitamin D. Goat
milk is also appropriate feed for dog and cat, but having no vitamin B12
content, a long-term drinking may result in anaemia. At the same time, the
fine physical structure of protein and fat in goat milk is more readily
digestible for puppies and kittens.
In the whole cow milk on an average 3.3% protein (with 26-29 g
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casein/l in it), 4.6% lactose, 3.8% fat, 1.18 g/l (29.5 mmol/l) calcium, 0.93
g/l (30.1 mmol/l) total phosphorus, 0.12 g/l (5.1 mmol/l) magnesium, 0.58
g/l (24.2 mmol/l) sodium, 1.4 g/l (35 mmol/l) potassium and 1.04 g/l
(30.2 mmol/l) chloride can be found. Biological value of the milk protein
and fat is high and they are efficiently absorbed, in contrast to the lactose,
which has only limited utilisation. Milk is a rich vitamin A, B and vitamin
C source. While producing skimmed milk, the milk fat is removed, conse-
quently the fat-soluble vitamins are not present, either. The iron and cop-
per concentration in the cow milk, compared to the bitch’s and queen’s
milk are lower (iron: 0.2-0.6, 6-8 and 3-4 mg/l; zinc: 4, 7-8 and 5-7; cop-
per: 0.05-0.2, 1.3-2 and 0.8-1.2 mg/l, cow, bitch and queen, respectively.
Average cow milk comprises 2.98 g/l Lys, 1.02 g/l Met, 0.26 g/l Cys, 0.53
g/l Trp and 1.22 g/l Arg, pH= 6.7 (JENSEN, 1995).
In fermented dairy products (e. g. yogurt) there is no lactose any
more, on the other hand, there are lactobacilli, lactic acid (in some cases
fungi and ethanol too), protein, minerals and vitamins. Their feeding help
maintain eubiosis in the intestinal flora, which is extremely important in
older animals, being prone to dysbiosis. Cheese has the entire valuable
component of the whole milk, except lactose and vitamins B, which, in
turn, are found in the sweet whey.
In both companion carnivores, especially in cats, lactose intolerance
may develop as aging progresses. In these animals drinking milk causes
osmotic diarrhoea. Until this time, the proposed daily lactose limit is 1 to
2 g/kg LW, i.e. 20-40 ml/kg LW (dog and cat, respectively). Since this
would not cover the fluid requirements, ad libitum drinking water should
also be assured. Some dog and cat may get allergic to the milk protein
(mostly to the casein), resulting in plasmacytic colitis and scratchy-itchy
dermatitis.
FISHES
From the point of view of carnivore feeding, fishes can be classified
as “fatty” (grassy) and “lean” fishes. The latter are some marine fishes (e.g.
cod, Gadus morhua and haddock, Gadus aeglefinus), having less than 2%
total fat, and predominantly storing lipid in the liver. The muscle tissues of
such fish contain 0.5-1.5% lipid in the form of functional lipids (lipoprotein
or phospholipids of the cellular membranes). On the contrary, “fatty” fish-
es, like herring, sardine, carp, salmon, and eel contains 5 to 25% fat; the
starlet, the sturgeon and trout situated between the two groups. They store
662
the lipids in form of triacylglycerides under the skin and in the muscle tis-
sues: this may achieve 15% of the muscle. The total fat level varies not only
according to the species, but also from the season, breeding cycle and feed
availability. Fat-free fish meat has a very similar chemical composition
than mammalian meat; its biological value is excellent. Both fresh and salt-
water fish carry significant amounts of polyunsaturated fatty acids, espe-
cially eicosapentaenoic (ω3-20:5) and docosahexaenoic (ω3-20:6 acids, the
so-called omega-3-fatty acids. The vitamin A and D concentration is low;
on the contrary, selenium and iodine are high.
Filleted fish meat is low in calcium, but the whole fish body is good
calcium and phosphorus source for dog and cat. Cat is fond of fish smell
and taste, therefore fishmeal generally is applied in commercial diets as
taste enhancer. Although carnivores prefer raw fish to cook one, it is advis-
able to feed them after heat treatment, because raw fish may contain thi-
aminase (see Table XXIX-3) and parasites, too.
EGG AND EGG PRODUCTS
Egg, whole egg powder, egg-white and egg yolk powder are valuable
raw materials. The white is usually about 58% of the weight of the whole
egg, the yolk 31% and the shell 11%. About three-quarters (74%) of weight
of whole eggs is water, 12.9% protein (1/8), 11.5% fat (1/9) and 1.0% ash.
The yolk is a more concentrated than the white, because egg yolks contain
51% water, whereas egg whites contain over 87%. Just all of the nutrient
and active ingredient required by dog and cat are present in egg, only the
amount carbohydrates, vitamin C and niacin is low. The biological value of
egg protein is excellent. Vitamins, A, D and B-vitamins are also present.
The vitamin A value of egg yolk is independent of colour. Sodium and cho-
lesterol are mostly located in egg yolk, too. In raw egg white there is a pro-
tein (avidin), capable of binding biotin in the gut lumen, preventing it from
absorption. Consequently, it is advisable to feed egg rather in hard-boiled
form. The average poultry egg weight is 50-63 gram.
CEREALS AND MILLING BY-PRODUCTS
Grains of (corn, wheat) provide energy to the animals mainly by their
starch components. Since dog’s and cat’s pancreatic amylase production is
low, the digestibility of cereal grains should be improved by previous heat
treatment (micronisation, extrusion, cooking etc.) The glycaemic index (GI)
should be taken into consideration (FIGURE XXI-13).Concentration and
the biological value of grain proteins are low; consequently they should be
663
complemented. The protein content of corn gluten, a by-product of starch

(distiller) industry, is relatively high (60 to 70%), but its biological value is
low. It is widely used as a component of medium to low quality dry pet
foods. Although pre-treatment improves the availability of phytic phospho-
rus, grains are generally low in minerals. On the contrary, brans of differ-
ent cereals are high in minerals and vitamins of group B, as well as in
dietary fibre. The latter may serve as source of ballast or indigestible organ-
ic matter, for the prevention of constipation. Sorghum starch practicallyis
resistant to digestion of carnivores, thereby can be used also, instead of
fibre in weight-reducing diets to dilute energy density.
FIGURE XXI-13:Glycaemic index of some typical ingredients
FATS AND OILS

Carnivores require unsaturated fatty acids in their diet, but since dogs are
able to transform the linoleic acid into polyunsaturated fatty acids, they
can be supplied through vegetable oils. Cats, having no D7-desaturase
enzyme activity, require at least arachidonic acid (ω-6-C20:4), basically
found in animal fats. Both dog and cat need exogenous ω-3 fatty acids (175
mg DHA/kg LW in dogs, HALL et al. 2006). Beyond the carrier of essential
fatty acids, fats and oil increase energy density, improve palatability and
help in preventing forming of dust clouds. Nutritional application of abused
frying oil and fat should be avoided, because they may contain dioxins and
664
peroxides.
VEGETABLES AND OTHER PLANT PRODUCTS
Vegetables are not typical carnivore feeds, but their fibre, minerals
and vitamins may be valuable. In some pet food they give the colour of end-
product. The following species can be used. Leafy vegetables: lettuce
(Lactuca sativa), cabbage (Brassica oleracea var.), savoy (Brassica oleracea
var. sabauda), field kale (Brassica oleracea var. acephala), red cabbage
(Brassica oleracea var. rubra), spinach (Spinacta oleracea), garden sorrel
(Rumex scutatus) and cauliflower (Brassica oleracea botrytis). Radish and
tomato are not advised, although some dogs eat them after boiling.
Roots and tubers, like carrot (Daucus carota), potato (Solanum
tuberosum), Swedish turnip or Swede (Brassica compestris rutabaga) can
be used only after boiling, because in raw form they are indigestible and
may cause diarrhoea. Boiled (roast, fried, mashed) potato is an excellent
source of energy, but it should be taken into account that its glycaemic
index is high, close to that of the sugar. Soluble fibre fractions of the car-
rot and Swedish turnip have outstanding dietetic effects, as a source of
FOS (fructose-oligosaccharides).
Leguminous seeds like pea, beans are high in protein and vitamin B,
but their preference at cats is low. Raw soybean is extremely high in anti-
nutritive substances (see Chapter VIII); therefore it cannot be used for dog
and cat feeding. Heat and chemical treatment may eliminate part of these
substances; thus some pet food producers are not reluctant to incorporate
soybean in the extruded products. Nevertheless, allergenic compounds are
resistant enough to survive these treatments, included deglycosylation
(TOOKER and STAHLY, 2006). Therefore it is not a miracle, that some dogs
and cats develop allergy to the soybean-containing feeds. All of the legumi-
nous seeds carry more or less non-starch carbohydrates (NSC), which in
turn, cannot be completely degraded in the small intestine and will arrive
intact in the large intestine, causing carbohydrate overload of the hind gut,
dyspepsia, bacterial fermentation and flatulence. On the other hand, the
glycaemic index of leguminous seeds is favourable, i.e. medium to low.
21.3.8.3. Feeding of table scraps

Nowadays, with the greater availability of high-quality dry and canned dog
and cat feeds, the practice of feeding a companion animal almost entirely
on table scraps has been drastically reduced over these years. In addition,
665
the trend to smaller families and particularly to pre-prepared and packaged

low-waste foods has cut down on the volume of scraps available for the
family dog or cat.
When using by-products of food industry or table scraps occasional-
ly, without a calculated feeding plan, it should be taken into consideration
that meats are low in calcium, sodium (according to the blood content),
iodine, vitamin A and vitamin E. Feeding raw meat, raw fish or by-product
of fishing industry, there is a danger of ingesting pathogenic agents
(Aujeszky disease, salmonellosis, TGE=transmissible gastroenteritis of
swine, BSE) or parasites. Feeding of too much bone and boiled liver leads to
obstipation. Large amounts of raw liver and kidney may result in diarrhoea
and particularly at cat, in addiction. Feeding exclusively on liver may cause
vitamin A toxicity. Egg and egg products (egg powder, eggshell) are excellent
feed ingredients, but peculiar attention should be given to the prevention of
possible salmonella infection. High amounts of milk, especially in older cat,
will cause diarrhoea on the basis of lactose intolerance or may contribute to
the development of a casein allergy. Cereals grains and vegetables can be
given only in cooked form. Vegetable dishes from bean and peas will cause
flatulence. Too hot, spicy and salty table scraps cannot be given to the dog
and cat. Although some dogs like them, feeding of tomato, mushroom,
fruits, sweets and cakes are not appropriate for them, because of the dan-
ger of diarrhoea.
Otherwise, there is very little to say against scraps as a major con-
stituent of your pet’s diet, always provided you eat more or less sensible bal-
anced diet yourself. If he has been raised on them, an animal will happily
eat all the leftover meat, fat, potatoes, vegetables and anything else that
comes in way. Chicken and fish are fine for him, too, if all the strong bones
are carefully removed before giving them to him. Since a scrap diet tends to
be high in fats and carbohydrates and low in lean animal protein, it is not
advisable to add fat to this sort of diet, but it is good idea to give the dog/cat
a meal of its own lean meat or boiled eggs and at least once a week by way
of supplement, plus a small daily ration of kibbled biscuit or even canned
dog/cat meal just to help keep things more or less balanced for the animal.
Scraps can of course be added as a supplement in them to any regular diet,
meat, dry feed, canned feed or whatever and will contain many essential
nutrients which will do pet nothing but good. Limestone or cracked eggshell
supplementation should be taken into consideration. While preparing
666
home-made diet or feeding table-scraps, the attention should be directed to

the characteristics of the candidate components, taking into account the
mineral and vitamin sources and the possible deficiency or toxicity dangers
(Table XXI-22. and XXI-23). For dogs and cats of rough usage (e.g. lactation),
the feeding of appropriate complete feed is advisable.
Table XXI-22: Mineral sources, consequences of deficiency and overdosage
667
Table XXI-23: Vitamin sources, consequences of deficiency and overdosage
668
21.3.9. Feeding frequency, feeding practice

Adult dogs may be fed once a day, but twice a day is also acceptable. Cats
practically need ad libitum access to feed and drinking water. If dry com-
plete feed is used alone, not only the continuous water supply, but also the
possibility of urination is essential in case of disciplined dog (danger of uri-
nary calculi). Dry and canned feed can be combined, namely the moist may
be given in the evening and the dry in the morning.
Puppies, kittens, heavy working dog, lactating bitches and queen
required at least three feeding a day, growing animals (from weaning till 6
months of age), up to five occasions.
FOR FURTHER READING

Hall, J.A., Picton, R.A., Skinner, M.M., Jewell, D.E. and Wander, R.C. (2006): The (n-3)
fatty acid dose, independent of the (n-6) to (n-3) fatty acid ratio, affects the plasma
fatty acid profile of normal dogs. J. Nutr. 136, 2338-2344.
Jensen, R.G. (1995): Handbook of milk composition. Academic Press. San Diego, New
York, Boston, London, Sydney, Tokyo, Toronto.
Kahn, C.M. (ed.) (2005): The Merck Veterinary Manual, Ninth ed. Merck & Co., Inc.
Whitehouse Station, NJ. USA.
McNamara, J.P. (2006): Principles of companion animal nutrition. Pearson. Prentice Hall.
Upper Saddle River, NY
Prola, L., Dobenecker, B. and Kienzle, E. (2006): Interaction between dietary cellulose con-
tent and food intake in cats. J. Nutr. 136, 1988S-1990S.
Tooker, B.C. and Stahly, T.S. (2006): Utilization of deglycosylated soy protein in mono-
gastrics. J. Anim. Sci. 84(Suppl. 1), 342.
W.C.P.N. (2000): Waltham FOCUS: Advances in Clinical Nutrition. Special Edition.
William Caple & Company Ltd. Melton Mowbray
Yamka, R.M., Friesen, K.G. and Schakenraad, H. (2006): The prediction of urine pH
using dietary cations and anions in cats fed dry and wet foods. Intern. J. Appl. Vet.
Med. 4(1), 58-66.
Zentek, J., Meyer, H. and Hess, M. (1994): Abhaengigkeit des Harn-pH-Wertes beim
Hund von der Mineralisierung des Futters und dietaetische Konsequenzen
(Dependence of urine pH in dogs in relationship with the mineral supply and its
dietetic consequences – in German, with English abstract). Kleintierpraxis, Issue
12.
669
Chapter XXII
FEEDING AND NUTRITION OF POULTRY
22.1. Digestive characteristics of birds
22.2.1. Principles of domestic fowl nutrition
22.2.1.1. Laying hen’s maintenance
22.2.1.2. Influence of feeding on laying performance and egg
composition
22. 2.1.3. Rearing of replacement pullets
22.2.1.4. Feeding of commercial hens
22.2.1.5. Moulting of commercial flocks
22.2.1.6. Nutrition of breeding hen flocks
22.2.1.7. Practical feeding of hens
22. 2.1.8. Feeding of cockerels in rearing and reproduction
22.2.1.9. Feeding of meat-type chickens (broiler)
22.2.1.10. Feed supplements applied in feeding of broiler
chickens
22.2.1.11. Concentrate mixtures
2.2.2.2. Principles of turkey nutrition
22.2.2.1. Feeding of growing turkeys
22.2.2.2. Feeding turkeys in the breeding period
22.2.2.3. Feeding of fattened turkeys
2.2.2.3. Principles of goose nutrition
22.2.3.1. Morphology of the digestive tract
22.2.3.2. Microorganisms in the gastrointestinal tract
22.2.3.3. Digestion in geese
22.1.3.4. Intestinal fermentation of polysaccharides
22.2.3.5. The requirement of growing and breeding geese for nutri
ents
22.2.3.6. Feeding of the breeding geese
22.2.3.7. Requirement of geese for vitamin and minerals
22.2.3.8. Practical feeding of geese
22.2.3.9. Feeding of young broiler and fattening geese
22..3.10. Production of fatty livers (foie gras)
22.2.4. Principles of ducks nutrition
22.2.4.1. Feeding of ducklings at rearing
2.4.2.2. Breeding ducks nutrition
22.2.4.3. Feeding of fattened ducks
22.2.4.4. Practical systems of the ducks feeding
22.2.5. Feeding and nutrition of the ostrich
22.3. The most important digestive and metabolic diseases of poultry
671
22.1. Digestive characteristics of birds

he anatomy and function of the avian gastrointestinal tract profound-
T ly influence the utilisation of feed and the nutrient requirements of

birds. In case of birds all aspects of the feeding are very versatile. This
is true for the way of the feed is taken in (see the wide range of different
beak types), the existence, measure and importance of function of crop or
caecum and the quantitative range (ratio) of the requirement of some nutri-
ents (e.g. a one-day turkey need 28% crude protein and 0.0025% biotin in
the air-dry feed). The digestive tract in avian species is (relatively) shorter
than those of mammals, and the times for retention of feedstuffs is shorter,
leading to the less efficiency in recovering the nutrients from feedstuffs.
Nutrient requirements are substantially determined by the physio-
logical cycle (hutching, growing, moulting) and the maturity at the age of
1 day (precocial = nest leaving or altricial =nest resting). The energy needs
– especially in an open-air keeping system – are highly influenced by the
environmental temperature and humidity. The latter makes it difficult to
extrapolate data of wild ancestor or equivalent to the domesticated birds, or
to the ones in the zoo.
Animal welfare considerations express further concerns: while the
free-living birds spend 80% of daylight time searching and ingesting of feed,
the domesticated or zoo birds receive their feed scheduled and the duration
of the search and intake is significantly shorter. Thus, one has to prevent
monotony, boredom and enrich the environment.
There are substantial differences in the digestibility of fibres, too:
many species can utilize fibrous feedstuffs by means of microbial fermenta-
tion in the caecum (goose, ostrich etc.), others (granivorous, nectarivore)
hardly benefit from fibre fractions at all. In some group of birds (like falcons,
parrots, woodpecker, sparrows, colibri) the caecum becomes rudimentary,
or even ceases to exist and its functions – except fibre fermentation – are
overtaken by kidneys and rectum.
In today’s organic husbandry even the domesticated poultry has
opportunity to catch insects and therefore ingest chitin. Chemically the
chitin is a mucopolysacharide compound; its original function is to
strengthen and to stabilize the structure of the insect’s body. Many birds
672
TECHAPTER XXII: POULTRY NUTRITION AND DIETETICS
possess chitinase produced by the proventriculus. This enzyme split chitin

into chitobiose, which is an N-acetyl-D-glycosamine dimer. The beta 1,4-
bond of the latter will be split by the chitobiase into N-acetyl-D-glycosamine.
Although, the energy released from chitin is less than that from starch, it
cannot be considered insignificant because its digestibility changes from 20
to 80%. The main goal of chitin decomposition is to expose the soft tissues
for the other digestive enzymes to act on. The indigestible part of the chitin
– as indigestible organic matter - enhances the gut movements (peristalsis)
and the passage of digesta. In the domestic fowl (Gallus gallus)) the chiti-
nase activity is low, whilst the African Grey Parrot lacks altogether. The
present review fundamentally concentrates on the production birds, but for
the sake of a comparative approach, other species are also mentioned.
Poultry (gallinaceous birds) and turkeys, guinea-fowl, pheasant and par-
tridges (galliformes) have practically identical digestive tract (FIGURE XXII-
1).
FIGURE XXII-1: Digestive tract and ovary of domestic fowl (picture of © Fekete S.Gy.)
Special features of the bird’s digestive physiology
The birds’ digestive system differs in many respects from that of the mam-
mals. Morphological differences include beak instead of lip, lack of teeth
and buccal muscles. Owing to these the way of feed intake will change and
birds swallow rough, poorly prepared pieces of feed. The mouth (or cavitas
oralis) consists of the beak (rostrum), tongue (lingua), salivary glands (glan-
673
CHAPTER XXII: POULTRY NUTRITION AND DIETETICS
dulae ois) and pharynx. Little or no reduction in particle size occurs in the
mouth area. The tongue is relatively rigid, containing only few taste organs
(buds). The salivary glands are scattered in groups around the lower jaw,
cheeks, tongue and pharynx. They secrete 10-30 ml per day of mucinous
saliva, with amylase and inactive lipase enzymes. The saliva and the mucus
of glands similar to the salivary glands in lining the oesophagus serve to
lubricate the feed bolus.
Either an enlargement of the oesophagus (goose, duck) or crop(s) (or
ingluvies) (fowls, granivorous) developed to facilitate the preservation and
moistening, (pre)soaking and softening of feed and the regulation of stom-
ach filling-emptying. The crop is primarily a storage organ and little or no
digestion occurs, except for some bacterial fermentation or minor amylase
activity. Geese and ducks (water fowl) does not have a distinct crop, but the
oesophagus is able to dilate along its length and provides an important
reservoir. In pigeons and female parrots part of the crop secretes a nutritive
substance for the young („crop milk“). Stomach is divided into two parts: the
glandular proventriculus and the muscled gizzard (ventriculus) stomach.
They are the site of the beginning of enzymatic digestion and grinding of
feed. The proventriculus does not serve as storage organ, but the thickened
mucosa contains many glands which secrete both hydrochloric acid and
pepsinogen, which are released when feed is present in the lumen of the
organ. The gizzard has a large and powerful circular and a rudimentary lon-
gitudinal muscle. Tubular glands secrete a material, which forms a thick,
horny layer lining the interior surface of the gizzard. This layer appears to
be of proteinaceous nature similar to keratin and protect the soft tissues
from the action of the hydrochloric acid and pepsin as well as mechanical
abrasion. The gizzard contracts rhythmically (three contractions per
minute) to crush the content and grind the edges of particles together, thus
reducing particle size. The latter is facilitated by the „grits“(small pieces of
stones, pebbles, sand and gravel, rough feed particles), which stand for the
teeth’s function.
When particle size has been sufficiently reduced, the chyme passes
through the gizzard-duodenal junction to the intestine. The intestine is a
multilayered tube containing a serosal layer, longitudinal and circular mus-
cle layer, submucosal layer and mucosal layer. The surface area of intes-
tinal lining (mucosa) is greatly expanded compared to that of a simple tube
due to extensive microscopic folding to form a carpet of somewhat flattened,
finger-like projections called villi. In the avian gut, villi exist throughout the
length of the small and large intestine, steadily decreasing in height along
the way. The luminal surface of each villus is, in turn, increased by many
microvilli to facilitate absorption. In the interior of the villi, beneath the
epithelial cells, is the lamina propria, which contains connective tissue, cap-
illaries, and smooth muscle and nerve fibres. No lacteal have been found in
the avian intestine, contrasting with the mammalian intestine. (Lacteal
674
=any of the intestinal lymphatics that transport chycle. Only two cell layer
separates the lumen of intestine from the blood, because the capillaries
bring the bloodstream to the base of epithelial cells. Between the villi are the
crypts of Lieberkuhn. The globlet cells of the villi surface produces
mycopolysaccharide. This mucous layer serves a protective function, but
may also assist in absorption, because it may contain calcium binding pro-
tein. The absorptive cells’ luminal surface is covered with the microvilli.
Disaccharidase enzymes are found on the luminal surface of the microvilli.
A tight junction attaches each cell to its neighbours so that chyme from the
intestinal lumen cannot pass between the cells.
Factors of intestinal integrity
MUCOUS is secreted by specialised epithelial cells of glands in the mouth,

oesophagus, by the individual globlet cells of the proventriculus and intes-
tine. The viscous mucous consist of glycoprotein and water. Its functions
include the protection of mucosal cells in the stomach and gut from autodi-
gestion and it is the first barrier to bacterial and fungal invasion. Some vir-
ulent strains have either mucinolytic enzyme (e.g. Candida albicans) or ure-
ase to break down this protective layer (Heliobacter pylori). The intestine
also secretes water and electrolytes to keep bacteria in suspension and
wash them downstream from the small duodenum and jejunum.
The other element of the defence system is the lamina propria, which
contains connective tissue that underlines the epithelial lining, the vascu-
lar and lymphatic channels and the immune system, called gut-associated
lymphoid tissue (GALT).
GALT Damages of the blood vessels by pathogenic agents
(e.g. Newcastle disease, avian influenza, invasive candidiasis. intestinal coc-
cidia and salmonella spp.) may cause ischemic injury of the mucosa (infarc-
tion) or leakage of blood from the vascular channel (haemorrhage). GALT
represents the largest secondary immunological organ of the birds’ body. It
is composed of B and T lymphocytes, plasma cells, the mobile macrophages
and the resident dendritic cells. GALT secretes IgA (secretary antibody) on
the mucosal surface in response to foreign antigens. In production environ-
ment the lamina propria is quite active due to the responsiveness of GALT
and is actually in a normal state of reactivity and mild inflammation.
Diseases like infectious bursal disease, chicken infectious anaemia,
Marek’s disease, haemorrhagic turkey enteritis and clinical coccidiosis may
impact secondary lymphoid tissues. Corticosteroids, lymphocytolytic myco-
toxins (e.g. T–2 toxin, diacetoxyscirpenol) are able to rapidly deplete GALT
(GLÁVITS and SALYI, 1998).
Two bile ducts enter duodenum just below the gizzard exit. One of
these ducts passes directly from the liver to the intestine and the other
675
passes from the liver via the gall bladder (vesica fellea) to the intestine. Bile
contains bile salts and other compounds necessary to emulsify the fat in the
diet. In columbines there is no gall bladder. Entering the duodenum adja-
cent to the bile duct are two or three pancreatic ducts. The pancreas con-
sists of two main lobes and two smaller lobes. They are the sources of many
digestive enzymes for the hydrolysis of protein, carbohydrate and lipid con-
stituents of the diet in addition to specialised enzymes that hydrolyse
elastin, nucleic acids and other minor dietary constituents. The alpha islets
of the pancreas produce glucagon and the beta islets insulin.
Quantitatively, most of the digestion and absorption takes place in the small
intestine under the influence of the digestive enzymes secreted by the pan-
creas and intestinal wall and the bile secreted by the liver. The venous sys-
tem of the gastro-intestinal tract comprises two portal veins. The right side
system receives blood from the mesenteric veins. The pancreatic vein enters
the right lobe of the liver. The posterior mesenteric vein is linked to renal
veins, and blood from the small intestine may enter the kidneys (see: patho-
genesis of visceral gout of vitamin A deficient poultry).
Approximately in the middle of the length of the small intestine is the
yolk sac diverticulum or Meckel’s diverticulum (diverticulum vitellinum).
During the last stage of embryonic development, it serves as a connection
between the egg yolk and the intestine. After exhaustion of the yolk, the
diverticulum regresses (after the day 5 of the life) and in the growing and
adult bird serves no further function. The end of the ileum is the ileo-cae-
cal-colic junction, which plays a role in the control of flow rate and in the
filling and emptying of the caecae. Except for the ostrich and goose where
two large caecum developed and is the main site for bacterial fermentation,
whereas generally only a small amount of digesta reach this gut section
(exception: goose). In the pigeons the caeca are far less important. The cae-
cae are paired tubular structures lying caudally along the ileum from the
ileo-caecal-colic junction. In the adult domestic fowl the length of the cae-
cae is approximately 20 cm. Three distinct areas are seen in the caecae: the
neck, the mid caecal and the apical area. The filling of the caecae occurs at
regular intervals in case of ad libitum feeding. Evacuation of the caecae
result from the strong contractions of its wall. Frequency of emptying is 4
to 8 times daily and depends on the degree of distension, the quantity of
protons present and their electrolytes.
As mentioned above, the large intestine is very short, it extends from
the ileo-caecal-colic junction to the cloaca, a distance of 4 to 9 cm.
Functionally the colon acts very differently to that in other animals with an
almost constant retro-peristalsis( other than during defecation and caecal
evacuation) which appears to redirect nitrogenous products from the kid-
neys back to the caeca where micro-organisms utilise the N for their meta-
bolic processes which may contribute to the birds’ metabolism. Lymph nod-
ules, with their number increasing with age, are present. The colon empties
676
into the cloaca, which has a greatly expanded diameter compared to that of
the colon. Through the cloaca urine and faeces mixed before elimination,
and this is also the route taken by sexual products, like eggs (by the
oviduct) and sperm (by two sperm ducts). The colon and cloaca are involved
in the recovery of water and electrolytes from the intestinal content (FIGURE
XXII-2). At the lips of the vent the epithelium becomes stratified squamous
type. The regularly occurring defecation is a consequence of the rapid con-
traction of the coprodeum. Owing to the morphological characteristics,
urine from the ureters can ascend up to the caeca where the main part of
the water and electrolytes may be absorbed. The urine get concentrated as
insoluble urates and eliminated in form of a paste covering the excrement
in a white-grey layer. The proctodeum is linked through its base to the
Bursa Fabricii which is a nucleoprotein-rich B-cell dependent lymphoid
organ, degrading with age.
FIGURE XXII-2. : Picture of male duck cloaca: 1. Coprodeum; 2. Rectum; 3. Urodeum; 4.

Ureteral opening; 5. Proctodeum; 6. Vent
In comparison with mammals, the functional differences include the

length and volume of the digestive tract being small and also the retention
time (approx. 24 hours) short. Consequently, birds require highly digestible
feed. There is no lactase activity but the sacharase production is also low,
thus some types of feedstuffs (which contain lactose and sucrose) cannot be
applied. In the first weeks of life the utilisation of saturated fatty acids is
low; therefore vegetable oils are used for increasing the energy concentra-
tion. The caecum in most animals is the site of fermentation of fibre frac-
tions but in chicken the caecum is believed to be capable of receiving only
very fine fractions. Breakdown of NSPs (non-starch polysaccharide) in the
gut may remove a valuable source of fermentable substrate for the resident
microbial population which are utilising N from the retrograde provision of
uric acid and other nitrogenous wastes from the urodeum. Thus, with the
677
exception of geese, the microbial activity in caecum is small, consequently

the fibre degradation is low and the synthesis of vitamin K and members of
group B does not satisfy the needs of the host organism, therefore birds
always require external supply in the feed and sometimes drinking water.
Feed selection (preference) of birds is controled by the shape, colour,
size and consistency and the taste has less importance. The preference
order of materials of different consistency is: cereals grain and others seed
> pulpy > meals > flours, that of the meals: wheat > barley > rye > pea >
corn (maize), and in form of whole grain: wheat > corn > oats > barley > rye.
To prevent selective eating of poultry, the different ingredients of the feed
mixture should be ground to the same particle size.
Feed intake is basically determined by the energy concentration of
feedstuffs, although, especially in pelleted form, luxury consumption may
occur. Some antinutritive substances, like water-soluble β-glycans and pen-
tosanes make the gut content more viscous slowing the transit of the diges-
ta, which in turn, decreases voluntary feed intake. This situation can be
helped by the adding of industrially produced enzyme preparations to the
feed mixture. During the period of eggshell production laying birds prefer
high calcium feedstuffs. The water intake is an acquired ability of the one-
day-old birds, which should be considered when designing the drinkers and
the feeding-drinking technology.
In case of poultry the energy requirements of birds and the energy
value of feedstuffs are generally given in mega joule (MJ) expressed appar-
ent metabolized energy, corrected to zero nitrogen retention (AMEn).
Later in the text metabolisable energy refers to the above described form. (In
France for growing birds the ME. corrected on 40% N-retention is also
used.) The protein requirement of different poultry species and the protein
concentration of the feedingstuffs are given in crude protein.
The interrelationship of protein and energy both in feeds and in rec-
ommendation data can be expressed by the “protein-energy ratio (P/E)”,
which means the amount of crude protein per MJ ME. This form of expres-
sion can be used for amino acids too (e. g. the lysine requirement of one-day
chicken is 0.8 to 1.0 gram Lys/MJ ME).
It is worth mentioning that the academic vitamin recommendations
(e.g. NRC, 1994) are the minimum requirements and in the practical poul-
try feeding – considering the needs for optimum production, environment,
health status, possible stress, vaccinations etc. – even 2-10 times higher
vitamin concentration may be applied.
678
© Dorota Jamroz and Janusz Kubizna
22.2.1. Principles of domestic fowl nutrition
22.2.1.1. Laying hen’s maintenance

The laying hens are animals characterized by high productivity. The mass
of eggs produced can be even 14-16 times higher than hen’s body weight.
Because of such high physiological demand on the layer’ organism, the pre-
cise balancing of mixtures and providing the perfect environment conditions
for the birds are essential. In comparison with cage keeping systems, alter-
native ones assured the psychological comfort, the chance of movement,
contact with surrounding, the possibility of scratching in the litter, the rest
on perches, laying of the eggs in the nests etc. In “pro-ecological” feeding
systems, the additional application of whole wheat grain is being taken into
consideration. Because of size, chemical composition and structure of the
grain, cereals are natural feed for birds. The use of whole grain allows
reducing the costs connected with the grinding process to be reduced. The
cereals must come from own, environment at friendly (ecological) produc-
tion, i.e. from cultivations carried out without application of mineral fertil-
izers and chemical plant protective preparations. For the production of
those “ecological” feed mixtures growth promoters, conservants, antioxi-
dants, pure amino acids and other substances can not be applied. The lay-
ers having free access to the open yards covered (or not) with grass are able
to receive the nutrients from plants and soil.
The eggs obtained from hens reared in free range systems are char-
acterized by higher weight (by 2-6%) and thicker, stronger shell than eggs
obtained from layers kept in cages. In this case the feed intake related to 1
kg of egg mass is growing considerably from 1.8-2.2 kg in the intensive
keeping even up to 3.5 kg. However, the amount of energy used for mainte-
nance of constant body temperature and for movement increases when the
birds are reared in free-range system but the mechanical injuries of the legs
are often observed. Difficulties in maintaining proper hygiene of open yards
and elimination of microbiological feed supplements create some health,
mainly parasitological, problems. The maintenance costs of animals exceed
by about 30-40 % the expenditure in traditional keeping of hens on litter or
in cages, so the productive effects could be about a dozen or so percent
lower. The requirement of hens for nutrients depends on many factors, such
as: type of using (laying hens producing consumable, hatching eggs or meat
type hens), hybrids, breed, body weight (light and semi-heavy laying hens,
meat hens), age and laying phase, quantity of the daily feed intake, keeping
system (in battery cages, housing on the litter, in aviaries, in free range sys-
tems etc.) but also on microclimate inside confinement rooms (intensity of
ventilation, heating of the poultry houses, coops or lack of it etc.), region,
season of the year.
679
In determining feeding intensity, with special regard to protein and

mineral substances, the problem of minimization of emission of unused
metabolites (particularly substances derived from nitrogen, phosphorus and
microelements metabolism) into the environment should be taken into con-
sideration. The hens consume about 2.3-4.1 g of nitrogen daily and excrete
0.8-1.8 g of this element, which means that the N utilization, depending on
age, laying phase and productivity, vary within 30-53 %. Year-long emission
of nitrogen to the environment from one layer can reach 290-657 g of N
excreted, mainly in the form of easily and fast degraded uric acid (40-70 %
of excreted nitrogen) and as a ammonia-N (4-20 % of total excreted nitro-
gen). Daily phosphorus intake vary within 530-840 mg, and excretion
reaches 213-428 mg/d/animal which amounts to 78-156 g of the phos-
phorus from the layer per year. These two values indicate the necessity of
accurate supervision of feeding systems.
22.2.1.2. Influence of feeding on laying performance and egg composition

The reproductive organ of hen and other birds as well, is made up of the
ovary and only one well developed oviduct (left). The egg is created in differ-
ent parts of the reproductive tract. The egg cell – the oocyte and the yolk
layer - is formed during many months in the ovary, the albumen (egg white)
is created in the main part of the oviduct (magnum) in over 3 hours, and the
sub-shell membranes are created in the isthmus within 1.5 hour. The cal-
cification of the shell is occurs in the uterine part of the ovary and lasts for
about 20-22 hours. The weight of egg of young hens usually does not exceed
45 g, at the age of 40 weeks it varies from 55-65 g and in older hens, after
60 weeks it reaches even 70 g (Table XXII-1).
Table XXII-1: Composition of eggs of commercial birds
680
Differences in egg size depend on the hybrid, hens producing eggs

with white or brown coloured shell, body weight and age. The average com-
position of the egg is presented in Table XXII-2.
Hens usually begin the production of eggs at the age of 19-22 weeks,
depending on the type and body weight. Peak of laying is reached after 6-8,
even 10 weeks from first egg laid. The flock is sorted out depending on lay-
ing rate within a period of 64-70 weeks, after 15-16 months, e.g. after 10-
12 months of laying. In meat type hens the laying period is considerably
shorter and lasts 6-7 months, depending on leading the flock. The number
of eggs obtained from meat-type hen amounted to 150-160 and 285-310
eggs from light hens. Feed intake in the laying period amounted in light
hens to about 35 kg, in semi-heavy - about 45 kg, and mortality and the
selection vary from 3.0 % up to 6.0%. The greater losses are noted in the
restless, nervousness light hens or in heavy meat-type ones. The egg pro-
duction process is regulated by the lighting program ensuring 14-16 hrs of
light daily in a continuous setup. The egg is formed during the night, but
usually is being laid between 6-8 a.m.
The curve determining the flock’ laying rate, expressed in % for 100
layers, is reflecting the physiological state of the bird, harmonious and even
production of eggs, as well as feeding, ventilation failures, room tempera-
ture variations and diseases of birds. The birds are reaching the laying peak
on the level of 98%. Towards the end of productive period the laying declines
to 60-65 %, even to 50 %, and the moment of liquidation of the flock
depends on the relation of expenses of the feed and the price of eggs. The
basis of the correct feeding of hens is applying of homogeneous feed mix-
tures, without sudden dietary changes. Composition of the egg – sex cell
(gamete) is quite constant (Table XXII-3), although quantities of some com-
ponents can be modified genetically or by feeding regime.
681
Table XXII-2: Chosen parameters of chemical composition of hen’ egg
682
Table XXII-3: The possibilities of changing of the egg composition: component of low or of
no fat
The labile components of the egg include vitamin A, E, B2, pan-

tothenic acid, folic acid, biotin given in large quantities in the feed, treated
as a “hatching” vitamins, can reduce the embryos’ mortality and improve of
hatchability of the eggs. The quantities of egg protein and amino acids, con-
tents of gross energy and the fat in egg are quite stable but the composition
of fatty acids can vary depending on the kinds of fats already present in the
diet. There are also some possibilities of manipulating the quantity of total
cholesterol in the egg (200-230 mg/egg) by applying polyunsaturated fatty
acids (Table XXII-4), by increasing the level of crude fibre in the feed that
can reduce the cholesterol absorption, by applying of different plant prepa-
rations and algae. The most effective method to decrease the cholesterol in
the egg is to increase the laying rate and thereby spreading the amount of
cholesterol produced daily by hens in higher number of eggs. Through
applying of feeds containing greater amounts of linolenic acid (n-3), CLA,
iodine and zinc, oil from fish derived from cold seas water, linseeds or the
683
linseed oil, rapeseed or rapeseed oil and other nutraceuticals is possible to

obtain enriched eggs, showing higher consumable value.
The quality of the shell of both hatching and consumable eggs is an
important technological feature (hygiene, transport). The strength of shell is
influenced mainly by its density and also by its thickness and mass. The
share of shell in the total egg weight depends on genetic value of birds and
on the feeding. In hen eggs, the shell makes up about 6-7 g, i.e. about 9-11
even 12 % of egg weight. The resistance against crushing applying 42-43
Newton indicates good qualities of the shell. Average thickness of it amount-
ed on average 0.33 mm. In the shell mineralization process the different fac-
tors, such as structural proteins, calcium, phosphorus, vitamin D3, vitamin
C, chlorine, and magnesium and electrolytes balance take part. In case of
insufficient quantity of calcium in feed the hen is able to use the limited
reserves of phosphates from long bones. The released but not built-in phos-
phorus is excreted from the organism. The calcium reserves are small and
are estimated to last for about 24-36 hrs, which causes that in the Ca defi-
ciency with the strength of shells lowered.
Gradual, slow release of Ca from macrogranular (coarse-grained) min-
eral supplements, calcareous grit, shells etc. assure a constant supply of
calcium necessary for good shell mineralization takes place mainly at night.
22. 2.1.3. Rearing of replacement pullets

The period of the growth of young breeding birds or hens that are intended
for commercial flocks may determine future laying performance. The growth
rate and the flock alignment should be consistent with the growth pattern
of the given hybrid. The acceleration of attaining of sexual maturity, accom-
panied by weak somatic development, should not be pursued. The birds
which prematurely start to lay, produce small eggs at the expenses of their
own body, thus the laying period can be shorter. The young hens with
excess body weight, slightly overfated in a period of the beginning of the lay-
ing, quickly reach the highs level of productivity, however in the peak of lay-
ing the advanced stages of the fatty liver degeneration was found in them.
Mortality of the best layers can be increased. Possible ways of rearing is
shown in FIGURE XXII-3.
684
FIGURE XXII-3: Optimal (C) and practical (A+B) growth intensity at replacement pullets
In the initial period of rearing, during first 6-8 weeks of life the pul-
lets are fed ad libitum with feed mixtures containing about 18% of protein
and energy value of 11.7-12.1 MJ ME/kg. The recommended level of lysine
amounted to 0.85-0.75%, methionie 0.33-0.35%. In order to obtain savings
in the field of the feeding, the level of crude protein can be lowered even to
15-16 % on condition of good balancing of exogenic amino acids and with
addition of pure ones. In this period the chickens fed smaller amounts of
feed. Daily intake varies within 10 and 40 g/d/head in light hens and
between 48-55 g in heavier birds.
During the second growth period, to 18-19 weeks (22 weeks in the
meat-type hens) the birds are growing slower. The feed mixtures intake
increases up to 100-120 g/d/bird. The control of the quantity of feed intake,
especially in heavier and meat-type birds, is necessary. The pullets of light
hybrids can be fed ad libitum, however to the hybrids of semi-heavy and
meat-type birds should be given the mixture in limited amounts. Controlled
feeding intensity ensure the equal growth rate of the whole flock and allows
to correct the pullets body weight gain according to assumptions of rearing
of concrete genetic material. The level of energy in feed mixtures is reduced
to 11.1-11.5 MJ ME/kg and crude protein to 14-15%. During this period it
is possible to make considerable savings in the choice of raw materials (feed
components) to the production of mixtures being guided by the lowest costs
of 1 MJ or/and 1% of feed protein, however the exogenic amino acids should
be precisely balanced.
During the whole rearing period it is possible to apply one kind of feed
mixture (about 14-15% of protein), phase feeding or the free choice system,
in which the pullets can freely consume protein concentrates or cereals-
685
mineral and vitamin mixtures. The continuously controlled body weight is

the proper way of checking of rearing of pullets. Totally in the rearing of light
pullets in the period up to about 21 weeks the feed intake amounted to 6.7-
8.3 kg and the body weight vary within 1350-1750 g. Losses (mortality)
should not exceed 4%. Content of the crude fibre in mixtures given to young
birds allows controlling its energy value, maximum fibre quantity should
not exceed 4.5%. The level of calcium should vary from 1.0 to 0.8%, while
the level of available phosphorus from 0.40-0.45 to 0.32-0.35%. Detailed
recommendations for feeding of pullets in the rearing period usually are pre-
sented in information characterized the genetic material.
In the period of growth the coccidiostatics are applied in mixtures in
order to protect the birds against coccidiosis. About four weeks before the
anticipated start of laying, that means at 17-19-21 weeks, it is necessary to
withdraw these supplements from the feeds, so that the birds are able to
develop their own resistance. The use of coccidiostatics prior to the start of
laying and later removing them from feeds can constitute a threat of disease
outbreak in a laying peak and threat of transferring of medicaments into
eggs. The level of the calcium in feed should be increased to about 3.0-3.5%
at the beginning of egg production. Too early use of mixtures containing
high amount of the calcium can lead to the loss of appetite in hens and to
kidney damage.
From 15-16 weeks the quantities of vitamins and the macro –
microelements in feed can be lowered. In the phase of transitional feeding,
i.e. between 19-21 weeks, the gradual exchange of feeds used in rearing
period to mixture intended for layers can be introduced.
In feeding of meat-type pullets the amount of feed mixtures should be
strictly controlled. The restricted feeding systems with reduced quantity of
daily diet by 10-20% can also be used. The control weighing of chickens,
usually about 5% of flock, is necessary. The economical feeding of pullets
by no means can lead to nutrient deficiency affecting the development of
reproductive organs, the correct ovulation and the good biological value of
eggs laid by young hens.
22.2.1.4. Feeding of commercial hens

Genetic progress, over the last ten years has caused substantial changes in
the genetic potential of commercial hens. The average body weight of hens
laying brown eggs diminished from 2.05 to 1.95 kg, white ones from 1.90 to
1.70 kg. The laying period lasts 10 or 11-12 months and the average weight
of egg mass up to 19.6 kg is obtained from one layer with the total feed
intake of 1.8-2.1 kg/kg of egg mass. Daily consumption of mixture, depend-
ing on hybrids, has been decreased to 95-110-113 g/d/head. These data
are pointing to the narrow range of reserves, which are usable to the birds
characterized by such high productivity and low body weight. Particularly
the first eight weeks of laying requires intensive feeding, so that young lay-
686
ers are able to reach full somatic maturity. The feed mixtures must contain
about 18% of crude protein and energy density of 11.6-11.7 MJ ME/kg,
3.5% Ca and 1.4-1.6% of linoleic and linolenic acids. All of these positively
affected the size of eggs.
About the magnitude of feed intake in a period of high laying to a
great extent the energy density is deciding. The energy requirement deter-
mines the quantity of feed mixture consumed and in the same way the
intake of all nutrients. The adjustment of protein contents as well as vita-
mins, mineral components to the energy value of the mixture will ensure the
correct feeding of hens. It could be stated that maintaining the balance of
nutrients to the energy value of diet is one of the most important subjects
of commercial layer hen nutrition. The quantity of mixture consumed is lim-
ited by capacity of the digestive tract resulting from the hens’ body weight.
Light, lively hens intake more energy in relation to their metabolic body
mass than calm, heavy birds. The layers kept on the litter or in free range
systems have a bigger chance of movement, scratching in the litter and in
the soil. The losses of energy are higher thus the demand of these birds for
energy is estimated to be about 10-15% higher than hens kept in cages and
almost deprived of space for movement. For light birds it is necessary to
foresee 1.45-1.55 MJ ME and 17.1 MJ ME/d/bird for semi-heavy ones. The
suitable quantity of the protein is foreseen usually on the level of 18-19.5
g/d/head. In the practical circumstances the light/ dark hours, the daily
exepted egg production and feed intake are given in strict technological pre-
scription (FIGURE XXII-4).
687
FIGURE XXII-4: A typical technology for laying hens
The level of energy in the mixtures for hens must be adjusted to the
conditions of birds, maintenance, especially to the temperature inside poul-
try houses. Requirement for energy is lowered drastically during a period of
heat. The feed intake by hens is decreased even to 60-80 g daily, which
means that birds consume not only less energy but also less protein and
other nutrients which are necessary for correct functioning of the organism
and eggs production. Decrease of energy concentration in the relation to the
stable quantity of nutrients is useful at high temperature inside poultry
houses and in the outside environment. It protects hens against nutrient
deficiency. In turn, within a period of great variations of the temperature
and in the situation of lack of heat in the laying houses, the increase of the
energy value of mixtures precisely adapted to the concentration of nutrients
and to the greatness of planned or predicted daily feed intake per bird is
necessary. So, the correct relation of the quantity of nutrients to energy
became the criterion of modern standardization of diets for poultry. It is
possible to calculate the requirement of hens for energy from the following
formulas:
688
For light hens MEn = 490×LW0.75+16**+ 9.6× O

For semi-heavy hens MEn = 460×LW0.75+20**+ 9.6×O
where:
AMEn – requirement for metabolisable energy when N bal-
ance is zero
LW 0.75 – metabolic body mass
** - deposition of energy in body weight gain (0.78)
O – daily weight of produced eggs (in g)
or
AMEn, kJ/d = [480 + (15–T) × 7] × LW0.75 + 23×ΔLW + 9.6×O
where:
AMEn= apparent ME corrected to N balance by “GfE” (DLG,
1999)
T – environmental temperature (correction to ET < 15°C)
LW - body weight (kg)
ΔLW - daily body weight gain (g)
O – daily egg production (g)
The energy value of feed mixtures depends on the contents of cereal

grain, which constitute the basic raw material in the mixture, on the quan-
tity and the kind of the fraction of carbohydrates (starch) as well as on con-
tents of the dietary fibre in the feeds, which mainly is composed of com-
pounds such as arabinoxylans, β-glucan, cellulose, pectins and oher non-
starch polysaccharides. In the dietary recommendations the higher, levels
of the crude fibre, determined traditionally in mixtures, are also presented.
The quantity of structural substances to a great extent determines the
degree of nutrient digestion, but also gives the proper feed structure that
has profitable effect on the motoric activity and functions of the digestive
tract. The level of crude fibre in the mixture offered to hens creates the pos-
sibility for manipulation with the quantity of feed that is eaten by layers
and, at the same time, is one of interventional factors in case of cannibal-
ism.
The contents of fat in feeds for layers usually varied between 3.0-
3.7%, and application of additional fat to the mixtures for layers is not com-
mon, because reaching the 11.1-11.7 MJ ME/kg energy value of mixtures
is possible without this supplement. Through application of fat, mainly of
the plant origin or from fish oil, it is possible to enrich the eggs with the
unsaturated fatty acids of n-3 and n-6 family. The best fatty supplements
for hens are: whale oil, oil from the livers of cods, the oil from the herring
menhaden. Supplement of the fish oil at amount of 1.5% increases the
quantity of the linoleic (18:3, n-6) and docosahexaenoic acid (DHA) (22:6, n-
3). The similar effect was obtained in the DHA range when 15% of whole lin-
seed or ground linseed was introduced into mixture, although the quantity
of the linoleic acid (n-6) was increasing at the same time. The valuable
689
sources of n-3 fatty acids are: oil from saltwater fish, linseed, rapeseed and
oils from these seeds.
The good sources of the linoleic acid (18:2, n-6) are the sunflower oil,
soya oil and oil from the evening primrose. Rape contains both linoleic and
linolenoic acid and these are in optimal proportions for the humans, i.e. in
the ratio of 2:1. Human organism is not able to synthesize the sufficient
amounts of polyunsaturated fatty acids, which are important for function-
ing of the cellular membranes, mitochondria, for maintenance of the enzy-
matic and receptor activity, for synthesis of prostaglandins, transformation
of cholesterol etc. The application of polyunsaturated fatty acids (PUFA) is
not generally recognized. Increased amounts of feeds that are good sources
of these acids can decrease productivity and the weight of eggs. In addition,
the stability of PUFA in animal products is not exactly known, especially
after heat treatment. The cholesterol level in egg is relatively constant and
reaches about 200-210 mg in normal-sized egg. The daily requirement of an
adult human reaches 250, maximum 300 mg. Main source of it is own syn-
thesis in the organism (60-80%). The threat for human health that results
from consumption of eggs as source of cholesterol is still debatable. The
great doses of vitamin E (50-400 mg/kg of feed mixture for hen), which is
perfect antioxidant, increase the stability of products enriched with PUFA.
The recommendations of the optimal concentration of crude protein
in mixtures for laying hens advised, depending on the size and the phase of
production, the amount from 15.0 to 17.5 even 18.0% in counting on the
air-dry feed mixture, although many results confirm the possibility of low-
ering the quantity of the protein in mixtures to 14 or 13%. In a deep reduc-
tion of protein level, the supplementation of mixture with pure amino acids
to the level of requirement for methionine (405-450 mg/d/bird), lysine (835-
870 mg), tryptophan (140-180 mg) etc is indispensable. Below 13% of crude
protein in the mixture the deficit of exo- and endogen amino acids occurs.
In precise balancing the level of ileal digestible amino acids from feed is
being taken into consideration and the proportion between each amino acid
consistent with the pattern of “ideal protein” is sought. In the pattern of
“ideal protein” the quantity of each amino acid is related to the lysine treat-
ed as 100% (Table XXII-5). The amount of protein balanced on the basis of
the amino acids composition and expressed in digestible components as
well as adjusting the quantity of amino acids to metabolisable energy con-
tents in mixtures assured obtaining optimal production of eggs and mini-
mization of feeding costs and of excretion of unused nitrogen to the envi-
ronment.
690
Table XXII-5: The amino acid patterns of hen’ egg protein and recommendations of their
ratio in feed
* estimated on the basis of N balances in the body, eggs and feathers
The requirement of hens for calcium varied, according to different

authors, between 3.5-4.5% depending on the production and on the laying
phase; more constant level of total phosphorus reached 0.5-0.7% (on aver-
age 0.53%), and the level of non-phytate, available phosphorus varied
between 0.33-0.40%. The size of particles of the supplement containing cal-
cium plays a significant role. Slow and regular release and absorption of Ca
from thick particles of the chalk or crushed shells, especially at night (in
darkness) when the shell of egg is formed, assure better mineralization and
solidity of it. The eggs with thicker and stronger shell are obtained from the
hens raised in backyards, where they can pick up small stones (sand) or cal-
careous grit. Also better structure and strength of the bones of jumps are
obtained.
In the mineralization of bones and shell of eggs the significant func-
tion is played by sodium (requirement varies between 0.14-0.16%), chlorine
(0.17%) and magnesium. These elements and some microelements, such as
Zn, Mn, Fe, Cu, Se, I are important for the metabolism of nutrients in organ-
ism, for the functioning of numerous enzymes and organs as well as for syn-
thesis of the egg’ components. Because the mineral components included in
the natural diet is insufficient for realising the genetic potential of animals,
the mineral-vitamin premixes should be used in the feed mixtures. These
premixes are adjusted to the requirement of hens and ensure to meet the
birds’ requirements for both microelements and vitamins (Table XXII-6).
Systematically increased amounts of vitamins in commercial vitamin
mixtures for poultry, including layers, is debatable. Presented recommen-
691
dations on the dosages of vitamins, determined in precise scientific

research, are often 6-10 times higher than the demands of birds. It is
because the recommendations also consider possible lowering of activity of
vitamins as a result of long storage of premixes or mixtures, different degree
of availability of vitamins from natural and processed feeds or vitamin
preparations as well as the risk of mistakes in standardization of these bio-
logically active substances. The results of research that indicate the neces-
sity of application of higher “anti-stress” doses of vitamins (especially vita-
min. E and vitamin. A – possibly in form of ß-carotene - at dose of max.
13,500 IU/kg) are also taken into consideration.
Recommendations of doses of vitamins according to standards of var-
ious countries show considerable diversity because of the various systems
of layers keeping, in the specificity of requirement of particular layer hybrids
for vitamins, in the assortment of applied natural feeds, in the tradition etc.
As an example, recommendations showed in Table XXII-6 are related to the
trends of production of consumable or hatching eggs, which has substan-
tial importance for the biological value of eggs.
692
Table XXII-6. Recommended amounts of vitamins and microelements in 1 kg of feed for

laying hens (88% of dry matter) (Polish Standards of Poultry Nutrition; IFiŻZ PAN, 1996)
* Supplement of the vitamin C is useful within period of summer heat and in the condi-
tions disadvantageous to hens (vaccinations, diseases)
** Larger quantities of manganese are necessary for meat-type hens
Radical representatives of the ecological conception in poultry feeding

are negating the necessity of introduction of mineral-vitamin premixes,
treating them as introduction of chemicals into diets. However, rely on the
abundance of vitamin in natural or processed feeds, such as dried cereal
grain, soya bean oil meal, rapeseed meal, animal meals is creating huge pro-
duction risk for greater flocks of poultry.
Feed supplements favouring maintenance of the good status of birds
health, improved the utilization of nutrients from feed mixtures, laying rate
and organoleptic features of eggs are being applied into the feeding of com-
693
mercial hen flocks. Too low a natural activity of cereal phytase requires
additional application of microbiological phytase preparations, facilitating
the phytates degradation in the digestive tract of birds. Apart from wheat,
wheat bran and barley, natural activity of phytase in grain and leguminous
seeds is low, but quantity of the phosphorus in these connections - high.
Application of this feed supplement can improve the utilization of the phos-
phorus from grain and seeds of leguminous plants and rape by about 15-
30% and on the same way can reduce excretion of the unutilised phospho-
rus into environment. Synthetic amino acids, probiotics, feed preservatives,
feed enzymes, antioxidants and other substances registered through WHO-
FAO and EU are permitted to be applied in the diets of commercial layers.
However, coccidiostatics should not be applied during the laying period (see
Chapter XVIII). When the flock is treated with therapeutic substances, the
withdrawal period in eggs distribution should be introduced.
In order to improvement yolk colour the preparations containing
carotenoids or pigments are being applied. These substances can be trans-
ferred into the yolk. Corn and dried green plant materials are rich in zeax-
antin, lutein, xantophylls. Other feeds hardly contain any carotenoids. Also
the dried flowers of marigold (Tagetes spp.) and paprika are good sources of
carotenoids, although the quantity of these active substances is decreasing
during storage, so the application of antioxidants is necessary. Synthetic
substances, derivative of apo-carotenic acid, cantaxantin and other
carotenoids make it possible to obtain yolk colour over 12 points in the 15-
point Roche scale.
In situations of the threat of microbiological contamination of feed the
application of inhibitors of fungus or mycotoxins created by them (detoxi-
cants) as well as preparations reducing the risk of Salmonella proliferation
is necessary.
22.2.1.5. Moulting of commercial flocks
High costs of the rearing of pullets intended for future laying hens can be
reduced by the prolonged productive period of hens as an effect of forced
moulting of the flock and by introduction the second laying season. Birds
periodically lose the plumage then are undergo in the period of rest. It is
possible to utilize these processes in intensive eggs production. In choosing
of methods of moulting the genetic traits, the body weight of birds, their
health status, the alignment of the flock and other factors should be taken
into consideration. The majority of methods is based on introducing the rad-
ical dietary restriction, i.e. on applying starvation for 8-10 days (withdraw-
al of feed) or 2-days with the reduced feeding every second day. The water
restriction through 1-2 days or making the access to water possible every
second day, limiting the quantity of Ca in the diet also is recommended.
Also the use of pharmacological and herbal preparations is possible. The
694
moulting process and the return to the complete feeding appeared after 30-
50 days, depending on the applied method.
22.2.1.6. Nutrition of breeding hen flocks
Breeding flocks in general are fed similarly to the commercial layers,

although there are numerous factors, which affects the hatching egg quali-
ty and needs to obtain healthy, vital chicks. The errors made in the feeding
of the parent flock are influencing the quality of progeny.
Substantial nutritional problems are resulting from the diversity of
the hens’ live weight, because apart from light and middle-heavy breeding
hens a great group are constituted the meat-type hens. With average body
weight of 2.9-3.7 kg they showed tendencies of consuming excessive quan-
tity of feed mixtures (150-170 g or even more) (Table XXII-7). Overeating
leads to fast fat accumulation in the bodies of hens in flock. The distur-
bances in egg production often lead to the sharp fall in laying or even to the
premature ending of laying after 5-6 months. In turn, too small portions of
feed in the relation to the possibilities of feed intake cause nervousness of
birds, aggression and can lead to feather picking or cannibalism. For meat
flocks, both in the period of rearing and in the period of laying it is neces-
sary to restrict the quantity of given feed consistent with the hen’s require-
ment. Meat-type birds usually consume about 10-15% more feed than their
dietary requirements. Breeding layers of light hybrids are usually fed ad libi-
tum, meat-type flocks – receive strictly controlled amounts of feed mixtures.
695
Table XXII-7. Recommendations for feeding of laying hens (Polish Standards of Poultry
Nutrition; IFiŻZ PAN, 1996)
Characteristic is the fact, that simultaneously to the increase in body

weight of breeding hens, the recommended content of protein in mixtures
decreases from 17.5-16.5% for light hens to 15.5-14.5% for meat hens. The
levels of exogen amino acids and of macroelements also decrease. The lower
concentration of some nutrients results from greater quantities of feeds con-
sumed by the meat-type layer. Breeding hens must receive larger quantities
of vitamins in the feed, especially vitamin. A, E, and so-called “hatching-
vitamins”- vitamin. B2, B6, the nicotinic acid, pantothenic acid, folic acid
and biotin protectively affecting the liver of layers and survival of embryos.
Heavy, meat-type layers shows higher requirement for manganese and zinc.
The level of Mn should be increased from 40-60 to 80, even to 90 mg/kg,
and the quantity of zinc from 40-50 mg to 60-70 mg/kg of complete mix-
ture. In feeding of breeding birds the application of different antimicrobial
supplements should be restricted. Their “protective” activity makes the
selection of the breeding material, characterized by great natural resistance
to the diseases, very difficult.
22.2.1.7. Practical feeding of hens
The systems of managements in hen are diversified not only by environ-

mental conditions, the possibility of the regulations of the temperature and
rate of ventilation, but also the kind and the intensity of the hens feeding
should be adjusted to. The most popular is the maintenance of hens on lit-
696
ter, to a smaller extent - in cages, in aviaries or in free-range system, with

or without regulated temperature.
Keeping the hens in cages requires carefully elaborated composition
of feed mixtures assuring to meet the requirement for all nutrients, because
birds do not have access to the other sources of nutrients. Restricted pos-
sibility of movement decreases the requirement for energy, which should be
reduced by about 10-15% in comparison to the standard needs.
Higher requirement for energy is shown by birds that have access to
backyards. Because of particular importance of good development of
plumage of birds exposed to the losses of body heat, the need to fill diets
with the higher amount of the sulphur amino acids, which are necessary in
synthesis of feather’ protein, occurs. Too fast a rate of intake of the granu-
lated feed increases the inactivity of hens and can lead to the feather pick-
ing or to cannibalism. Although, giving of mixture in meal form is
unfavourable, because the layers are busy for a longer time because selec-
tively consume the ingredients of feed mixtures. Giving of the whole wheat,
not-crumbled grain and grit positively affects the bird’ behaviour.
In practice the layers are fed with complete mixtures in which cereal
grains (the whole or grounded wheat, barley, maize) shared about 60-65%.
Moreover, the oil seed meals and seeds of leguminous plants, the chalk and
phosphates, salt and the premixes also are used. The share of maize in mix-
tures should not exceed 30% because of the high level of energy in this
grain. Great quantity of rapeseed oil meal or leguminous seeds can cause
the enteritis and pancreatitis in hens or make the taste of eggs worse.
Some hybrids of hens with the determined genotype, laying brown
eggs, are not able to metabolize (oxidize) the trimethylamine (TMA) in the
liver. The level of this compound is rising as a result of microbiological
degradation of choline in intestine. TMA is transferred into eggs giving the
“fishy” taste and smell. This limits the application of large amounts of rape
seeds containing sinapine, i.e. ester of the sinapic acid and choline. Large
amounts of leguminous seeds or the fish meal also unfavourably affect the
taste of eggs. Commercial hens are being fed with mixtures that do not con-
tain the compounds of animal origin. The amount of exogen amino acids
could be completed by applying of good quality soya bean meal and pure
amino acids.
22. 2.1.8. Feeding of cockerels in rearing and reproduction
The number of cockerels in the breeding flocks usually varies from 8 to

about 10% of the whole population. Thus the application of separate feed
mixtures must be realized through specially constructed feeders. Feeding
adjusted to the requirement of growing cockerels is possible only when the
young males are reared separately. During the first three weeks of life the
cockerels should be fed ad libitum with mixtures containing 18-19% of good
697
quality crude protein and with energy value of 11.5-11.9 MJ/kg.

Restrictions in the amount of feed given to cockerels should begin after the
third or fourth week of life, adjusting the restrictive system of feeding to
their condition, live weight and alignment of the flock. It is possible to
replace part of the mixtures with cereals (wheat, oats) or to apply mixtures
and grain alternatively or apply the feeding every second day. Quantity of
mixtures given daily per cockerel, depending on the body weight and at the
age of 18-20 weeks varies from 95-110 to 115 g, in the second phase of hens
productivity it varies from 110 for light to 165 for meat-type cockerels.
Different systems of mixture dosage are being recommended distinguishing
the week’s portion of the feed on 6 or 5 days, with 1-2 days of starving.
Cocks are introduced to the hen flocks (1:9, 1:10) at 18-21st week of
life. They are higher than females and have broader heads thus putting
feeders at the height where they can be reached by cocks rather than small-
er hens, allow the males to be fed separately. It is also possible to put delim-
iters depending on the size of the heads of birds. During the period of repro-
duction males require smaller quantity of crude protein 10-11%; 0.3%
methionine, 0.6% lysine, only 0.9-1.0% Ca, while the layers have to receive
3.2-3.5% of calcium. Excess of calcium lowers the quality of the semen and
is dangerous for kidney functions. The precise balancing of microelements
– iodine, copper, cobalt, selenium and from vitamins - vitamin. E (above 25
mg/kg), A - 15,000, vitamin. B12 etc. is especially important for breeding
animals. Correct composition of mixture for cocks is difficult because of the
desired low level of protein, thus in practice males are being fed with mix-
tures containing about 14% protein. The dosage of feed should be strictly
controlled.
22.2.1.9. Feeding of meat-type chickens (broiler)
In contrast to the controlled feeding and body weight of future reproductive

flocks, the purpose of broiler feeding is to obtain the maximum of body
weight gain in the shortest possible time with good feed utilization, high
slaughter yield and maximum best quality of meat. During the last 30 years
the significant genetic progress has taken place and, at present, the broiler
chickens are reared only to week 5. 6. or 7 of life depending on the market
conditions and production costs. It is also noteworthy that now the feed uti-
lization index varies between 1.7-1.9 kg per kg body weight.
Fattening performance of different flocks can be compared by means
of the Dutch Index (DI).
Raising % × Sold weight, kg

DI = ———————————————————— × 100
FCE, kg/kg × Raising time, day
698
For example, if a flock was sold on day 42 of raising with an average

live weight of 2.3 kg, having 8% losses (mortality +culling) and a feed con-
version efficiency of 2.0, the DI is (92×2.3/42×2.0×100=252. The higher is
the Dutch Index, the better is the performance.
Different feeding systems are applied depending on the genetic poten-
tial of the chickens and economic condition of market. These depend on the
application of different feeding periods, such as feeding with pre-starter
mixture (1-7 days), starter (to 21 day) then grower and finisher (last 5-7
days before slaughter) or other systems. The mixtures with high concentra-
tion of nutrients in the intensive broiler rearing are proposed, but there are
also “economic” mixtures containing lowered amounts of energy, protein
and other components which are proposed. The “ecological” feeding systems
do not allow the use of too many feed supplements, which also contain syn-
thetic amino acids necessary to correct deficiencies, especially when the
low-protein mixtures are given to the birds.
In standard feeding of broiler chickens, the mixtures should contain
21-24% of crude protein, 0.45-0.52% methionine, 1.1-1.4% lysine, 11.9-
13.5 MJ ME/kg, Ca 0.9-1.1% and P available 0.38-0.45%. In heavy broilers
(roaster) rearing, up to 8-10 weeks the grower and finisher mixtures with
lowered to crude protein 16-17% are applied. These mixtures contain also
lowered amounts of other compounds, such as methionine 0.36-0.30, 0.84-
0.70%, lysine and expanded energy to protein ratio. Uncontrolled increase
of energy concentration can lead to the excessive fat deposition in the body.
Thus the optimal level of protein in feed, density of energy and contents of
other nutrients in feed mixtures should result from applied rearing system.
Meat-type (broiler) chickens are able to consume huge amounts of
feed and their appetite, which depends on the energy value of their diet,
result mainly from the fast growth. The energy level in the diets is the basis
of balancing all nutrients, because, according to our present knowledge, the
ratio of energy to the amino acids digested in the intestine or to the avail-
able minerals determines the efficiency of chicken nutrition. In modern sys-
tems for balancing of concentrate mixtures the index of energy to protein
ratio is still not used. Nowadays, the relation of metabolisable energy to the
amino acids degraded in the intestine is considered to be a crucial factor of
mixture quality. Considerable are the relations of exogen amino acids to the
lysine used as a reference amino acid (100%). “Ideal protein” pattern should
be adjusted to the genetic value of birds, their age and sex. This way of bal-
ancing protein is still not common because of the necessity to control the
actual amino acid composition in the feeds applied in the chickens feeding.
It is the energy value of mixtures for broiler chickens that determines
to a great extent body weight gain, rate of feed intake, its conversion as well
as fat deposition in the carcass. Depending on the planned intensity of pro-
duction and the time of slaughter (day 35, 42 or 56) the energy level vary
between 11.9-13.5 MJ ME/kg. According to the international standards
699
minimal energy value of mixtures should not be lower than about 12.0 MJ
of metabolisable energy in 1 kg of feed. The high energy value of fed mix-
tures can cause a deposit of large amounts of fatty tissue in the body and
can increase the frequency of outbreak of some metabolic diseases, such as
sudden death syndrome. The ratio of 1 MJ ME to 0.6-0.9 g of lysine and 0.3-
0.4 g methionine is considered to be optimal.
The fast-growing birds strongly need minerals. The minimal level of
minerals is usually established in the national recommendations. The
amounts of some macroelements can not be lower than e.g. 0.8-1.0% for Ca;
0.6-0.7% for P total and 0.12-0.14% for Na. Deficiency of macro and
microelements are rare because in composition of complete mixtures the
special mineral-vitamin mixtures (premixes) adjusted to the age and feeding
period of chickens, are used. Important is the balancing of the phosphorus
in the mixtures; phytic phosphorus is available to the birds to a very small
extent only because of low activity of phytase present in raw feeds or secret-
ed by intestinal microflora. The knowledge of the amounts of available phos-
phorus from natural feeds and the use of microbial phytase allow decreas-
ing the amount of phosphorus coming from phosphates, to be decreased.
Great variability in the contents of microelements in natural com-
pounds and incomplete data on their availability make the precise balanc-
ing of their level in mixtures almost impossible. The premixes that contain
microelements such as chelates, oxides, sulphates, chlorides or other salts
are significant supplements added to mixtures. Because of environment and
human health protection, the application of uncontrolled, high levels of Cu,
Zn or Mn in feed mixtures or utilizing of the “pharmacological” effects of
high doses of those chemicals (micro-elements) are prohibited.
In standardization of vitamins application for broiler chickens in the
fattening period, body weight of birds, energy value of mixtures and other
parameters should be considered. Recommendations, according to the stan-
dards from different countries, varies considerably. For some vitamins the
high, “anti-stress” dosages are also recommended, e.g. for vitamin A even
up to 20.000 IU/kg, however, according to the EU Directive, the permissi-
ble level of this vitamin for broiler chickens can not exceed 13.500 IU/kg
mixture and vitamin D3 maximum 3.000 IU except the turkey broiler (to
5.000 IU/kg feed). Increased dosages of vitamin E, even to 400 mg/kg can
enrich the final product – broiler’ meat – with this vitamin that show antiox-
idant properties.
Parallel to the goal of producing of functional food for human, in the
nutrition of meat-type chickens many different supplements are introduced.
These included increased amounts of vitamin (A, E), but also microelements
– I, Se, polyunsaturated (in them also condensated) fatty acids (CLA) of fam-
ily n-3, which are present in fish oil, in rape oil, in oil from common evening
primrose (Oenothera biennis) or from linseed oil (Linum sativum). The levels
of these supplements do not exceed the maximal amounts that are allowed
700
in the poultry feeding in EU countries.
22.2.1.10. Feed supplements applied in feeding of broiler chickens
The non-starch polysaccharides included in cereal grain, such as arabi-

noxylans, β-glucan, cellulose and other can constitue from about 10%
(corn) through 12-14% (wheat, triticale, rye) up to almost 30% in dry mat-
ter in oats. These substrates contain energy which is unavailable to birds.
They are unable to degrade these chemical bounds because of the lack of
endogenous enzymes. The pool of energy that is obtained from the degra-
dation of these carbohydrates by intestinal microflora is not sufficient for
the birds’ organism. Moreover, these polysaccharides, especially their
hydrophylic fractions can cause negative effects in the lumen of gastroin-
testinal tract – increase of the digesta viscosity, difficulties in the digestive
enzymes secretion and in absorption of nutrients through the intestine wall
that is covered with the hydro gels. It results in the disturbances of water
management in the intestine and in the microbial stability of digesta.
In order to improve the use of potential energy from NSP and decreas-
ing of antinutritional activity of them, the enzymatic preparations are intro-
duced to the mixtures. They are characterized by higher activity of β-glu-
canase, cellulase, xylanase and other carbohydrases. The both quantity and
quality of the enzymatic preparations must be adapted to the kind of sub-
strate i.e. structural carbohydrates prevalent in the cereals or in legume
seeds that are used in feed mixtures (e.g. β-glucanase – barley; arabinoxy-
lanases – wheat etc.) and to the enzymatic activity declared by the produc-
ers of these. Also phytase belongs to the group of enzymatic supplements.
This enzyme is used for the purpose of improving the availability of phytic
phosphorus from grain, legume seeds and their derivatives.
Great interest of scientists and producers is focused on the nutrients
and feed supplements that can stimulate the immune system of animals
and indirectly that of humans. Some of them are high quality proteins that
act in hormone synthesis and the creation of blood cells, polyunsaturated
fatty acids from family n-3 and especially conjugated linoleic acid (CLA),
vitamin A, E, C as well as microelements – Zn, Se and original active plant
substances – plant extracts in which the flavonoids plays a most important
role. The purpose of replacement of banned antibiotics with the probiotics,
prebiotics and organic acids is to stabilize the intestinal microflora, to
decrease the colonization of intestine with the pathogens and to improve the
chickens’ health status.
The deficiency of exogen amino acids originating from natural materi-
als can be corrected with synthetic amino acids (methionine, lysine, threo-
nine and tryptophan). Deficiency of other amino acids is relatively rare.
Coccidiostatics that are included in feed mixtures for broiler chickens
decrease the possibility of coccidiosis outbreaks, although their application
701
must fully comply with the recommendations that guarantee effective

amounts of ionophores or other substances to prevent Eimeria from prolif-
eration. Introduced prophylactic vaccinations may eliminate the necessity of
using coccidiostatics in the future.
22.2.1.11. Concentrate mixtures
The broiler chickens are fed on concentrate mixtures as an only source of

the nutrients. These are made from the ground cereal grains, mainly corn,
in the countries of Middle and North Europe from wheat, barley, soy bean
oil meal, legume seeds and/or rape seeds oil meal from the 00 varieties. The
application of fish meal is permitted, although there is a ban on the appli-
cation of the meat or meat-bone meals in mixtures. The mineral compo-
nents are completed through the introduction of chalk (limestone), feed
phosphates and mineral-vitamin premixes into mixtures. The purpose of
aggregation of the mixtures (pelleting) at a temperature of 70-85°C is to
maintain their homogeneity, to make the selection of feed particles impos-
sible, to put the structure into shape close to the natural size of feed parti-
cles - cereal grain, to improve the digestibility of nutrients (conditioning with
water steam can improve the starch digestibility) and also, to promote inac-
tivation of antinutritive substances.
2.2. Principles of turkey nutrition

Morphological characteristics of the digestive tract in turkeys are similar to
those of hens. Turkeys have short digestive tract in relation to the length of
the body (5-6:1), actively functioning crop, inhabited by lactic acid bacteria.
Fast passage of feed (digesta) through the digestive tract (4 to maximum 8
hours when the large quantities of structural substances are given). Since,
in relation to large live weight, secretion of digestive enzymes is medium,
therefore maintenance of high rate of nutrients digestion careful balancing
of diets and feeds preparation is necessary for these high need birds.
Turkeys are characterised by high rate of growth up to about 11-12
weeks, although very good body weight gain is obtained in the turkey hens
to up to the age of 15-18 weeks, and in turkey cocks even up to 24 of week.
Smaller than in other species quantities of subcutaneous and abdominal
fat, meatier carcass and great amount of protein in it cause turkeys to have
the highest requirements for protein in comparison with other commercial
birds.
Daily weight gains in young fattened turkeys reach 150-160 g and are
the highest in the group of fattened poultry. The huge muscular mass cre-
ated in the modern hybrids results from the intensive metabolism, but also
is the reason for many diseases and development anomalies of young
turkeys, such as: deformations of poorly mineralised bones, mainly legs,
702
increased pressure of extended muscles on the blood vessels which causes

dysfunctions of blood circulation, deaths as an effect of sudden death syn-
drome (SDS) and ruptures (cracking) of the aorta and other diseases.
22.2.2.1. Feeding of growing turkeys

The feeding of young turkeys should consider the future use of birds - as
reproductive flocks or as fattened turkeys. This diversity of future use deter-
mines the intensity of feeding, genotype of birds, and energy concentration
in feed mixtures etc.
In the first period of intensive growth the turkeys both hens and cocks are
fed on the same mixture given in quantities prescribed in recommendations
prepared by the producers of the animal material and adjusted to the rate
of growth. The quantity of protein in the starter mixture usually varies with-
in 28-26% and energy values vary between 11.7-12.2 MJ ME/kg (Table
XXII-8).
703
Table XXII-8: Feeding of the parental flock of semi-heavy turkeys (BUT-8; and BUT-9)
and heavy turkeys (Big-6) in the rearing period (BUT, 1995) (JANKOWSKI, 2001)
704
Page 704
15:59
2008.09.03.
* minimum and maximum val-

JAV_22_VND_FSGy:MINTAÚJ.qxd
ues are given

** quantitative limitation of diet
should be introduced
The mixtures for young, growing turkeys contain high quantities of vitamins and mineral components than those
for other species of poultry and depend on genotype of turkeys (Table XXII-9 and Table XXII-10).
Table XXII-9. Nutrient requirements of turkeysUnitGrowing turkeys, males and femalesBreeders

2008.09.03.
15:59
Page 705
705
2008.09.03.
15:59
Page 706
706
Legends to the TABLE XXII-9
a - The age intervals for nutrient requirements of males are based on actual chronology from previous research. Genetic improvements
in body weight gain have led to an earlier implementation of these levels, at 0 to 3, 3 to 6, 6 to 9, 9 to 12, 12 to 15, and 15 to 18
weeks, respectively, by the industry at large.

b - The age intervals for nutrient requirements of females are based on actual chronology from previous research. Genetic improvements
in body weight gain have led to an earlier implementation of these levels, at 0 to 3, 3 to 6, 6 to 9, 9 to 12, 12 to 14, and 14 to 16
weeks, respectively, by the industry at large.
c - These are approximate metabolisable energy (ME) values provided with typical corn-soybean-meal-based feeds, expressed in kcal
MEn/kg diet. Such energy, when accompanied by the nutrient levels suggested, is expected to provide near-maximum growth, par-
ticularly with pelleted feed.
d - Requirement may increase with wheat-based diets.
e - The levels of Mn (60 ppm), I (0.4 ppm), Se (0.2 ppm), vit. A (5.000 IU), D3 (1.100 ICU), thiamin (20 ppm) are provided in the same
niveau in all growth periods.
In the composition of premixes applied to the feed mixtures the special-

ized coccidiostatics permitted for application in turkey diets occurs. They
must be consistent with the preventive application of veterinary services.
However, sometimes they are removed from planned diets, when an immu-
nization against coccidiosis is possible. During this period the prevention
against histomonadiosis is also preferable. Size of feed particles should also
be adjusted to the age of birds. Young turkeys receive crumbled feed (the
diameter 2-3 mm) then granules with the diameter of 4-5 mm. The concen-
tration of nutrients in mixtures is reduced in relation to the age of birds,
and the constant control of the body weight and the alignment of the flock
are necessary. The amount of the feed portion is rationed. The growth rate
and development of the growing birds should be consistent with the tech-
nology given by the producer of the animals. In case of too intensive growth
the quantity of the given feed should be limited (reduction by 10, 15 or
20%), the whole, not-crumbled grain of the wheat or the oat is given.
Table XXII-10: The levels of microelements (mg/kg) recommended to growing turkeys
707
* Recommended supplement to the mixtures

** Total amount in mixture
22.2.2.2. Feeding turkeys in the breeding period
The fundamental principles of feeding turkeys in the breeding period are

similar to those given for hens. The preparation for the reproduction should
begin at the age of 26-28 weeks, in accordance with the management’s rec-
ommendation. Along with introducing the light program stimulating physi-
ological processes of beginning of laying the concentration of nutrients suit-
able for the anticipated energy value of mixtures and temperature inside
rooms should be increased.
Ad libitum feed intake by the turkey layers should be accompanied by
periodic control of the live weight because there the accumulation of fat in
the carcass should be prevented. Great amount of fat tissue leads to an
increase in the frequency of liver diseases reduces the hatchability of eggs,
can cause oviduct prolapsed, kidney diseases etc. Periodically it is possible
to limit feed rations or use whole cereal grain.
Turkey cocks are usually kept and fed separately, because higher
quantity of calcium in mixtures is necessary for females to produce eggs
(2.2-2.4%). In the cock’ organism the high Ca level can lead to serious dam-
ages of the excretory system - of kidneys and ureters. The excessive increase
in the body weight of turkey cocks drastically reduces their reproductive
potential and also leads to overloading of the limbs and leg diseases.
22.2.2.3. Feeding of fattened turkeys

Intensity of feeding of young fattened turkeys has to be adjusted to the
genetic potential of breeds or hybrids, and at the same time the economic
aspects should also be considered, because the expenditure of feeding fat-
tened birds could constitute as much as 60% of production costs.
Recommendations for feeding of fattened turkeys are much more
diversified than e.g. those of hen broilers. It results from the fact that there
are greater differences in the characteristics of various hybrids, in their dif-
ferent rate of growth and the body weight gain as well as in the recommen-
dations elaborated by various breeding companies. Thus the requirements
for nutrients are always related to a certain hybrid, and differ in the sex.
Depending on the concept the phase feeding of fattened turkeys is applied
with changing of the composition of mixtures usually every 4 weeks. The
turkey hens are raised to the 16th, maximum 20th, tom turkeys to the 24th
week of life.
In order to obtain good efficiency in rearing and the best possible
quality, fatless carcasses and at the same time savings in utilization of mix-
tures, in the rearing of turkeys periods of restrictive feeding are introduced,
mainly in the early phases of the growth. The whole wheat grain and cheap-
708
er feeds, various feed supplements, mainly enzyme preparations, probiotics

and organic acids are also applied successfully. Also the concentration of
vitamins and some mineral components in mixtures are increased which
prevent diarrhoeas (zinc compounds), ruptures of the aorta (Cu, Mn, Co),
fatty liver degeneration (biotin, selenium) and limbs diseases.
In optimization of the composition of mixtures the amino acids pro-
file, according to the “ideal protein” pattern for growing turkeys, should be
taken into consideration. Also, contents of polyunsaturated fatty acids,
CLA, selenium, iodine and many other components that are makes it pos-
sible to obtain the improvement of “pro-health” quality of final product –
turkey meat, are being increased. Introduction of the feeding programmes
into practice in the purpose of production of “programmed and functional”
food is obtained by modifying the chemical composition and components of
complete mixtures.
2.3. Principles of goose nutrition

The goose (Anser domestica) was domesticated around 5000 years ago (FIG-
URE XXII-4).
FIGURE XXII-5: Wall picture fron the Ancient Egypt
It is still strongly connected to the habitat and plant production, mainly

because it consumes large quantities of green matter and/or root vegeta-
bles. These birds, better than other species, can utilize the home-made fod-
ders containing high amounts of the crude fibre, thus the goose is often
named “ruminant” among poultry.
22.2.3.1. Morphology of the digestive tract
709
The adult goose is able to consume large quantities of fodders, about 20%
of its live weight, as much a 1000 g green matter. Very strong beak with
sharp brinks, the flexible and extensive oesophagus, the big muscular giz-
zard (45-60 g/1 kg of the body weight) covered with the thick and tough ker-
atinous layer indicate the adaptation to taking in, crumbling and removing
of excess water from green matter and roots. The internal pressure in the
gizzard of hens amounts 10.2-15.3 Pa, in ducks 17.8-18.3 and it is highest
in the goose reaching 27.0-28.6 Pa. The goose is characterized by the
longest intestine among domestic birds. Also relation of the length of intes-
tine to the length of the body is broadest in the comparison with hen or
duck. The length of the digestive tract in the adult goose may reach even 4
m.
Variations of the length of each part of the digestive tract are espe-
cially clear in young birds. In 4-5-week-old geese, the length of the digestive
tract reach about 70-80% of the adult size, at the age of 2 months it reach-
es 85-98%. In older geese the length of small intestine is increasing slight-
ly. The development of the digestive tract ends before the age of 34-37 weeks
(7.5-8 months), i.e. after reaching the sexual and somatic maturity.
Intensity of the birds feeding during the growing season, as well as intro-
duction of feeds rich in structural carbohydrates significantly affect body,
the length and the weight of intestine. The use of greater portions of feeds
that contain high amounts of pectins and hemicelluloses lead to an increase
in the length of intestine by about 3-8% (oats), by 9-12% (dried green fod-
der) and by 27-49% (dried beet root pulp) in comparison with control ani-
mals fed standard mixture.
The kind of feeds offered to birds and the amount of structural poly-
saccharides in them are not left without effect on the intestinal wall’ mor-
phology and on the digestion of nutrients by geese. Increased share of oats
(>50%), dried beet root pulp (about 31%), dried alfalfa and grass (20%) in
mixture positively affects the thickness of the small and large intestine wall
(by 16-23%). The feeds containing pectins which binds water and causes
light dehydration of the intestinal digesta, mechanically affect the intestin-
al villi causing the shortening and swelling of them, and even morphologi-
cal changes - active congestion of the network of vessels of mucous mem-
branes with abundant lymphocyte infiltration and enlarging of lymphatic
follicles. It is interesting and noteworthy that the negative changes in the
intestinal wall, in kidneys and in the liver, observed when the dried alfalfa
was given to young geese, did not occur when the birds were fed on dried
grass, and i.e. feed containing less saponins and phytohormones.
22.2.3.2. Microorganisms in the gastrointestinal tract
Numerous micro organisms are colonizing the digestive tract of poultry. The
number of this population varies in the limits of 106-1011 per gram of the
710
intestinal content (Table XXII-11). Eubiosis - the balance of the population

of Lactobacillus, Gram+ and Gram- bacteria, E. coli and other groups of
microorganisms - depends on pH, the rate of passage of the intestinal con-
tent, concentration of the protein (the buffering, NH3 formation) and on the
amount of mineral components.
Table XXII-11. The pH value and number of microorganisms in the goose digestive tract
In the caecum and large intestine the microbiological degradation of

the components of dietary fibre and formation of large quantity of short-
chain fatty acids occurs as a result of bacterial cellulase, xylanase and other
enzymes activity. In these parts of the digestive tract the presence of
Bacterioides succinogenes and Ruminococcus flavefaciens is comparable
with their quantities in the rumen. The digestibility of amino acids in the
intestine depends not only on the kind of protein fed but also on participa-
tion of different bacteria in the process of degrading these bonds.
22.2.3.3. Digestion in geese
The specificity of digestion of nutrients in waterfowl results from the fact

that in these birds the crop function is fulfilled by the dilatation of the
oesophagus. In this part of gastrointestinal tract the saliva containing
mucin that facilitates swallowing of the feed is secreted. The quantity of gly-
cosidase and other enzymes and ptyalin is very low and it is also debatable
whether they are secreted by the organism or are of exogenic origin deriving
feeds. The main digestion process is starting in the glandular stomach, in
which the hydrochloric acid and proteolytic enzymes (pepsin) will degrade
the proteins and peptid bonds. Beta-galactosidase secreted in mammals,
does not occur in the liver of poultry. Thus the birds are not able to digest
711
lactose. It is degraded during the bacterial fermentation in subsequent parts

of the intestinal tract. The characteristic feature of poultry, and that of
geese, is the relatively small pancreas containing of a many islets of
Langerhans. Insulin, but mainly glucagons are secreted by them. In poultry
the basis of alpha-amylase activity is performed by an enzyme of pancreat-
ic origin. The alpha-amylase is not produced in the liver and the activity of
this enzyme depends on the flow of blood and the transport of this enzyme
into liver. The low alpha-amylase activity in blood usually indicates good
utilisation of carbohydrates from the diets.
Digestion of the starch’ fraction of the diet is particularly essential in
the feeding of birds because these carbohydrates constitute a fundamental
component of feed mixtures for poultry. It was found that the different lev-
els of starch digestion depend on its physico-chemical composition and on
its origin. The starch from grain is degraded mainly by the pancreatic alpha-
amylase, however the weak utilisation of potato starch – feed used in geese
feeding, is caused by the low activity of alpha-amylase secreted by salivary
glands, which in poultry are less active than in other animals. High activi-
ty of proteases is determined in the stomach. It decreases in further parts
of the digestive tract, although the amylase activity is reaching its maximum
value in the content of the ileum. High levels of celulolytic activity is detect-
ed in the content of the caecum, which is accompanied by fatty acids for-
mation, what proves the dynamic of microbiological carbohydrates fermen-
tation at a level similar to the rumen of sheep. Cellulolytic activity of the
caecum content is almost two times higher; also the activity of dehydroge-
nase is three times higher than in the rumen of sheep. However these issues
are still not well recognized.
The long and capacious digestive tract in geese creates favourable
conditions for good digestion of bulky feeds – roots, green matter, grains and
leguminous seeds. Corn and the wheat are the best digested grain (Table
XXII-12). Attention should be given to the good digestion of the crude fibre
in geese (12-40%).
Root-feeds are very well digested by geese. However, the weaker
degradation of N-free extractive substances coming from raw potatoes
(about 44%) and from the carrot and beets (77-83%) should be underlined.
Dried feeds obtained from these fodders to a high extent are degraded in the
digestive tract. Thermal processing of potatoes (potato flakes) considerably
improves the apparent digestibility of the starch and other saccharides, it
reaches even 91%. The use of silages in geese feeding, which is recom-
mended in practice, is going to be less justified. The values of digestion coef-
ficients of nutrients, determined for these feeds and resulting from the
change of pH of the intestinal content and the decreased activity of enzymes
digesting carbohydrates, were generally low. However good digestibility of
the nutrients was determined for green matters. Apparent digestibility of the
crude fibre from mixtures given to geese varies between 10-33%, of the
712
hemicellulose between 55-76%. 15-34% of cellulose and 66-74% of pen-

tosanes is digested by geese. Quite high digestibility of pentosanes has a
significant importance, because the quantity of these substances e.g. in
barley and oats amounts to 8.2-11.1%, while in corn and wheat the quan-
tity of pentosanes is below 6% of the dry matter.
Table XXII-12. Apparent digestibility of some feeds in geese, %

(JAMROZ and BIELIńSKI, 1985)
1 root feeds 300-400 g; cereals 20-120 g; complete mixtures 65-150 g; dried grass 70-
110 g
The digestibility of pectins present in the feeds for poultry in consid-

erable quantities is also debatable. About 8% of pectins is present in the rye,
dried green matter contains about 8-10%, dried beet pulp 12-16%. High
713
digestibility of these substances was found in the geese (58-68%) and in

pigs. The relatively good digestion of the crude fibre fractions in geese is
explained by the microflora activity and by specific function of the great cae-
cum.
22.1.3.4. Intestinal fermentation of polysaccharides
The cereal grains are the basis of mixtures given to geese. The energy val-
ues of them are dependent on the starch, crude fibre as well as on non-
starch carbohydrates: beta-glucans (4-17%), pentosanes (5.8-8.7%), pectins
and other polysaccharides content. The bacterial digestion (degradation)
and fermentation of these substances in the intestine happens in presence
of Lactobacillus, Streptococcus, E. coli, Bacterioides. Similarly to other birds,
as a result of polysaccharides degradation and glucose fermentation inside
the goose intestine, short chain fatty acids (SCFA) are created. The quanti-
ties of these acids vary in geese from 20-25 mmol/l in the small intestine
up to 190 mmol/l in the content of caecum. The share of acetic acid in total
VFA amounts to 50-60%, of propionic acid - utilised in gluconeogenesis -
19-32%, and butyric acid 18-21%. It should be noted that considerably
larger quantities of fatty acids are formed in ducks than in intestine of geese
or chickens, despite the fact that the lower coefficients of apparent
digestibility of N free extract and crude fibre are just observed in ducks. It
could also be stated that the large quantities of these undigested carbohy-
drates is strongly degraded by bacteria.
The fermentative processes are most intensive in the caecum, in
which the digesta contains a small quantity of bigger particles of structural
NDF and ADF substances. Presumably these are not passed to this part of
intestine. The question, whether the SCFA formed in birds’ caecum are sig-
nificant source of energy for these animals still remains unexplained.
Volatile fatty acids can be metabolized in the intestinal mucosa, which is
necessary for the correct functioning. Absorbed, can cover up to 15-20% of
energy for maintenance in rabbits and pigs. The degree of VFA absorption
from geese intestine is not yet exactly known, although sometimes it is esti-
mated on the level of 15-45%.
22.2.3.5. The requirement of growing and breeding geese for nutrients
The future productivity of breeding geese to a great extent is determined by

the intensity of rearing of young goslings. A wide range of requirements for
substantial nutrients depends on the growth phase and future use of geese.
For crude protein it varies from 22.5 (0-3 weeks) to 14 % (13 weeks to the
maturity); for metabolisable energy 11.7-10.0 MJ ME/kg; Ca 1.0-0.6%; P
total 0.7-0.5% (P available 0.4-0.3%); lysine 1.1-0.73%; methionine 0.48-
0.32%.
714
Moderate intensive, controlled rearing of goslings apply feed mixtures

containing the following components in subsequent phases of geese devel-
opment: crude protein from 20-21-18 to 14%; metabolic energy from 12.0-
11.5 to11.0 MJ ME/kg; Ca from 1.0-0.9 to 0.6%; P total 0.6%; lysine 0.9-
0.65%; methionine 0.45-0.33% positively affects the correct growth of
goslings and makes it possible to obtain good laying in the first and the fol-
lowing years of geese production. Intensive feeding in the rearing period
indeed makes the earlier start of laying and higher production of eggs in the
first season possible compared to birds fed economically, although in the
next years the productivity of layers fed semi-extensively during rearing is
considerably higher. Total feed consumption during the rearing period
reaches about 4.5 kg (1-4 week), 8.5 kg (5-8 week) and about 9.0 kg (9-12
week), which in total reached about 21-22 kg per one reared goose.
22.2.3.6. Feeding of the breeding geese
The amount of the basic nutrients and their energy content in complete-
mixtures given to breeding geese, according to different authors and
sources, vary at greatly (Table XXII-13). These data concern different breeds
of geese and different management systems. Application of green grass in
the geese diets (200-700 g/d/bird) cause that the composition of addition-
al feed mixtures should be adjusted to the combined feeding system.
Diversification of mixtures composition should also be determined by the
laying phase and the intensity of production. The concentration of energy in
mixtures given within a period of rest must be lowered to 9.2-10.0 MJ
ME/kg, and the crude protein to about 12%. There is necessity of clear
diversification of energy and crude protein requirements of the different
phases of production period (Table XXII-14).
715
Table XXII-13: The examples of the requirement of breeding geese for nutrients
716
Table XXII-14. Some data on the requirement of breeding geese for nutrients (Polish
Recommendations of Poultry Feeding, 1996) and NRC (1994)
The effect of adaptation of the geese to the different quantity of the

protein in feed is considerable. Positive N balance is still possible to obtain
when the laying geese are fed on a diet containing 10% protein. By feeding
diets containing 13.5% and energy density of 10.5 MJ ME/kg, the geese are
able to achieve good production (over 70 eggs). The number of fertilized eggs
and the number of goslings per goose are similar to the case noted when
higher concentration of nutrients in feed was given to birds (16% of protein
and 11.6 MJ ME/kg). Geese do not require the presence of animal protein
in the feed. However, because of correct moulting, particularly after feather
plucking, when they are likely to be fed on low quantity protein in mixture,
the periodical supplement of synthetic DL-methionine seems to be prof-
itable.
The energy value of mixtures for geese should not exceed 11.5 MJ
ME/kg in order to prevent excessive fat deposition and impairments of liver
functioning. Overfatening of geese, particularly males, negatively affects
their sexual activity and in the same way the fertilizing of eggs. However, it
was often underlined that the higher number of eggs with higher weight was
717
obtained if better quality mixtures were given to birds.

The ganders more greedily and faster consume the feed than females
and easily become overfed and obese. Efforts to prevent decrease of egg fer-
tilisation after 3.0-3.5 months of continued reproductive by intensification
of the feeding, application of additional vitamins; particularly A and E (to 40
mg/kg) have not given explicitly profitable result.
Introduction of two laying cycles (in total 8-9 months of the repro-
duction) makes it possible to obtain more than 90 eggs from one goose.
Managing the reproductive process is done through the light program and
proper feeding in the period of rest (24 g of crude protein, 2.09-2.30 MJ ME
daily per head), in the preparatory period (30-35 g of protein, 2.50-2.80 MJ
ME/d/head) and in the a period of eggs production - about 3.80 MJ ME and
52-56, even to 60 g of the crude protein daily per layer. Concentration of
nutrients in mixtures should be modified depending on the planned size of
daily ration of mixture. The quantity and the kind of other domestic (farm)
feeds given to geese should also be taken into consideration.
22.2.3.7. Requirement of geese for vitamin and minerals
Demand for vitamins in goose is still weakly understood and the numeric
values given by different authors or standards are characterized by a great
variability (Table XXII-15). The current NRC standards do not give the rec-
ommendation of vitamin application for this species.
718
Table XXII-15. Some examples of the goose requirement for vitamins
* standards for ducks
719
Wild geese or geese kept on the pasture can consume large amounts of
green fodder and in the same way great quantities of natural xanthophylls.
However, the amount of lutein, zeaxantin and other carotenoids in feed mix-
tures is not high. It usually varies between 22-24 mg/kg of mixture. These
components come mainly from corn or dried green grass. Carotenoids, β-
carotene and vitamin A play a significant role in the organism, inter alias
stimulate the synthesis of progesterone - the hormone important for breed-
ing.
The yolk of goose egg contains more vitamin A (3720-7400 IU/100 g)
than the yolk of hen egg (3210 IU/100 g). This quantity depends on the age
of goose, the contents of vitamin in the feed and on the successive month of
reproductive period. The view that these birds have a particularly demand-
ing requirement for retinol is prevailing, although our own long term exam-
inations do not confirm this suggestion.
The level of 10.000 IU vitamin A per kg of the mixture is ensuring the
proper biological value of hatching eggs. Higher quantity of this vitamin
improves, though ambiguously, the efficiency of laying and hatching. The
addition of 16-24 mg of ethyl ester of β-carotenoic acid to the standard level
(10.000 IU vitamin A/kg) in the geese feed or 8 mg/kg together with 20.000
IU vita. A/kg considerably increases the number of goslings obtained from
a layer, chiefly as a result of reducing of embryos’ mortality during the
hatching of eggs. Applying of this carotenoid improves the survival of
goslings coming from the layers fed in this way. In our own experiments it
was observed that the layers towards the end of laying season had a good
appetite, were lively and in good health.
Some examinations indicate the usefulness of increasing the quanti-
ty of vitamin E in mixtures for geese even up to 40 mg/kg of the feed.
Tocopherol improves the index of eggs fertilization to 91-96% and their
hatchability as well. The necessity of increasing dosages of vitamin A, D3
and E when the big quantities of beets are fed, also is being underlined. The
oxalic acid included in the beets reduces absorption of these vitamins, as
well as Ca and P in the digestive tract of the goose. Some authors are of the
opinions that giving vitamin B1, niacin, folic acid and biotin in the diets for
waterfowl is not necessary because their quantities in raw, natural feeds
seem to be sufficient. However, observations in practice contradict this view.
Given to geese fed only on concentrate mixtures, additional amounts green
matters or yeast rich in these nutrients improved the health status of
goslings, in which the rickets was incorrectly recognized. Cholin in 1000
mg/kg quantities protects the reared geese against perosis. The require-
ment of geese for nicotinic acid is being determined at 60, and for the pan-
thotenic acid at 20 mg/kg of feed. However, these suppositions are poorly
documented with data from the small number of experiments.
The requirement of geese for minerals are characterized mainly by the
quantity of calcium necessary (to 2.7%) and for the phosphorus (to 0.6%).
720
The data from research on the requirements of these birds for microele-
ments are scarce in the available literature. Determining of geese the needs
for minerals through the control of the level of these components in the
blood serum is extremely difficult because in these birds there is a great
variability of many biochemical indices depending on the season and the
sex. The most significant variations concern the concentration of the calcium, phospho-
rus, lipid compounds of blood, iron and transferrins in females. Variability of these indices
in ganders depends on season is small.
22.2.3.8. Practical feeding of geese

Many problems of the practical feeding of geese are related to the applica-
tion of different components in mixtures in an attempt to reduce the feed
costs. Instead of corn, barley, wheat and soya bean oil meal – the standard
components of concentrate mixtures - there are the ground horse bean
seeds and peas (to 20%), sweet lupin (to 15%), rape seed and its derivative
feeds (by-products) (10-23%), dried beet pulp (to 35%), dried grass, rye,
CCM (corn cob mix) and many other components could be successfully
introduced into geese diets. The dried grass can be applied up to 35-40% of
mixture, but above 30% the laying rate and hatching of eggs could be affect-
ed. For young goslings the share of dried grass should not exceed 20% of
mixture. High contents of saponins in dried green matter from legumes,
reaching 1% (e.g. in dried Lucerne), can decrease the geese productivity and
the hatching of eggs. The share of dried Lucerne in the diet must be signif-
icantly lower than the amount of dried grass. The goose, like no other
species of poultry, well utilizes the green matter from pasture effectively.
After the reproductive season i.e. in the period of rest, the geese can use the
pasture, eating 500-700, even 1000 g of green matter. It also willingly con-
sumes other green feeds, such as - winter wheat, rye grass from durable
grasslands, clover, Lucerne. After harvest geese perfectly search for the crop
residues. At the same time geese have excellent freedom of movement.
Extensive keeping of birds in the autumn months to a great extent reduces
feeding expenditures. During the 4-month long pasture season, an average
of 120-150 m2 per one goose should be planned, depending on the kind of
pasture and its efficiency.
During of the 5-6 month-long of reproductive period the adult goose
eats a total of 65-75 kg of feed mixtures. The bird can excrete 1.3-4.0, even
4.5 g of nitrogen daily, which in a year-long scale amounts to 260-1500 g N
per one bird. The amount of unutilised, excreted nutrients is strictly
dependent on the composition of diets or mixtures.
It is possible to utilize the droppings of goose for feeding purposes.
The amount of protein in 1 kg dry matter of excrements may vary from 14
- up to 28%. In the arid form obtained from the drying facilities they can be
used as a good component of mixtures for ruminants and other animals.
However, because of sanitary consideration such utilization of droppings is
721
prohibited in EU countries.
There is no doubt that the goose is one of these bird species, which
are useful especially in the “ecological” backyard poultry keeping. This bird
gives the chance of applying and using different cheap feedstuffs, produced
in every agricultural farm.
22.2.3.9. Feeding of young broiler and fattening geese
Currently the raising of young broiler-geese up to 9-10 weeks of life is rarely

practiced because of the unprofitable relationship between feed prices and
the final product. The goslings should receive complete mixtures with care-
fully balanced composition and contents of the protein amounting to 22-
20% (during 1-3 weeks of life). In the second rearing phase the quantity of
this component is being gradually lowered to 17.5% (4-8 week) and to 16-
15% during the last weeks of raising. The energy value of mixtures should
not be higher than 11.7-11.9 MJ ME/kg, the quantity of Ca should be low-
ered from 1.0 to 0.6 %, P should vary between 0.6-0.5%; vita. A lowered
from 10000 to 5500 IU/kg, wit. D3 from 1500 to 500 IU/kg. Young geese
are sensitive to the deficit of the niacin, cholin and vitamin B2 deficiencies.
The coccidiostatics should not be applied in raising young and fattened
geese.
Intensively fed young fattened geese are kept for 15-16 weeks. To the
fourth week of life the birds should be fed on diets containing 19-20% of
protein (11.3-11.4 MJ/kg, given ad libitum), during the next 5-12 week - 17-
16%, of protein and from 13th week until the end of rearing period about
15% of protein. From the 3rd-6th week geese can consume green matter of
very good quality. In 7-8 week intake of green matter varied between 500-
1000 g/d/head. At this period the daily intake of mixtures reaches 200-230
g/d/head. In the 3rd dietary period the geese eat about 1000-1500 g green
matter and in the 14th-16th week only barley and the oat (to 300 g), in the
last week - merely oat (to 550 g/d/head). The green matter can be partly
replaced by carrot.
The total intake of green matter in rearing up to 14th-15th week of life
reaches 65 kg/head. Feathers plucking occur in the12th week and the peri-
odic application of premixes containing methionine – a valuable component
for the regeneration and the re-growth of pens - is recommended
Application of fodder yeast, enzyme preparations containing carbohy-
drases (especially cellulase and hemicellulase – used when large quantities
of barley and oats are applied), sprouted cereal grain that stimulates the
appetite is particularly profitable, because during the last few days of the
fattening period the geese are not active and show weaker appetite.
22..3.10. Production of fatty livers (foie gras)
722
Fattening of geese for fatty livers is banned in the majority of EU countries;

however some countries still permit this kind of production. For fattening of
this kind well developed geese, usually at the age of 4 months, are used.
However, 12-week-old birds are also useful animal material. Formerly
applied intensive fattening through 5-6 weeks, was shortened to a maxi-
mum of 3 weeks (depending on age, shape of birds and the degree of somat-
ic development). Initially 300-500 g of corn is being given to geese, and then
it is increased up to about 800 g.
In order to improve the digestibility of carbohydrate compounds of
corn grain, it is thermally prepared. There are also supplements of the salt,
enzymes and yeast applied. The quantity of fat in the fatty liver exceeds
40%, and its mass reaches 500-700 g depending on the length of the fat-
tening period. Final parameters of the mass and the composition of the liver
depend on the kind of feeds given to the birds (small supplement of animal
origin feeds is being applied in order to deliver the structural proteins), on
the age of geese and on the length of fattening period.
2.4. Principles of ducks nutrition
The digestive tract of ducks in relation to the length of the body is signifi-
cantly shorter in the comparison with goose (5-6:1). In the rate of passing
of the content and the degree of nutrients digestion the considerable simi-
larity is found between the young duck and chicken, however the digestive
system of adult duck is more similar to the goose. The ducklings not always
better than chickens or goslings digest structural components, although in
the caecum of ducks there are greater quantities of short chained fatty acid
(SCFA) formed. FIGURE XII-6 shows the influence of energy concentration
(practically: fibre level)on feed utilization. Value of metabolisable energy of
feeds used in poultry feeding, determined mostly for hens, can be estimat-
ed for ducks, according to different authors, 5-6% higher than for chickens.
FIGURE XII-6:
Feed conversion
efficiency in
function of ME-
density at the
growing duck
(TAMAS, 1995)
723
In the feeding of ducks the specific feature of this species - the high-
est gain rate - should be taken into consideration. Ducklings in 5-7 day of
life doubled the body weight determined directly after hatching. Young fat-
tened ducks easily becomes overfattened, which should be considered in
balancing the energy in diets or mixtures.
22.2.4.1. Feeding of ducklings at rearing
The duckling’s requirement for nutrients depends on their genotype, future

use, the system of management and on many other factors. Substantial dif-
ferences are found in the genotype; therefore ducklings of different species
(duck, Muscovy, their F1 generation) should be reared separately.
The Muscovy ducks requirement for nutrients is higher in all stages
of rearing. It is especially high for the protein and for the exogenic amino
acids (Table XXII-16). It results mainly from the better muscles mass of
these ducks. In the subsequent phase of rearing it is possible to give good
quality green matter, carrot, although it requires the changes in the com-
position of mixtures given to ducks.
Specific feature of growing ducklings is their high requirement for the
niacin - nicotinic acid. For Pekin ducks the dosage of 70-50 mg/kg is being
recommended. For Muscovy ducks these recommendations are quite gener-
al and determine the needs for niacin at the level of 25 mg/kg. The weak-
ness of legs, sitting of ducklings on legs - typical symptoms of the niacin
deficit are often mistakenly treated as a symptoms of rickets, in course of
which the level of Ca, P and vita. D3 in the diet is being increased, what in
turn is intensifying the symptoms of niacin deficit. The ducklings that have
problems in movement or are sitting down on jumps should be fed with mix-
tures containing yeast, good quality green feed or supplement of the vitamin
PP. The rate of growth of ducklings should be strictly controlled and should
be consistent with recommendations of the producer for the genetic materi-
al.
724
Table XXII-16: Chosen data regarding the requirement of ducks (1 kg feed, 88 % dry
matter) (The Polish Standards of Poultry Feeding) and NRC (1994)
2.4.2. Breeding ducks nutrition

The basic principles of breeding ducks and geese are similar. Between 25-
27 weeks of life the duck start the period of reproduction. Depending on the
laying phase and the production level 16-18% of protein and energy densi-
ty (concentration) from 11.3 to 11.9 MJ ME/kg mixtures are recommended.
At high laying the level of Ca should reach 3.0-3.1% and available phos-
phorus should vary between 0.28-0.45%.
Within a period of rest or moulting the quantity of the crude protein
should be reduced to 11-12%, level of calcium to 1.0-1.2% (Table XXII-17).
Table XXII-17 . Feeding of the meat-type reproductive ducks
725
In 1 kg: vit. A 5.000-10.000 IU; D3 1.250-2.500 IU; vit. E 12.5-25 mg; vit.
K 1-2 mg; choline 400-800 mg; B2 2.7-5.5 mg; panthotenic acid 2.5-5.0
mg; B12 0.07-0.014 mg, folic acid 0.2-0.5 mg; biotin 0.12-0.25 mg; niacin
25-50 mg; B1 1-2.0 mg; B6 1.5-3.0 mg; Mn 30-60 mg; Fe 46-80 mg; Cu 4-
8 mg; Zn 30-60 mg; Se 0.15-0.3 mg; I 0.2-0.4 mg
22.2.4.3. Feeding of fattened ducks
Fast-growing fattened ducks need to be intensively fed with precisely bal-

anced mixtures during the first three weeks of life (Table XXII-18). The duck-
lings are very sensitive to the quality of protein thus applying of feeds of ani-
mal origin (fish meal) that are rich in exogenic amino acids or supplemen-
tation with the pure amino acids is recommended. An important condition
for good feather development in ducks is the high level of sulphur amino
acids in mixtures. The ducklings for intensive growth require a particularly
large amount of vitamin A (8.000-10.000 IU/kg). In case of its deficiency the
lubrication of duckling eyes can be observed. Other necessary vitamins are
D3 (about 2.500 IU/kg), niacin (60 mg/kg) and some biologically active sub-
stances. They react faster than chickens or goslings on deficits of vitamins
or mineral components.
Young Muscovy males grow remarkably fast and their requirement for
nutrients is found to be about 5-8% higher than that of females or Pekin
ducks as a species.
Similar levels of the feeding are required for the interspecies cross-
breeds (Carina moschata L. – Muscovy male x Anas drake platyrhynchos L.
– Pekin duck female), that are characterized by a good muscle mass and
smaller fat accumulation than Pekin ducks.
Table XXII-18: The variants of broiler ducks feeding (JEROCH, 1998)
726
In 1 kg: vit. A 8.000 IU; D3 2.500 IU; E 20 mg; K 1,5 mg; choline
800 mg; B2 4 mg, panthotenic acid; 12 mg; B12 0,01 mg; folic acid 0,5
mg; niacin 60 mg; B1 2 mg; Mn 60 mg; Fe 80 mg; Cu 8 mg; Zn 60 mg; Se
0,2 mg; J 0,4 mg.
22.2.4.4. Practical systems of the ducks feeding

The majority of flocks of ducks are fed on complete mixtures. Breeding
flocks in the resting period and ducklings in semi-intensive fattening can
receive green fodders (grass, Lucerne, clover), carrot, crumbled beets,
steamed potatoes and feeds, which are usually applied for hens and geese.
The duck can eat, depending on age, 80-250 g of succulent feeds, 200
or even 300 g of the mixture or different concentrates (mixture of the cere-
al grain, leguminous seeds). Because of the ban on feeding ducks for fatty
liver, in our opinion, the information given in one of the previous chapters
is enough and give substantial information on this kind of use and feeding
of waterfowl.
727
CHAPTER XXII: POULTRY NUTRITION AND DIETICS
22.2.5. Feeding and nutrition of the ostrich

The first ostrich farm of the World was established in South-Afrika, mainly
to produce ornamental feathers for the theatre and music-hall dancers.
Ostrich skin is equivalent to that of lizard suitable for producing hand bags,
boots and footwear. The meat of the ostrich contains 19 to 20% protein and
only 1 to 3% fat, which is very suitable for manufacturing well preservable
smoke-cured meat produces. More breeds exists, but the so-called blue-
necked is the most popular.
Interestingly enough, the digestive characteristics and nutrient
requirements of the ostrich partly resemble to that of the turkey, partly
those of the goose. Notwithstanding that there is no caecum in the digestive
tract, the fibre digestion of the adult bird is very good, like in the goose. The
growth rate of chicks is very high with a turkey-like protein, mineral and
vitamin requirements. The ostrich’s egg, relative to the live weight is small,
weighing 0.9 to 1.5 kg. In the daytime eposited eggs are hatched by the
cocks, from the afternoon till the next morning the hens is sitting on them.
The first laying period is of low productivity (only 10 to 12 eggs), which con-
tinuously increases from cycle to cycle and in the sixth-seventh year achieve
the top egg production with up to 70 eggs. The emerge of the “newborn”
birdie from the egg is very difficult, being the egg-shell very strong, this is
why the hatching should be supervised and if necessary, helped. This is the
first slaughter possibility: gourmet carcass.
After the successful hatched out, the one-day-old chick of 30 cm
height can be fed on turkey grower I first in crumbled, later in small pellet
form till the age of 12 weeks. Ostrich chicks achieve a live weight of 15 kg
at the age of 3 months. The intensive growth of the first 12 weeks requires
high protein diets (18 to 22% of crude protein). The energy concentration
may be low (approximately 10 MJ ME/kg) because the fibre utilization of
birds continuously increases. The feed conversion efficiency in this period
is very good, with a value of 1.8. The calcium and phosphorous require-
ments are 20% higher than in case of the growing turkeys, because of the
very intensive growth and the high amount produced bone. The magne-
sium, vitamin D and biotin needs are higher, compared to the other broiler
birds. Vitamin C supplementation is supposed to be beneficial.
728
After the age of 12 weeks more and more cheap roughage can be fed:
first chopped green grass and alfalfa, whole corn plant, corn stalk and final-
ly even straw of good hygienic quality. This basic feed should be completed
by/with 1 to 2 kg of corn grain or cereal concentrate in form of large pellets
(diameter: 16 mm) till the slaugher.
At the age of one year the breeding animals are selected and the
remaining birds are slaughtered with a live weight of 100 to 130 kg. The
breeding maturity is achieved at the age of 18 to 30 months. Breeding birds
are living in “trio”, one cock with two hens. Adult live weight of hens is 110
to 175 kg, that of the cocks 300 kg. The were able to adapt to the European
climate, the cold weather does not mean a great stress unlike the high air
humidity.
729
22.3. The most important digestive and metabolic

diseases of poultry
ACUTE DEATH SYNDROME (FLIP-OVERS)

It is the disease of intensively growing, 3-4-week old growing chicken and
turkey. Death is caused by the breakdown of heart functions and the col-
lapse of the blood circulation. Dead birds can be found on supine position.
The pathological findings are not characteristic. More cases can be found in
males than in females. Among the predisposing factors are considered the
high, easy fermentable carbohydrates (especially sugars), ionophore coc-
cidiostats and heat stress.
In growers and layers, acid-base disturbances caused by heat stress
highlighted a role for accumulation of plasma lactate as an extrarenal buffer
to combat severe alkalosis. However an increase in blood lactic acid dam-
ages the cardiac system and could predispose broilers to sudden death syn-
drome. Some day-long feed restriction (minus 10-15%) around the day 20
of the age may moderate mortality, and the losses in weight gain can be
compensated until slaughtering.
AORTIC RUPTURE OF TURKEY

A disease of the fast growing (12-16 weeks old) turkey of susceptible genet-
ic line. Affected birds die from exsanguinations into their body. Besides
genotype and gender (toms are more prone ) too high a protein and fat level
in feed during the first 3 months of age, and lack of copper seems to be
among the main factors causing the disease. The latter, being active part of
the lysosyl oxidase enzyme, is essential in the formation of collagen and
elastic fibres of vessels’ wall. Toxic constituent of the vetch (Lathyrus odor-
atus), the beta-aminopropionitril has very similar damaging effect on the
vessel’s wall. High blood pressure facilitate the manifestation of the ailment.
ASCITES SYNDROME (OEDEMA DISEASE)

The syndromes have been described in intensively developing broiler chick-
ens. It is basically a cardiopulmonary failure, where owing to the high
venous pressure large amounts (up to 500 ml) of serum-containing liquid
enter the abdominal cavity of the affected birds. The right ventricle of heart
is enlarged. The main cause is the poor oxygenation of the tissues, which
may derive from several factors, like keeping, genotype, relatively small lung
capacity etc.
The selection for high protein accretion and good feed conversion may
also facilitate the outbreak. Of the nutritional factors one have to mention
the high energy concentration and protein to energy ratio (P/E), as well as
luxury consumption due to pelleted form of the feed mixture. Each of them
730
enhances protein synthesis and the oxygen needs of tissues. These circum-
stances may be aggravated by the unfavourable microclimate of the stall
like increased ammonia or carbon dioxide concentration of the air. Males,
having higher protein content in their body, are more susceptible. High salt
level or the presence of mycotoxin in the feed may facilitate the outburst.
In endangered flocks all factors should be eliminated, which may
inhibit the balanced corporal growth and protein retention. Thus, use mash
form, energy-low protein diet, consider limit.time feeding and keep the level
below 0.475% (LEESON et al. 1995)
CAGE LAYER FATIGUE

Osteoparalysis in battery keeping is caused by disorders of calcium metab-
olism. It is often accompanied by laying eggs of thin eggshell. The numeri-
cal egg production and the plasma calcium level generally are maintained
but the ash content of bones drastically decreases which may lead to spon-
taneous fractures. The increased calcium supplementation itself is not suf-
ficient for recovery, but together with extra ascorbic acid intake – by assur-
ing the activity of vitamin D and increasing the bone’s mineralization – may
often help.
EARLY MORTALITY OF ONE-DAY CHICKEN

High percentage mortality of the first week of life IS generally not infectious,
but it is due to inadequate micro element and vitamin supply of the parent’s
generation, because the one-day-old chicks meet the majority of their
requirements from the yolk (vitellus).
GOOSE STRESS LAMENESS SYNDROME

Under practical farming condition, one-day-old goslings and growing geese
suffer multiple stressors like overpopulation (crowdedness, cold, hyperven-
tilated stall, wet bedding, too dry or humid air, increased ammonia concen-
tration, troubles in water and feed distribution, meteorological fronts,
serum application, vaccinations, plucking of feather, heat stress etc.). This
physiological state would require more ascorbic acid as that of the capacity
of endogenous synthesis in kidneys.
The relative vitamin C deficiency retards growth, impairs immune
functions and causes fatty infiltration in fibroblast cells of collagenous
fibres of conjunctive tissue. The lack of formation of fibroblasts’ filaments
and their polymerisation primarily occur in the Achilles tendon. The tendon
will degrade and as a clinical sign, a special lameness develops: the walking
becomes uncertain and painful, the tarsal joint cannot be bent, and the
birds try to keep balance of their body by the flapping of the wings and they
lean on their beaks for support. The treatment is possible only in the early
stages, and is not enough to supplement feed with ascorbic acid (1-10
mg/kg LW/day for a week, but also the stressors should be eliminated.
731
FIBRE DEFICIENCY AT GEESE

Gees are partly water, partly grazing herbivore birds. Fresh grass intake of
an adult goose may achieve 2 kg per day. which is equivalent to 100-140
gram crude fibre. The utilisation of nutrients in dry form is lower 50% than
in fresh form.
These birds require the fibre for the optimum functioning of gizzard
and caecum, whereas the real energy, gained from the caecal fibre fermen-
tation is relatively small. The minimum crude fibre needs are between 10 to
12% in air-dry matter; the commercial goose feeds, in contrast, has an aver-
age of 6% crude fibre. Lack of fibre – especially during the growing phase –
causes the so-called “bulky feed hunger” (“desire for indigestible organic
matter”) syndrome, which is a special form of stress. Birds become nerv-
ous, vitamin C requirement may increase and the development and function
of caeca is disturbed. First an atrophy occurs, which, in turn, may lead to
the gees eating the bedding, to a subsequent inflammation of the inner
mucous layer of the caecal lumen. Across the inflamed walls of vessels plas-
ma enters the lumen, which results in a fibrinous (cheesy) inflammation of
the caeca (“fibrinous typhlitis.”). The absorbed degradation products of the
fibrin act as antigens, inducing the production of antibodies and the devel-
opment of an amyloidosis. The only way of prevention is to assure a suffi-
cient fibre supply from the first days of life.
HINDGUT ACIDOSIS
Metabolic acidosis damages the gut wall reducing the feed conversion effi-
ciency. Metabolic acidosis restricts calcium absorption from the small intes-
tine and increase osteoclastic function causing bone resorption. In layers
the kidneys stabilize pH by adjusting cation and anion excretion. Acidosis
inhibits carbonate formation and vitamin D activation and increases bone
solubility and urinary calcium. Eggshell formation involves severe metabolic
acidosis, which may be partially buffered by the P fraction of orthophosphate.
Accumulation of lactic acid occurs in the hindgut of layer type birds
when they are exposed to new (novel) single-cereal-based feeds. This accu-
mulation is influenced by the quantity of feed consumed at different phas-
es of the production cycle. Critical periods: a greater “meal” feeding pattern
may occur after the dark period with birds not eating until after oviposition.
Feed intake also can increase rapidly at post-point of lay and at peak pro-
duction, which may involve maladaptation of the gut to larger feed volume
in the short-term. This is certainly a consideration in replacement pullet
systems where the birds, after long-term heavy feed restriction are exposed
to larger quantities of feed (or ad libitum) than they have been conditioned
to eat.
Any adaptation to a higher starch loading (e.g. whole grain inclusion
in the diet) requires a relatively long period of gut adaptation, sometimes as
732
long as three weeks. The total organic acid load (the VFA’s plus lactic acid)
is higher on ground grain diet than on whole one. Simple measurement of
pH change in fresh excreta may allow to be monitored of complex alteration
in digestive process and microbial activity. (The pH of the healthy broiler
chicken’s excreta between 6.5 and 7.3).
Furthermore, any increased water intake, i.e. due to heat stress or
disease, may result in a similar increase in fermentation metabolites as
digesta transit time may be reduced.
MALABSORPTION SYNDROME (STUNTING SYNDROME, HELICOPTER DISESE)

Malabsorption syndrome consists of growth depression, diarrhoea, pale-
ness, leg weakness and incomplete feather development of broiler chicken
and sometimes of turkey. The diseases basically have viral origin and the
first clinical signs at day 4-7 can be seen. The crop of the affected birds is
enlarged; they eat and drink a lot, which is followed by diarrhoea. Till week
3-4 the sick birds – owing to the unequal growth of their feather – have a
helicopter appearance.
The dietary, predisposing factor is the defective transition from the
own vitellus-supply to the external feed. The digestive enzyme activity is
insufficient, and the development of an active immunity is slow. Bile acid
concentration in the digesta is also diluted, possibly impairing lipid absorp-
tion. The decreased bile concentration is associated with an increase in
luminal aerobic and anaerobic bacteria.
Clinically, malabsorption is the output of voluminous, bulky faeces
with increased osmolarity owing to unabsorbed nutrients (mostly excessive
fat). It occurs because of defective intraluminal digestion due to ineffective
enzymes or lack of enzymes. Defective absorption could be accompanied
with the loss of mature enterocytes that have been replaced with immature
cells lacking full absorptive function. Again, NSP may be involved with mal-
absorption because gelatinous luminal content blocks the access of
enzymes to otherwise digestible nutrients. This problem is also associated
with increased mitotic activity and enlarged crypt depth in the gut mucosa,
indicating an increased turnover of enterocytes and a relative increase in
secretory enterocytes or a decrease of mature enterocytes.
Extra vitamin supply in the drinking water and use of antibiotics of
broad spectre moderate economical damage. Among the most obvious non-
infectious factors are the high tannic acid, trypsine inhibitor, nitrite-nitrate,
biogenic amines and mycotoxin content of the feed, or the too high salt level.
Heat stress also decreases the absorption of nutrients, as well as the over-
feeding after feed restriction (rationing), because in the latter case the diges-
ta transit time accelerates. Infectious and non-infectious causes may com-
bine with each other.
733
OSMOTIC DIARRHOES
It involves excessive osmotic forces exerted by luminal solutes. Poultry diets
high in salt is one of the causes andmay occur with diets formulated with
lots of bakery by-product meal. Osmotic factors may be involved in diges-
tive problems associated antinutritional factors (non-starch polysaccha-
rides, NSP) in barley, rye and wheat and leguminous seeds. These complex
carbohydrates, typically hexoses and pentoses, are resistant to digestive
enzymes, create a viscous environment within the intestinal lumen,
increase the mass of luminal digesta and produce moist sticky droppings.
(For more details see Chapter VIII).
SKELETAL DISORDERS
The intensively growing broiler chicken and especially the turkey is loaded
with many skeletal disorders: rickets, juvenile osteoporosis, perosis (FIG-
URE XXII-5), tibial dyschondroplasia and torsion of the tibia. In the back-
ground of these diseases there are genetic predispositions of the too high a
growth rate and the primary or secondary lack of specific (micro)nutrients
(Mn, niacin, folic acid, cholin, biotin). Eating of feedstuffs wit antinutritive
substances (soybean, rapeseed, sorghum) may also facilitate the manifesta-
tion of these skeletal troubles.
FIGURE XXII-7: Perosis in growing turkeys (© FEKETE S.Gy. 1998)
Practically a perosis-like lameness may occur in growing turkey and

caused by zinc deficiency (the basic problem is the degeneration of the tibio-
734
tarsal chondrocytes), but – opposed to the perosis - the Achilles tendon typ-
ically slip out of the condyle on the medial side.
FATTY LIVER HAEMORRHAGIC SYNDROME (FLHS)

It is a manifestation of a disease of high producing laying hens, mostly
housed in cages. The functional trouble of the liver cells results in fat accu-
mulation in the hepatocytes and subcapsular haemorrhages.
Pathogenesis: the first sign of it is the significant (30-40%) drop in egg
production. The collapse of blood circulation and internal haemorrhages
may lead to death. After obesity becomes excessive with fatty infiltration of
the liver and fat accumulation in the abdominal cavity occur. The basic
cause is the intensified de novo lipid synthesis of the liver. Rations rich in
carbohydrate (elevated insulin concentration in the blood) and the oestro-
gen in blood related to egg production facilitate the process. The poor phys-
ical activity in cage, in its turn, inhibits the catabolic hormone secretion of
the adrenal cortex. Significant improvement can be achieved by a partial
replacement of carbohydrates by oils (soybean, sunflower, rapeseed or corn
oil) in the ration.
The FATTY LIVER AND KIDNEY DISEASE (FLKD) is an ailment of young
chickens, caused by biotin and cholin deficiency. It is characterised by
swollen, pale and friable fatty liver with or without haemorrhages,
hydropericardium and pale swollen kidneys. Field outbreaks are most com-
mon between 10 days and 4 weeks of age and the mortality varies from 2
to10%. The addition of 100 mg biotin and 19 mg cholin for 1000 birds in
drinking water for some days will prevent new cases appearing.
ABOUT VITAMIN DEFICIENCIES BRIEFLY

Vitamin A deficiency causes the keratinisation of pharyngeal epitheli-
um and small white pustules in the mouth, pharynx and oesophagus.
Clinical signs include lacrymation, conjunctivitis, accumulation of cheesy
material in the conjunctival sac, mouth and oesophagus, in severe cases
hyperkeratosis of corneal epithelium resulting in blindness. A serous dis-
charge is also seen from the nostril
The kidneys are pale, the tubules and ureters are distended with var-
ious amount of urates. The urate deposits may be found on other visceral
organs, too. Decrease of the excretion capacity in pullets, young laying hens
and birds of prey causes visceral gout (BOKORI, 1955). The process is facili-
tated by the (partly) portal venous circulation of the bird’s kidney, and also
the allergic effect of some soybean causing nephrosis (CSAKY et al, 2004).
The first sign of the gout in growing Canaries is the distorsion of legs. Lack
of vitamin A in young cockatoo, parrots and budgerigar makes the birds
susceptible to candidiasis. In birds of prey kept in captivity, the third eyelid
become opalescent, grey-white is the first clinical sign of vitamin A defi-
ciency.
735
Vitamin D deficiency in rapid growing birds exhibits signs of retarded

growth and rickets: the beak, claws and bones become soft. In hens there
is an increase in thin shelled eggs, followed by the decrease in egg produc-
tion. Affected hens tend to sit like a penguin and the sternum may be bent
and ribs turned inwards (“S”-shaped sternum). There is a poor calcification
in chickens, especially at the epiphyses of tibia and femur, the parathyroid
glands are enlarged. In laying hens well defined knobs appear at the costo-
chondral junction (painless hard swellings on limb joints and ribs: “rickety
rosary”) and the sternum may show lateral bending. In birds of prey in cap-
tivity, fed predominantly with meat, the legs will become thin and longer,
bending inwards, owing to vitamin D and calcium deficiency (osteomalatia
in growing and osteoporosis in adult individuals).
The vitamin E and selenium deficiency will cause exsudative diathesis
(oedema of subcutaneous tissues associated with abnormal permeability of
capillary walls), pancreas necrosis. As a secondary effect, the lack of antiox-
idant protection encephalomalacia develop in 2-4-week old chicken, char-
acterised by ataxia, backward or downward retraction of the head (“crazy
chick disease”). On necropsy, the cerebellum shows hyperplasia and
petechial haemorrhages and necrosis on the surface, with oedematous
meningies. (Clinically it may be similar to the carriage of the head seen in
Mn deficiency of one-day-old chicks, but in the background the incomplete
development of the neck muscles is to be found. In chicken and turkey, if
vitamin E deficiency is accompanied by deficiency of sulphur amino acids,
lesions may appear in breast and thigh muscles (light coloured streaks
showing the hyaline degeneration. In growing ostrich fed one-sidedly by
alfalfa pellets of corn meal myodegeneration may also develop in the mass
muscle, similar to the lambs’ “white muscle disease”.
It should be emphesised that the deep breast muscle myopathy
(Oregon disease or green muscle disease) of turkey is not a vitamin E defi-
ciency syndrome, because it is caused by the intense and frequent flapping
of the wings. In ducks the muscles of the gizzard will suffer from myode-
generation.
Vitamin K deficiency leads to an anaemic appearance (pale comb, ball)
caused by the haemorrhages in various organs, including subcutaneous tis-
sues, muscles and internal organs. The prolonged administration of bacte-
rocidal chemical may destroy intestinal bacteria responsible for vitamin K
synthesis. Since vitamin K is fat-soluble, there is no storage of vitamin K3,
and thus the continuous supply is indispensable. The increased blood-clot-
ting time and the haemorrhages in the subcutaneous tissues make it nec-
essary to differentiate it from the Newcastle disease and some mycotoxico-
sis.
The relative deficiency of ascorbic acid (vitamin C) (heat stress, bat-
tery keeping system) makes the bird susceptible to tibial dyschondroplasia.
The reason is that the activation of vitamin D – and its mineralization, too
736
– may be facilitated by ascorbic acid. A special deficiency syndrome of vita-

min C and calcium can be regarded as “Cage Layer Fatigue” and the “Goose
Stress Lameness” (see above).
The clinical signs of thiamine (B1) deficiency may be characterised by
opisthotonus, polyneuropathy, cyanosis, bradycardia and paleness.
Chickens sit on their flexed legs drawing back their heads. Atrophy of gen-
ital organa also occurs. Inadequate supply of one-sided meat feeding (birds
of prey) is one of the most frequent causes. Besides the described symp-
toms, prey birds turn the neck round or keep it loxotic.
Lack of riboflavin (B2) through the lesions of the peripheral nerves
may cause a “curled toe paralysis”. The clinical signs include the chickens
sitting on the hock keeping their toes bended (FIGURE XXII-8).
FIGURE XXII.-8: Typical, curled- toe’ on growing chick (© KATONA-ZSOMBOR, 2008)
At more developed stages, the chickens lie down with extended legs,
swinging flaccid and droping the wings. In case of slight B2-deficiency
chicks withdraw their necks, keep their swings loose and their tail feather
are directed towards the soil. After diagnosing subclinical riboflavin defi-
ciency in chicken and turkey it may be remedied by giving the vitamin and
measuring the glutathione-reductase activity. If the treatment increases
enzyme activity the deficiency can be confirmed. Feeding cereals exclusive-
ly, especially in cleaned, hulled form, may lead to outbreaks. (It should be
differentiated from the “hawk paralysis” of birds of prey, kept on a badly
formed roost, without sufficient physical activity.) The paralysis of 2-3-week
old chicken are caused by oedema and secondary degeneration (demyelini-
sation) in the peripheral nerves (nervus ischiadicus and nervus brachialis)
between the neurofibrils, as well as in the myelin sheath of axons and in the
737
nucleus of Schwann cells without any inflammatory alteration (GLAVITS et al.

1994). The embryos have defective downs referred to as “clubbed”.
The pyridoxine (B6) has an important role to play in transamination
and its absence decreases the oncotic pressure of blood. One of the first
clinical signs is the inflamed oedema of the eyelid, rough, deficient plumage
and extrapyramidal symptoms (weakness, inco-ordination of movements,
with heads lowered and droopy wings). Lack or low availability of niacin
(nicotinic acid) is shown by poor feathering and squamous dermatitis (first
on the axillar region), inflammation of the buccal cavity, including the
tongue (the tip of the tongue may appear black) and diarrhoea. It may occur
on corn and wheat based diet.
The pantothenic acid deficiency causes faded, rough and brittle
plumage. It may aggravate the clinical manifestation of the genetic failure,
the French moult of growing parrots and budgerigars. Dermatitis of descen-
dent type develops with inflammatory changes at the corner of the beak, at
eye-lids and partly on the toes, crusty scab like lesions, eyelids may be
sealed together. In contrast, dermatitits of descendent type, starts with
encrustation and cracking, developing into fissures. (Similar symptoms can
be caused by severe zinc deficiency too.) The occurrence of biotin deficien-
cy is possible, because its availability in the cereals is low and it is sensitive
to the oxidising effect of rancid fats. Lack of biotin in fast growing chicken
and turkey may cause perosis.
The perosis is a complex syndrome, meaning many development trou-
bles of the bones. Owing to an incomplete cartilage synthesis the otherwise
normal bones suffer abnormal changes. The longitudinal growth of the long
tubular bones become unequal, the tibia bends inwards or outwards, or
undergo spiral torsion. Changes and enlargement of the tibiotarsal (hock)
joint may cause the displacement of the Achilles tendon (owing to the later-
al distorsion, the tendon of the gastrocnemius muscle frequently slips from
the medial condyle: “sliped tendon, enlarged hock disorder”, in German:
“Fersenkrankheit”). Besides biotin deficiency manganese, cholin, niacin and
folic acid deficiency also often give rise to these symptoms.
SHORTLY ABOUT MYCOTOXICOSIS

The target organs of the most important mycotoxins are the liver, spleen,
bone marrow, mucous membranes, kidneys. and the genitali.Toxins, pro-
duced by the fungi Aspergillus flavus and parasiticus (aflatoxin B1, B2, G1
és G2) can primarily cause ailments in water fowls. In the aflatoxicosis the
main site of the damages is the liver. In acute form of the disease the liver
has a distinct yellow colour with haemorrhages on the surface. The struc-
ture of the liver suffers disintegration and in the hepatocytes vacuolar
degeneration and necrosis can be seen. In subacut cases the proliferation
of bile vessels and nodular hyperplasia of the hepatic cells can be seen.
738
All poultry species are susceptible to the damaging effect of F–2 tox-
ins (zearalenon) – except the domestic fowl –. Due to the effect of F–2 fusar-
iotoxicosis the egg production will not change, but the sperm production
and the spermatogenesis are affected. The T–2 toxin (and related other tri-
chothecene skeleton toxins. the HT–2 toxin, DAS, NIV, FX, DON) cause
growth depression, necrosis covered with crusted deposition on the beak, on
the skin in the angle of the mouth, on the tongue and palate. In acute cases
haemorrhagic hepatic dystrophy and tubulonephrosis as well as atrophy of
lymphoid organs (like bursa Fabricii, thymus and gut wall) can occur. In the
tissues strong lymphocyte depletion occurs. Egg production is affected even
by the intake of feed with relatively low (0.5 mg/kg) toxin concentration.
Birds – unlike the mammals –. are rather resistant gainst the fumon-
isine mycotoxin produced by Fusarium moniliforme.
The disease caused by Stachybotrys alternans (syn. S. atra and S.
chartarum), named stachybotryotoxicosis causes necrotic inflammation
on the ball, proventriculitis and enteritis, necrosis and depletion of cells of
lymphoid and myeloid-haematopooetic organs. Given the fact, that this
mould proliferates mostly on roughages and bedding straw, in poultry flock
the outbreak is rather rare. The Aspergillus and Penicillium species produce
ochratoxin A (and the less toxic ochratoxin B) may cause interstitial fibro-
sis and degeneration of the kidneys. The latter may lead to uricosis and vis-
ceral gout. Hatchability of eggs is affected by the intake of feed containing
as little as 1-2 mg/kg toxin. Higher doses develop skeletal changes similar
to rickets.
FOR FURTHER READING
Degussa (1992): Ideales Protein beim Broiler. Ein neues Konzept zur Optimierung der
Aminosäurenversorgung. Feedback Special,3/36.
DLG-Futterwerttabellen (1984): Aminosäurengehalte in Futtermitteln. DLG Frankfurt am
Main.
Elkin, R.G. (1987): A review of duck nutrition. Research World’s Poultry Sci. J.,43(2), 84-
106.
European Table of Energy Values for Poultry Feedstuffs, WPSA (1989): Spelderholt Centre
for Poultry Research, Beekbergen.
Farrell, D.J. and Stapleton, P. (1986): Duck Production Science and World Practice.
Univ.New England, Armidale
Firman, J.D. and Boling S.D. (1998): Ideal protein in turkeys. Poult. Sci. 77,105-110.
Glavits, R. and Salyi, G. (1998): Mycotocisoses in poultry. Poult. Intern. 37(12), 26-40.
Jamroz, D., Wiliczkiewicz, A., Orda, J. and Skorupińska, J. (1994): Ileale und postileale
Fermentation von Getreidekohlenhydraten bei Jungmastgeflügel. Wien.Tierärztl.Mschr.
81, 80-84.
Jamroz, D. (1992): Empfehlungen zur Gänseernährung. Wien.Tierärztl.Mschr.,79, 47-58.
739
Jankowski, J. and Faruga, A. (1996): A note on performance of medium type turkeys fed
on practical diets containing different levels of supplemental vitamins and trace min-
erals during the rearing stages. J. Anim. Feed Sci. 5, 157-162.
Jeroch, H. (1996): Faustzahlen zur Geflügelfütterung. Jahrbuch für die Geflügelfütterung
. Verlag Eugen Ulmer, Stuttgart.
Kirchgessner, M., Eder, K., Müller, H. L. and Jamroz, D. (1999): Zur energetischen
Bewertung von Nicht-Stärke-Polysacchariden beim Geflügel. J. Anim. Physiol. a.
Nutr. 81, 51-55.
Klasing, K.C. (1998): Comparative animal nutrition. CAB INTERNATIONAL. Oxon
Larbier, M and Leclerq, B. (1992-original): Nutrition and feeding of poultry, Ed. Wiseman,
J. (1994-English version). Nottingham University Press.
Leeson, S., Diaz, G. and Summers, J.D. (1996): Poultry metabolic disorders and mycotox-
ins. University Books. Guelph. Ontario.
Mikulski, D., Jankowski, J., Faruga, A. and Mikulska, M. (1997): The effect of enzyme
supplementation of triticale-barley feeds on fattening performance of turkeys. J.
Anim. Feed Sci. 6, 391-399.
Mineral Requirement s for Poultry. Mineral Requirements and Recommendations for
Growing Birds,1985. Working Group No.2 Nutrition, of WPSA Report, World Poultry
Sci. J. 41(3), 252-288.
National Research Council (1994): Nutritional Requirements of Poultry: 9th Ed. National
Academy of Sciences, Washington DC
Noble, D.O., Muir, F.V., Krueger, K.K. and Nestor, K.E. (1996): Effect of altering the early
protein level on males from two strains of commercial turkeys. Poult. Sci. 75, 1334-
1344.
Noy, Y., Frisch, Y., Rand, N. and Sklan O. (1994): Trace mineral requirements in turkeys.
World’s Poult. Sci. J. 50(3), 253-268.
Rivas, F.M. and Firman, J.D. (1994): The influence of energy and protein on turkeys dur-
ing the finisher period. J. Appl. Poult. Res. 3, 327-335.
Scott, M.L. and Dean, W.F. (1991): Nutrition and Management of Ducks. Cornell Univ.,
Ithaca, NY
Sen. J.L., Jeffrey, M.J. and Kerr, B.J. (1994): Influence of amino acid supplementation of
low protein diets and metabolizable energy feeding sequence on performance and car-
cass composition of toms. Poult. Sci. 73, 1867-1880.
Waldroup, P.W., Adams, M.H. and Waldroup, A.L. (1997): Evaluation of National
Research Council amino acid recommendations of Large White turkeys. Poult. Sci.
76, 711-720.
740
Chapter XXIII
RABBIT NUTRITION AND DIETETICS
23.1.Digestive physiological characteristics

23.1.1. Regulation of feed intake
23.1.2. Digestive functions in the caecum
23.1.3. Intake of the soft faeces (caecotrophy)
23.1.4. Role of caecotroph in the gastric carbohydrate fermen
tation
23.1.5. Digestive physiology of weaning
23.1.6. Weaning systems
23.1.7. Mineral metabolism
23.1.8. Vitamin requirements
23.2.1. Energy metabolism and nutrition in rabbit does
23.2.1.1.Special features of voluntary feed intake
23.2.1.2.Maintenance requirements
23.2.1.3. Growth requirements in young rabbits
23.2.1.4. Pregnancy requirements
23.2.1.5. Lactation requirements
23.2.1.6. Effect of nutritional strategies
23.2.7. Effect of management strategies
23.2.7.1. Parity order
23.2.7.2. Reproductive rhythm
23.2.7.3. Litter weaning age
23.2.2. Protein and amino acid nutrition
23.2.2.1. Protein requirement
23.2.2.2. Amino acid requirement
23.2.3. Minerals, vitamins and additives
23.2.4. Nutrition and feeding of breeding bucks
23.3. Practical rabbit nutrition
23.3.1.Optimum composition of feed mixtures
23.3.1.1. Supply of energy
23.3.1.2. Role of indigestible organic matter
23.3.1.4. Minerals and vitamins
23.3.1.5. Young females and breeding does
23.3.1.6. Consequences of the deviations from the
recommendation
23.3.2. Feeding practice
23.3.2.1. Form of feed mixture
23.2.2.2. Feeding technique, amount of daily ration
23.3.3. Characterization of important rabbit feedstuffs
23.4.1. Troubles with digestion
23.4.2. Feed toxicity
23.4.3. Mycotoxicosis
741
T
he raising of rabbits is dynamically spreading all over the world. This
fact is testified too not only by the size of stock and the increasing
amount of products (meat, wool, coat), but also by the number of
publications dealing comprehensively with the breeding and feeding of rab-
bits. The reason for this phenomenon are as follows: the meat of the domes-
tic rabbit is outstanding in terms of both dietetic effect and its chemical
composition and does not compete with man and pig as a consumer of valu-
able cereals and leguminous proteins, or with poultry consuming concen-
trated feeds. It was proved by comparative examination that the rabbit
incorporates more protein in its body than broilers do and that its feed pro-
tein utilization is also more favourable. Authors have shown that owing to
its extremely low cholesterol and sodium levels, rabbit meat may play an
important role in the prevention of vascular diseases. Nor do religious rules
prohibit its consumption, in general.
The species not only allows large-scale meat production, but it is also
quite suitable for organic, or backyard production (by housewives, old age
pensioners or schoolchildren. Coordinated by large-scale units, the two
types of breeding can fit favourably. Thanks to its excellent properties rab-
bits can convert roughage with high fibre content (including numerous non-
traditional feedstuffs and tropical plants) without the deterioration of the
quality of the carcass. Since the rabbit is biologically capable of living con-
tinuously in the stage of reproduction, each unit of the product carries con-
siderably less burden – the cost of maintenance, stock replacement – than
sheep or cattle.
Another characteristics that holds out good promise is the great
genetic variety of the species manifesting itself in the intensity of growth,
fertility, maternal ability, milk production, resistance to diseases, heat tol-
erance, etc., which are all prerequisites of the success of selection pro-
grammes aimed at improving these properties. In view of the above, the FAO
launched programmes in several developing countries to introduce or inten-
sify rabbit raising. At the same time, rabbit is also a wool (angora) and fur
(rex) producing animal, and it is also a preferred pet (especially the dwarf
breeds) and one of the most important subject of the in vivo teratological
studies.
742
CHAPTER XXIII: RABBIT NUTRITION AND DIETETICS
23.1. Digestive physiological characteristics

23.1.1. Regulation of feed intake

„Feed intake according to energy” is a characteristic feature of the rabbit. It
means, that between certain limits, the daily dry matter intake is deter-
mined by the actual energy need of the animal (LEBAS, 1975). When the
energy concentration (MJ DE/kg) of feed mixture decreases, the voluntary
dry matter intake decreases. (FIGURE VI-1) This is why rabbit can be fed ad
libitum in practise. This self-regulating capacity is still underdeveloped in
young animals and can be disturbed by some antinutritive substances of
feeds. Consumption of mycotoxin (T–2 containing) feed, and higher than
17oC environmental temperature also decrease feed intake.
For the sake of logical unity the regulation of caecotrophy should be
described here. It depends on three main factors, i.e. the stimulation of rec-
tal mechanoreceptors, the perception of the specific odour of the soft faeces,
and the inner motive determined by the blood level of metabolites and hor-
mones. For the latter we have only indirect evidences. Namely, in case of
energy deficiency of the organism the rabbit consumes the total quantity of
the produced caecotroph. During ad libitum feeding the dry matter intake
depends on the protein requirement of the animal or, more precisely, on the
protein and fibre level of the ration FEKETE and BOKORI, 1985). This means
that caecotrophy is higher if the feed mixture contains less protein and/or
more fibre (FIGURE XXIII-1). The production of the soft faeces is practical-
ly independent of the amount and composition of the ration. It is the com-
position of the caecotroph that varies according to the diet. The time of feed
intake is a regulating factor of drinking, kinetic activity and excretion of
urine and faeces.
FIGURE XXIII-1:Regulation of the caecotrop intake
743
The relative volume of the different sections of the digestive tract is different
from that of other domestic animals. From the functional point of view it is
worth mentioning that the rabbit has the largest stomach and caecum (36
and 43% of the total wet gastrointestinal content) among the monogastric
animals. The pH value varies in the different parts of the digestive tract: on
average, it is 2.0-2.2 in the stomach, 6.1 in the duodenum, 7.3 in the
jejunum, 7.2 in the ileum, 5.7-6.1 in the caecum and 6.1-6.5 in the colon.
The total germ count in the caecum is 109-11/g, in the colon and rec-
tum 108/g, and in the jejunum even less. Bacteria of the genus Bacteroides
occur in the largest number (102-109/g); the number of aerobic microbes is
0-102/g, that of anaerobic one 0-106/g. Lactobacilli and clostridia (0-103/g)
appear only owing to feeding. Originally rabbits do not contain E. coli in
their gut, but in most cases, according to the environment, it can be found.
In the caecum of the healthy rabbit in a concentration of 106/ml protozoa
and a rabbit-specific saccharomyces (Cyniclomyces guttulatus, 106/g) do
occur. According to the original composition of the intestinal flora, some
broad spectrum antibiotics (e.g. ampicillin, lincomycin) can disturb the bal-
ance, causing dysbacteriosis and subsequent mortality.(FIGURE XXIII-2)
FIGURE XXIII-2:Rabit digestive tract, with the pH conditions
Rabbits chew thoroughly; the chewing movement per minute is 12.

Caecotroph is swallowed in intact form. The degradation of the proteins
begins in the stomach under the influence of pepsin-hydrochloric acid com-
plex, which is capable of hydrolysing peptide bonds of most proteins, mucin
744
being one important exception. The microbial protein of the caecotroph can
be digested only after pre-treatment of a specific bacteriolytic factor, pre-
sumably equal to the colonic lysozyme which is incorporated in the soft fae-
ces and by means of the caecotroph it is transferred to the stomach. The
digestion of proteins continues in the small intestine owing to the activity of
pancreatic trypsine, chymotrypsine, and carboxypeptidases A and B. The
intestinal mucosa contains a wide range of aminopeptidases which com-
plete the process of protein digestion. Amino acids enter the rabbit jejunal
brush border membrane vesicle and later into the blood. The residue of
dietary protein and digestive enzymes gets into the caecum, where it will be
utilized by the microorganisms. The digestion and absorption of lipids take
place similarly to other monogastric animals by means of pancreatic lipase,
sterol ester hydrolase, phospholipase A, bile salts and a co-lipase. Bile acids
are excreted in a form conjugated mainly with glycine and in a smaller pro-
portion with taurine. Approximately 70% of the rabbit’s bile pigment is
biliverdin, because the synthesis of bilirubin is limited.
The readily available carbohydrates (sugars, starch) can be digested
completely and their absorbed end products are glucose, fructose and galac-
tose. Heat and chemical treatment influences the digestibility of starch.
Blood glucose and insulin concentration does not increase in rabbits fed
with crude starch. Heat treatment (100oC) enhances the hydrolysis. The
quickest hydrolysis and absorption can be obtained by feeding acetylated
distarch adipate.
The hydrolysis of fibre components (pectin, hemicellulose, cellulose)
is accomplished by the intestinal bacteria, mostly those of the caecum. The
end products of that fermentation are short-chain fatty acids. The produced
VFAs (formic, acetic, propionic, butyric) and the other organic acids,
remaining from the feed (malonic, fumaric, succinic acid) are actively
absorbed through the caecal and colonic wall into the blood. The proportion
of the three main short-chain fatty acids, acetic, propionic and butyric acid
in the caecum is 10:1:1.53.
23.1.2. Digestive functions in the caecum

The concentration of the bacterial flora in the flora caecum is l09-11/gram,
with the predominance of the strictly anaerobic Bacteroides colonising in
the first week of life. The main N source for the bacteria is NH3 produced by
their desaminase and urease activity. Cellulolytic activity is low, so the
energy required for the assimilation of ammonia will be obtained from the
degradation of sugars, pectin and xylose. In case of an incomplete protein
supply the blood urea will be excreted in the caecum. A certain amount of
urea derives from the feed. The biuret can be more efficient N source than
urea. Urea will be hydrolysed mainly by bacteria adhering to the mucous
membrane.
As a function of the protein level of feed and the NH3 concentration of
745
caecal content, the produced ammonia will be built into the bacteria and
will be utilized by caecotrophy. (Certain absorption of microbial lysine by
non-caecotrophic routes in rabbits, but it represented only 14% of the total
microbial lysine absorption, much lower than that obtained by the re-inges-
tion of soft faeces (BELENGUER et al. 2005). Besides the protein concentration
of the feed mixtures used in practice the ammonia-assimilating capacity of
the caecal flora is trifling, so supplementation of the rabbit’s feed with urea
does not result in any benefits. On the other hand, feeding of urea may
cause emphysema of the gut wall and load the detoxifying capacity of the
liver.
23.1.3. Intake of the soft faeces (caecotrophy)
The word „coprophagy” means simply the ingestion of faeces, independent-
ly from its forms and causes. Some animals practise coprophagy in case of
disturbed behaviour or severe (micro) nutrient deficiency. For other group
of animals this phenomenon is normal, even necessary for the digestive
physiology and in these cases it is justified to call it re-ingestion rather than
coprophagy. The caecotrophy is the most specialized form of the physiolog-
ical coprophagy (re-ingestion), developed by taxonomically distant mam-
mals (some rodent species, beaver, herbivorous monkeys etc.), where a par-
ticular mechanism allows the separate excretion of the caecal content rich
in nutrients and the normal, hard faeces poor in valuable material. The re-
ingestion of caecal content (caecotroph) makes the better utilization of
nutrients possible. The caecotroph will be swallowed without chewing. The
typical composition of the two types of faeces is the following: dry matter 53
vs. 39%, and in the dry matter crude protein 15 vs. 34, ether extract 3 vs.
5, crude fibre 30 vs. 18, ash 14 vs. 15, N-free extract 38 vs. 19%, gross ener-
gy 18 vs. 19 MJ/kg dry matter for hard faeces and caecotroph, respective-
ly. (Table XXIII-1).
746
Table XXIII-1: Chemical composition of the rabbit’s two types of faeces

(FEKETE and BOKORI, 1984)
*Dry matter as % of fresh matter; all the other data as % of DM
The caecotroph and the hard faeces will be produced in the proximal
colon, at different times. During the production of hard faeces a separating
mechanism, by the contractions of the colonic haustrae, removes the fluid
and small particles back in the caecum (FIGURE XXIII-3).
FIGURE XXIII-3: Separating mechanism of the rabbit’s proximal colon

(after CHEEKE, 1987)
747
The process is helped by the active water secretion of the colon. The
fusus coli is supposed to be the pacemaker of the contractions. The
described separating mechanism does not act during the excretion of the
caecotroph, which is fundamentally a caecal content, enclosed by mucin
produced by the colonic gland cells. The mucilaginous membrane inhibits
the diffusion of ions and absorption of electrolytes. The hard faeces contains
more dry matter, but its protein level is low. The soft faeces mostly consists
of bacteria of high protein and vitamin B concentration. The physical form
of the hard faeces is well-known; the caecotroph is a cluster, composed by
5-10 small globules, fixed together by the mucous membrane. The form of
clusters changes, from the compact vine grape to the strawberry (FIGURE
XXIII-4).
FIGURE XXIII-4: A caecotroph (left) and two pieces of hard faeces (© FEKETE, 1984)
During the formation of hard faeces the caecal content will thorough-
ly be transformed in the colon. The wall of the proximal colon produces a
lytic factor (protein of 180,000 Dalton) which dissolves the bacteria, espe-
cially those of low carbohydrate content. Bacteria of high carbohydrate con-
tent, produced during the soft faeces phase are resistant against this lyth-
ic factor. Apart from the described separating mechanism this lythic factor
is responsible for the fact that the bacterium concentration 1/5-1/10, the
protein content ½-1/3 that of the caecotroph. The distal colon produces,
mainly in the soft faeces phase, a species-specific lysozyme, which will get
into the stomach with the caecotroph and there it has a bacteriolytic role.
During the formation of soft faeces the plasma level of aldosterone is the
lowest, does not stimulate the re-absorption of sodium from the caecotroph.
Consequently, the sodium concentration and the sodium to potassium ratio
are higher that in the hard faeces. Stress (or ACTH injection) has a similar
effect.
748
23.1.4. Role of caecotroph in the gastric carbohydrate fermentation

The starch of the feed will be hydrolysed under the action of amylase into
glucose molecules. Amylase can derive from three sources: feed, saliva and
caecotroph bacteria. The third (with is a molecule of 40,000 Dalton) is the
most important since it is less sensitive to acidic milieu. The produced glu-
cose diffuses into the caecotroph balls and will be transformed into lactic
acid and carbon dioxide. Lactic acid rediffuses into the lumen: its absorp-
tion begins in the stomach but is completed only in the small intestine. The
mucous membrane surrounding the caecotroph pellets plays an important
role: it protects the bacteria from being attacked by gastric juices. (FIGURE
XXIII-5).
Figure XXIII-5: Fate of the caecotroph globules in the stomach
The importance of this phenomenon is not the energy surplus by lac-

tic acid, but rather the lightening of the hind gut’s carbohydrate overload
(see 23.4.) as pancreatic amylase production of the rabbit is modest, and
the acidification of the stomach content activating pepsinogen.
23.1.5. Digestive physiology of weaning

After birth the first, exclusive feed for bunnies is the mother’s milk. The rab-
bit’s milk is rich in dry matter (26-31%), fat (10-15%) and protein (12-13%);
the only carbohydrate is the lactose (0.9-1.0 %). The transition period from
milk to solid feed begins at the age of 20-21 day. The doe generally suckles
only once a day. The long breaks facilitate/stimulate the bactericide
processes in the stomach. The suckling time is early morning, and lasts 3-
749
5 day. The bunnies have no fixed teat.

The function of the mucosal barrier is immature in the newborn rab-
bit: intestinal antigen intake and endotoxin binding are greater than in
adult animals. The feed antigens are absorbed more readily into the circu-
lation during the first weeks of life. (Artificial raising with cow’s milk: dan-
ger of „sudden cot death”). During the suckling period in the digestion of
carbohydrates the lactase is predominant; at the 3rd week, parallel with a
decrease in lactase activity, those of the saccharase, maltase and amylase
grow. The colonisation is very early, after 24 hours it reaches the 109/g
digesta concentration in the caecum and colon. Germ-free raised pups do
not practise caecotroph. The formation of the normal enteral microflora is a
very important factor in the development of ileal villi. The protein digestion
is done by means of the rennin of the stomach and the pancreatic trypsine
and chymotrypsine. The activity of pepsin-hydrochloric acid activity is
important to the age of 4 weeks, but full maturity cannot be reached earli-
er than 5-6 weeks. The fat digestion is helped by the mother’s milk lipase
and partly by the pancreatic one, but the majority of the milk fat is absorbed
intact. The adult lipase activity develops until the 5-6th weeks.
23.1.6. Weaning systems

Weaning at the age of 27-30th days. At 18-20 days the bunnies begin to eat
from the feed of their mother. In the middle of week 4 the dry matter of the
ingested feed is equal to that of suckled milk. The pancreatic amylase activ-
ity of an average bunny is now significant, the intestinal flora is stable and
the caecotrophy is practised. To have a security margin, the weaning on day
27-30 is proposed.
Weaning at the age of 35-42 days of life. From bunnies’ biological and
animal health point of view the weaning 5-6-week old is proposed to
decrease mortality and to minimise the drop in the rate of body weight gain.
Before and after weaning (1-2 weeks) a high fibre, low in starch and
camomile (Anthemis nobilis) hay (with flower) containing feed is proposed
(FEKETE and VETESI, 1985) to decreased enterophathies. At the same time the
average daily gain of young rabbit is smaller, but later, owing to the com-
pensatory growth the deficit in live weight will disappear.
1.7.3. Raising of orphan pups.
The best way of course is the giving to nurse doe. If it is not possible,
until the opening of the eyes (approximately at 10-day), the most practical
way of feeding the bunnies is the eye dropper method. Cow’s or goat’s milk
should be made more concentrated by mixing cream or egg yolk into it. The
mixture should be heated and concentrated with a small amount of corn
meal or maltose. The proper temperature can be controlled by dropping it
on the back of the hand. Although in natural conditions the pups suckle
only once a day, in artificial raising, it is better to give the mixture in two
equal portions (12 -12 h). This helps to avoid diarrhoea and undernutrition,
750
because the artificial milk normally does not reach the nutrient concentra-
tion of the mother’s milk. To facilitate urination the inguinal region should
be stroked for 1-2 minutes before each feeding. For the 12-14-day-old bun-
nies a small amount of oat flakes and some blades of grass are offered. At
that time they are able to learn to eat from a teaspoon. From 15-18 days
special rabbit starter can be given besides the milk replacer.
By mixing trypsine and chymotrypsine to the milk replacer the sur-
vival rate of orphan pups increases. The following milk replacer (ARTIFICIAL
RABBIT MILK) proved to be effective: in 1 litre of cow’s milk 62.5 gram deep-
frozen milk protein, some teaspoons oil and 10 gram vitamin supplement
should be dissolved. Each gram of the vitamin supplement contains 4 mg
calcium-panthotenate, 1 mg niacin, 0.4 mg pyridoxine-hydrochloride, 0.2
mg thiamine-hydrochloride, 0.2 mg riboflavin, 25 µg pteroil-monoglutamic
acid, 25 µg biotin, 10 µg cyanocobalamine, 60 µg retinol, 2.5 µg kolecalcif-
erol and 0.99 g glucose.
23.1.7. Mineral metabolism

Calcium, phosphorus and magnesium. The calcium metabolism of rabbit
(and guinea pig) differs from that of other mammals. The surplus calcium of
the feed will be absorbed, later – mostly in the form of calcium-carbonate –
excreted in the urine. If the daily ration has a high Ca level, the urine is con-
centrated, white, and similar to limewater. Owing to the damage of the cap-
illaries, this urine may contain yellow-red pigment. In the majority of mam-
mals the intestinal absorption is the major site of calcium metabolism, and
the surplus is eliminated in the faeces.) In contrast, the extra intake of cal-
cium will be absorbed independently of the needs of the organism which
causes the plasma calcium level to increase and as the threshold is exceed-
ed it is excreted in the urine. The metabolic route of the excretion of the sur-
plus magnesium is also the urine, but this mechanism is independent of the
calcium intake.
Too high a level of dietary calcium (>0.75%) has no effect on the repro-
ductive performance of does, whereas too much (1.1%) phosphorus has. The
magnesium requirement of pregnant does is at least 20 mg/kg LW/day. The
deficiency does not influence the blood concentration, the litter size, but the
kindling time will be elongated and the Mg content of mothers’ femur
decreases.
Potassium. The requirement is 0.6-0.9%. The surplus (>0.5 g/kg LW)
potassium is excreted by the kidneys, the plasma concentration remains
unchanged, but enhanced diuresis and polydipsia can be found (for 1 g K
86 ml extra urine). Extra potassium antagonises with the absorption and
retention of Mg.
Eating feed of too high potassium level (4.2 or 6.4%) cause
histopathological damages in the testicles and suprarenal gland. In practi-
cal circumstances the forages grown on soil intensively fertilized with liquid
751
manure, may have a high potassium level. The continuous long-lasting

feeding of a daily ration of 1.0% potassium causes nephritis, the potassium
deficiency (>3%) may develop muscle dystrophy.
Sodium and chloride. There is no sufficient data concerning the real
salt requirement of rabbits. The NRC (l977) recommended 0.5% NaCl sup-
plementation is sufficient in most cases. Separately taken, the growing rab-
bit requires as a total of 0.25% sodium and 0.32% chloride in the air-dry
diet.
Sulphur. Anaerobic bacteria of the rabbit’s intestinal tract are able to
reduce sulphur. This chemical form can be built into their amino acids,
which – by means of the caecotrophy – can be utilized by the host organism,
the rabbit. The angora wool is extremely high in S (38 mg/gram, cf. sheep
wool 27-33 mg/gram). The apparent absorption of sulphur is 65-77%; the
urinary excretion is 27-35%. The daily sulphur retention is 34-47% of the
ingested amount. The average daily wool production of the Angora rabbit is
2-3 gram; therefore the feed should contain 0.3% sulphur in the air-dry
matter. In case of deficiency, the supplementation of the feed mixture with
0.1-0.2% Na2SO4 may be useful. The needs of growing meat rabbit are met
by the sulphur content of the natural feedstuffs.

Vitamin A, carotene. The daily 8 µg/kg LW vitamin A (approx. 500-600 UI/kg
feed) is sufficient for the maintenance and growth. The reproductive func-
tions require more: 14-20 µg/kg LW. The natural rabbit feedstuffs general-
ly contain the precursor carotene. The transformation to vitamin A occurs
predominantly in the gut wall during absorption.
Symptoms of vitamin A deficiency are similar to those other mam-
mals. Characteristic for rabbit are congenital hydrocephalus, anorexia, con-
junctivitis, blur of cornea and a decrease in the elastin content of lungs and
aorta. Classically, characteristic symptoms of A-hypervitaminosis are: a dis-
position for convulsions, loss of hair and osteomalatia. Feeding of pregnant
does with a ration of higher (190,000 IU/kg feed) vitamin A level may cause
foetal resorption, abortion, hydrocephalus of newborn, reduced litter size
and weight. In the blood of DOES WITH VITAMIN A TOXICOSIS the level of retinol-
, retinyl-palmitate and the total vitamin A are increased. The vitamin A con-
centration of the liver may be 100-200 times higher. The ingestion of the
same amount vitamin A by the growing rabbit did not have a negative influ-
ence other than decreasing the feed intake and daily weight gain by a few
percentage.
Vitamin D. The vitamin D deficiency can cause rickets in growing rab-
bits. The exact daily need is controversial, 23,000 IU vitamin D3/kg feed is
already toxic and it causes the calcium and phosphorus concentration of
the blood plasma to increase. The animal will be emaciated; in the soft tis-
sues, the mucous membrane of the stomach, in lung, trachea, kidneys,
752
blood vessels and in females ovarian calcification occurs with high mortali-
ty.
Vitamin E. The minimum daily vitamin E (alfa-tocopherol) require-
ment is 1 mg/kg LW. This is equivalent to l5-20 mg/kg air-dry feed, but in
practice more than 40 mg/kg air-dry feed is offered. In liver coccidiosis the
tocopherol level of tissues drastically falls. The most typical deficiency
symptom is muscle dystrophy (Zenker muscle necrosis) accompanied by
fatty infiltration of the liver and regressive processes in the ovaries and tes-
ticles. The concomitant deficiency of cholin and/or potassium facilitates the
development of myodegeneration.
Vitamin K. The amount of vitamin K, produced by the caecal bacteria
and utilized by the caecotrophy, is sufficient for maintenance and growth.
Pregnant does require 2 mg/kg feed supplementation to prevent bleeding in
the placenta and the consequent abortion. The higher vitamin K supply can
be given in form of alfalfa (hay/pellets), too.
Water soluble vitamins. By means of the caecotrophy the rabbit can
meet most of its requirement to the group B. The rabbit is capable of syn-
thetising vitamin C, thus the supplementation is not necessary.
As a generally accepted practice the recommended vitamin concentra-
tions in feed mixtures are the following: 10,000 IU vitamin A/kg, 900 IU vita-
min D3/kg, 50 mg vitamin E/kg, 2 mg vitamin K3/kg, 2 mg vitamin Bl/kg,
4 mg vitamin B2/kg, 20 mg panthothenic acid/kg, 2 mg vitamin B6/kg,
0.01 mg vitamin Bl2/kg, 50 mg niacin/kg, 5 mg folic acid/kg and 0.2 mg
biotin/kg.
Microminerals like iron, zinc, manganese, copper, iodine, selenium are
recycled by the caecotrophy, and the supplementation is only recommend-
ed when (semi)synthethic diets are used.
It is stated that the poorer growth rate of rabbits in larger litters as

opposed to smaller litters is more likely to be due to insufficient maternal
nutrition during pregnancy than to foetal competition. As a reference, a
thorough knowledge of the changes of the major chemical components tak-
ing place in pregnant rabbit and her uterus is extremely important in tera-
tological studies, in which the rabbit is the most frequently used model ani-
mal (FIGURE XIII-6).
It can be concluded that in contrast to other animal species (BRODY,
1945), in rabbits the development of foetuses does not only present a con-
siderable demand in the last trimester of gestation but already from the sec-
ond half of the pregnancy and the changes of fat to protein ratio are similar
to that of humans (FEKETE et al. 2005).
753
FIGURE XIII-6: Changes of the maternal (BW-Doe) and foetal (BW-F) weights, maternal
(GE-Doe) and foetal (GE-F) energy content and the fat to protein ratio
(Fat/Protein) in the foetus (FEKETE et al. 2005).
The above described facts explain well, why the breeding doe received
special emphasis in this manual.
754
© Gerolamo Xiccato and Angela Trocino
T
he scientific and related technical advances in rabbit production dur-
ing the last 20 years have radically changed farm management. Main
changes in rabbit farm management firstly affect the reproduction
sector. Both research and genetics offer commercial hybrid lines selected for
high reproductive efficiency with high prolificacy and high milk production.
The spread of artificial insemination allows controlled reproduction
rhythms, programmed reproductive events and therefore it optimises
labour. On the other hand, however, the same conditions that contribute to
the innovation of rabbit production bring about some negative conse-
quences. High producing hybrid rabbit does is not capable of ingesting a
sufficient energy level to support their nutritional requirements and is
excessively exploited during their life, with a consequent reduction of their
reproductive career length (longevity). The use of too intensive reproductive
rhythms may accentuate these negative figures.
23.2.1. Energy metabolism and nutrition in rabbit does

Energy metabolism and energy requirements in young and breeding does
may vary according to several factors. The most important ones are the body
size (depending on breed and age), the physiological state: maintenance,
growth, pregnancy and lactation; the environmental conditions; tempera-
ture, humidity, speed of air flow and management (i.e. different reproduc-
tive rhythms, litter size, weaning age). An intense body energy deficit has
been proved in rabbit does during lactation, especially in highly reproduc-
tive hybrid lines whose voluntary feed intake is often unable to cover nutri-
ent requirements for lactation and concurrent pregnancy (PARTRIDGE et al.
1986). The doe’s energy deficit may be controlled by means of increased vol-
untary feed and energy intake; reduced body energy output by anticipating
weaning age as well as increased body energy recovery by prolonging the
kindling-to-kindling interval.
23.2.1.1. Special features of voluntary feed intake

The appetite regulation in reproducing rabbits is mostly controlled by
chemostatic mechanisms, for which reason the total energy quantity ingest-
ed daily tends to be constant when expressed related to live weight (LW) or
even better to metabolic body mass (kg W0.75). While in growing rabbits (0.6
to 2.5 kg LW) voluntary intake is about 0.95 to 1.00 MJ DE/day/W0.75,
young breeding females ingest a lower amount of energy decreasing from
0.95 to 0.70 MJ DE/day/W0.75 in the period from weaning to first insemi-
nation.
During the first pregnancy daily energy intake further decreases from
755
0.60-0.65 MJ DE/day/W0.75 in the first 25 days until 0.40-0.45 MJ

DE/day/W0.75 in the last 5 days, due to the increasing volume of foetuses
in the abdomen. On day of kindling, the doe ingests only a little amount of
feed or even nothing (FIGURE XXIII-7).
FIGURE XXIII-7. Evolution of voluntary feed intake (kg/day) (full line) and energy intake
(MJ DE/day/W0.75) (dotted line) from mating to the end of the third lacta
tion in breeding does submitted to a semi-intensive reproductive rhythm
(insemination 12 d post-partum). (I: insemination; K: kindling; W: weaning.)
Energy consumption is higher in lactating females, which can ingest

up to 1.5-1.8 MJ DE/day/W0.75 at the lactation peak and 1.1-1.3 MJ
DE/day/W0.75 on average. The highest values are recorded for multiparous
does. After weaning of the litter, does quickly decrease their feed intake in
a week at about 35-45% of the lactation level, that is 0.5-0.6 MJ
DE/day/LW0.75 (XICCATO et al. 2004).
The chemostatic regulation in growing or non reproducing rabbits
appears higher than 9 to 9.5 MJ DE/kg. Below this limit, a physical-type
regulation linked to filling the gut by feed is prevalent. In lactating does, the
energetic limit of chemostatic regulation is higher than the above figure,
around 11 and 11.5 MJ DE/kg. Between the range of9.5 and 11 MJ DE/kg,
both physical and chemostatic regulations act in a combined way.
Therefore, an increase in the DE concentration over 10 and 11 MJ DE/kg
permits a further increase of daily energy intake. The chemostatic regula-
tion depends also on the dietary energy source and is higher in fat-added-
diets than in high-starch diets.
23.2.1.2. Maintenance requirements

Like in other animals, DE requirements for maintenance (DEm) in rabbits
depend on metabolic body mass and physiological state. Since no experi-
756
mental information on DEm requirements is available for young males and

females (2.5 to 3.5 kg LW) kept for reproduction, the value of broiler rabbit
(0.43 MJ DE/day/W0.75) can be used (Table XXIII-2). In breeding does, esti-
mations of DEm often are different, because body energy changes during
lactation are not always considered. When body protein and fat changes are
considered in the different physiological condition (growing, pregnancy and
lactation), the proposed maintenance energy equal to 0.40 MJ DE/day/W0.75
for non-reproducing does, 0.43 MJ DE DE/day/W0.75 for pregnant or lactat-
ing does and 0.46 MJ DE/day/W0.75 for concurrently pregnant and lactating
does. The value of 0.40 MJ DE/day/W0.75 recommended for non-reproduc-
ing does can also be used for adult bucks.
Table XXIII-2. Maintenance energy requirement and efficiency of DE utilization and body
energy retention (PARIGI- BINI and XICCATO 1998).
23.2.1.3. Growth requirements in young rabbits

Growth requirements and efficiency of energy utilization in young males
and females kept for future reproductive activity can be considered similar
to those of rabbits kept for meat production. During the first period of
growth (from weaning to 10-12 weeks of age), body increment (retained
energy=RE) is mainly retained as protein, while a lower amount is retained
as fat. The energetic values for body protein are 23.0-23.5 MJ/kg. The cor-
respondent value for rabbit fat is lower than in other mammalians and dif-
ferent from young and adult rabbits (35.6 vs. 36.5 MJ/kg).
The efficiency of DE utilization for body energy retention (RE/DE) is
0.50-0.55, depending on the proportion of retained protein and fat (Table
XXIII-2). The efficiency of energy utilisation for growth is influenced by the
composition of growth, because energy as protein is retained less efficiently
(0.40) than as fat (0.65). In the second period of growth (from 10-12 weeks
to first mating), the feeding plan should avoid fattening which could impair
reproductive efficiency and career length. An increase in the dietary energy
level can affect the composition of body gain and the partition of energy
757
retained as protein and fat. The body composition changes are not linearly
correlated with DE intake (DEI), because some constituents (i.e. fat) tend to
increase more than proportionally.
23.2.1.4. Pregnancy requirements

Primiparous does undergo wide variations in body composition, tissue dep-
osition and energy retention. During early and mid gestation (0-21 days),
LW increases like in non-pregnant does. During late pregnancy (21-30
days), empty body weight (LW minus gut content) decreases as a conse-
quence of protein and fat losses and energy transfer to the rapidly growing
foetuses (FIGURE XXIII-8.
FIGURE XXIII-8: Live weight and energy changes of maternal body and foetuses in the peri-
od around first pregnancy and kindling.
The efficiency of DE utilisation for maternal tissue accretion in preg-

nant or non-pregnant does has been estimated equal to 0.49. The efficien-
cies of dietary DE utilisation for foetal growth are 0.31 in pregnant nulli-
parous does and 0.27 in lactating and pregnant does. Explanation for the
high energy cost of foetal growth, observed also in pigs (0.20-0.30), may
come from the very high protein content of the foetal body and its quick
turnover.
23.2.1.5. Lactation requirements

The energy output in the milk (Emilk) during lactation is exceptionally high
in rabbits, compared to other species, due to the intense milk production
(200-300 g/day) and its high dry matter concentration (30-35%); protein
758
(10-15%) and fat (12-15%). The average milk energy output per kg of meta-
bolic body mass is higher in rabbits than in cows: a 4-kg doe producing 250
g/day of milk excretes 0.75 MJ Emilk/day/kW0.75, while a 600-kg cow pro-
ducing 25 kg/day of milk excretes only 0.6 MJ Emilk/day/W0.75.
Chemical composition of rabbit milk changes substantially during
lactation. In particular, dry matter content decreases in the first 1-3 days
when colostrum becomes milk, then it remains constant for 2-3 weeks, and
finally increases as the milk yield decreases. During the third decade of lac-
tation, a contemporary increase in fat concentration is recorded FIGURE
XIII-9). The average energetic value of milk is 8.4 MJ/kg, about 2.8 times
higher than that of cow milk (3.0 MJ/kg).
FIGURE XXIII-9. Evolution of the chemical composition of milk (LEBAS, 1971).
The utilisation of DE for milk production is estimated at 63% in both

lactating and concurrently pregnant and lactating does (Table XXIII-2). The
efficiency of energy utilisation retained in the doe body (RE) for milk pro-
duction is calculated 81% in lactating does and 76% in lactating and preg-
nant does similar to other animal species (bovine and pigs) and in accor-
dance with theoretical calculations.
For the calculation of energy requirements and body balance for lac-
tating non-pregnant does, see Example.
Example. Estimate of DE requirements and body energy and fat balance in lactat-
ing non-pregnant does (assuming protein balance in equilibrium) (from PARIGI BINI
and XICCATO, 1998)
Reference data:
Average live weight (LW) = 4.25 kg
Days of lactation = 30 days
759
Milk production = 220 g/day

Gross energy content of milk = 8.4 kJ/g
Gross energy content of fat in the maternal body = 36.5 kJ/g
Efficiency of DE utilisation for milk energy (Emilk) = 0.63
Efficiency of body energy (REd) utilisation for Emilk = 0.78
Dietary DE density = 11 MJ/kg
Maximum DE intake (DEI) = 1250 kJ/day/kg LW0.75
Protein balance = 0
Calculated data:
Average W0.75 = 2.96 kg
Emilk = 220 x 8.4 = 1848 kJ/day
DE requirement:
DEm = 430 kJ/day/kg W0.75 = 1273 kJ/day
DE requirement for milk production (DEmilk) = 1848 / 0.63 = 2933 kJ/day
Total DE requirement = DEm + DEmilk = 4206 kJ/day (1420 kJ/day/W0.75)=4.2
MJ DE/W0.75
Body energy and tissue balance

Maximum DEI = 1250 x 2.96 = 3700 kJ/day
Maximum feed intake = 336 g/day
DEI deficit = -506 kJ/day
Emilk from dietary energy = 1848 - 506 x 0.63 = 1529 kJ/day
Emilk from body energy = 1848 - 1529 = 319 kJ/day
Body energy (REd) retained = -319 / 0.78 = -409 kJ/day
Fat loss = -409 / 36.5 = -11.2 g/day
Total body fat loss during lactation = -336 g
Total body energy loss = -12.27 MJ/animal
23.2.1.6. Effect of nutritional strategies

FEEDING THE YOUNG DOES
Young does should face the first mating, pregnancy and lactation with an
adequate body condition to support high nutritional requirements for
reproduction at their best. The growing period can be divided into two
phases:
- first period, from weaning (30-35 days) to puberty (10-12 weeks) and LW
from 0.8 to 2.4 kg, during which feeding programme and growth perform-
ance are quite similar to those of rabbits kept for meat production;
- second period, from puberty to first mating (17-18 weeks) and LW up to 3.2-
3.5 kg, during which period daily growth rate lowers and feeding program
aims to induce a correct morphologic and reproductive development. In the
first period, the growth models developed for meat rabbits can be used: daily
gains (g/day/W0.75) of live weight (LWG), empty body (EBG = 0.87×LW),
water (WG), protein (PG), fat (FG) in growing rabbits from weaning (0.8 kg)
to slaughter (2.4-2.5 kg) can be estimated according to digestible energy
intake (DEI). The partition between energy retained as protein (REp) and fat
760
(REf) is different according to the dietary energy intake. When DEI=DEm (i.e.
0.43 MJ/d/W0.75), the energy equilibrium is reached as a consequence of a
gain of energy (36 kJ/day/W0.75) as protein (+1.5 g/day/W0.75) and an
equivalent loss of energy as fat (-1.0 g/day/W0.75). At the same time, EBG
is positive (6.5 g/day/W0.75) primarily due to water retention. With increas-
ing DEI, LW and EB gain increase and protein gain (in weight) always
remains higher than fat gain.
The voluntary DEI of young rabbits kept for reproduction is an aver-
age of 0.90 MJ/day/W0.75, slightly lower than that of in growing rabbits for
meat production (0.95-1.00 MJ/day/W0.75). When fed ad libitum, RE as
protein and fat is equal (REp = REf = 0.122 MJ/day/W0.75) and rabbits
reach about 2.4 kg at 11 weeks of age. If feed restriction is applied in the
first period of growth (80-85% of ad libitum), or even less (70.75% of ad libi-
tum), daily weight gain is lower and at the same weight (2.4 kg) the body
composition is characterized by lower fat and energy concentration (FODOR
et al. 2002).
In the second period of growth, the somatic and physiologic develop-
ment of the future reproducing female advances slower. Body protein and
ash concentrations tend to remain stable at 20% and 3%, respectively (state
of chemical maturity), while fat and water (inversely each other) levels
depend on age at first mating and feeding regime. Reproducing females are
usually mated the first time at 16-18 weeks of age, or at 75-80% of adult
weight, i.e. 3.3-3.5 kg for an adult weight of 4.2-4.4 kg. From 10-12 weeks
to first mating, voluntary feed intake slightly decreases from 0.80 to 0.70
MJ/day/W0.75 and daily weight gain reduces from 35 to 20 g/d.
At 17 weeks of age, breeding rabbits given ad libitum access to a diet
containing 10 MJ/kg DE may reach about 3.4 kg LW and about 18% body
fat concentration. This condition could be excessive if further fattening dur-
ing pregnancy or quick overfattening (more than 20%) in 2-3 weeks in case
of failed pregnancy is considered. Overfattening can provoke subsequent
distocia and impaired reproductive performance. An earlier insemination
(e.g. 15 weeks) would avoid overfattening, but it could be unfavourable due
to the still incomplete development of endocrine and reproductive systems.
To avoid overfattening by postponing the first insemination, feed is
often given at a restricted level. The choice of restriction level and feeding
programme are of great importance to permit a correct morphologic and
physiologic development of future breeding rabbits, prepare them for the
reproductive career with suitable body energy reserves and to stimulate
their feed intake capability. Restricted feeding regimes delay the moment at
which the target weight is reached (3.4 kg), thus decreasing daily weight
gain (Table XXIII-3).
761
Table XXIII-3. Theoretical performance and body composition in young females submitted
to restricted feeding from 11 weeks of age until first mating (3.4 kg LW)
Feed restriction at 90% of ad libitum level (0.675 vs. 0.750 MJ

DE/day/W0.75) decreases daily weight gain by 20% and allows to reach the
target weight at 18 weeks with a body fat concentration of 16%, which can
be considered as an optimum. Restrictions at 80% or even at 70% further
delay the age of first mating and do not permit body fat storage in a suffi-
cient quantity. To avoid reduction in sexual receptivity at the first insemi-
nation in restricted does, a FLUSHING with a lactation diet given ad libitum is
usually performed during 4-7 days before the first insemination (ROMMERS
et al. 2001 and 2004).
One of the main criticism of feeding restriction is the reduction of vol-
untary feed intake in the successive reproductive phases (pregnancy and
lactation) with the accentuation of risking a negative energy balance
between reproductive cycles (FORTUN-LAMOTHE, 2003). For this reason, feed-
ing restriction (80-90%) could be applied in the first growth period (5-11
weeks), followed by unrestricted feeding. This regime should result in an
optimal body condition at 17-18 weeks of age and a voluntary feed intake
higher than following traditional restricted programmes. Feed restriction
can also be continued in the first part of pregnancy, especially when the live
weight exceeds the target one, while ad libitum feeding with lactation diet is
recommended during the last 2 weeks of pregnancy to take into account
increasing pregnancy requirements and decreasing feed intake nearby kin-
dling.
If ad libitum feeding is preferred, a growing diet containing a moder-
ate energy level (9.5-10 MJ/kg) rich in fibre and well provided with protein,
should be given throughout most of the growing period until the first mat-
ing, in order to allow good somatic development in the doe and increase vol-
untary feed intake during subsequent reproductive career. Since the rich-
762
fibre diet administration to growing rabbits increased feed intake and gut
content (FEKETE and BROWN, 1993), some studies were performed to stimu-
late voluntary feed intake during the growth of young does and increase dry
matter and energy intake during subsequent reproductive activity. The
administration of high fibre-low energy diets to young females before the
first mating increases voluntary feed intake during growth and pregnancy,
while decreases body fat and energy deficit at the end of the first lactation
(XICCATO et al. 1999). A similar feeding regime does not affect reproductive
performance at birth, while stimulates feed intake during lactation and milk
production with consequent higher litter size and weight at weaning.
FEEDING REPRODUCING DOES
Insufficient feed intake of primiparous does and the consequent inability to
cover lactation and pregnancy requirements are widely recognised as the
main reasons for their body energy deficit. During early pregnancy, increas-
ing dietary energy concentration by fat/oil addition usually reduces dry
matter intake and does not determine significant variations of digestible
energy intake. During the last week of pregnancy, voluntary feed intake is
limited by physical intake capacity (PASCUAL et al. 2003).
The effects of feeding level on reproductive performance are contro-
versial: higher pup mortality at birth has been found with high digestible
energy intake during pregnancy, which has not been confirmed in long-term
experiments. During lactation, feeding high digestible diets increases DE
intake. This effect is even more accentuated when fat-added diets are used
in comparison with high-starch diets both in primiparous and multiparous
does. Body energy balance in does, however, is always negative and not sta-
tistically different from dietary treatments (FIGURE XXIII-10).
FIGURE XXIII-10 Body energy balance in an empty body in does at different physiological
state was fed on diets with increasing DE concentration (XICCATO et al.1995).
In fact, a higher dietary energy supply determines an increase in milk
763
production, limiting its beneficial effect on body condition (empty body=EB)

both in primiparous and multiparous does (Table XXIII-4).
Table XXIII-4: Effect of dietary energy level on reproductive performance and energy balance
in rabbit does at the 28th d of the second lactation (PASCUAL et al. 2000)
When analysing energy balance in reproducing does fed on diets at 17

to 18% CP and 10 to 11 MJ DE/kg in different experiments, the increase of
DE intake (net of the requirements for simultaneous pregnancy) leads to a
proportional increase in milk energy output. The production of milk energy
(Emilk) and the energy retained in the doe body (REd) are functions of the
DE intake (DEI).
The gross efficiency of DEI utilisation is 43.4%, or rather, each addi-
tional kJ of ingested DE determines an increase in milk energy production
of 0.434 kJ. This efficiency is different from the net efficiency of DEI utili-
sation for milk production, which is 0.63, because part of the available ener-
gy is used to reduce energy deficit. This deficit can be decreased by 0.203
kJ per each additional kJ of DEI. This trend, linear throughout the intervals
tested, shows a DEI at the energy equilibrium (REd=0) equal to 1.585
MJ/day/W0.75. A similar voluntary intake is very rare (on average) in prim-
iparous and secondiparous does. At the equilibrium point, the energy milk
output is 0.71 MJ/day/W0.75 which corresponds to about 250 g/day of
milk in a 4.25 kg rabbit, assuming a milk energy concentration of 8.4
MJ/kg.
23.2.1.7. Effect of management strategies

23.2.7.1. PARITY ORDER. Unlike primiparous rabbits, multiparous does are
usually considered capable of ingesting high amounts of feed required to
achieve body energy and protein equilibrium. Substantial body fat and ener-
gy mobilisation, however, is observed also in multiparous lactating does.
Various authors describe significant increases (5-15%) in feed intake from
the first to the second and from the second to the third kindling, followed by
lower but not significant increases for successive parities. DE intake rises
by 9% between the first and the second lactation and only by 3% from the
second to the third lactation, while milk production increases by 10% and
8%, respectively. The unchanged substantial gap between dietary energy
intake and milk energy output account for the body deficit also maintained
at higher parities (FIGURE XXIII-9). In does submitted to semi-intensive
764
rhythm and traditional weaning, however, body energy deficits do not longer
appear in does after their 3rd kindling. Different results may be ascribed to
the rabbit strain (commercial hybrids or selected pure breeds) and/or the
body balance measurement methods (comparative slaughter, ultrasound
technique, total body electric conductivity).
FIGURE XXIII-11. Evolution of total body energy concentration (TOBEC measurements) in

rabbit does from 1st to 3rd kindling and according to their physiological
state (BOLET and FORTUN-LAMOTHE, 2002).
23.2.7.2. REPRODUCTIVE RHYTHM

The spread of artificial insemination has allowed programming reproduc-
tion activity inside the herd. Reproductive rhythms are currently defined
intensive, when scheduling insemination in does 1-7 days post partum (p.
p), semi-intensive, with insemination 7-20 days pp, or extensive with
insemination of 20-35 days pp.
Under practical conditions, the most frequent systems perform re-
mating at a fix day in the week that is 3-5 days, 10-12 days, 17-19 days or
24-26 days after kindling, which determine exact theoretical intervals
between the two kindling of 5, 6, 7, or 8 weeks, respectively. New solutions
are now being investigated (post-weaning insemination associate to early
weaning, alternate post-partum and post-weaning insemination), but the
most preferred re-mating programme remained the semi-intensive rhythm, a
compromise between the doe’s needs of recovering body reserves between
one reproductive cycle and the next (FIGURE XXIII-12) and the economic
demand of the increasing the number of bunnies weaned per year. In fact,
post partum insemination implies an excessive exploitation of the doe,
765
which finally results in a strong reduction of reproductive performances and

life span. On the other hand, extensive rhythm allows only a few number of
kindling per year and may cause mothers getting fat and subsequent
impairment of reproductive performance.
FIGURE XXIII-12. Theoretical energy requirements, digestible energy intake and energy
balance in multiparous rabbit does re-mated 12 d post partum (XICCATO, 1996)
Milk production is largely affected by the reproductive rhythms and

the different overlapping periods of pregnancy and lactation. Does submit-
ted to intensive reproductive rhythms begin to decrease milk production
after 15-17 d of lactation and a sharp drop in the last week of pregnancy
appears (FRAGA et al. 1989). In fact, nutritional requirements increase con-
sistently after the third decade of pregnancy due to exponential foetal devel-
opment and the hormonal changes caused by the imminent kindling that
contrast milk production (FORTUN-LAMOTHE et al. 1999).
One of the main reasons for lengthening the interval between kin-
dlings is prolonging the dry period in order to permit complete body energy
recovery. In primiparous does, a severe body energy deficit was observed
within the first and second kindling with inseminations 12 d post partum (-
26% of initial body energy content) but a less serious deficit (-15%) with
insemination 28 d post partum (FIGURE XXIII-13a). When multiparous
does were submitted to early weaning (21 or 25 d) body energy deficit dis-
appeared in those submitted to semi-intensive (insemination 12 d post par-
tum) and extensive (26 d post partum), rhythms while deficit was found to
be severe only in rabbits submitted to intensive reproductive rhythm (2 d
post partum) (FIGURE XXIII-13b). Better body conditions were observed in
766
the latter study, where only the intensive rhythm provoked a substantial
energy deficit, which might be ascribed to the concurrent action of high par-
ity order and early weaning age.
FIGURE XXIII-13a and FIGURE XXII-13b: Effect of reproductive rhythm on body protein,
fat and energy balance at kindling in reproducing does: (a) primiparous does with litter
weaned at 28 d (PARIGI-BINI et al. 1996); (b) multiparous does, average of litter weaning a t
21 and 25 d (XICCATO et al. 2004).
High reproductive efficiency has been reported in does submitted to

extensive rhythms, even if high embryonic mortality was observed. The role
of high prolactine and low progesterone levels in lactating does in reducing
the foetal survival rate has been elucidated, while the effect of nutritional
deficit caused by concurrent lactation is less clear. An alternating re-mat-
ing rhythm (alternatively 1 d post partum and post weaning associated
with weaning at 26 d) may improve both receptivity and fertility when com-
pared to a fixed re-mating rhythm (11 d post partum) (CASTELLINI et al.,
2003).
23.2.7.3. LITTER WEANING AGE. Under field conditions, bunnies are usually
separated from their mothers around 32-35 d of age or even later. Previous
research, in fact, reported a negative correlation between weaning weight
and post-weaning mortality and induced breeders to increase weight by
delaying weaning age. On the other hand, more recent studies demonstrat-
ed the possibility of successfully anticipating bunnies weaning age.
The greatest interest in early weaning technique lies in the possibil-
ity of reducing the doe body energy deficit, by shortening the lactation
length (period of energy deficit with body energy utilisation for milk synthe-
sis) and prolonging the dry period (period of energy surplus with body ener-
gy restoration). Milk production requires a great energy effort also in its final
period, because milk dry matter and lipid concentrations increase from the
day 20 onwards when production begins to fall. Moreover, the natural pro-
gressive weaning does not permit the complete recovery of the body energy
lost during lactation because of the very short dry period.
In does at their first, second and third parturition, reducing weaning
age from 32 to 21 d of age improved body energy balance (from –19% to –8%
767
of the initial energy concentration) but was unable to achieve equilibrium

(Table XXIII-5). Also in multiparous does, weaning at 25 d did not prevent
body energy deficit (–8% of the initial energy content), while weaning at 21
d only provided a balance that approached equilibrium (-3%).
Table XXIII-5.Body balance in lactating and pregnant does between initial (1st) and final
3rd) kindling: changes of empty body composition compared to initial kindling (XICCATO et
al, 2004)
Early weaning failed to definitively balance the energy deficit in these

does because of the substantial decrease in feed intake after weaning (about
40-50% of lactation period). With natural weaning, in fact, feed intake
remains at the highest levels also during the last decade of lactation, thus
partially permitting the recovery of body energy lost during the first 20 days.
Sudden decrease in feed intake that occurs after early weaning reduces
daily energy surplus and delays complete restoration of body reserves.
Nevertheless, a negative effect of very early weaning on rabbit repro-
ductive performance was also recorded in terms of the lower litter size and
bunnies born alive in multiparous does whose litters were weaned at 21
days of age). This result was ascribed to the marked effect of the weaning
technique on the metabolic and hormonal pattern at the time of foetus implan-
tation from 7 to 11 days of pregnancy. The occurrence of mastitis may also
be observed in the field as a consequence of the abrupt interruption of lac-
tation at 21-25 days. When early weaning (25 d) is associated with early
mating (4 d post partum), high prolificacy and litter size at weaning are
observed in comparison with does subjected to a traditional mating-wean-
ing system (mating 11 days post partum and weaning at 35 days of age).
23.2.2. Protein, amino acid and micronutrient nutrition

23.2.2.1. Protein requirement
When speaking about nitrogen nutrition in rabbits, the first question yet to
be solved is to determine the best unit for expressing requirements. Despite
the fact that true ileal amino acid digestibility would be the best unit, crude
protein (CP) and apparent digestible protein (DP) are the most commonly
used units, for which both requirements and raw material compositions are
available. Finally, due to the chemostatic regulation of voluntary feed intake
in rabbits, the digestible protein to digestible energy ratio (DP/DE) should be
considered to evaluate correctly the nitrogen intake in rabbits.
As for energy, protein requirements may be calculated on the basis of
768
maintenance, growth and production needs as summarised in Table XXIII-

6.
Table XXIII-6: Maintenance digestible protein (DPm) requirement and efficiency of dietary
and body protein utilization (FRAGA, 1998)
YOUNG RABBITS
In young rabbits, maintenance requirements are estimated at 2.9 g
digestible protein (DP)/day/W0.75 (FRAGA, 1998). Protein supply must also
consider growth requirements of young does, which vary in accordance with
the growth rate. The efficiency of dietary digestible protein utilization for
growth is 0.56. Overall DP retention (RP/DP intake) decreases linearly with
increasing live weight from 0.40 to 0.10 (TROCINO et al., 2000 and 2001).
During growth, empty body protein concentration changes from 12% (61%
DM) at birth to 17% (68% DM) at weaning (35 d) and about 20% (59% DM)
at 10-12 weeks of age. Afterwards, body protein concentration is quite con-
stant (20% of empty body weight; about 18% LW, after achieving chemical
maturity). Lacking specific information, the same figures for growing rabbits
may be used for protein maintenance requirements in non-reproducing
adult females and bucks.
PREGNANCY
During the first pregnancy, rabbit does retain protein in their body in the
early gestation period (0-21 days), while they transfer some protein from
their body into the rapidly growing foetuses in the late period of pregnancy
(21-30 days) (Table XXIII-7). This is due to the exponentially growing protein
requirements of the foetuses and the intense foetal protein turnover, which
was demonstrated to be approximately 5 times higher than that of the
maternal tissue.
LACTATION AND SIMULTANEOUS PREGNANCY

Protein requirements for maintenance have been estimated by using the-
comparative slaughter technique equal to 3.73 and 3.76-3.80 g
DP/day/W0.75 in lactating and lactating and simultaneously pregnant does
769
(PARIGI-BINI and XICCATO, 1993). In lactating does, the coefficients for utilisa-
tion of digestible protein (DP) and doe’s body protein utilisation for milk pro-
duction and foetal growth are calculated at 0.77 and 0.59, respectively. In
concurrently lactating and pregnant does the coefficients for digestible pro-
tein (DP) and doe’s body protein utilisation for milk production and foetal
growth are calculated at 0.76-0.80 and 0.60-0.61, respectively.
Table XXIII-7. Composition and partition of weight gain in pregnant does (PARIGI-BINI et al.
1990)
×EB=empty body
High milk production and the high milk protein concentration (10-
12%) accounts for high protein requirements for milk synthesis. When lac-
tation and pregnancy overlap each other, as already described for energy,
also protein requirements increases to a different extent depending on the
reproductive rhythm. Limited body protein losses (5 to 10%) usually occur
only in concurrently pregnant and lactating does subjected to intensive
reproductive rhythm (Table XXIII-8). Sometimes, however, a negative protein
balance has been reported in multiparous does having only lactation.
Table XXIII-8: Protein balance during the first lactation of does according to their physio
logical status (XICCATO et al. 1995)
PROTEIN TO ENERGY RATIO (DP TO DE RATIO)

In practice, dietary protein levels recommended for young females and
bucks range from 15 to 16% crude protein and 10.5-11.0% digestible pro-
tein. In reproducing does, crude protein from 17.5% to 19.0% and digestible
protein from 12.5% to 13.8% are recommended. These values correspond to
a ratio of 10.5 to 11.0 g DP/MJ DE for young rabbits and 11.5 to 12.5 g
DP/MJ DE for reproducing does. The higher values are recommended for
does under intensive breeding rhythms.
Dietary DP to DE ratios lower than recommendations significantly
decrease milk production and reproductive performance. In pregnant lac-
770
tating does, a reduction of the DP to DE ratio may decrease litter weight and
size, while has less effect on protein body balance (Table XXIII-9).
Table XXIII-9. Effect of DP/DE ratio on reproductive performance and composition of empty
body gain between the first and second kindling (XICCATO et al. 1992)
Even if abundant protein levels in reproduction diets are usually rec-

ommended, protein excess with unbalanced DP to DE ratios (>13 g DP/MJ
DE) can cause productive and sanitary failures in terms of inefficient regu-
lation in the energy voluntary intake of primiparous does and consequent-
ly negative effects on the suckling litters and doe body conditions (BARGE et
al. 1991).
23.2.2.2. Amino acid requirement
Recent literature on rabbit amino acids requirements is scarce, therefore,
amino acids levels actually recommended are still those provided in 1989
(Table XIII-16). Total amino acids requirements have been studied on the
base of dose-response trials or using the amino acid pattern in rabbit milk
and body in growing rabbits, giving less importance to the requirements for
maintenance and reproduction (Table XXIII-10).
Table XXIII-10. Amino acid composition of the whole body in growing rabbits and in rabbit
milk relative to lysine concentration (FRAGA, 1998).
The amino acids supply through caecotrophy, which occurs in rab-

bits, has been considered adequate for a long time to support all essential
771
amino acid requirements in rabbits, but is true under natural circum-

stances. The most limiting essential amino acid for breeding does is methio-
nine (and/or cystine), immediately followed by lysine and threonine. A min-
imum level of 5.4 g/kg of total sulphur-containing amino acids (4.0 g/kg of
digestible sulphur-containing amino acids) is required to obtain adequate
productivity in growing as well as in non reproducing rabbits, while a high-
er level (6.3 g/kg of total and 4.9 g/kg of digestible sulphur-containing
amino acids) is recommended for reproducing females to increase milk pro-
duction and to reduce the interval between parturitions, and improve feed
utilisation efficiency. Recommended levels of lysine (for lactation diets with
10.5-11 MJ DE/kg) are 6.8 g/kg total lysine (5.2 g/kg digestible lysine) for
the maximum reproductive performance and 8.0 g/kg total lysine (6.4 g/kg
digestible lysine) for maximum milk production (TABOADAB et al., 1994,
1996).
During the lactation peak (10 to 20 days), a minimum dietary con-
centration of 5.8 g/kg total threonine or 3.8 g/kg digestible threonine is
necessary to maximize feed intake and milk production. Optimum supply is
6.4 g/kg total threonine and 4.4 g/kg digestible threonine, while higher or
lower values impair both the number of weaned bunnies and feed efficien-
cy (DE BLAS et al. 1998).
23.2.3. Minerals, vitamins and additives

23.2.3.1. Macro-minerals
Lactating does have higher requirements for some minerals (Ca, P, Mg, Zn)
than growing rabbits or non-lactating does, related to their high output in
milk (FIGURE XXIII-14), besides their numerous structural and physiologi-
cal functions (Table XXIII-11).
Table XXIII-11: Macromineral recommendations (g/kg as fed) for lactating does according
to different authors
Even if high quantities of calcium (1.5 to 2 g/day) and phosphorous

(0.6 to 1 g/day) are excreted through milk, the mineral balance during lac-
tation is substantially in equilibrium or slightly positive by the end of lacta-
772
tion. The most critical period for calcium supply should be late pregnancy and
lactation. When lactation and pregnancy fully overlap each others, a slight
mineral deficit can arise. On the opposite, mineral excesses can induce sig-
nificant alterations in both doe fertility and prolificacy and may negatively
interact with other mineral absorption processes and the environmental
impact (e.g. P, heavy metals).
In reproducing does, 1.2% calcium supply is recommended. As
described above, the calcium absorption varies according to the amount of
dietary calcium and is not affected by metabolic demand. A high dietary cal-
cium percentage allows great calcium and magnesium absorption and
reduces phosphorous utilisation (KAMPHEUS, 1991). Calcium absorption
depends primarily on the apparent digestibility (absorption rate) of raw
materials, ranging from 81% for calcium carbonate to 53-54% for other
inorganic (calcium diphosphate, calcium oxalate) or organic sources (alfal-
fa, clover).
Finally, dietary calcium content is important for the potential inter-
action of this element with the toxic action of heavy metals such as the
reinforcement of the negative effect of cadmium on L-threonine absorption
(MESONERO et al. 1995), which holds importance for the teratological tests,
too.
FIGURE XXIII-14. Evolution of the main mineral milk composition during lactation (LEBAS
et al. 1971)
The too low level of phosphorous (>0.45%) does not affect reproductive
performance (number of pups born alive, number of pups weaned, etc.) or
female live weight. On the contrary, high phosphorous concentration
(0.76%) reduces the litter weight at weaning. Based on these results and to
limit the P environmental excretion, the recommended levels were reduced
from 0.70% to 0.55%. The origin of phosphorous is less important than it is
for calcium. In fact, phytic phosphorous coming from vegetal raw materials
773
shows the same utilisation efficiency of mineral phosphorous or even bet-

ter. Calcium to phosphorous ratio between 1.5-2 to 1 is recommended (In non
pregnant does, its variations do not affect feed intake, while in pregnant
does a low ratio (1:1) stimulates feed intake and a high ratio (2:1) improves
calcium retention, reduces phytic phosphorus hydrolysis, increases litter size
and reduces mortality at birth. As stated, calcium and phosphorous levels
should vary according to the reproductive rhythm: slightly higher levels are
advisable (1.3% Ca and 0.6% P) when intensive systems are used.
As far as other macrominerals are concerned, there is no new exper-
imental information that allows us to modify the practical feeding recom-
mendations proposed before (see Table XXIII). Only sodium (from 0.30% to
0.20%) and chlorine (from 0.35 to 0.30%) reductions has been proposed
(MAERTENS, 1995). Regardless from the high K milk concentration, no K sup-
plementation is required due to the relatively high supply from raw materi-
als currently used in rabbit feeding.
23.2.3.2. Micro-minerals
There is an absence in recent data in this field (Table XXIII-12). Available
contributions evaluated the effects of dietary supplementation with iron,
copper and the interaction between selenium and vitamin E on the repro-
ductive performances in rabbit does.
Table XXIII-12. Micromineral recommendations (ppm as fed) for lactating does
Despite the low iron concentration in rabbit milk, young bunnies are
not usually susceptible to iron deficiency; they show a good body iron con-
centration at birth, thanks to a suitable passage of this micro mineral by
placenta, which satisfy their needs until the beginning of solid feed intake.
Iron supplementation (80 mg/kg Fe as FeSO4) in diets at 129 mg/kg Fe pro-
duced positive effects on the reproductive performance with higher litter size
at birth and weaning and also decreased the number of lower stillbirth,
which, however, require further investigation. Recommended iron levels for
lactating does may vary from 30 to 100 mg/kg.
Recommended levels of copper vary between 5 and 15 mg/kg with
higher values for reproducing does. Copper deficiency is not likely to occur
in rabbits, since raw materials used for feed formulation are sufficiently pro-
vided with it. High copper levels such as sulphate (75 to 450 mg CuSO4/kg
of feed) used to be proposed as a growth promoter, like in poultry and pigs,
774
improving digestive utilisation of feeds, thus protecting young rabbits from

enterotoxaemia and reducing mortality. Copper has been demonstrate to
accumulate in the liver and kidney and not in the muscles, and excreted
through the bile in the faeces. Studies in growing rabbits do not always con-
firm the positive effect of Cu as a growth promoter (FEKETE et al., 1989), while
no experimental evidence exists for reproducing does. In the past Cu sup-
plementation caused great environmental pollution due its large supply in
pig feeding and now its use as an animal growth promoter is forbidden in the
EU.
A positive effect on foetal growth and litter weight at birth due to
dietary supplementation with selenium (0.1 mg/kg) was observed by STUK-
LEC et al. (1994), whereas the supplementation with 0.3 mg/kg determined
low reproductive performance, which could be a sign of approaching the lev-
els of toxicity. A supplementation of 0.7 mg/kg DM of Se and 40 mg/kg DM
of vitamin E in females under heat stress conditions lead to a higher con-
ception rate and litter size (EL-MASRY et al. 1994).
Others microminerals are considered less important to the extent
that they are usually provided with diets and caecotroph and no problems
linked to their excess or deficiency which are detected under normal rear-
ing conditions. In fact, minerals and vitamin premix are usually added to
commercial diets, which adequately satisfy mineral and vitamin require-
ments in both growing and reproducing rabbits.
Nowadays, studies on supplementation with microminerals or vita-
mins mainly deal with their effect under sub-optimal environmental and
rearing conditions (i.e. stressing thermal conditions, limited availability of
raw materials, low-quality raw materials). Under intensive and good-con-
trolled rearing conditions, rabbit requirements are usually satisfied and
major problems about mineral supplementation regarding the environmen-
tal safeguard and control pollution of animal origin.
23.2.3.2. Vitamins
Vitamin A supplementation in the diet is essential for both growth and
reproduction, but at high dosages this substance reduces reproductive effi-
ciency and produces teratogenic effects with the decrease of foetal growth,
bone abnormalities, early foetal resorption, late abortions, reduced litter
size, increased natal and post-natal mortality and hydrocephalus.
Regardless of its transformation into vitamin A, a specific and positive
action has been hypothesised for ß-carotene on reproductive performance
by some authors, which however has not been confirmed by others. (In the
background of the benevolent action the antioxidative effect can be hypno-
tized.) The rabbit is capable of protecting itself against ß-carotene excess by
decreasing the conversion efficiency on vitamin A in the gut wall, but is
unable to react against an excess of vitamin A added to the diet in the form
of retinol (esters, acetate, or retinol palmitate). The addition of vitamin E (50
775
mg/animal/day) or vitamin C (50 mg/animal/day) or ethoxyquin (25

mg/animal/day) in does fed on diets with an excess of vitamin A reduces
the frequency of reproductive dysfunctions.
Recommended levels of vitamin A for a lactation diet (10-12,000
IU/kg) are far below the toxic level (about 90,000 IU/kg) (Table XXIII-13).
Table XXIII-13: Vitamin requirements for lactating does
The optimum levels of vitamin D in breeding rabbits have not been

established yet and a range of 900-1000 IU/kg is recommended. Vitamin D
excesses (3,250 to 13,200 IU/kg) provoke the mineralization of the soft tis-
sues, anorexia, weight loss, and foetal mortality. Tissue calcification (aorta,
kidneys, ovaries etc.) is worsened by the contemporary dietary excess of cal-
cium. Under practical conditions, an excess of vitamin D is considered to be
more dangerous than its deficiency.
Recent information on vitamin E requirements in reproducing does
is missing, whereas the supplementation of 50 ppm is usually recommend-
ed, to be increased in case of impaired immunity or infections. No toxicity
due to the excess of vitamin E has been described, while its deficiency is
usually associated with the occurrence of muscular dystrophy as well as
impaired reproductive performance, in terms of increased infertility, abor-
tions and mortality at birth.
In intensive reproductive system the lack of vitamin K may lead to
haemorrhage in the placenta and consequent abortions.
The capability of intestinal microflora to synthesize sufficient quanti-
ties of water soluble vitamins (B-group, C, etc.) and their availability
through caecotrophy usually allow rabbits to satisfy their requirements. No
776
recent research about requirements of water soluble vitamins in doe breed-

ing conditions has been conducted and recommendations are mostly based
on current diet formulation and practical responses. Under intensive rear-
ing conditions, a positive response to a dietary water soluble vitamin sup-
plementation is sometimes observed. The supplementation of vitamins C
and E improves the reproductive response in rabbits subjected to stress
conditions (like high temperatures), reducing post-natal mortality and
slightly increasing the litter size and weight. Therefore a dietary supple-
mentation of 50-100 mg ascorbic acid can be recommended under stress
conditions.
23.2.3.4. Additives
In both reproducing does and weanling rabbits, live yeast supplementation
proved some positive effects on performance and health status, especially
when animals were kept under sub-optimal environmental conditions with
high stocking density and low hygiene control.
Various strains of Bacillus spp. were also tested with contrasting
results. The supplementation of B. cereus var. toyoi at 2x105 spores/g diet
in rabbit does and suckling rabbits tended to improve litter weight and lit-
ter size at weaning at 25 d. A similar positive effect on reproductive per-
formance was detected when diets for reproducing does were added with B.
subtilis and B. licheniformis (3×109 spore/g).
Little information is available on the use of enzymes in rabbit feed-
ing, especially when emphasis is put on reproducing does, showing no clear
positive effects.
777
23.2.4. Nutrition and feeding of breeding bucks
ubstantially less studies are available on nutrition and feeding of
S breeding bucks than those of does. The spread of artificial insemina-

tion has further reduced the ratio of breeding males to females ratio
(as far as 1 to 100) and interest in developing specific diets for bucks has
also declined. At the same time, increasing attention is given towards the
semen quality, both fresh and refrigerated, with special emphasis on the
effects of dietary supplementation with minerals, vitamins and fatty acids
(ALVARINO, 2000).
Feeding the bucks must satisfy their maintenance and growth,
requirements from the sexual maturity (around 5 months) until 7 months
of age, around which most breeds and hybrids complete their corporal and
sexual development. From this age onwards, only maintenance require-
ments have to be assured, while nutritional requirements for semen pro-
duction are not relevant.
Voluntary feed intake in males increases until 5 months of age then
reduces by 30% (MAERTENZ and LUZI, 2004). Restricted feeding (35 g/kg LW),
however, may be used to avoid over fattening in heavy breeds which could
cause sore hocks, thus reducing their productive life.
In terms of reproductive performance, an excessive restriction may
reduce libido and sperm output (ejaculation), but semen characteristics are
not affected (FODOR et al. 2003). Few evidences exist on the effect of reduced
dietary protein on libido and semen quality which are not in accordance
with the reproductive performance of the inseminated doe (Tables XXIII-14
and 15).
In adequately balanced diets, neither the supplementation with

essential amino acids (lysine and methionine), nor with vitamin premix
778
(vitamin A, D and E) may substantially modify quantity and quality of fresh

semen or relating reproduction performance (fertility, born/litter, born
alive/litter)
Over-nutritional levels of an association of vitamin E and vitamin C
reduce oxygen molecules and oxidative products and improved antioxidant
capacity both in the fresh and stored semen (CASTELLANI et al. 2000), but
have limited effects on semen quality and its fertilizing ability (CASTELLANI et
al. 1999; MINELLI et al. 1999).
The use of diets containing high levels of 18:3 (omega-3) fatty acids
(and supplemented with over-nutritional levels of vitamin E) improves the
quality of rabbit spermatozoa in terms of higher live cells, speed and move-
ments of spermatozoa
Under thermal stress conditions, dietary supplementations with vita-
min E (40 mg/kg DM) and selenium (0.7 mg/kg DM) increase blood testos-
terone and total spermatozoa concentration in males, while a further sup-
plementation with zinc (35 mg/kg DM) also stimulates semen volume, live
spermatozoa concentration and motility. Under heat stress conditions the
addition of 0.7 mg/kg DM of Se and 40 mg/kg DM of vitamin E leads to
the improved chemical and physical semen quality in bucks (EL-MASRY et al.,
1994).
The compilation of all the nutrient requirements of replacement
young female rabbits and breeding, adult does can be read in the part of
“Practical feeding and nutrition (Table XXIII-20.)
779
23.3. Practical rabbit nutrition
or the practical farming the nutrient requirements of four „rabbit cat-
F egories” have been formulated: adult, non-producing (buck, non-preg-

nant doe, culling breeding animal), pregnant, simultaneously preg-
nant and lactating and “growing“ (from month one till the slaughtering at
the age of about 2.5 month). There is also a special recommendation for the
replacement of young and adult breeding does. In extreme climatic condi-
tions (especially in hot and wet climate) values are not applicable.
23.3.1. Optimum composition of feed mixtures

Comparing the nutritional needs of the above mentioned four categories, it
is evident that the lactating (eventually also pregnant) doe requires the most
concentrated feed. The reason for that is that the daily milk production is
100-300 gram of milk, which is three times concentrated than the cow’s
milk (for more details see previous Section 2), whilst the energy reserves of
her body are limited. As the second on the „requirement list” plays the grow-
ing, and only after come the pregnant and the resting animal do. The adult,
non producing laboratory rabbits should receive feed according the values
of the pregnant ones. Though the doe do not produce but the experimental
procedures (e. g. blood sampling, weighing, body temperature measurement
etc.) requires some extra (micro) nutrients.
Table XXIII-16. gives the requirements of energy, major nutrients and
amino acids, Table XXIII-17 the macro minerals, the, Table XXIII-18 the
micro minerals and Table XXIII-19 the vitamin needs of the given four cate-
gories, related to air-dry feed mixture. Table XXIII-20 summarizes the spe-
cial values for young replacing females and adult breeding does. It should
be emphasizing that there is a separate item for minimum fibre requirement
(for details see later).
23.3.1.1. Supply of energy

One of the most important feeding behaviour of the rabbit is that the daily
voluntary feed intake is adjusted to the energy concentration of the feed. .
Of course the prerequisite of this regulation is a balanced ration (protein,
mineral and vitamin levels). The average daily digestible energy intake per
kg metabolic body mass (W0.75) is 0.9-1.0 MJ. Lactating does eat more: 1.2-
1.3, even 1.5 MJ/W0.75 (in peak lactation, d 15-20). The anatomy of the rab-
bit’s digestive tract allows this regulation to function within the limits of
feed energy density of 9.2-13.4 MJ DE/kg air-dry feed.
Parallel to the high energy density the protein, mineral and vitamin
level should be increased. Rabbits require essential fatty acids (linoleic,
780
linolenic and arachidonic acids) too. An average of 3-4% mixed fats and oils
of the usual diets satisfy the fatty acid requirement, therefore the fat/oil
supplementation can be justified only for energy enrichment. The measure
of the latter depends upon the energy density of the basic feed.
Paradoxically, in spite of its high fibre requirements, rabbits poorly
digest fibre. Consequently, the fibre does not play an important role in the
energy supply of the rabbit.
23.3.1.2. Role of indigestible organic matter

The indigestible organic matter (IOM) or ballast (bulk) has a great part in
the feeding and dietetics of rabbit. The indigestible organic matter is in fact
the indigestible part of the fibre fractions of the cell wall material. The form
of their expression (crude fibre, NDF, ADF, lignin) depends on the analyti-
cal procedure used. The recommended crude fibre level for growing 13-15,
for lactating rabbits 10-11% is. If the digestibility of fibre were too high. it
cannot fulfil its role as „indigestible organic matter”, then the crude fibre
concentration should be increased (e. g. in case of high dried sugar beet
pulp in the feed mixture). As a „rule of thumb” an approximately 10% „indi-
gestible fibre” level should be assured.
The amount and physical form of the indigestible organic matter does
influence the utilisation of the other feed components: parallel to the high-
er UOM the transit time of the digesta fasten up which causes a decrease in
the digestibility of energy supplying compounds. The decrease in protein
digestibility– owing to the compensation by caecotrophy – is significantly
smaller than in other monogastric animals (bird, pig). The optimum particle
size falls between the limits of 0.1 to 0.5 mm, both the finer and the larger
being disadvantageous. Lack of indigestible organic matter leads to diges-
tive troubles, diarrhoea and higher mortality.
23.3.1.3. Proteins and amino acids (AAs)

Investigations of the last decades elucidated the rabbit’s protein require-
ment. From the 21 AA 10 are essential and the glycine semiessential for this
species. One third of the sulphur containing AA may be cystine, the remain-
ing 2/3 should be methionine. (The same is true for the phenilalanine and
tyrosine). The average lysine and sulphur containing amino acid require-
ment is 0.6 while that of the arginine is0.8% in the air dry feed mixture. In
case of lysine and arginine the over dosage is not dangerous; the „thera-
peutic index” of the methionine is small, and higher dosages are harmful. If
the dietary protein contains all of the essential amino acids in available form
(„ideal protein”), than 15-16% crude protein level is sufficient for the grow-
ing rabbit. Using proteins of medium or low quality, however the total pro-
tein concentration should be set higher.
The minimum protein requirement of breeding does, expressed in
ideal protein is 17-18%. Giving more protein, the milk production can be
781
stimulated; using less than 16% protein decrease the litter weight at wean-
ing. It is worth mentioning that the protein deficiency or the amino acid
imbalance decreases the voluntary feed intake. Although theoretically – by
means of the caecal flora – rabbits are able to utilize NPN materials (urea,
ammonium salts, biuret etc.), at the protein levels of the practical nutrition
their application is useless, even harmful.
23.3.1.4. Minerals and vitamins

The calcium and phosphorus needs of the growing rabbit are significantly
less than those of the lactating does. Too high a ratio of sodium and potas-
sium to chloride (see cation anion balance) may cause kidney damage and
reproductive failures. There are controversial data concerning the antidiar-
rhoeal effect of the copper-sulphate supplementation.
By means of the caecotrophy rabbits can ingest sufficient amount of
vitamins (K and group B) for maintenance and medium intensive produc-
tion. For the fast growing young rabbits 1-2 mg/kg vitamin B1 and B6, 6
mg/kg vitamin B2 and 30-60 mg/kg niacin supplementation may improve
performance. Addition of ascorbic acid, except in heat and oxidative stress,
has no effect. Over dosage of vitamin A leads to toxicosis and that of vita-
min D to hypervitaminosis, thus the concentrations given in Table XXIII-19
should be considered!
23.3.1.6. Young females and breeding does

In Table XXIII-20 the special nutritional needs of the replacing young
females and adult breeding does can be found.
23.3.1.7. Consequences of the deviations from the recommendation

Lack of indigestible organic material may cause diarrhoea. Too much (more
than 12%. 9.3 MJ DE/kg air-dry feed) IOM is also disadvantageous diluting
energy density to such an extent that the rabbit cannot eat sufficient
amount of feed, worsening the average daily gain. Very high (over 20% crude
fibre may cause impaction of the caecum.
Table XXIII-21. shows the expected modifications of the production
parameters, if the nutrient concentration of the feed differs from those of the
recommendations. It is worth mentioning that a decrease in production
intensity (e. g. organic husbandry) is not necessarily uneconomic; it
depends on the actual financial environment. In case of a multiple deficien-
cy feed, the production consequences can be elucidated only by means of
feeding trial.
23.3.2. Feeding practice

23.3.2.1. Form of feed mixture
In order to insure homogenous mixing, the ingredients of the commercial
rabbit feed previously are grinded. However, rabbits have a strong aversion
782
toward the feeds of meal (flour) form. Thus, the small grinded and thor-
oughly mixed rabbit feed should be pelleted. The ideal pellet dimension for
rabbit is 3-5×10-12 mm in column shape. Table XXIII-7 shows the effect of
diameter of the 10-12 mm long pellets on feed intake, daily weight gain and
feed conversion efficiency.
23.2.2.2. Feeding technique, amount of daily ration
The classic feedstuffs keeping backyard rabbit keeping are the cereals, the
wheat bran and the roughages (in summer chopped grass, in winter mead-
ow and alfalfa hay). In wintertime the feeding of fodder beet and carrot is
also usual. More and more small breeder uses the combination of the com-
mercial pelleted feed and natural ingredients (green grass, lettuce, cabbage,
carrot, cereal grains etc.) In case of intensive reproductive cycle all types of
animals – except the breeding bucks – are fed ad libitum. In semi-intensive
reproductive technology the breeding does are recommended to be fed on a
restricted ration (30-35 gram dry matter/kg LW/day) from weaning till the
next kindling using the mixture of the pregnant animals (category II in Table
XXIII-16-19). Growing broiler rabbits are continuously allowed to eat ad libi-
tum. The number of drinking sites should allow the sufficient water intake,
i.e. twice the dry matter intake. (If the water supply shortened, rabbits
decrease their voluntary dry matter intake.)
For practical predictions see the following AVERAGE DATA OF PELLETS
INGESTION:
-growing (broiler: week 4-11): 110-130 gram/day;
-suckling doe (together with her litter till day 28): 350-380 gram/day;
-adult, maintenance: 120 gram/day and
-“pen—mother“ unit: 1-1.4 kg/day.
To produce 1 kg LW (included the maintenance of breeding animals)
requires approximately 4 kg air-dry pellets.
23.3.3. Characterization of important rabbit feedstuffs

The energetic value of the rabbit feeds are given in digestible energy (DE),
expressed in megajoule (MJ). The protein needs – supposed to be at least
70% digestible – is given in crude protein. In the following the recommend-
ed incorporation ratio of the different feedingstuffs is shown.
The amount of OATS is not limited, even the whole cereal ratio may
consist of oats without any harmful side-effect. BARLEY is used for the
replacement of oats. Proposed amount of WHEAT in the feed mixture is 10-
40%, its starch is more digestible for rabbits that that of the corn. CORN may
enhance fat accretion worsening the carcass quality. It is poor in fibre, pro-
tein, essential amino acids and calcium, thus, requires supplementation.
Maximum inclusion level for growing is 30-35, for breeding rabbits 20-25%.
RYE grains have low fibre and contains antinutritive pentosane compound,
therefore they have obstipation effect. Maximum incorporation level is 10-
25%. The SORGHUM can be used similar to barley. High tannic acid contain-
783
ing sorghum sorts can cause constipation in rabbits. MILO grains can be
used in 10-15%. The DRIED SUGAR BEET PULP is recommended in 5-25%;
owing to its soluble fibre content has an excellent dietetic effect (prebiotics),
minimizing the occurrence of diarrhoea.
PEA and BEAN may provoke bloat or flatulence, the upper incorpora-
tion level is 10-15%. The SWEET LUPIN seed having balanced protein and fibre
content, may be fed up to 30%.The ratio of RICE BRAN may be a maximum of
15-20% because its silica content is limiting, decreasing protein digestibili-
ty. It is prone to get rancid. Extracted SUNFLOWER MEAL – according to its fibre
level – may vary between 3-35%. It is rich in sulphur containing amino
acids; its protein has a high biological value. The SOYBEAN MEAL can be mixed
up to 30% in the feed. The main reason for using of LINSEED MEAL in some
percentages is its excellent dietetic effect. The ALFALFA MEAL – according to
its protein and fibre level – usually gives 20-60% of the pellets. Yeast (if reg-
ulatory is not prohibited: milk powder or fish meal) may also be used in 1-
3%.
As a source of indigestible organic matter, the good quality STRAW,
CORN STALK and WHOLE CORN PLANT meal, or the wheat of rye GREEN MEAL is
proposed. The 1-3% of MOLASSES or sugar, besides its good taste and energy
content, improves the consistency of the pellets. For this goal one can use
methyl-cellulose or lignin-sulphonate too.
Small breeders may use for lactating does in 20-40 grams/day the
fresh BIER and APPLE BY-PRODUCT (HUSKS O POMACE) and for each age category
of the dried BREAD AND ITS CRUST (fibre supplementation is necessary!!!!).
After certain adjustment (transition/habituation) the feeding of BOILED POTA-
TOE is also applicable.
In case of GREEN FEEDSTUFFS (acacia foliage, different grasses, young
green corn plant, green alfalfa chops), HAYS, FODDER BEET AND CARROT the
upper limit is given by the intake capacity of rabbits.
784
785
786
787

23.4.1. Troubles with digestion

Carbohydrates in rabbit feed can be divided into two groups: the readily
available carbohydrates (RAC. e. g. sugar, starch) and those, participating
in fibre structure. Feeding too much RAC increases the incidence of entero-
toxaemia. This can be explained by the carbohydrate overload of the hind
gut, which is in actual fact, a fermentative dysbiosis. The latter will devel-
op, if the feed mixture contains high amount of starch (e. g. corn grain) and
part of it cannot be digested in the small intestine and will serve as nutri-
ent for the caecal bacteria. Too small (fine) particle size facilitates the
process. Pathologic fermentation in the caecum favours the multiplication
of pathogenic bacteria (among others for the iota toxin producing
Clostridium spiroforme). The absorbed iota-toxin will cause lethal entero-
toxaemia (hemorrhagic inflammation of the caecum, watery diarrhoeas
causing exsiccation, paralysis of the central nervous system). Increasing the
fibre level up to 12-15% minimized the occurrence of the disease. It is not
clear, that the effect derives from the energy diluting (starch concentration
decreasing) influence (“per se” effect), or from the specific characteristics of
the fibre. Nevertheless, it is highly probable that it is not sufficient to mini-
mize the presence of starch, but also important are the fibre fractions.
(Efforts to prevent the enterotoxaemia by buffer – sodium bicarbonate, zeo-
lites, MgO – supplementation had trifle or negative results.)
The sufficient “indigestible fibre” is important not only to prevent diar-
rhoeas, but also in maintain optimum growth rate and to exclude fur chew-
ing. It is highly important to differentiate the effect of fibre on the digestibil-
ity of nutrients and its general, positive dietetic influence on the whole
organism. The sum of the dietetic effects (maintenance of the healthy peri-
stalsis of the gastro-intestinal tract, adsorption of toxin in the gut lumen,
assuring the natural feeling of fulfilment, the so-called feeding comfort, giv-
ing substrate for the microbial fermentation and volatile fatty acid produc-
tion in the caecum) up to a certain fibre level (20% crude fibre) may be high-
er than the negative consequences of the decreased digestibility of nutri-
ents. Over 22% of crude fibre the incidence of caecum impaction and con-
stipation is higher.
Fibre deficient diets – especially at angora rabbits – facilitate the for-
mation of gastric hair ball (zootrichobezoar) (FIGURE-XIII-15).
788
FIGURE XXIII-5: Rabbit stomach with the, from anthral part originated hairball
(FEKETE and BOKORI, 1985)
To prevent mortality due to diarrhoea the following minimum fibre

levels are recommended: 15.3% ADF (acid detergent fibre), 11.7% crude
fibre and 11.7% indigestible fibre. The required fibre level should be set by
combining the natural ingredients or by adding straw meal.
Above 24.6% ADF concentration the live weight gain decreases. Each
per cent ADF in the feed mixture decreases the digestibility of dry matter (-
1.17% unit), organic matter (-1.32% unit) and crude protein (-0.64% unit). at
higher fibre level even the digestibility of crude fibre will be lower. feed com-
position also influences the volume and mass of the stomach (parallel to the
high fibre-low starch diet the intake will be larger); lower than 11.7% ADF
causes the enlargement of the caecum, because caecotrophy is not prac-
tised. The starch has no separate dietetic effect: in case of a sufficient fibre
supply even 25% starch has no harmful effect on the digestive processes.
After fasting the normally 67-72 mmol/kg caecal content VFA level dramat-
ically declines to 10 mmol/kg.
The pathogenesis of the diarrhoea of weaning rabbit may also be
explained by the high alkali value of the solid feed. (ALKALI VALUE OF A FEED
is equivalent to the amount of HCl (in mmol/kg) which is able to set the pH
of 1 kg feed on pH 3.) Too alkaline diet means that the bicarbonate overload
is increasing the blood. i. e. HCO3-concentration, which, in turn, will be
excreted into the caecum. The developed high caecal pH (6.5 instead of the
6.0.-6.2) attenuates the local antibacterial effect. The final result may be the
multiplication of pathogenic bacteria (e.g. E. coli), diarrhoea, death
(PROHASZKA and BARON, 1982). Instead of the optimum alkaline values most
of the feeds have a value of 500-700. The bacterial imbalance in the caecum
and the intake of feed high in protein may lead to the putrid dyspepsia.
The supplementation of the drinking water with 20 ml/l acetic acid,
789
or 20 ml/l lactulose (LACTULOSE=1,4-fructos-galactosid, a non-fermentable

and indigestible disaccharide, able to bind the NH3 in the gut lumen) may
decrease mortality rates; the lactic acid proved to be ineffective. Animals,
drinking water with acetic acid, had a lower pH in the caecum (instead of
6.47 – 6.19). The described protective effect was even sufficient in case of
ampicillin provocation. Actually, the supplementation of drinking water with
18.75 mg/100 ml ampicillin (or other broad spectrum antibiotics) for some
days will cause diarrhoea and mortality among the weaning rabbits. The
intake of antibiotics was approximately 10 mg/rabbit/day. Ampicillin acts
by upsetting the intestinal balance of the bacteria.
23.4.2. Feed toxicity

The pathogenesis of the peroral ANTIBIOTIC-ASSOCIATED ENTEROTOXAEMIA (ampi-
cillin, linkomycin, clindamycin etc.) (diarrhoea) can be explained by the
reduction of number of the predominantly Gram positive anaerobic bacteria
(in the majority Bacteroides). Parallel to their lower number the concentra-
tion of the produced non-dissociated volatile fatty acids, and the antimicro-
bial effect falls. (This antibacterial effect is the strongest in pH range
between 6.0-6.5, with a volatile fatty acid concentration of 100-2000
mmol/kg caecal content.) Thus, the use of antibiotics (especially penicillin
family) can change the intestinal flora and disrupt their natural balance.
Reduced competitive protection of indigenous intestinal flora permits the
overgrowth of pathogenic E. coli, S. typhimurium, Proteus, Shigella, Klebsiella
and clostridia) and proliferation of their toxins, resulting in fatal enteritis
and diarrhoea.
To the antibiotic NARAZIN (ionophor type coccidiostatics) rabbits are
extremely sensitive, the LD50–value is low, 11.9 mg/kg LW (OSZ et al. 1988).
In practice toxicosis may occur when poultry premix is added to the rabbit
feed: After the compound is taken in, uncertain, uncoordinated movement
and the flaccidity of the limbs develop. In some cases neurological signs
(tonico-clonico cramps, spastic contractions, salivation and the side-posi-
tion of the head) appear and some of the animals will die. Necropsy reveals
pale fields in the heart and muscles (Zenker-type myodegeneration, necro-
sis of the myofibrils) and lympho-histiocytic infiltration.
23.4.3. Mycotoxicosis
Mycotoxicosis are diseases, caused by toxins (“mycotoxins”) produced by
fungi (moulds) and ingested mostly by the feed. The best known toxins are
the AFLATOXINS (B1, B2, G1 and G2), produced by the Aspergillus flavus and
A. parasiticus fungi. Intake of feed, containing 350 ppb (ppb=µg/kg) alfa-
toxin B1 may cause chronic aflatoxicosis in growing rabbits. The decline in
feed intake is 30, in weight gain 70%. After 3 weeks the first mortality cases
occur, and at day 45 it reaches 20-30%. Sick animals lose weight, practise
fur chewing and the number of leukocytes fall. Necropsy revealed hepatosis
790
and splenomegalia. Subacute B1-alfatoxicosisra (0.06-0.09 mg/kg LW/day

per os) can be characterized by diffuse hepatic necrosis, disturbed blood
clotting and disseminated intravascular coagulation. Phenobarbitals, CoCl2,
piperonil-butoxid and cysteine have some preventive, the oxitetracyclin some
curative effect.
Ingestion of CITRININ, produced by Penicillium and Aspergillus spp.
Develops diarrhoea and nephrosis in rabbits.
The T–2 TOXIN (and its metabolites) is produced by the Fusarium
sporotrichioides, F. poae and F. tricinctum mould species. The damage of
brain tissue is characteristic for the fusariotoxicosis of rabbit (and horse).
Ingestion of feed of 0.024% VOMITOXIN (DEOXINIVALENOL, DON) causes 100%
embryonic mortality; T–2 toxin is excreted by the milk, causing death of
suckling pups. GLAVITS and co-worker (1988) in acute and subacute peroral
test have found the necrosis of lymphoid cells in the mucous membrane of
the small intestine and the depletion of “B” and “T” dependent zones of lym-
phoid organs (spleen, lymph nodes, bone marrow, sacculus rotundus). The
necrosis of MPS (mononuclear phagocyte system) cell in the liver is propor-
tional to the amount of ingested toxin. The LD50-value of T–2 toxin in rabbit
is between 1 and 4 mg/kg LW. It means a high sensitivity towards trichote-
cen-structured mycotoxins, thus the rabbit may serve as a good model for
their research. Long-lasting subtoxic peroral intake (1 mg/kg LW/day)
attenuates the capacity of the liver to metabolize and the sensitivity of the
organism increases. Feeding pellets of subtoxic (12.5 and 25 mg T–2
toxin/kg DM) decreases the voluntary feed intake by 60-70% (similar to the
pig’s feed refusal). The water content of the faeces from animals under toxin
influence increases by 10%. The T–2 toxin (and its metabolites) concentra-
tion in the urine, faeces and caecotrophy are proportional to the intake;
consequently, from the analysis of the mentioned biological samples the
mycotoxin state of a group of animal can be evaluated. (FEKETE et al. 1988).
By reingestion of the caecotrophy the toxin gets once more into the stomach of
the rabbit, which may explain the high sensitivity of this species to the myco-
toxins. The intake of feed with a very low amount (0.19 mg/kg feed) of T–2
toxin during some weeks, in female rabbits will decline the progesterone
production of the yellow bodies in the ovaries (FIGURE XVIII-1.) nd may also
lead to the collapses of the immune functions (FEKETE and HUSZENICZA,
1993).
FOR FURTHER READING

Besenfelder, U., Solti, L., Seregi, J., Muller, M. and Brem, G. (1996): Different roles of β-
carotene and vitamin A in the reproduction of rabbits. Theriogenology 45, 1583-1591.
Castellini, C., Dal Bosco, A. and Mugnai, C. (2003): Comparison of different reproduction
protocol for rabbit does: effect of litter size and mating interval. Liv. Prod. Sci. 83,
131-139.
791
Cheeke, P.R. (1987): Rabbit nutrition and feeding. Academic Press, Onc. Harcourt Brace
Jovanovicch Publisher. Orlando – New York – London – Tokyo
DeBlas, C. and Wiseman, J. (1998): The nutrition of the rabbit. CABI Publishing,. Oxon
Fekete, S. and Bokori, J.(1985): The effect of the fiber and protein level of the ration upon
the cecotrophy of rabbit. J. Appl. Rabbit Res. 8, 68-71.
Fekete, S. and Gippert, T. (1986): Digestibility and nutritive value of 19 more important
rabbit feedstuffs. J. Appl. Rabbit Res. 9, 103-108
Fekete, S. and Huszenicza, G. (1993): Effects of T–2 toxin on ovarian activity and some
metabolic variables of rabbits. Lab. Anim. Sci. 43:646-648.
Fekete, S. Gy., Hullar, I., Romvari, R., Andrasofszky, Emese and Szendro, Zs. (2005): Study
of the energy and protein balance of pregnant rabbit does using two comparative
methods. Acta Vet. Hung.
Fekete, S.-Tamas, J.-Vanyi, A.-Glavits, R. and Bata, A.(1989): Effect of T–2 toxin on feed
intake, digestion and pathology of rabbits. Lab. Anim.Sci. 39, 603-606.
Fodor, K., Fekete, S. Gy., Zoldag, L., Bersényi, A., Gáspárdy, A., Andrásofszky, E., Kulcsár,
M. and Eszes, F.(2001): Influence of feeding intensity on corporeal development,
body composition and sexual maturity in female rabbits. Acta Vet. Hung. 49, 399-
411.
Fortun-Lamothe, L., Prunier, A., Bolet, G. and Lebas, F. (1999) Physiological mechanism
involved in the effects of concurrent pregnancy and lactation on foetal growth and
mortality in the rabbit. Liv. Prod. Sci. 60, 229-241.
Fraga, M.J., Lorente, M., Carabaño, R.M. and de Blas, J.C. (1989): Effect of diet and remat-
ing interval on milk production and milk composition of the doe rabbit. Anim. Prod.
48, 459-466.
Harcourt-Brown, F. (2002): Textbook of rabbit medicine. Butterworth-Heinemann. Oxford
– Auckland – Boston – Johannesburg – Melbourne – New Delhi
Lebas, F. (2000): Vitamins in rabbit nutrition: literature review and recommendations.
World Rabbit Sci. 8(4), 185-192.
Nizza, A., Di Meo, C. and Taranto, S. (2000): Effect of lysine and methionine on libido and
semen characteristics of bucks. World Rabbit Sci. 8(4), 181-184.
Parigi Bini, R., Xiccato, G., Cinetto, M. and Dalle Zotte, A. (1992) Energy and protein uti-
lization and partition in rabbit does concurrently pregnant and lactating. Anim.
Prod. 55, 153-162.
Partridge, G.G., Lobley, G.E. and Fordyce, R.A. (1986): Energy and rabbit metabolism of
rabbits during pregnancy, lactation and concurrent pregnancy and lactation. Br. J.
Nutr. 56, 199-207.
Pascual, J. J., Cervera, C., Blas, E. and Fernández-Carmona, J. (2003) : High-energy diets
for reproductive rabbit does: effect of energy source. Nutrition Abstracts and
Reviews, Series B: Livestock Feeds and Feeding 73: 27R-39R.
Rommers, M.J., Meijerhof, R., Noordhuizen,J.P.T.M. and Kemp, B. (2004): Effect of feeding
program during rearing and age at first insemination on performance during sub-
sequent reproduction in young rabbits. Reprod. Nutr. Dev. 44, 321-332.
Vetesi, F. (ed.) (1990): Rabbit pathology and health (in Hungarian). Mezogazdasagi Kiado.
Budapest.
Xiccato, G., Parigi Bini, R., Dalle Zotte, A., Carazzolo, A. and Cossu, M.E. (1995) Effect of
dietary energy level, addition of fat and physiological state on performance and
energy balance of lactating and pregnant rabbit does. J. Anim. Sci. 61, 387-398.
Xiccato, G., Trocino, A., Sartori, A. and Queaque, P.I. (2004): Effect of doe parity order and
litter weaning age on the performance and body energy deficit of rabbit does. Liv.
Prod. Sci. 85, 239-251.
792
Chapter XXIV
HORSE NUTRITION AND DIETETICS
24.1. Digestive characteristics of horses

24.2. Nutrient requirements of horses
24.2.1. Dry matter intake
24.2.2. Energy and protein
24.2.3. Minerals
24.2.4. Vitamins
24.2.5. Essential fatty acids
24.2.6. Water
24.3. Practical nutrition of healthy horses
24.3.1. Principles of ration formulation
24.3.1.1. Roughage portion
24.3.1.2. Concentrated feeds
24.3.2. Feeding technique
24.3.2.1. Feeding frequency (incidence of feed distribu
tion)
24.3.2.2. Feeding sequence (order of offered feeds)
24.3.2.3. Feeding length (duration of feed intake, eating
interval)
24.4. Realisation of the practical nutrition of the healthy horse
24.5. Horse dietetic
24.5.1. Dietetic of special (patho)physiologic states
24.5.1.1. Raising of orphan foals
24.5.1.2. Geriatrics: feeding and nutrition of the old
horse
24.5.1.3. Insufficient voluntary feed intake.
24.5.2. Feeding Horses with Specific Disease
24.5.2.1. Starvation
24.5.2.2. Hoof defects
24.5.2.3. Recurrent myopathy
24.5.2.4. Recurrent airway disease
24.5.2.5. Diarrhoea
24.5.2.6. Hepatic dysfunction
24.5.2.7. Chronic renal failure
24.5.2.8. Urinary calculi
24.5.2.10. Intestinal colics (FIGURE XXIV-4)
24.5.3. Dietetics of miscellaneous diseases
CHAPTER XXIV: HORSE FEEDING AND DIETETICS
24.1. Digestive characteristics of horses

hile compiling a feeding evaluation system for horses, the digestive
W characteristics should be deeply considered. The horse is one of the

caecal-colon fermentor herbivore and as such combines some of
the advantages of the strict monogastric animals and those of ruminants.
Failure to understand the anatomy of the gastro-intestinal tract and basic
digestive physiology is the root of many feeding errors. The digestive process
begins with seizing feedstuffs, which are supplied as fresh pasture, long hay
or variation of stored forage and as concentrate feed containing processed
or unprocessed cereal grains. In order to properly start the digestive process
the horse needs a sound, functional mouth. This means that the teeth
should be in good conditions with functional occlusive surface. The adult
female horse has 18 upper and 18 lower teeth consisting of six upper and
six lower incisors and 12 upper and lower molars. In stallion and gelding
there are also the 2 lower and 2 upper dens caninus and sometimes one
upper and one lower extra praemolar (the so called “wolf’s tooth”, P0) too.
The function of the teeth is grasping, gathering and chewing of feed. The
teeth participate in the digestive process primarily by reducing the particle
size of feed. Chewing also stimulates the flow of saliva which may initiate
chemical digestion of feed and lubricates feed prior to its passage to the
remainder of the tract. Dental problems resulting from wear and genetic
defects interfere with dental function. Most of the time horses that waste
feed, bolt their feed or are finicky eaters have dental anomalies, which once
corrected, allow the horse to utilize feed more efficiently. Horses should
have annual dental exams and should have their teeth floated to sharp
edges be detected on the upper (outside) or lower (inside) molar arcades.
Older horses which through normal wear have lost functional dentation or
younger horses with dental problems may benefit from use the of pelleted
concentrate rations since the particle size has been mechanically reduced
in the pelleting process.
The next section, the oesophagus is a muscular tube connecting the
mouth and the stomach. Feed is forced down the oesophagus by peristaltic
waves or muscle contractions. Saliva produced by secretory glands in the
794
mouth lubricates the feed and prevents the feed from becoming lodged in
the oesophagus. Therefore, it is actually rare for the horse to choke or for
oesophageal obstruction to occur. The horse’s stomach is relatively small
when compared to the capacity of the entire tract. The horse really evolved
as a continuous grazer and its better equipped to utilize small frequent
meals rather than large meals of concentrates. Rate of passage through the
stomach is such that the ingesta remains in the stomach only for a very
short time. The pH of the stomach is quite acidic. Although the major
impact of the stomach on the digesta is acid hydrolysis and enzymatic
digestion of protein, there is a significant amount of lactic acid produced
from the fermentation of soluble sugars by microbes located in the fundic
region of the stomach (pH=5.4 vs. pyloric pH=2.6). The chyme (ingesta)
passes from the stomach into the small intestine.
The small intestine is composed of the duodenum, the jejunum and
the ileum. The small intestine receives a continuous flow of pancreatic juice
which increases due to nervous and hormonal factors associated with the
meal. The small intestine is, or should be if the feeding program is proper-
ly designed the major zone of absorption for simple sugars derived from
starch digestion, for amino acids from protein digestion, for free fatty acids
resulting from digestion of the lipid component of the diet, or fat soluble
vitamins, A, D and E and some minerals. The indigested part of gut content
passes through the ileo-caecal orifice into the caecum.
The caecum and large colon are analogous to the rumen and reticu-
lum of the cattle and contain milliards of bacteria and protozoa which serve
in a symbiotic relationship with the horse enabling the digestion of cellulose
and other fibrous fractions of the feed. In addition, the caecal and colonic
microbes synthesize B vitamins and vitamin K which are available to the
horse to help meet the requirements for these nutrients. The produced
acetic acid will be absorped through the gut wall contributing to the energy
supply of the host animal. There is a significant amount of microbial pro-
tein synthesized in the horse’s hindgut. Whether the microbial protein is
available to horse to contribute to meeting amino acid requirements is still
somewhat controversial. Anyway, in some circumstances (suckling foal,
protein deficiency) by means of the coprophagy the microbial protein can
really be utilized by the horse. The rectum is the posterior part of the diges-
tive tract and serves primarily as a storage area for faecal products which
have not been digested. This residue is held in the rectum until sufficient
material accumulates which then results in nervous stimulation and void-
ing of faeces through the anus (defecation). FIGURE XXIV-1 vizualizes the
main digestive processes of horse
795
FIGURE XXIV-I: Schematic presentation of the horse digestive system and basis process-
es.
The feed intake is strongly regulated. In case of a balanced ration,

the average daily dry matter intake is 2 kg/100 kg body weight.
Nevertheless, this figure can be modified by a series of circumstances, such
as the roughage-concentrate ratio, the age, physiological and reproductive
status of the animal, the rearing intensity and the type of work. The daily
dry matter intake ranges from 1.5 to 3.0% of the live weight (LW).
Generally (except the French net energy system, the “unité fourragère
cheval”, UFC) the nutrient requirements of horses and the nutritive value of
feedstuffs for horses are expressed in digestible energy (DE), expressed in
megajoule or Mcal (1 Mcal= 4.184 MJ) and digestible crude protein (DCP).
For the purposes of the official feed control, the use of the crude protein (CP)
is also allowed (HUNGARIAN FEED CODEX, 1990).
796
24.2. Nutrient requirements of horses

© Annette Zeyner
utrient requirements are derived from the needs for maintenance
N plus growth, reproduction, lactation and physical activity. The rec-

ommendations include an additional charge for extra needs that can-
not be surely calculated (e.g. from different climate conditions or sponta-
neous activity of growing horses). Recommendations are given for energy,
protein, selected minerals and vitamins. In some cases (lactating mares,
growing horses) it is recommended to supply individual essential amino
acids. Above this, horses need water and essential fatty acids, too.
24.2.1. Dry matter intake

To create an appropriate ration it is necessary to be informed about the vol-
untary feed intake. Feed intake patterns should be expressed on dry matter
basis. Table XXIV-1 informs us about the mean and maximal dry matter
intake of horses. This depends on the animal’s body weight (LW) and ener-
gy need. Horses from smaller sized breeds consume relatively more dry mat-
ter than horses of large-sized breeds. As a tendency, feed consumption is
positively related to the energy need. Sucking foals consume an especially
high amount of feed, including milk. Pregnant mares in the last weeks prior
to birth are able to consume always up to 2 kg/100 kg LW/day. In situa-
tions of extremely high energy needs, e.g. for strenuous exercise, the feed
intake may temporarily not be able to meet the increased demand. In prac-
tice, the realised feed intake will be limited by additional patterns like
palatability and physical characteristics of the feed as well as the mental
state of the horse.
Table XXIV-1: Mean and maximal dry matter intake in horses (in kg per 100 kg of LW an
day (BOULOT, 1987)
1 quite more in sucking foals
797
24.2.2. Energy and protein

Recommendations for energy and protein may be expressed as ‘digestible
energy’ (DE) and ‘digestible protein’ (DP). To maintain adult horses 0.6 MJ
DE per kg of metabolic body size (W0.75) are recommended. For horses of
extensive breeds (e.g. Shetland pony, Exmoor pony, Dartmoor pony), an
energy recommendation that is approximately 10% lower may fit better
because these horses exhibit energy-sparing attributes like a calm tem-
perament as well as an especially thick coat and firm hair coat. Related to
the metabolic body weight, growing horses have age-dependently a higher
energy need for maintenance than adult horses. Regarding the protein need
for maintenance, 3 g of crude protein per kg of W0.75/day are recommend-
ed in adult horses. Recommendations for DE and DP are given in Table
XXIV-2.
Table XXIV-2: Recommendations for the daily supply of energy and protein in horses
(according to GfE, 1994)
To supply horses properly, an optimal relationship between protein

and energy in the total ration is necessary. This can be expressed as PEQ
(protein-energy quotient, in g DP/MJ DE). In rations for horses at mainte-
nance level the PEQ should amount to 5. Because a lot of feedstuffs contain
relatively more protein, the PEQ tends to be higher. This may be tolerated
798
to a large extend, but horses should be prevented from receiving excessive

protein supply because this may place a huge burden on the kidneys and
liver and consequently on the energy balance. The excretion of water, min-
erals and nitrogen will increase, too. In this way, more ammonia may be
released to the environment and can irritate the respiratory tract.
It is highly complicated to measure the work load of horses.
Therefore, it may be more practicable to subdivide it according to a scheme
into light, moderate and heavy. In general, the energy need for physical
activity rose with increasing velocity and proportion of vertical activity (gal-
lop and jump). The need over maintenance may amount up to 10, 25 to 50
and more than 50 to 100% for light, moderate and heavy work, respective-
ly. An extremely high work load (e.g. distance riding) may involve an energy
need that lies above 100%. The needs for energy and protein in working
horses increase in parallel. Thus, the PEQ for working horses remains con-
sistently at 5 like at maintenance level.
Pregnancy in broad mares can be subdivided into early pregnancy
until the 7th month and an advanced stage of pregnancy starting from the
8th month. However, the mare gains on weight significantly always from the
5th month of pregnancy. This weight gain seems to be mainly induced by
the retention of water and nutrients within foetal membranes and placental
fluid. Until the end of early pregnancy the foetus itself reaches only 17% of
the birth weight (MEYER and AHLSWEDE, 1976). First when the mare becomes
heavily pregnant, pregnancy induces a relevant extra need for nutrients and
energy. The mares’ body weight at birth should amount to 17 to 18% above
the barren status (13 to 14% are due to the development of the foetus,
uterus, mammary gland, foetal membranes and placental fluid. The remain-
ing 4% extra-genitally stored nutrients are at a physiologically necessary
level (MEYER and STADERMANN, 1991). To realise this, the energy need
increases by 25 and 40% between the 8th and 11th month of pregnancy,
respectively.
Marginal energy supply to the heavily pregnant mare will not influ-
ence the body weight of the foal at birth, but may elongate pregnancy.
Opulent energy intake during advanced pregnancy can contribute to the
foals’ body weight, but may also cause obesity in the mare resulting in dis-
turbed birth, conception rate and feed intake. Pony mares should be strict-
ly prevented from obesity developed throughout pregnancy because obese
mares ingest lower feed during late pregnancy and ponies are especially
endangered to develop fat mobilisation syndrome and to suffer from hyper-
lipidaemia. During advanced pregnancy the protein need increases by 42
and 80% in the 8th and 11th month, respectively, and thus faster than the
energy need. In this way, the PEQ rose from 5.7 during the 8th month of
pregnancy until 6.4 prior to birth. Practical feeding conditions seem to
ensure the necessary supply of essential amino acids to pregnant mares.
However, mixed feed for lactating mares that mostly contains soybean meal
799
to add lysine can always be given in heavily pregnant mares because feed
change is to be prevented in the peripartal state.
Lactation caused an especially high extra need for energy and pro-
tein. The milk yield increases until the 3rd month post partum. In contrast,
the content of energy, fat and protein in the mare’s milk decreases consis-
tently after birth. When lactation reaches its maximum, the need for ener-
gy and protein attains twice and 3.5-times, respectively, of the maintenance
level. Furthermore, the PEQ is especially high, 9.4. Until the 5th month of
lactation the energy and protein need amounts to only 1.65- and 2.40-times
the maintenance level (PEQ 7.4). In most cases foals were weaned at this
time (they are 5 to 6 month old). Lactating broad mares do not only have a
specific protein need but also a special need for lysine that can often not be
met by common rations. The lysine need per litre of milk amounts to 3 g
(BOCHRÖDER und SCHUBERT, 1994). This must be added to the need for main-
tenance (0.19 g/W0.75: NRC, 1989). When lactation reaches its maximum,
a warm-blooded type mare (600 kg LW) may give 20 litres of milk and, thus,
need 83 g of lysine per day. This can only be supplied by applying of lysine-
rich feedstuffs like green grass, soybean meal or directly lysine as feed addi-
tive. High amounts of skimmed milk powder are contraindicated in adult
horses because it may induce osmotic diarrhoea. Marginal amounts of
lysine in the feed do not reduce the lysine content of the milk, only the milk
yield.
In growing horses, the need for energy and protein depends strictly
on the growth intensity as well as the type of housing and, thus, the ability
to move spontaneously. Recommendations given in Table XXIV-2 refer to a
medium growing intensity and a keeping system where ample spontaneous
activity is allowed. Medium growing activity means that the young horse
reaches 60% of its final body weight when it becomes 12 month old. While
the horse is growing, the retention of energy per unit of body weight increas-
es while that of protein decreases. Therefore the energy need rises in
absolute terms, but there is a drop relative to the metabolic body mass.
Excessive energy does not increase the animal’s final withers height, but
weight may increase faster than height. Intensive growth combined with low
spontaneous activity obviously one of the triggering factors of developmen-
tal orthopaedic diseases, the DOD (VAN WEEREN and BRAMA, 2003). Growing
horses start with a considerable protein need after birth that decreases as
the young animal becomes older (Table XXIV-2). Besides protein a special
need for essential amino acids should be taken into consideration (Table
XXIV-3). During the sucking period these may be supplied mainly via the
mare’s milk. When the foals become older essential amino acids must be
provided in increasing proportions and after weaning totally by the solid
feed. For this reason, the foal must be trained to consume significant
amounts of solid feed and the mixed feed should contain enough essential
amino acids (lysine, sulphur-containing amino acids, and threonine). A pre-
800
liminary ratio of lysine to methionine+ cystine to threonine can be recom-

mended to be 1:0.6:0.6. Grain protein is relatively rich in sulphur-contain-
ing amino acids, but especially poor in lysine. Thus, grower feeds contain
commonly soybean meal or skimmed milk powder.
Table XXIV-3: Recommendations for the daily supply of essential amino acids to growing
horses (NRC, 1989; SAASTAMOINEN and KOSKINEN, 1993, GfE, 1994)
24.2.3. Minerals
Minerals can be subdivided into macro and trace elements. Regarding
macro elements, recommendations exist for the supply of calcium, phos-
phorus, magnesium, sodium, potassium and chloride. In the same way data
for the following trace elements are given: iron, copper, zinc, manganese,
cobalt, selenium, and iodine.
Macro elements. Calcium and phosphorus are especially important
for the function of the skeleton. Magnesium is essential for neural conduc-
tion and muscle contraction. Sodium, potassium and chloride are exten-
sively excreted via the sweat. The latter elements are particularly important
for thermoregulation as well as to maintain the balance of body fluids, acids
and bases. At maintenance level true digestibility end endogenous losses
are relevant to formulating recommendations for the supply of macro ele-
ments (Table XXIV-4).
Table XXIV-4: Recommendations for the daily supply of relevant major elements to horses
at maintenance level (GfE, 1994)
Table XXIV-5 shows the recommended amounts of the macro ele-

ments to meet the needs for maintenance plus growth, reproduction, lacta-
tion and physical activity in horses.
801
Table XXIV-5: Recommendations for the daily supply of relevant macro elements in horses
(GfE, 1994)
Beside the total amount of calcium and phosphorus, the ratio of both
elements should be observed. For adult horses it is recommended to fall
between 1.5 and 2.0:1. Long lasting calcium deficiency and excess phos-
phorus can induce secondary nutritive hyperparathyroidism resulting in
osteodystrophy fibrosa generalisata. Under practical feeding conditions
horses have excessive potassium mostly from roughage, but native feed-
stuffs are low in sodium and chloride. This explains the need for free access
to sodium chloride in all horses except for young foals, which are still not
able to consume proper amounts from the free choice salt.
Work causes sweat loss and so an increased need particularly for
sodium, potassium and chloride. At moderate environmental temperatures
(18-20°C) the sweat loss of horses depends on the work load and may vary
between 0.75 and more than 5.00 litres per 100 kg of LW/day (MEYER et al.
1990). One litre of sweat contains approximately 3.1 g sodium, 1.6 g potas-
sium and 5.5 g chloride, but the loss of further elements is not so signifi-
cant: 0.12 g/l calcium; 0.05 g/l magnesium, <10 mg/l phosphorus, 11 mg/l
zinc, 5 mg/l iron, 0.3 mg/l copper and selenium in traces (MEYER et al.
1990). The need for sodium, potassium and chloride depends on physiolog-
ic activity and weather conditions (environmental temperature and relative
humidity) and may therefore extremely vary within a short time frame.
Sodium and chloride should be provided via free access to salt. However,
the intake of loose sodium chloride may be higher than that of salt from a
solid block. Because forages in most cases provide excess potassium a
severe deficit of this element is only temporarily in extremely sweating hors-
es. Immobilization of a horse over a long period causes decreased retention
of calcium and phosphorus within the bone. Before such a horse is exer-
cised again, the supply of these elements should be temporarily increased
802
above normal.
Pregnancy first gives rise to an extra need for selected minerals at
when the mares are in an advanced stage because it is not until the eighth
month of pregnancy that a significant storage built in the foetal skeleton.
Until birth, the need for calcium and phosphorus increases up to 1.5-1.7
times the maintenance levels, and rises therefore at a higher rate than the
pregnancy-induced extra need for energy and protein. The need for sodium
is not clearly elevated. However, practical experiences indicate that an
unsatisfactory supply of sodium to mares in an advanced stage of pregnan-
cy may support the retention of meconium in foals. Broad mares should
have free access to a salt block, but this should not be attainable by the foal.
The extra need for minerals during lactation results essentially from
the excretion of these elements via milk. For this reason, the need for mag-
nesium and sodium increases only marginally and for potassium and chlo-
ride to a slightly higher extent. In contrast, lactation causes a need for cal-
cium and phosphorus that reaches the twice to three times of the mainte-
nance levels.
The need for calcium and phosphorus for growth remains nearly
consistent during the development of the animals. These minerals are need-
ed for fast bone retention in the young foal. With increasing age, the need
for retention decreases relatively while the need for maintenance rises.
Other elements are not used in such high amounts for growth.
Trace elements
Recommendations for the supply of trace elements are given in relation to
the intake of dry matter (Table XXIV-6). To calculate this, a daily dry matter
intake of 2% of the body weight is assumed. In rations containing high pro-
portions of concentrates the need may be somewhat higher.
Table XXIV-6: Recommendations for the daily supply of relevant trace elements to horses
(mg/kg DM1; GfE, 1994)
1) related to a daily dry matter intake of 2% of LW
803
Most native feedstuffs for horses are rich in iron and therefore the
need is commonly met. Feedstuffs rich in phytate (e.g. grain, extracted oil
seeds), cadmium, manganese, zinc and copper can decrease the utilization
of iron. The iron status may be marginal in horses with severe blood loss,
parasitic infestation and diarrhoea. Contrary to other neonates, a lack of
iron is not a common problem in newborn equines. Only the situation of
immature born foals is complicated because the foetus retains about 50%
of the iron’ in the last third of pregnancy. Parenteral iron substitution is not
well tolerated by foals and the safety of oral doses (e.g. by iron fumarate or
iron citrate) is at least unsure.
Contrary to colostrum, the mare’s milk contains clearly not enough
copper to meet the foals need (MEYER, 1994). However, foals may be able to
use copper that was stored within the liver before birth. This may work quite
well provided that the heavily pregnant mare received the amount of copper
she needed. Inadequate copper supply in horses may cause disturbed car-
tilage formation, haematopoiesis, development of nerves and vessels with
vascular rupture as well as contracted tendons, osteochondrosis and pig-
mentation disorders like the Arabian Fading Syndrome. In relation to the
beginning of developmental orthopaedic diseases the relationship of copper
to the energy supply in growing horses may be of special interest. To ensure
that foals are provided with enough copper it is recommended to double the
dry matter-related copper recommendation (Table XXIV-6) in mixed feeds for
suckling foals (20–24 mg Cu/kg DM). In general, the utilization of copper is
limited by high amounts of iron, zinc, molybdenum, cadmium and phytate
in the feed. Contents of 50 mg/kg dry matter in the total ration should not
be overwhelmed, especially regarding the antagonism to zinc and potential
hepatotoxicity.
The utilization of zinc may be limited by high amounts of calcium,
copper and phytate in the feed. When using practical horse rations the need
for zinc is mostly met. Extensive wound healing may be supported by the
addition of zinc to the normal need (1 mg Zn/kg LW/day). In heavily preg-
nant mares and growing horses it should be remembered that the action of
zinc is antagonistic to that of copper.
The utilization of manganese can be impaired by excessive iron and
calcium. Green plants from calcareous soil contain commonly less man-
ganese, contrary to the feed from soils with a comparatively low pH value.
Manganese is not very toxic. Excess supply may support anaemia because
of its antagonism to iron.
Cobalt is used by gut microbes to synthesize vitamin B12. Contrary
to ruminants, health problems in horses caused by deficient cobalt supply
are not known from practice. As part of the glutathione peroxidase seleni-
um helps to protect biologic membranes from oxidative destruction. Signs
of selenium deficiency can mostly be observed in connection with a lack of
vitamin E. However, white muscle disease in foals is obviously a selenium
804
deficiency disease. The injury may be caused always during pregnancy. In

Europe, soils provide in most cases only little selenium and this is especially
so when the pH value is low. Selenium is a highly toxic element. Signs of
chronic selenium poisoning in horses include the loss of hair in mane and
tail as well as the deformation of the hoof due to poor quality horn, unspe-
cific lameness and exungulation. Such signs have been observed above 2
mg selenium/kg of dry matter over a long period of time. The LD50 is said
to amount to 3.3 mg/kg of LW.
Near the sea coast green plants contain usually enough iodine to
meet the horses’ need. In contrast, the feed from mountainous areas pro-
vides commonly less iodine. Brassica spp. as well as feedstuffs containing
cyanogenous glycosides (linseed, legumes, and manioc) may have a detri-
mental influence on the utilization of iodine and thereby, induce a higher
need. Iodine in the ration can be equalized by commercial feeds in which
iodine-containing salts may be used as additives. The lack of iodine in preg-
nant mares can handicap the foetal thyroid function and cause goitre and
skeleton deformation in neonates, but excess iodine is detrimental, too. It
should be noted that algae of marine origin contain abundant amounts of
iodine.
24.2.4. Vitamins
Recommendations exist to supply selected fat-soluble vitamins (A, D, E) and
water-soluble vitamins (B1, B2, and biotin) to horses (Table XXIV-7). Except
when mixed feeds are given, vitamin A is provided for in native feedstuffs via
ß-carotene in horses. It can be assumed that about 400 IU of vitamin A can
be received from 1 mg of ß-carotene. Green fodder, artificially dried grasses
and legumes, grass silages and carrots may provide enough ß-carotene to
meet the horse’s need for vitamin A. Haylage and hay contains lower
amounts of ß-carotene and the content decreases further during storage.
Table XXIV-7: Recommendations for the daily supply of relevant vitamins to horses
(GfE 1994)
1 when fat supplements are given; 2 for heavily working horses until 4; 3 for heavily work-
ing horses until 5
805
Whether ß-carotene has a vitamin A-independent role to influence

fertility of broad mares is controversial. However, to provide the mares with
some ß-carotene during the winter season commercial preparations as well
as carotene from green meal are just as effective (KADEN, 2000). For this rea-
son, 200 to 400 mg of ß-carotene per day is the usual amounts. Inadequate
supply of vitamin A or ß-carotene may cause decreased fertility, growth and
resistance to bacteriological and parasitic infections. Runting syndrome and
corneal opacity with overflow of tears are also possibly due to vitamin A defi-
ciency. Contrary to ß-carotene, an excessive supply of vitamin A may induce
health problems. Excess vitamin A can cause alopecia, ataxia, sagging of
the limb at the fetlock joint due to overextension of flexor tendons and pos-
sibly disturbed bone re-formation with an increased tendency to fracture
(SCHRYVER, 1990). Such an excess results mostly from the abuse of com-
mercial mixed feeds.
Relevant amounts of vitamin D (vitamin D2) can always be found in
sun-dried hay. Vitamin D3 is usually provided by commercial mixed feeds.
Contrary to most other animals, in horses the vitamins D2 and D3 are obvi-
ously not significantly converted into the active vitamin D hormone
(HARMEYER et al. 1992). However, as long as solar radiation is ensured it has
not been demonstrated yet that horses need oral vitamin D administration.
On the other hand excessive vitamin D may cause severe health problems
including calcinosis of the duodenal wall, myocardium and kidney.
Young green plants and the appropriate artificially dried conserves
contain significant amounts of vitamin E, provided that the latter had been
produced carefully. The production of hay and pre-dried grass silage as well
as the storage of forages is connected with significant losses of vitamin E.
Mean concentrations of vitamin E are to be found in grains, but mostly as
ß-, g- and d-tocopherols which show a limited biologic value. The yellow fat
disease in foals is a vitamin E deficiency disease. Health problems related
to vitamin E deficiency seem to occur seldom in adult horses or may exhib-
it unspecific symptoms and possibly reduced immune competence. ZENTEK
(1990) described signs similar to equine rhabdomyolysis syndrome in hors-
es of different age which were fed on rations especially low in selenium and
vitamin E. Because of its function within the anti-oxidative system, the need
for vitamin E is particularly high in heavily exercised horses as well as hors-
es that ingest a fat-enriched feed. For the latter, the extra need can amount
to 1.5 to 3.0 mg of a-tocopherol acetate per gram of poly-unsaturated fatty
acid. Excess vitamin E seems not to be a relevant problem in feeding prac-
tice.
In horses, vitamins of the B complex are synthesized by gut
microbes. Since it is assumed that most of these vitamins of microbial ori-
gin are manufactured in the terminal gut, it is not yet fully understood how
the required amounts are absorbed. However, synthesis can be limited in
case of disturbed gut microflora caused by high concentrate - low fibre feed-
806
ing and oral antibiotic intake, or may be relatively low if the need is elevat-
ed e.g. caused by high carbohydrate metabolism. Some toxic plants like
marsh horsetail (Equisetum palustre) and bracken (Pteridium aquilinum)
contain a thiamine-splitting enzyme. As one of the first signs, thiamine-defi-
cient horses are conspicuously nervous. In case of unsure supply, the
essential vitamins of the B complex can be added by dried yeast or the
respective additives as part of mixed feeds.
Vitamin C (ascorbic acid) is a part of the anti-oxidative system, too.
Horses are able to synthesize this vitamin in the liver. Under extreme con-
ditions (heavy exercise, transportation, heat stress, infection diseases and
other kinds of injury) the synthesis may be relatively low. However, a poten-
tial oral need is difficult to quantify and may strongly depend on the indi-
vidual situation. Crystalline and amorphous ascorbic acid as feed additive
show unsatisfactory biological properties, contrary to more protected com-
pounds like ascorbyle phosphate and ascorbyle palmitate (ZEYNER and
LENGWENAT, 1993).
24.2.5. Essential fatty acids

It is known from practice that the ingestion of poly-unsaturated fatty acids
(PUFA), e.g. from plant oils or linseed, results in smooth and bright hair coat
indicating significant effects. Thus, it is preliminarily recommended to sup-
ply 5 g linoleic acid per kg of dry matter in horses (MEYER, 1984). This
amount is not to be found in the feed in every case, especially if the ration
contains high proportions of straw, beets and beet products. However, rele-
vant practical problems caused by marginal amounts of essential fatty acids
need not be expected because the body fat of horses is able to store and
release fatty acids over a long period of time and may ensure the fluidity of
membranes. The potential ability of n3-PUFA (e.g. from linseed oil or fats of
marine origin) is another aspect is to down-regulate inflammatory process-
es. This may become important in dietetics to prevent inflammatory disor-
ders like laminitis and chronic airway obstruction.
24.2.6. Water
The supply of proper amounts of tap water (Table XXIV-8) to horses is
important to maintain their health with special regard to digesta passage,
thermoregulation as well as the balance of electrolytes, acids and bases. The
need for water is influenced by the loss via sweat and milk and the incor-
poration into reproductive products. It can be assumed that horses con-
sume 3.5 and 3.0 kg of water if the ration is rich in roughage and roughage
plus concentrates, respectively. This difference is due to the higher water-
bonding ability of fibres within the digestive tract.
807
Table XXIV-8: Recommendations for the daily supply of water to horses
808
24.3. Practical nutrition of healthy horses
© Annette Zeyner
ractical horse feeding includes the appropriate supply of drinking
P water, energy, major nutrients, minerals and vitamins, which can be

achieved by a targeted combination of individual feedstuffs. The best
solution is a continuous access to good quality water assured by an auto-
matic drinker. The feed-related diseases and metabolic troubles can be pre-
vented by means of appropriate feed selection, ration formulation as well as
feeding techniques and technology. This requires the use of feedstuffs of
impeccable hygienic quality and free or low in antinutritive substances. In
case of horse special emphasis should be given to the ratio of high-fibre and
high-starch feedstuffs, practically the proportion of roughage to concen-
trate, and also to the feeding technique (frequency and duration of feedings
and sequence of individual feedstuffs).
24.3.1. Principles of ration formulation

In the majority of cases horse rations consist of a basal diet combined with
concentrates. Concerning the supply of high in structural fibre feedstuffs
(roughages, haylage), the minimum requirements, should be taken into
account in case of high-starch feedstuffs (concentrates, cereals) the maxi-
mum tolerable limits as well as their effect upon the digestive tract and
digestive processes (Table XXIV-9).
809
Table XXIV-9: Effect of predominantly high-fibre or predominantly high-starch feeding on

the digestive processes in horse
1) 1 h postprandial after given 1 kg feed; 2) independently on the small intestinal digestibil-

ity of cereal starch; 3) very high starch flow in the caecum
24.3.1.1. Roughage portion

The effect of the fibre-rich feed on the parameters in the digestive tract can
810
be summarized as follows (see also Table XXIV-9 above):
The sufficient supply with high-fibre feeds (roughages) is indispensable

for the undisturbed running of the digestive processes. Besides, assuring
the sufficient amount of fibre to offer effective fibre is an essential prerequi-
site. Fine grounded roughages are not able to sufficiently stimulate chewing
and may lose most of their favourable dietetic feature (see Chapter I). To
prevent the possible digestive troubles the following minimum amounts are
recommended:
≥1.0 kg roughage of effective fibre content per 100 kg LW daily.
The optimum allowances are generally higher than in case of other
monogastric animals. The primary high-fibre roughage is the hay, but in
many cases one third of that may be replaced by straw of good hygienic
quality. Feeding haylage has a similar effect on intake and gut environment.
At the same time, silages are faster ingested than hays, but related to dry
matter basis, the speed of hay and silage ingestion are similar to each other.
While using ensilaged green fodder the calculation of the minimum
roughage amount should be done on dry matter basis. In all cases the use
of straw of good hygienic state for bedding will help in assuring the supply
of fibre-rich feed. In fact, the roughage supply of horses, fed on silage of low
dry matter content and possibly kept without straw bedding, may prove
insufficient. In these cases the following minimum fibre concentration
811
should be considered for adult horses:

≥18% of dry matter crude fibre in the daily ration.
Deficient supply of structural fibre has higher health danger (dysbio-
sis, caecum acidosis) than the intake of too high amount of roughages. The
very high crude fibre concentration in the ration decreases the digestibility
of nutrients. Having a one-sided high intake (> 1 kg/100 kg LW) of micro-
bial hardly degradable roughage (e.g. straw) there is a danger of constipa-
tion, especially in case of lacking exercise.
24.3.1.2. Concentrated feeds
In many cases the energy requirements of horses (sport horse, breeding
mare) cannot be covered by feeding solely the basis feed. Another reason of
using concentrates may be the aim of preventing the „hay or straw stom-
ach“ (the so-called cosmetic reason). High energy feeds, such as concen-
trates (cereals, mixed feed), containing high starch and feeds high in fat, are
given.
Too much starch ingestion may overload the gut milieu and upset
digestive processes in the stomach and small intestine as well as in the cae-
cum. The effects of too high starch intake on the digestive processes in hors-
es were summarized in Table XXIV-9, namely:
812
The disproportionately high concentrate intake significantly increases

the risk of digestive troubles and their consequences (colics, damage of gas-
tric mucosa, and rupture of the stomach wall, acute laminitis, endotoxin
shock and diarrhoea). Consequently, offering a great amount of readily fer-
mentable carbohydrates (from starch) and fructans of meadow grass are
high risk factors for horses that are not accustomed to it. During the patho-
genesis of feed-related acute laminitis (founder) the high protein supply
obviously is an additional trigger factor. FIGURE XXIV-1 visualized the
model of enteral and extraenteral consequences of overfeeding with prae-
caecally indigestible, but in the caecum readily fermentable carbohydrates,
resulting in the predisposition to acute laminitis (founder).
FIGURE XXIV- 2: Model of the etiologic role of postileal readily fermentable carbohydrates
in the development of acute laminitis (founder).
813
The ileocaecal passing starch load depends upon the amount and the small
intestinal digestibility in a given period of time. The small intestinal
digestibility of feed starch, according to the origin and pre-treatment is very
variable (Table XXIV-10).
Table XXIV-10: The small intestinal starch digestibility in horses deriving from different
sources and in function of pre-treatment
814
Table XXIV-10: continued
A, a-amylase; aDpc/pi, apparent digestibility of the starch, pre-caecal/pre-ileal; LW, live

weight; CF, fistula at the caecum; e, evening meal; E, enzymes (a-amylase, xylanase, ß-glu
canase, pectinase); gr., ground; hp, hydrothermal processed; IF, fistula at the ileum; JF,
fistula at the jejunum; L, lactose; m, morning meal; MF, mixed feed; C+AM, pell.,
corn+alfalfa meal, pelleted; p. m., post mortem; S, starch; 1) extremely high digestibility in
one animal only; 2) donkey; 3) additionally pelleted
Among the untreated starch sorts before all the oats starch has the
highest precaecal digestibility. Grinding of cereal grains to coarse meal may
increase the ileal digestibility. At the same time, feeding of too fine-ground
or even powder-like flours could cause digestive troubles and by means of
inhalation during the feed ingestion causes the irritation in the higher parts
of the respiratory tract. Besides the positive effect of amylase addition to the
corn grain the small intestinal digestibility may be significantly improved by
the hydrothermic pre-treatment. The glycaemic and insulinaemic response
following the ingestion of starches from different origin and pre-treatment
does not generally agree with the expected ileal digestibility (Table XXIV-10).
Thus, the amount of starch given per feeding is considered to be of immense
importance. Starch sources for horses may be classified according to their
ileal/precaecal digestibility and to the related possible health risks as fol-
lows.
A) Starch of high ileal digestibility
oats, hydrothermic treated cereal grains, finely ground cereals, occasional-
ly steamed potatoes [[ in case of a high intake there are only occasional risks
of limited extent for the caudal section of the digestive tract (damages of the
stomachal mucosa, increased gas formation in the small gut)
815
B) Starch of low ileal digestibility

barley, corn (maize), manioc (cassava), raw potatoes [[ in case of a high
intake possible risks for the hind gut (caecal acidosis with its consequences
like acute laminitis). The risks of starch-rich feeding both for the foregut
and hind gut may safely be decreased by the restriction of the amount of
feeds of high starch content. In this respect the following recommendations
are valid:
< 0.5 kg/100 kg LW/feeding from oats and hydrothermic pre-treated
cereals or < 0.3 kg /100 kg LW/feeding from barley and corn.
The reason for the stronger limitation of the barley and corn is the
low small intestinal digestibility of their starch content. In case of
high concentrate feeding the number of daily feeding (frequency)
should be increased. The feeding of fats and oils decreases the rel-
ative starch concentration, preventing therefore the incidence of
digestive troubles and their consequences. Despite the lack of gall
bladder the capacity of horses to digest fats is high. The apparent
faecal digestibility will increase very steeply with the growing lipid
proportion of feed mixtures and finally attains a plateau above 80%
(FIGURE XXIX-3).
FIGURE XXIV-3: Apparent digestibility of ether extract depending on the fat concentrtionof
the feed mixture (ZEYNER, 2002)
The drop in crude fibre digestibility caused by high starch ingestion
816
may be alleviated by a partial replacement of starch by fat in the feed mix-

ture. At the same time, giving of high fat portions has the risk of reducing
fibre digestibility through the alteration of the caecal microflora by means of
the ileal indigestible fats. This risk can be expected at the level of above 8%
ether extract in the dry matter or while giving the high-fat concentrates not
often enough. Therefore adult horses may receive no more than 1 g fat/kg
LW daily and this amount should be distributed at least to three times.
24.3.2. Feeding technique

The concept of feeding technique comprises the feeding frequency, the feed-
ing sequence and the duration of feed intake. Although the above individual
factors have their specific effects, they act together as a unity and their
assessment should be adapted to the actual situation, like formulation of
the daily ration, technological opportunities etc.
24.3.2.1. Feeding frequency (incidence of feed distribution)
Horses, kept on the meadow between May and September are occupied with
a14-to-16-hours with the feed intake daily. Breaks between grazing, both
during the day and the night, are not longer than 2 hours. This type of feed
intake is very similar to that of the wild living ancestors of the horse. The
frequent ingestion of small bites prevents the overloading of the digestive
tract. In stall keeping systems this type of feeding frequency cannot be real-
ized. The feeding frequency for concentrates is determined by the required
energy supply from the concentrates and the necessary limitations of the
feeding time. At least the, in the following described minimum in the feed-
ing frequency should be respected. The higher the amount of concentrates
in the daily ration is, the more frequently should be fed and the character
of starch should also be taken into consideration. This type of calculation is
reasonable only if the fibrous feed supply of horses is simultaneously
assured. The distribution of roughages is determined by the principle that
possibly for a long period of time chewing opportunity should be given to the
horse. That helps in stabilizing the milieu in the digestive tract and may pre-
vent behaviour anomalies. The roughage feeding should be divided at least
into two daily occasions. In addition, it is favourable to make a continuous
access to straw bedding of good hygienic state possible for the horses unless
there is a contra-indication.
By means of applying feeding automats the daily concentrate
amount may be divided into many small feeding portions and it therefore
817
significantly contributes to the species-specific keeping of horses. In this

system horses may receive up to 12 small feed amounts daily. These fre-
quent stimuli are physiological and they rationally occupy the animals and
the development of bad stall behaviour can be prevented. Once the horse got
accustomed to the feeding automat, the rare, sporadic distribution of great
amount of concentrates may develop excitation behaviour (stamping, paw-
ing the ground, scratching, and champing).
24.3.2.2. Feeding sequence (order of offered feeds)
Besides the feeding frequency the sequence (i.e. the order of distribution of
feeds) has a great influence on the gut milieu stability, the balanced fer-
mentative processes and the digestive functions.
In case of stall feeding systems the main energy source is the con-
centrate, which should be given preferably, only after the distribution of
roughages. In this way, under the influence of fibre-rich roughages, stable
milieu circumstances could be developed in the gut canal. If the ratio of for-
ages is significantly overwhelming, the amount of concentrates serves only
as an additive to the badly fermentable, fibre-rich feedstuff, and then the
concentrates should be given together with the forages, to be able to stimu-
late the microbial activity. When using straw as single forage, the concen-
trates should be distributed before offering the straw to assure the balanced
caecal fermentation processes. Nevertheless, in the meantime, this type of
ration is not frequently used and under common stall feeding conditions the
greatest part of the energy is offered in form of concentrates. Accordingly,
the generally advised feeding frequency is the forage then concentrate dis-
tribution order. In grazing systems the distribution of hay before the morn-
ing driving out helps in preventing alterations in gastric digestion. Feeding
concentrates, if it is necessary, is optimal after arriving to the pasture or
during the day on the pasture.
24.3.2.3. Feeding length (duration of feed intake, eating interval)
The duration of feed intake in horses is in close correlation with the sort and
physical form (chopped, pelleted) of the feedstuff (Table XXIV-11).
818
Table XXIV-11: Influence of sort and physical form of feed on the mastication number in
large horses
w.d. = without data; 1) rich in leaves; 2) duration of feed intake is hardly changed by crush-
ing or coarsely grinding; 3) 20% chopped straw or 30% hay, *briquette
Large horses require approx. 40 minutes for the ingestion of 1 kg of

roughages. On the contrary, the same amount of concentrates will already
be eaten within 10 minutes. After chopping roughages will be ingested fast
and less chewed. The duration of concentrate intake may just be doubled
by mixing 20% of chopped straw or 30% of chopped hay into the concen-
trate for fast eating (gobbling) horses. To prevent the constipation colics the
length of chops should be at least 3, or rather 5 cm. The parameters of the
feed intake in ponies require a different evaluation to that of for large and
medium size horses. The duration of roughage intake and the concentrate
ingestion is approximately doubled and up to 4 times more in ponies than
in large horses, respectively. Feed intake and the related chewing activity
are linked to different functions. The most important of these are as follows:
a) saliva production and soaking of the swallowed feed pieces, b) cutting of
the feed particles, c) mashing and crushing of the feed and release of nutri-
ents (passing into the liquid phase).
a) Saliva production and soaking of the swallowed feed pieces
The sufficient insalivation of the feed is important to assure the phys-
iological passage of the oesophagus as well as for a quick and satisfactory
soaking of the stomach content with the gastric juice and to prevent unde-
sirable fermentation processes. At feed intake duration of 40 minutes/kg for
819
roughages and 10 minutes/kg for concentrates the saliva production may

be assumed in large horses at about 3.0 to 3.5 kg/kg roughages and 1.0
kg/kg concentrates. Owing to its high swelling capacity, the dry sugar beet
pulp should be soaked with 4-times more water before feeding to prevent
the oesophagus obstruction. The dry matter content of the swallowed feed
bites is approximately between 11 and 15%, while it is much higher:
between 33 and 38% in case of roughage and concentrate intake, respec-
tively. Accordingly, there is a high stomach dry matter after fast filling rate
the concentrate ingestion. The high rate of gastric filling means a significant
risk for the induction of fermentation troubles in the stomach area.
b) Cutting of the feed particles
The intense chewing and grinding of feeds, especially those of fibre-
rich ones, is of high importance for the healthy passage through the whole
intestinal tract and for preventing the constipation colics. The swallowed
feed bites in horses the diameter of 30 to 50% of the feed particles is below
0.1 mm and only 30% of all the particles (supposed a length of 12 mm) are
above 1.5 mm. In case of grass intake or ingestion of dried sugar beet pulp
the number of large particles is higher. The distribution of particle size in
the ileum content and in the faeces is similar to that of the oesophagus, cer-
tainly with an increasing tendency for large particles. Young, long clover
and agrostis (bent-grass) apparently cannot be sufficiently ground by hors-
es and is of a predisposal factor to constipation. Too shortly chopped (< 3
cm) roughages and the so-called golf lawn and lawn mower grass, as well as
very large amounts of hardly fermentable roughages (> 1 kg/100 kg
LW/day) are similarly insufficiently chewed and after swallowing it may lead
to constipation. To prevent these types of alterations in the passage rate
caused by the insufficient breaking into small pieces, sufficient time should
be given to horses for the undisturbed feed ingestion (Table-XXIV-11). The
peaceful feeding time will not only improve the grinding of feeds, but also,
support the physiological gastric digestive processes, especially the secre-
tion of gastric juice. Large amounts of roughages can be given while the
horses are allowed more time for their eating, e.g. in the evenings. The dis-
tribution of roughages before the concentrates, or the addition of chopped
hay (eventually straw) may slow down the concentrate intake. Critical feed-
ingstuffs (agrostis, large amounts of clover, short chop, golf lawn, lawn-
mower grass) should be neglected.
Feedstuffs of critical hygienic status (e.g. mouldy) are capable of alter-
820
ing the gut motility and thereby facilitating the formation of diarrhoea or
constipation. In addition to the objectively short available feeding time, the
different stressors (excitement during feeding time, meal jealousy) may
accelerate feed ingestion, eliciting the subsequent insufficient grinding and
unsatisfactory gastric juice production. Horses are allowed to be ridden or
used for at least two hours after feeding.
c) Crushing of feeds and release of nutrients
By means of the chewing process the individual feed bites become
ground and crushed, therefore their nutrients go into liquid phase already
in the oesophageal fluid in a measurable extent. The release of the readily
soluble nutrients makes digestion easier in the stomach and gut by means
of the produced enzymes of the horse organism. Since the crushing and the
grinding of feeds depend upon the undisturbed chewing process, the suffi-
ciently long and undisturbed feeding time is highly significant for horses.
The necessity of supporting the digestive processes by means of preliminary
grinding or crushing is discussed especially in case of oats. While feeding
rations rich in fibre, the previously crush of oat grains improved approxi-
mately by 4 percentage units of the digestibility of organic matter, in com-
parison to ration containing the whole grains. Considering this small ener-
getic gain, the crushing of the oats grains is relevant only for horses with
damaged dentition, for very old animals and for horses with inappropriate
closure in dentition due to developmental failures.
821
24.4. Realisation of the practical nutrition of the

healthy horse
he horse is a herbivorous, caecum and colon fermentor animal.
T Therefore grazing is the most natural form of feeding. If the grass on

the meadow is not sufficient, one has to supplement it with cut grass.
The horse shows its unsatisfied fibre needs by chewing the environment or
eating the beddings. By means of the fermentation in the hind gut the
dietary fibre utilization is quite good. The fibre requirement is high, about
14-16% of the dry matter. The grazing adult horse consumes 30-50 kg grass
a day. This daily portion consists of cut grass, green Italian millet, rye, bar-
ley, oats, and Sudan grass. (If the need arises, one can feed green sorghum
and maize too.) Of the leguminous plants, such as green alfalfa, saintfoin
(Onobrychis sativa), red clover and the common vetch are included in dif-
ferent mixtures, the best of which is oats with vetch. In the latter case, how-
ever only half of the mentioned dosage is recommended, in order to avoid
protein overfeeding.
The basis of the winter feeding is hay (6-10 kg), some straw and
rarely beet, silage or other succulent feedstuff. (For race horses the average
daily hay portion is as much as 3-4 kg. The most commonly used hay is the
meadow hay. The hay of Italian millet, common vetch with oats and Sudan
grass are also preferred. The feeding of alfalfa and clover hay is also com-
monly used, but the danger of the loss of leaves and the consecutive nutri-
ent losses are great. Most of the concentrate feed is made up of alfalfa meal,
too. The bulk need can be covered by healthy straw too. In small farms peo-
ple give for night corn stalk (with leaves). The spring straws (oat and barley)
are better than the hays of poor quality. In small farm one mixes the
chopped straw (size: 2-3 cm) with salted water and molasses, and feeds
them after 1-2 h of standing. In wintertime the use of succulent feed is also
possible, i.e. 2-3 kg of carrots, without cutting, potato: 3-5 kg/day. Sugar
beet is fed together with chopped straw. The use of silage of very good qual-
ity (corn, corn-alfalfa) in a daily portion of 15 kg proved to be excellent, espe-
cially in fattening foals and heavy draught-horses.
The feeding of concentrates is limited from veterinarian and eco-
nomical point of view. In Middle Europe the oats, barley and corn and wheat
822
bran are widely used. For race horses one use the extracted (solvent) linseed
meal and molasses (sugar) too, and the feeding of concentrated horse feeds
(in form of pellet) are also available. The concentrate portion generally used
is 0.5-1.5 kg; 2-5 kg for sport and race horses.
The drinking water requirement is at least 3 litre/kg feed dry mat-
ter. The average salt (NaCl) consumption is 10-15 gram/100 kg LW. During
the feeding of the pregnant mare there is an extra need for carotene (70-240
mg/day).
Foals. For the suckling foals generally a creep feed (oats, hay) is
given. It is recommended to give vitamin A and D-supply for the winter-
foals. The oats consumption of the suckling foals equals to their age in
month in litre. The average weaning time is about 5-7 months of age. The
ponies receive ¼ of the above mentioned quantities.
The commonly used weaning time is 5 months. The basic feedtuffs
for the foals according to their age are as follows: suckling: mare’s milk and
oats (“The foal is born by the mare, but its mother is the oats”.) Weaned will
receive concentrates plus hay or pasture, the growing: concentrates plus
hay of good quality or pasture and the older growing (till 1.5 year-old) good
pasture, later hay, pasture, a few concentrates. The main “loss” of the foal
at weaning is 4-5 litre of mare’s milk. Because the maximum hay intake at
that time equals to 2-2.5 kg, one should feed concentrates. The concen-
trates can be oats, barley, in an average quantity of 3-4 kg. (The average
“creep” feed consumption until weaning is 200 kg oats and 100 kg alfalfa
hay (50 % can be meadow hay).
The foal can reaches 70-75% of its mature withers height’s at the age
of 6 months; 85-88% at the age of 12 months; the 60-65% of mature body
weight in 1-year-old (thoroughbred) and 65-70 % in case of the heavy bred.
For the typically autumn weaned foal of 6 months one gives besides pas-
ture, 3 kg of hay and 3 kg of concentrate. The use of milk powder is advis-
able, because the pasture at that time may be very poor. During winter hay,
concentrates and 0.5-2 kg beet are fed or 3-5 kg silage of good quality is also
possible. The next spring and summer: 10 kg of grass, in the evening con-
centrates (cereals). While determining the nutrient requirement of foals 3
categories are used: Arabian type (mature weight 400 kg), half-bred (mature
weight 500 kg) and draught horse (mature weight 600 kg). A typical exam-
ple for the 7 month-old foal’s daily ration is as follows: hay 2.5 kg, oats or
barley 3.5 kg, wheat bran 0.5 kg (optional), beet or carrot 0.5 kg.
823
The total daily needs of working horses are different depending on the
intensity: light, medium and heavy work. E.g. for a horse of 500 kg, in case
of a medium intensive work 100 MJ DE is given, which can be met by 5 kg
meadow hay, 2 kg alfalfa hay, 0.5 kg barley, 0.5 kg corn, 0.5 kg molasses
and 1 kg straw. In case of an intensive work one has to supplement with 2
kg of concentrates. In normal circumstances, the appropriate energy supply
includes the covering of the protein needs.
824
24.5. Horse dietetics
24.5.1. Dietetic of special (patho)physiologic states
24.5.1.1. Raising of orphan foal

The mare’s milk is less concentrated and sweeter than the cow’s milk. The
suggested proportions for milk replacer are: 600 ml dairy milk, 400 ml
water, 35 g sugar. Daily portion: 4, later 9 to 10 litres/day, given each 2-3
hours. In addition, 2-3 raw (crude) eggs/day to improve the essential amino
acid and vitamin supply can be given. For shorter period of time the
skimmed milk can be simply used. In great studs some nursing mares or
goats can be applied.
In case of danger of dehydration a total parenteral nutrition (TPN)
may help. The iv. applicable solution consists of 1000 ml 5% amino acid
solution, 500 ml 50% dextrose, 30 mEq potassium chloride, 30 mEq sodi-
um bicarbonate and complex vitamin injection. The daily dose is 2.7-3
litre/45 kg live weight.
24.5.1.2. Geriatrics: feeding and nutrition of the old horse

Horses could be expected to live well into their 30s or beyond, depending on
many factors, including the accurate feeding. A horse older than 20 years of
age should be considered a geriatric animal. The practice of geriatric equine
medicine, the body weight-body condition scores, dentistry, disease resist-
ance and arthritic conditions should be emphasized. The most important
differences of young and old horse’s blood chemistry are in the elevated
median corpuscular volume and the decreased plasma ascorbic acid con-
centrations. The later may reflect a more intensive glyconeogenesis, due to
increased cortisol production. The digestion and utilization of protein, fibre
and phosphorus may decrease, those of the calcium does not change in
older animals.
In case of body condition loss the horse should be fed a very palatable,
easily masticated and digested, dust-free diet that has a slightly higher pro-
tein content (12-16%), maintenance levels of calcium (less than 1%), and
slightly elevated phosphorus content (0.4-0.65%), maintaining 1.5 to 1 Ca:P
ratio. Use of soybean meal helps in meeting the protein needs. The addition
825
of some yeast culture may improve nitrogen retention and pre-digestion or

extrusion may aid in feed digestibility. Adding fat or oil (1-2
cups/horse/day) to the diet of thin animals will help in maintaining appro-
priate body condition. As choke (oesophageal obstruction) appears more
common in aged horses that do not salivate or chew well, pelleted feeds may
need to be avoided, unless pre-soaked in water. Geriatric horses tend to
have a lower fibre digestibility, may be prone to choking, and appear to have
a higher incidence of colic caused by impactions. Older horses may have a
depressed immune system. The addition of ascorbic acid (10 grams, twice a
day, will increase antibody response.
There is also a higher incidence of renal calculi in aged mares and
geldings fed alfalfa hay. In geriatric horses with renal disease, beet pulp,
legume hay, should be avoided. Grass hay with protein content of 8-10%,
vegetable oils (1-2 cups/day), if the animal is of low body condition score,
and supplemented B-vitamins (brewer’s yeast) may all be beneficial. In
arthritic conditions first the suffering should be released through the use of
nonsteroid anti-inflammatory drugs and chondroitin sulphate. With chron-
ic founder (hyperglycaemia, due to reduced insulin response), which may
occur associated with pituitary adenomas, starch intake should be cur-
tailed, and both feed and fibre intake increased.
24.5.1.3. Insufficient voluntary feed intake.

Hypophagia due to a decrease ability or desire to eat (anorexia) occurs with
many diseases. The absence of feed intake even for a few days, particularly
in conjunction with trauma or a disease will adversely affect all body sys-
tems, making it more difficult for the animal to respond to therapy and
recover from the disease. Nutritional support is an adjunct therapy. For
example: without antibiotics, all the feed or nutrients possible will not cure
a foal with septicaemia. Nutritional support can make an important contri-
bution towards the goal of all therapy: rapid and complete recovery of the
patient with minor complications. The sick horse should be encouraged to
eat if it is able to do so and no contra-indications exist for oral feed intake
such as oesophageal lesions. If it is painful for a horse to eat, besides giv-
ing it an analgesic, feeding green grass or a feed mash (like water mixed with
bran, complete pelleted feed or meal) may decrease the pain associated with
eating sufficiently so that the horse will consume feed voluntarily.
Fever and pain, not just oral but anywhere may decrease feed intake
826
in which case giving analgesics and antipyretics may be helpful. Analgesics

may help feed intake in horses with laminitis, orthopaedic problems or
other localized sites of pain. They are less effective in horses with systemic
diseases. Feed intake in the sick horse should be encouraged by feeding
small amounts of feed frequently, removing feed that is uneaten for more
than 2 hours. One should also try and offer a variety of fresh feed (e.g., leafy
alfalfa hay, grain, sweet feed or bran). Lush green grass, grazed or freshly
cut is quite palatable for most horses. Bran mashes are popular for feeding
to sick horses but for most horses bran is poorly palatable.
Classical MASH: mixing bran with equal amounts of grain (particu-
larly steam flaked oats, barley or sweet feed) may improve palatability.
Including alfalfa meal or pellets will make it a more adequate complete diet.
A bran mash can be made up by boiling 2 litres of this complete ration and
feeding it warm but not hot to the sick horse. By including 1 cup of
molasses and one teaspoon of salt it may be more palatable to some hors-
es. Be careful with the hygienic status of bran, it may frequently be mouldy.
Occasionally sick horses will have peculiar feed preferences, so try different
feeds. Some sick horses reject feeds normally quite palatable and will con-
sume poor quality hay or bedding. The concern for anorectic horses is not
what they eat but how much they are eating.
Although some horses may eat while lying down, many won’t.
Therefore, getting a recumbent horse to stand may be beneficial to its
appetite. For this reason it may be helpful to sling a horse that is down
(apart from the medical benefit). Horses may also eat more if they see and
hear other horses eating. Some horses like to eat from the ground rather
from a manger, so try different feeding locations. Many people have the
impression that giving B vitamins may have the effect of stimulating a sick
horse’s appetite. Even if they don’t, giving B vitamins will be helpful in pro-
viding B vitamins in a horse with anorexia. Giving diazepam (valium, sedux-
en) in a low dosage (10 mg to a 500 kg horse) may stimulate eating for 15
to 20 minutes if feed is within easy reach and there are minimal distractions
in the environment. Response to repeated injections is not consistent.
Diazepam is metabolized by the liver and should not be used in horses with
liver dysfunctions. Anabolic steroids may increase feed intake several days
after administration but not immediately. It should be reserved for horses
in convalescence and used in the recommended dosage. Higher dosages
may have a detrimental effect on reproduction in stallions and mares.
827
TUBE FEEDING ADULT HORSES

Several enteral diets have been fed (via nasogastric tube) to sick
horses, ranging from slurries of pelletized feeds or alfalfa meal to commer-
cially available liquid diets designed for use in human patients. These liq-
uid diets have been classified into three categories: a) Blender diets - final-
ly ground whole feed suspended in water, b) Composition diets – composed
of highly digestible protein, carbohydrates, and fats, and c) Elemental diets
– diets containing small peptides and/or free amino acids rather than whole
protein. Elemental diets are very expensive and rarely used in horses. The
ingredients for preparation of a “Alfalfa-casein-dextrose slurry” are 454 g
alfalfa meal, 204 g casein (or dehydrated cottage cheese), 204 g dextrose, 52
g electrolyte mixture, 5 litres water; that of a “Pellet. vegetable oil slurry” 454
g pelleted complete feed, 50 ml corn oil, 2.5 litre water. Approximately 6
batches from the first and 9 from the second mixture are required to meet
the maintenance energy requirements of a 500 kg adult horse.
In practice, diets for feeding a horse through a large diameter tube
may be simply prepared by soaking a complete pelleted horse feed in at least
enough warm water to make it sufficiently fluid to pass through a stomach
tube. Dehydrated alfalfa pellets or meal may also be use. Grinding the pel-
lets or meal in a kitchen blender is not essential but may make the result-
ing slurry flow more easily. Water should be added just before the slurry is
administered; otherwise it will be too thick and viscous. Vegetable oils are
high density feeds and may be added to a healthy horse’s diet up to 20% of
the solid feed given or administered. A home prepared liquid diet for tube-
feeding to a 500 kg horse is given below: water 21 litres, dextrose 300 gram
(start with 300 gram and increase daily by 100 gram to a maximum of 900
gram), dehydrated cottage cheese 300 gram (start with 300 gram and
increase daily by 100 gram to a maximum of 900 gram), alfalfa meal 2 kg,
vegetable oil 150 ml, electrolyte mixture 230 g.
Add the alfalfa meal immediately before the diet is given otherwise it
will swell, making it impossible to give via a stomach tube. If the amount of
alfalfa meal is decreased, the low fibre may cause diarrhoea in the horse.
This diet may occasionally cause diarrhoea and laminitis and should not be
fed to horses with a previous history of laminitis. The electrolyte mixture
can be made up of the followings: 10 gram NaCl, 15 gram NaHCO3, 75 gram
KCl, 60 gram K2HPO4, 45 gram CaCl2, 4 gram MgO. Another favourite tube
feed is to boil oats 1.5 kg /day and split it into 3-4 meals per day. This will
828
meet about 25% of the daily energy requirements of a 500 kg horse. Often
a day or two of forced feeding with Jungle Oats is all that is required to get
a hypophagic horse to start eating normally.
Regardless of the diet used, the first day only one third of what is
needed should be given. If no significant signs of diarrhoea occur, increase
the amount by one third until the daily requirements are met (usually after
3 days). Start by tube feeding every 2-4 hours, gradually increasing the vol-
ume and decreasing the frequency. A hypophagic horse should be tube-fed
at least every 6 hours (4 times daily). Do not exceed 6 litres of fluid per any
feeding as a larger volume may be associated with reflux and signs of colic.
Always check for the presence of fluid in the stomach before the next feed
is given and if more than 50% of the previous feeding can be aspirated, dis-
continue the tube feeding.
Low fibre diets are often associated with diarrhoea, but it is often mild
and not associated with dehydration. It is not necessary to stop the tube
feeding with mild diarrhoeas, but the horses should be closely monitored for
signs of laminitis.
24.5.2. Feeding horses with specific disease

Most hospitalized horses do not need a special diet. They do not require the
high amount of feed given to working horses. Most can be well maintained
on a diet consisting of free choice, good quality hay. One to two kg of con-
centrates can be fed to these horses to help maintain appetite and adapta-
tion to grain once the disease is cured. For some specific diseases a spe-
cialized diet may speed up recovery and aid in the healing from the disease.
Some specific examples are given below. Besides cases of injury-induced
inactivity and instances where a horse does not or cannot eat enough on his
own, there are other specific disorders where therapeutic nutrition can ben-
efit the horse.
24.5.2.1. Starvation
Starvation per se occurs more commonly from late fall to spring and likely
reasons include financial constraints, neglect or ignorance and very seldom
cruelty. A horse with a long winter coat may look all right from a distance
and the poor condition can only be appreciated by palpation of the spine
and ribs. Fortunately most people notice and seek help if the starvation pro-
ceeds to recumbency. In horses with a condition score of 3 on a scale 1 to
9 (HENNEKE et al. 1983) i.e. transverse processes cannot be palpated but
829
spinous processes and ribs easily visible, tailhead prominent but individual
vertebrae cannot be identified) or more should be allowed free access to
water and salt and a diet of both hay and grain (13-16% protein) divided in
4-5 feedings daily. Once a horse is close to 40% below its optimum weight
it will become recumbent. Initially in sternal and later in lateral recumben-
cy. Survival is unlikely once a horse reaches 50% of its optimal weight,
regardless of the treatment. It takes a horse in good body condition at least
60-90 days without feed to become recumbent.
A chronically starved horse will have a long, dull coat that tends to
mask the extensive loss in body condition. A horse recumbent due to star-
vation will have a condition score of 1 (on a scale of 1 to 9, i.e. ribs, dorsal
spinal processes and tuber coxae will be easily palpated and lack of muscle
mass obvious). Most will die if they are unable to get up within 72 hours.
Therefore, they should be assisted to get up manually or in a sling.
Treatment should include intravenous and oral fluids without dextrose (they
are usually hyperglycaemic). If these horses are not eating they can be tube
fed as described previously. If they are eating, provide frequent small
amounts of grain (0.5–0.7 kg in a 500 kg horse) to which 10-20% vegetable
oil is added. Start slowly and increase the amount gradually over a period
of 3-4 days. A vitamin supplement containing vitamin A, D, E, K, thiamine
(B1), riboflavin (B2), pyridoxine (B6), cobalamine (B12), niacin, pantothenic
acid, biotin, folacin, and choline can be included in the daily feed. If too
much is fed too rapidly in a starved horse, acute laminitis, diarrhoea and colic
may occur, in which case, the amounts fed should be decreased. Starved
horses can attain their optimum body weight in 60 to 90 days.
24.5.2.2. Hoof defects

can occur from a multitude of causes and can be minimized with proper
hoof care. Biotin supplementation may be helpful in enhancing the repair of
hoof defects and preventing their recurrence. Horses with thin brittle
hooves, cracks in the weight bearing border of the coronary horn with crum-
bling of the lower edges of the walls or open white lines may benefit from
prolonged biotin supplementation. Biotin is widely marketed and a number
of commercial products are available. It will take more than 6 months for
the improvement to be noticeable in the case of hoof cracks and more than
19 months in horses with white line disease. Thoroughbreds need 15 mg
biotin per day and draught breeds twice as much. Horses with stratum
830
externum (on the outside) defects respond to biotin supplementation alone,

whereas, horses with stratum medium defects need calcium and protein to
respond if they are on a diet deficient in these two nutrients. It is not pos-
sible to differentiate clinically between stratum medium and externum
defects. Excess biotin intake by the horse has not been shown to be detri-
mental.
24.5.2.3. Recurrent myopathy

(exertional rhabdomyelitis) or tying up (or “set fast syndrome”) is common
in some thoroughbred fillies, although it can also occur in other breeds and
geldings. The relationship with carbohydrates and myopathy has been
known for decades. By lowering the carbohydrate intake and supplement-
ing with vegetable oils (250 ml per feeding) the incidence of tying up can be
lowered in most horses with the problem.
24.5.2.4. Recurrent airway disease

Recurrent airway obstruction (chronic obstructive pulmonary disease,
COPD or heaves) is an allergic condition to fungal spores commonly found
in hay and bedding. It is a disease of older (7 years and older), stabled hors-
es characterized by chronic coughing, exercise intolerance, laboured
breathing and nasal discharge. Once the allergy develops it will remain
present for the rest of the horse’s life. It is a disease that can be controlled
but not cured. It is vital to change environment of horses with Recurrent
Airway Obstruction’s and often it is essential to keep them outside and min-
imize exposure to the fungal spores.
If a hay cube cannot be fed, the hay should be soaked in water at least
5 minutes before feeding. The same applies for dusty concentrates. Wet feed
not immediately consumed should be replaced to prevent mouldy feed. All
feed should be fed close to the ground and not in a deep container so that
dust particles will tend to fall away from the horse’s nose and not be
inhaled. Straw, shavings and wood products should not be used for bedding
and the best bedding (if the horse must be stabled) is river sand, which is
more difficult to clean.
ELISA test are available (e.g. Equine Elisa™) for the following poten-
tial allergens: grasses (blue June grass, red top, bromegrass, orchardgrass,
timothy, ryegrass, fescue, Bermuda grass, Johnson grass, grain mix,
quackgrass), weeds and shrubs (alfalfa, dock sorrel, plantain, pigweed,
831
lamb’s quarter, ragweed mix, golden rod, sagebrush, Russian thistle, cock-
lebur, saltbush, kochia, marshelder), trees (cottonwood, juniper or cedar,
boxelder, willow, oak mix, pine, live oak, elm mix, bayberry or wax myrtle,
basswood or linden, sycamore, mesquite, cypress, pecan), mould epidermal
(Aspergillus, Alternaria, Helminthosporium, Hormodendrum, Penicillium,
Rhisopus, Grass smut mix, Stemphylium, mixed feather, house dust),
insect (black fly, cadois fly, horse fly, black ant, mosquito) and feed (alfalfa,
barley, corn, oat, molasses, wheat).
24.5.2.5. Diarrhoea
Horses with or recovering from diarrhoea should not be starved as lack of
oral alimentation for more than 2 days will cause intestinal atrophy and loss
of integrity which may delay the recovery even more and can result in sep-
ticaemia in young foals. If the diarrhoea is caused by small intestinal dys-
function, grain and concentrates should be withheld. On the contrary, with
a large intestinal dysfunction, concentrates can be fed but only in small
more frequent portions. Excess grain (especially corn) should be avoided
because indigested starch will pass through the small intestine into the
large bowel where fermentation may disrupt the normal large colon and cae-
cal function and make the diarrhoea worse.
Regardless of the cause of diarrhoea, fibre ingestion is beneficial as
a source of volatile fatty acids, which along with certain amino acids are the
primary source of nutrition to the cells lining the intestine. Fibre may also
stimulate intestinal segmental contraction (which will slow passage of
ingesta) and add bulk to form faeces. Because of these benefits, horses with
diarrhoea should be fed good quality hay. If horses receive oral antibiotics,
they may benefit from the oral administration of caecal contents, yogurt or
a commercial probiotic to inoculate normal intestinal flora.
Diarrhoea of young foals generally is caused by infection; conse-
quently, besides the dietary support the basic disease should be eliminat-
ed.
Sand induced diarrhoea can be seen in horses taking in sand inad-
vertently with feed or purposely by some horses particularly foals purpose-
ly. Inadvertent intake of sand is increased in horses on over grazed and
sandy-soil pastures and when feed is consumed from the ground. An insult
to the gastrointestinal tract that alters motility may also prevent a horse
832
from clearing normal small quantities of sand allowing it to accumulate and

causing clinical signs of weight loss, diarrhoea and in severe cases colic. The
sand accumulates in the ventral portion of the large intestine, irritating the
mucosa and causing diarrhoea and weight loss. Sand accumulation can in
some horses be heard on auscultation, seen in the sediment of a faecal
emulsion or abdominal radiographs may in some horses be required to
show up the sand.
Treatment includes feeding a soluble fibre containing psyllium (125
grams metamucil from Plantago ovata seed given twice daily) together with
a good quality hay and preventing additional intake of sand. Although the
diarrhoea may respond quickly to this treatment it may be necessary to give
a longer course of treatment to clear all the accumulated sand. Bran, oil and
other laxatives will have very little effect on clearing the sand. Having trace-
mineralized salt mixed with equal parts of bone meal may prevent some (but
not all) horses from consuming sand voluntary. A good deworming pro-
gramme is also essential to prevent intestinal damage and altered gut motil-
ity.
24.5.2.6. Hepatic dysfunction

In horses with hepatic dysfunction the plasma concentration of branched
chain amino acids (BCAA, i.e. leucine, isoleucine and valine) may be
decreased and that of aromatic amino acids (AAA, i.e. phenylalanine, tyro-
sine, tryptophan) as well as ammonia is increasing leading to clinical signs
of hepatic encephalopathy. These alterations can be minimized by feeding a
diet adequate to meet the energy and protein requirements but not excess
protein and with a protein high in BCAA and low in AAA with a high
BCAA:AAA ratio. The diet should also be high in starch to limit the liver’s
need for glucose synthesis. If the horse with liver disease is anorectic it
should be tube fed as described previously.
A diet that meets the requirements for hepatic disease can contain 0.5
kg corn meal/100 kg body weight per day divided into 5-6 equal portions.
Legume forage such as alfalfa may be best due to the high BCAA:AAA ratio
if the high protein content is tolerated by the horse (this can be determined
by measuring the plasma ammonia level). If the plasma ammonia is too high
(more than 80 μmol/l) grass forage should be fed instead. If the hepatic
damage is severe or the course of the disease is more than a few weeks, a
vitamin supplement containing vitamin A, D, E, K, thiamine (B1), riboflavin
833
(B2), pyridoxine (B6), cobalamine (B12), niacin, pantothenic acid, biotin,

folacin, and choline can be included in the daily feed. BCAA/AAA ratio of
some important horse feedstuffs: legumes (1.7-2.0), mature grass (1.2-1.5),
oats (1.65), barley (1.65), corn (2.15), wheat bran (2.0-2.2), linseed, canola,
sunflower grain (1.9-2.2), soybean meal (1.8) and casein (2.0).
24.5.2.7. Chronic renal failure

Chronic renal failure occurs more commonly in older horses and is charac-
terized by weight loss, high water intake, and increased urine production
due to an inability to concentrate urine. They will also have electrolyte dis-
turbances (mainly a low sodium and chloride level in the plasma) and vari-
ous degrees of anorexia caused by an inability to excrete metabolic waste
products like urea and reabsorb electrolytes. A diet for a horse with renal
failure should be high in carbohydrates and low in protein (to decrease urea
production), phosphorus and calcium. Excess calcium should be avoided as
this is also excreted by the kidney in horses (this differs from many other
species). A diet low in calcium, phosphorus and protein can be provided by
feeding grass forage. Concentrates may be fed up to one half of the daily feed
intake. Feeds to avoid include legumes like alfalfa (too high in calcium and
protein) and bran (too high in phosphorus). A vitamin supplement contain-
ing vitamin A, D, E, K, thiamine (B1), riboflavin (B2), pyridoxine (B6), cobal-
amine (B12), niacin, pantothenic acid, biotin, folacin, and choline can be
included in the daily feed to compensate for urinary loss of the B vitamins.
Salt and water should be provided ad libitum.
24.5.2.8. Urinary calculi

Urinary tract calculi are not common in horses and if present are usually
seen in the bladder of males (females can pass them through the relatively
short urethra). Diet modification to decrease urine concentration of calculi
constituents and to alter urine pH to make it unfavourable for calculi for-
mation has been shown to be an effective way of preventing calculi forma-
tion in horses. The dietary manipulation differs according to the calculi com-
position which should be analysed before starting a course of diet supple-
ments. To prevent a recurrence of calcium carbonate a low-calcium diet and
urinary acidifiers should be fed. The diet should ideally meet but not exceed
the daily requirements of phosphorus and calcium. This can be accom-
plished by feeding mature hay-grain mixture with no added calcium. Early
834
growth grass especially legumes are high in calcium and should be avoid-
ed. Adding ammonium sulphate at 75 mg/kg to the diet will be sufficient to
decrease urine pH from 8 to 5 in horse. Ammonium chloride will also lower
the urinary pH, but is less palatable to the horse. Urinary acidifiers may
cause more harm than good in that they decrease the dietary cation-anion
balance (see Chapter XI), which may increase the horse’s calcium excretion
in the urine. Increasing the salt intake has been advocated to increase the
urine production. Nevertheless, increased sodium may enhance the reab-
sorption of calcium in the gastro-intestinal tract and may not be beneficial in
preventing calcium carbonate crystals in the horse.

Severe burns, trauma, sepsis cause the horse’s metabolic rate to rise. Also
pushed upward are fat mobilization and utilization, protein catabolism, and
utilization and excretion of vitamin C and the B vitamins. Because of this,
the horse’s requirement for energy, protein, fluid, and water-soluble vita-
mins might be as much as 100% above maintenance needs. In these cases,
it is recommended to feed the horse a high-protein (14-16%), high-fat (7-
10%) ration, supplemented with B-vitamin complex, vitamin C, and possi-
bly vitamin E. Feed should be easy for the horse to consume. Suitable
rations might include legume-mix hay, pellets or extruded feeds in slurry
form, and supplements given orally, by nasogastric tube, or intravenouosly.
FIGURE XXIV-4: Typical signs of colic (abdomel pain)
835
24.5.2.10. Intestinal colics (FIGURE XXIV-4)

a) Intestinal impaction. Impaction of the large colon with ingested
feed is one of the most common causes of colic and is characterized by dry,
doughy contents. Colics caused by impactions are treated with analgesics
and some form of laxative tube by nasogastric tube. It is essential to keep
feed away from these horses until the impaction is passed. To assist in pre-
venting impactions, ensure proper dental care, supply fresh palatable water
at all times, and make sure that the horse gets regular exercise and a diet
consisting of good quality hay. Following the successful treatment of
impaction make sure that the hay fed to the horse is not too high in fibre
and not of poor quality. Instead low fibre highly digestible forage like grow-
ing grass or legumes will help in keeping the faeces soft. If necessary, 90-
120 gram of magnesium sulphate (MgSO4×7H2O or Epsom salt) may be
added to the daily diet.
b) Intestinal calculi. Enteroliths cause impactions and colic and must
generally be removed surgically. They form in the large intestine by miner-
al deposition around a node consisting of metal, cloth or nylon and most
often consist of magnesium ammonium phosphate (struvite). Diets high in
wheat bran and alfalfa (and thus high in calcium, phosphorus, magnesium
and protein) have been incriminated but not proven in the formation of
enteroliths. Some alfalfa is high in magnesium and will also promote alka-
line intestinal content, two factors implicated in the formation of struvite. In
areas where enteroliths are more common, they can be minimized by feed-
ing a diet consisting of grass forage and grain and avoiding legumes and
wheat bran (unless required). Feeding apple cider vinegar may help as it will
lower intestinal pH to below 6.6. The daily dose required is 110 ml per feed-
ing of grain in a pony and twice as much in a 500 kg horse.
c) Right dorsal colitis. Right dorsal colitis is a specific inflammation
of a small segment of the right dorsal colon associated with phenylbutazone
administration. Clinical signs include weight loss, intermittent diarrhoea
and recurrent colic and is directly associated with the intake of roughage.
This condition may be difficult to diagnose and will require very specific
dietary management. The aim is to feed a complete ration containing not
more than 30% fibre. This can be achieved by feeding concentrates with
coarse alfalfa meal (low bulk diet) to which 20% vegetable oil is added. The
diet should be split in 5-6 smaller portions per day to decrease the mechan-
ical and physiologic load on the colon. Horses that start eating their bed-
836
ding due to a fibre hunger can be allowed to graze small amounts of green
grass for short periods during the day. Short chain fatty acids (e.g. butyric
acid or sodium butyrate) may aid in the mucosal repair of the large colon
and including psyllium (metamucil) in the diet for 4-6 months (125 gram
per day in a 500 kg horse) will increase the short chain fatty acid ratio in
the colon and promote healing.
d) Post-colic surgery
A horse that has had relatively simple colic surgery that didn’t involve intes-
tinal resection (removal) should be started back on his normal feed as soon
as possible. However, the horse should be fed small amounts frequently,
gradually increasing the serving size until achieving his usual routine. (This
assumes that his original diet was not a cause of the colic.) If a resection
was required, feeding can get complicated. Soaked hay cubes or complete
pellets, or liquid nutrition products might be recommended. Requirements
for fat, protein, fibre, and various vitamins might change from normal. It is
highly recommended that any time major changes are needed in the horse’s
diet; the switch should be made gradually. Start by swapping out just 20-
25% percent of his usual feed for the new feed. Then slowly increase the
amount of new feed while decreasing the amount of the old feed over the
course of two to four days, until the change is complete.
e) Large colon resection or dysfunction. Horses with extensive large
colon or ceacal resections should receive a diet high in protein (> 12%),
phosphorus (0.4-0.6%) and low in fibre (less than 28%) to compensate for
their decrease in apparent digestibility. A diet of this type can be provided
by feeding a good weanling-type diet. In addition water must always be
available as their needs for water is increased due to the small area of water
absorption. In horses with only left colon resection the intestinal function of
the large colon will quickly return to normal and they will not require spe-
cial diet modification.
f) Small intestinal resection or dysfunction. The duodenum and the
jejunum are the primary sites for digestion and absorption of starch, pro-
tein, vitamins and most minerals with the exception of phosphorus. The
ileum is the primary site of fat and fat-soluble vitamin absorption. The large
intestine can compensate to some extend for the lack of absorption of pro-
tein, carbohydrates and B group vitamins. With extensive small intestinal
resection, a diet consisting of good quality forage and only small amounts of
grain should be fed. Good quality alfalfa and/or growing forage are the best.
837
Calcium absorption is primarily from the proximal small intestine, but sup-
plementation is not necessary if alfalfa is included in the diet. With ileum
resection the parenteral administration of fat soluble vitamins A, D and E
may be necessary.
g) Rectovaginal surgery (or laceration). Decreasing faecal volume,
pressure and straining to defecate may be helpful following repair of recto-
vaginal laceration. This can be accomplished by feeding a diet low in fibre
and high in energy. A complete pelleted feed with 25 ml of oil/litre of pellets
will accomplish this. If a complete pelleted ration is not available, alfalfa or
green growing grass will also help to keep the faeces soft.
24.5.3. Dietetics of miscellaneous diseases

The horse dietetics other, hereby non treated field would be the nutrition
and performance and productivity (feeding of feeding for athletic perform-
ance, that of broodmares and stallions), the possibilities to avoid mineral
and vitamin deficiency syndromes (e.g. white muscle disease) and some typ-
ical locomotion-related syndromes, like orthopaedic disease (development
orthopaedic disease or DOD), (meta)physitis, osteochondrosis, Wobbler syn-
drome, acquired flexural deformities and nutritional hyperparathyroidism
will be discussed in the following edition. The diagnosis and treatment after
the intake of a natural or synthetic toxin is also an important field of horse
dietetics.
FOR FURTHER READING

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Nutr. Physiol. 2, 111.
Donoghue, S., Kronfeld D.S, Berkowitz, S.J. and Copp RL (1981): Vitamin A nutrition of the
equine: Growth, serum biochemestry and hematology. J. Nutr. 111, 365-374.
Davies. M.E. (1971): The production of B12 in the horse. Br. Vet. J. 127, 34.
Dill, S.G. and Rebhun,. W.C. (1985): White muscle disease in foals. Comp. Cont. Ed.
Practic.Vet. 7, S627-S635.
Edens, L.M., Robertson. J.L. and Feldman, B.F. (1993): Cholestatic hepatopathy, throm-
bocytopenia and lymphopenia associated with iron toxicity in a thoroughbred gelding.
Equine Vet. J. 25, 81-84.
Ellis, N.W. and Lawrence T.L (1978): Energy undernutrition in the filly foal. Br. Vet. J. 134,
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su plemental lysine and threonine on growth and development of yearling horses. 13th
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839
Chapter XXV
CATTLE FEEDING AND NUTRITION

25.1. Characteristics of the digestive physiology
25.1.1. Schematic overview of the ruminant digestive functions
25.1.2 Regulation of feed intake and possibility of prediction
25.1.3. Characteristics of rumen digestion
25.2. Calf nutrition
25.2.1. Digestive characteristics of newborn calf
25.2.2. Drinking of milk replacers
25.2.3. Deficiency syndromes and metabolic troubles of calves
25.3. Feeding and nutrition of dairy replacement heifers
25.3.1. Antecedents
25.3.2. Concept of isometric and allometric growth
25.3.3. Practical recommendations
25.4. Feeding and nutrition of lactating dairy cow
25.4.1. The energy and protein balance
25.4.2. Synthesis of lactose, milk protein and fat
25.4.4. The body condition scoring system
25.4.5. Homeorrhetic control
25.4.6. Principles of mineral and vitamin supply
25.4.7. Biologically active substances of feedstuffs
25.4.8. Principles of practical dairy cow nutrition
25.4.8.1. Composition of the ration
25.4.8.2. Practical classification of feedstuffs
25.4.8.3. Feeding systems
25.4.8.4. Preparation of yearly feed balance
25.4.8.5. Practical veterinary interventions
25.5. Beef cow and replacement heifer
25.5.1. Feeding of replacement beef heifers
25.5.2. Nutrient requirement of the beef cow
25.5.3. Yearly production-reproduction cycle of beef cow
25.5.4. Feedstuffs of breeding beef cattle
25.6. Fattening beef
25.6.1. Energy requirements
25.6.2 Prediction of voluntary dry matter intake
25.6.3. Modifying effect of environmental temperature
25.6.4. Protein (nitrogen) metabolism of beef cattle
25.6.5. The compensatory growth
25.6.7. Methods of fattening
25.6.8. Feedstuffs for beef cattle
25.6.9. Growth promoters in cattle fattening
25.6.10. Deficiency syndromes, metabolic troubles and poisonings of
beef cattle
25.7. Veterinary technical advice in cattle herds
25.7.1. Determination of the locally valid requirement data
25.7.2. Veterinary feed evaluation in spot
25.7.3. Evaluation of the real supply
25.7.4. Profile investigations
25.7.5. Special field of the veterinary nutritional consultation
CHAPTER XXV: DIGESTIVE CHARACTERISTICS OF CATTLES
25.1. Characteristics of the digestive physiology

Knowledge concerning calves should be dealt with separately, because the
newborn calf functionally is still a monogastric animal (called also in the lit-
erature as “preruminant”). The digestion in abomasum and small intestine
of ruminants is more or less similar to the monogastrics. The fermentative
processes in the reticulo-rumen, however, are a very specialized type of
digestion.
The basic difference is that the proportion of the separate digestive
sections, related to the body weight is higher, if compared to the monogas-
tric: 12-20%. It is not the only explanation of the excellent feed utilisation
of the ruminants, because in case of some monogastric, like horse may even
be higher (1:16). The real key is that the proportion of the individual sec-
tions of the GIT is also different; reticulum, rumen and omasum may be
treated as one functional unit. This is the reason why ruminants are more
efficient from the point of view of nutritional strategy, than monogastric
herbivores, because they can utilize fibrous feedstuffs. The omnivores take
a medium place, carnivore having the less effective group in this respect.
The effectiveness of nutritional strategy is an economic and not an energetic
notion (i.e. transformation of gross energy of feedstuff), but it means that
ruminants are able to transform raw materials, which cannot be used as a
monogastric feedstuff (e.g. straw). Ruminants in this way are, not nutrient
competitors with human being, too. Therefore the role of ruminants in the
food chain is indispensable.
25.1.1. Schematic overview of the ruminant digestive functions

During carbohydrate (incl. fibre) degradation in the forestomach volatile
fatty acids (VFAs) are deliberated, providing energy for ruminants. Up to
90% of the total energy requirements of exclusively grazing cattle may derive
from these VFAs. Ruminal bacteria do not use fatty acid for energy, even
842
there is a bacterial fatty acid synthesis, too. Another basic difference, com-
pared to the monogastric animals is the nitrogen metabolism: part of the
ingested crude protein may be degraded to ammonia, which, in turn, may
be used by the rumen microbes. This process makes possible the use of NPN
(non protein nitrogen) compounds in the production of amino acids and
bacterial protein.
The described special features of the forestomachs are due to their
hosted micro organisms. Thus, the developing rumen should be inoculated
and populated by appropriate bacteria, protozoa and fungi. Tables XXV-1
and XXV-2 enumerate the main rumen bacteria and protozoa. The typical
substrates, the most important fermentation products and the special
nutrient requirements are also given. Above the fundamental building com-
ponents (VFAs, CO2, ammonia, some vitamins etc.) microbes require, as
vitamin-like substances, branched chain fatty acids. Most of the protozoa
cannot split fibre and there are some, which are consuming bacteria. Their
main usefulness is not the VFA production, but the protein, provided by
their own body in the abomasum, the bypassing of the essential, unsatu-
rated fatty acids through the rumen and the continuous mixing, homogeni-
sation of the rumen content.
Table XXV-1: Characterisation of the most important rumen bacteria

(after VAN SOEST, 1982)
--------------------------------------------------------------------------------------------------------------
Genus, species Substrate Product
--------------------------------------------------------------------------------------------------------------
Bacteroides amylophylus starch F, A, S
Bacteroides succinogenes cellulose F, A, S
Ruminococcus albus, cellulose and F, A, E,
flavefaciens xylane S, H2, CO2
Butyrivibrio fibrosolvens xylane, starch F, A , B, H2, CO2
Lachnospira multiparus pectin F, A, L, E, H2 ,CO2
Selenomonas ruminantium lactic acid, starch A, P, L, CO2, H2
Methanobacterium ruminantium formic acid, H2 methane
--------------------------------------------------------------------------------------------------------------
Legend: F = formic acid, A = acetic acid, P = propionic acid, B = butyric acid, E = ethanol,
L = lactic acid, S = succinic acic.
843
Table XXV-2: Characterisation of the most important rumen protozoon

(after VAN SOEST, 1982)
--------------------------------------------------------------------------------------------------------------
Group, genus Substrate Product
--------------------------------------------------------------------------------------------------------------
Holotrich
Isotricha, Dasytricha starch, sugars A, B, L, H2
Entodiniomorph
Entodinia starch F, A, P, B, (L)
Epidinium starch, A, B, H2, (F),
hemicellulose (P), (L)
Ophryoscolex starch A, B, H2, (P)
Diplodinium ? H2, fatty acids
------------------------------------------------------------------------------------------------------
Legend: see Table XXV-1. Parentheses mean that the quantity of produced compound is
tiny.
Besides their nutritional effects, protozoa have important influences

on host ruminant health status. Protozoa are more efficient in detoxification
of a variety of ingested compounds (mycotoxins, the toxic amino acid, the
mimosine, nitrites and nitrates) than bacteria. They help in disappearance
of lactic acid; even the active protozoa population reduces the rate of lactate
formation. Bloat and recurrent mild diarrhoea are more frequent in defau-
nated growing calves and drylot cattle on high concentrate ration. Finally,
the presence of rumen protozoa population decreases the chronic copper
toxicity by its sulphide production during protein breakdown (WILLIAMS and
COLEMANN, 1992).
An important part of the general overview of rumen fermentation is
the description of interrelationship of energy (in form of VFAs) and ammo-
nia release; in other words, the relation of rumen carbohydrate and nitro-
gen metabolism (FIGURE XXV-1).
844
FIGURE XXV-1: Scheme of ruminal ammonia and VFA release
Knowledge of timing of the carbohydrate and protein degradation is

important in understanding the feed intake regulation, too. Which types of
combinations may occur in function of time between the ruminal ammonia
and energy (VFA) release ? The ideal is when carbohydrate breakdown rate
is the same as ammonia production. In this case there are neither nitrogen
losses (economic profit) nor loading of detoxifying capacity of liver (physio-
logical advantage). If the energy release is slower than that of protein degra-
dation, an excess of ammonia will be present. This ammonia, after having
absorbed, loads liver and means metabolic and economic losses. This later
situation should be prevented, especially, when NPN compounds are fed. To
achieve this goal, either the rumen should be supplied by easily fermentable
carbohydrates (e.g. sugars, molasses), or part of the protein should be given
in rumen undegradable form. Therefore, never is the question that general-
ly a ruminant can be fed on urea or not, but the good approach is, that in
the given feed mixture and feeding technology is useful to give NPN sources,
or not. The same is true for the use of by-pass proteins: the UDP content of
the intake protein cannot limit the ammonia supply of the bacteria through
the remaining RDP part of the feedstuff.
845
25.1.2. Regulation of feed intake and possibility of prediction

In a wild living ruminants (e.g. moose) the real feed intake can be exactly
predicted by the knowledge of net energy and sodium requirements, the edi-
ble water plant offer as sodium source and the rumen capacity. Moreover,
moose is a concentrate selector during grazing; on the contrary, cattle a
grass/roughage eater, with less frequent feed rhythm. Since the possibility
for feed selection of domesticated ruminants is more limited than in the
above described situation, therefore knowledge of feed intake regulation is
essential.
In ruminants, the regulation of feed intake is much more complicat-
ed, than in monogastric, in which the “feeding to energy” principle is func-
tioning. It means that the animal ingests according to its real energy needs.
The same basically is true also for ruminants, but the rumen fermentation;
the wide range of fibre concentration may modify the basic picture. The
daily ration of ruminants consist of a significant amount of roughage, with
its varying fibre composition: lignin (totally indigestible, the polyphenyl
ester compound of total fibre), the living cell wall content (pectin, hemicel-
lulose and cellulose), partly digestible for the ruminal and intestinal
microbes and the well digestible cell cytoplasm. The relative proportion of
the mentioned compounds depends on the vegetative state of plant.
Therefore, the feed intake of ruminants is strongly influenced by the rumi-
nal degradability and intestinal digestibility of roughages of different matu-
rities. The actual composition of the ruminal flora may also have a great
impact on the fermentation. According to the previous processes, different
percentages of the ingested material appear in the faeces: the lignin is near-
ly 100%, the other fibre fractions, according to their degradability and
digestibility range from 10 to 60-70%, and the cell plasma may totally be
absorbed from the gut. (This later is not valid for the part of the plasma,
embodied by indigestible fibre.)
How the described factors can determine the ruminants feed intake?
The high lignin and cellulose content slows digestion down, giving animal a
feedback of satiety (fill full). The tool of feedback is the physical fulfilness of
the rumen. If the release of energy in the rumen is not parallel to that of the
ammonia, the number of cellulolytic bacteria decreases, because the ammo-
nia utilisation requires substantial amount of energy. The lower fibre degra-
dation slows down the rumen outflow rate, intensifying the feeling of fulfil-
ness. Summarizing, the low digestible roughage decreases the feed intake
846
by physical filling and by providing less available substrate for the rumen
microbes. The ammonia assimilation in bacteria and the number of cellu-
lolytic bacteria drops, degradation of fibrous material slows down, worsen-
ing further appetite. Thus, ingestion of feedstuff, high in indigestible fibre
fractions appears lower than the dry matter-based prediction.
As an illustration, let’s compare the required rumen degradation time
of some cattle feedstuffs to be 75% of potentially digestible cell wall broken
down: corn silage 1-1.5 days, green alfalfa 1 day, green grass 30 hours and
meadow hay 1.5 days (VARHEGYI and VARHEGYINE, 1992). These can be
explained by knowing the percentage of fibre fractions present in the feed.
PREDICTION POSSIBILITY OF FEED INTAKE (see also Chapter VI)

One of the approaches based on the physical control mechanism, and on
the knowledge of rumen volume (V, litre) gives the potential roughage intake
in grams:
FIroughage= 36 × V + 540.
The dairy cow related equation uses data of metabolic body size
0.75
(W , kg) and the amount of 4% fat corrected milk (FCM) production
(kg/day) giving the daily dry matter intake (DMI, kg):
DMIdairy cow= 0.01 W0.75 × (0.225 × FCM + 8.796)
The simplified form of this is: 0.025 W+ 0.1 FCM. It means that above the
live weight, the actual milk production should also be taken into consider-
ation.
In case of beef cattle the daily dry matter intake (DMI, kg) can be cal-
culated on the basis of metabolic body size (W0.75, kg) and the net energy
concentration of dry matter (NEm, MJ/kg DM):
DMIbeef = W0.75 Ñ (0.0355 NEm - 0.026 NEm2- 0.0196)
25.1.3. Characteristics of rumen digestion

SALIVA PRODUCTION. The appropriate saliva production is indispensable to the
normal digestive functions of ruminants. The reason for that is that the
slightly alkaline saliva effectively contributes to the maintenance of physi-
cal and chemical homeostasis in the rumen. In ideal cases the pH of rumi-
nal content is between 6.2 and 6.8. The range between 6.0 and 7.0 is still
physiological, but at about pH 6.0 most of the protozoa and cellulolytic bac-
teria cannot grow. Under pH 6.0 there is practically no effective fibre degra-
dation in the rumen, therefore even the slight local acidosis (pH 5.9-6.0)
847
may be harmful and undesirable.

The main stimulus of saliva production is the chewing, therefore
while feeding concentrate or high moisture feedstuffs, salivation decreases.
Concentrate feeding also increases the osmolarity of rumen fluid, which in
turn, inhibits saliva production. The insufficient amount of saliva may
cause both production and digestion problems. The mostly used prevention
is feeding buffers, like sodium bicarbonate. There are also parasympathico-
mimetic compounds (e.g. slaframine), stimulating saliva
CARBOHYDRATE METABOLISM. Main groups of carbohydrates, arriving in
rumen are as follows: cellulose, hemicellulose, pectin, starch and sugars. In
starch, the glucose units are linked by alpha-bonds, therefore the pancre-
atic amylase is able to split them. In the cellulose, the beta-bonds of glucose
units can be split only by bacterial cellulose. Hemicellulose consists of pen-
tose subunits; and the pectin is a glucuronic acid complex. Contracting the
chemical reaction steps, practically each compound get to the state of pyru-
vate.
Practical examples are as follows: hays are high in cellulose, sugar
beet pulp or apple pomace in hemicellulose and pectin, molasses in sugar,
cereals and other concentrates in starch. From the pyrovate, depending on
the original raw material, acetic acid, propionic acid, butyric acid and
methane are produced. Methane production is indirect; it is formed from
formic acid and hydrogen, by the contribution of methanobacteria. Slightly
simplified, propionic acid is produced from starch, therefore it is energeti-
cally advantageous. On the contrary, from the roughages acetic acid, CO2
and CH4 are released, in which the exhaled methane represents important
energetic losses. Small amount of butyric acid is produced from both types
of feedstuffs (FIGURE XXV-2).
848
FIGURE XXV-2: Volatile fatty acid pro-

duction in rumen, in relation to the
time elaps from feeding
In special feeding conditions (e.g. concentrate or sugar overfeeding)

the above described reactions may modify, and lactic acid and alcohol are
produced, too. The ruminal degradability of individual feed components sig-
nificantly differs. The undegradable part of the starch (bypass or protected
starch) passes over the rumen fermentation intact. For example after 8
hour-stay in rumen the degraded part of protein-free dry matter in wheat,
barley, corn and rye grain is 80-90%, that of oats 67% and only 37% in case
of sorghum grain (SCHMIDT et al. 2000). It is not advisable to give more that
1.5 kg bypass starch during a day.
The function of released metabolites is different: propionic acid hav-
ing gluconeogenetic property, promotes the glucose and glycogen formation
in the liver. Acetic acid is the precursor of milk and body fat. Butyric acid
is ketogenic, however, in low concentration it is the physiological stimulus
of proliferation of rumen epithelium (GALFI, KUTAS and NEOGRDY, 1986).
Methane, which is eliminated by eructation (belching) means energetic loss-
es. The significance of absorbed VFA can be illustrated by the faact that 70-
89% of the ruminant total energy requirements can be covered by the
volatile fatty acids, released in the rumen.
849
Based on the above described, ruminants are able to use feedstuffs,

which are inappropriate or of low value for monogastric animals. Moreover,
ruminants allow for the use of by-product ingredients that would otherwise
have to be as handled wastes for landfills. At the same time, ruminants even
require fibrous feedstuffs (roughages), to cover their energy needs and to
maintain the healthy rumen fermentation.
ROLE OF LIPIDS IN RUMINANTS. This question only recently received more
attention, because the typical ruminant feedstuffs (e.g. grasses, hays,
silages, beet etc.) have relatively low ether extract content. The fate of lipids
in rumen basically depends on their melting point, being lower or higher
than the body temperature. From this respect, oils have low, mammal fats
and especially tallow, high melting points. High melting point lipids do not
disturb rumen fermentation, bypassing into abomasum and small intestine,
more or less get digested and a proportion of them absorbed, the indi-
gestible part excreted by faeces. Low-melting point oils first get saturated
and all the cis-bound becomes trans one. The rumen microbes transform
unsaturated fatty acids in which hydrogen addition via microbial enzymes
removes double bonds in fatty acid acyl chain converting it to saturated
(biohydrogenation). Intermediates in the ruminal milk fat biohydrogena-
tion are the trans-11 CLA isomers, which are known to arise from transfor-
mation of linoleic acid (C18:2). The key enzyme of the production of cis-9,
trans-11 CLA is the ∆9-desaturase of mammary tissue. This compound has
a number of benefits to human health, including anticancerogenic proper-
ties.
Further on fatty acid are neutralized into potassium and calcium salts
(soaps), which may coat fibre particles, preventing their degradation and
consequently, methane production. The hydrogen accumulation inhibits
the action of methane-producing bacteria and the whole rumen fermenta-
tion is shifted towards the propionic acid production. Utilisation of oil, arriv-
ing through the abomasum into the small intestine is good, mostly by the
absorption in emulsified form. Emulsification of lipids in ruminant is good,
because it is facilitated not only by the bile, but also by the pancreas origi-
nated lysolecithin. Neither fats nor oils serve as a source of VFA in the
rumen.
Producing protected fats and oils, industry looked for combining the
advantageous features of the two different melting points lipids. Protected
means that these preparations protect themselves from saturation and they
850
also protect the rumen from disturbing its fibre degradation. Nevertheless,
at the level of small intestine, they are utilised. One of the processes of get-
ting protected lipid is the slight denaturation of oilcake proteins, which in
turn, protect embodied oil. Another approach is the saturation (hydrogena-
tion) and micro crystallisation. Tiny (1-4 µm of diameter) crystalline parti-
cles (prills) do not disturb rumen fermentation, i.e. they are rumen inert and
after having arrived in the small intestine can be emulsified and absorbed.
The calcium and potassium soaps of fatty acids also are considered as pro-
tected lipids. One of the best of them proved to be the calcium salts of palm
oil to resist biohydrogenation. Amides of unsaturated fatty acids also were
developed: specific fatty acid and amide linkage may protect them from bio-
hydrogenation, and e.g. oleamid feeding substantially enhanced oleic acid
concentration in milk, which is advantageous in enhancing the plasticity
and softness of milk fat for processing.
Summarising, carbohydrates and lipids are energy sources for rumi-
nants and, among physiological ranges, they are interchangeable.
PROTEIN METABOLISM OF ADULT RUMINANTS. The protein metabolism of
ruminants should properly be called nitrogen metabolism. The further
metabolic route of ingested protein and NPN depends upon the organism N-
supply: in case of insufficient protein intake, the majority of liver-produced
urea re-cycles by the saliva, back into the rumen. If the N-supply is over-
whelming, the surplus urea excretes through the urine, milk and uterine
fluid. Different by feedstuffs part of protein bypass undegraded the rumen
(UDP=undegradable protein or RUP=rumen undegradable protein), the
remaining part is degraded up to amino acids and ammonia. If the required
energy is available (see FIGURE XXV-1 about the timing of ammonia and
energy release), ammonia is built in bacterial body.
Basically the tissue amino acid supply of ruminants derives from two
sources, namely from, in the small intestine digestible bacterial and by-pass
proteins. Protein, digestible at the level of small intestine is the appropriate
notion to express the real protein value of feedstuffs and the protein require-
ments of ruminants. In the forthcoming physiological steps the protein and
amino acid metabolism of ruminant do not differ from those of monogastric
animals. Rumen bacteria, by means of glutamine synthesise enzyme, are
able to fix ammonia, but this chemical process requires considerable
amount of energy. The adjustment period of urea supplementation serves to
help the multiplication of rumen bacteria able for ammonia assimilation.
851
The main question of the modern ruminant protein evaluation sys-

tems is that what is the portion of the rumen degradable portion from the
ingested crude protein and how much digestible protein get available in the
small intestine as a total. These systems basically differ only in the way of
expression, prediction and calculation. Accordingly, the absorbed amino
acids derive not only from the feed protein, but also from the bacterial and
protozoa bodies. In the composition of the average bacterium body the pro-
portion of nucleic acid is important (up to 30%), which is not discriminated
using the Kjehldal crude protein determination (NÑ×6.25). The level of true
protein is approximately 50% in bacteria. The amino acid composition, as
well as the digestibility (76-85%) of protozoa is better than those in bacteria
(71%).
The other source of the protein, digestible in the small intestine is the
digestible part of the rumen undegraded protein. It is called by-pass protein,
escape protein, undegradable dietary protein (UDP), while the RDP (rumen
degradable protein) comprises also the NPN-compounds.
Rumen degradability (dg) means that from 100 units of protein how
many do degrade in the rumen (incorrectly/unjustified it is often called
`rumen digestibility`). HIGH DEGRADABLE PROTEINS (dg: 70-90%) contain aver-
age grass and meadow hay, corn silage, barley and wheat grain, wheat
gluten, extracted (solvent) groundnut, sunflower and soybean meal, as well
as beans. Adding of `”average” attribute is important, because if the hay or
silage get stuffed and get caramelised, or during the preparation of brown
hay and warm silage the Maillard-reaction occurs, proteins pass through in
the medium or low degradable group.) Proteins of forage grasses, legumi-
nous green plants, green forages, haylage, whole plant corn, corn grain,
corn gluten, heat treated soybean, lupine and sorghum grains, wheat bran,
brewer marc, cottonseed, linseed and pea are of MEDIUM DEGRADABILITY (dg:
50-70%). LOW DEGRADABLE (dg: 20-50%) proteins are in millet, grains, meat-
and-bone meal, blood meal and (selected) fishmeal. (Use of feedstuffs of ani-
mal origin while feeding ruminants in Europe is strictly prohibited.)
Degradability of feed protein cannot be accurately characterised by a
single number, because degradation depend also on the feed intake and on
production level of ruminant. The major modifying factor of the degradabil-
ity in these cases is the stay(ing) in rumen time. If a feed particle spends
less time in rumen, its undegradable portion increases. When more time is
available for the fermentation of the same feed protein, its degradability
852
increases. For example, the UDP portion of an average soybean protein is

50% in high yielding dairy cow, 35% in beef cattle and only 15% in suckling
beef cow. The explanation of this phenomenon is implied in the difference of
feeding levels (intensity) of the three types of animals. The appropriate feed-
ing level (one feeding level is equal to the maintenance requirement) for high
yielding dairy cow is 3-4, 1.5-2 for growing beef cattle and only 1.15-1.3 for
suckling beef cow. The rumen outflow rate increases parallel to the higher
feed ingestion. Average feed particle spends 24 hours in dairy cow rumen,
36-48 hours in growing beef cattle and in suckling beef cow or in dry dairy
cow up to 72 hours. The longer is the time, spent in rumen, the higher is
the protein degradability.
Above mechanisms may play a role in neutralisation of mycotoxins.
Trichothecen-structured mycotoxins get degraded in rumen during 2-3
days. Nevertheless, high concentrate intake accelerates rumen exchange
(outflow) rate, further decreases the pH. In such a condition the toxins may
remain intact, like the T—2 toxin in the ewe and heifer study (HUSZENICZA,
FEKETE et al. 2000; FIGURE XVIII-2). According in vitro study using rumen
fluid, during 24 hours only 37-56% of zearalenone can be degraded (MACRI

et al., 2005).
PROTECTED PROTEINS
One of the possibilities to increase the by-pass value of the daily
ration is the combination of protein sources of different degradability. The
other way is the technical treatment of the feed. Heat treatment, by causing
slight denaturising of protein, prevents rumen degradation, but does not
alter digestibility in the abomasum and small intestine. Chemical treat-
ments (e.g. using formalin) may also be effective, but being corrosive, in
many countries it is not allowed to be applied. Tannic acid is also useful,
but can change the faeces consistency by increasing its water content.
The FIGURE XXV-1 visualizes the time-related ammonia and energy
release in the rumen. The urea fermentation potential (UFP) gives the quan-
titative aspects, too. UFP expresses the amount of urea (in grams), which
can be assimilated, if the given feed or feed mixture is offered ruminants. It
depend on the protein degradability (dg, g/kg) and the in site available ener-
gy (expressed in total digestible nutrient, TDN, %) In formula:
UFP, g = (1.044 × TDN-dg)/2.8
This equation can be applied both for individual feedstuffs and for
853
CHAPTER XXV: DIGESTIVE CHARACTERISTICS OF CATTLESS
feed mixtures. If the UFP value is zero, it means that the available energy is
just enough to assimilate the released ammonia. Positive values mean ener-
gy surplus, therefore urea supplementation is possible. Negative UFP means
that part of released ammonia cannot be assimilated by the rumen bacte-
ria; it will be absorbed and converted by the liver into urea and either recy-
cled or excreted. In these cases feeding any NPN-sources has no usefulness,
it can be harmful.
The UFP, as a matter of fact, is the inverse of the rumen protein
(N) balance, used in the Metabolizable Protein systems (MPN-MPE). To
monitor UFP value is relatively simple: blood urea reflects rumen ammonia
concentration and the milk urea nitrogen has a very high positive correla-
tion with the blood urea (or BUN=blood urea nitrogen. 1 mmol/l urea equal
to 60 mg/100 ml BUN). The optimal range for milk urea is 2.5-5.0 mmol/l
(15-30 mg/100 ml BUN). It can serve as a quick test to predict the UFP
value of a feed mixture: the higher is the milk urea concentration; the lower
is the UFP value of the ingested feed (FIGURE XXV-3).
FIGURE XXV-3: Correlation between milk urea (y, mmol/l) and the UFP (x, gram) fed
dairy cows on different rations MAGDUS, FEKETE et al. 1988)
854
CHAPTER XXV: CALF NUTRITION
25.2. Calf nutrition

The average live weight of the newborn calf depends upon the adult weight
of the cow, namely 6.275% of the mature weight (NRC, 2001). From the
point of view of digestive physiology the newborn calf (lamb, kid) are a
monogastric animal and the forestomach can be found only by histological
examination (Table XXV-3, FIGURE XXV-4).
Table XXV-3: Relative measure of reticulo-rumen, omasum and abomasum, %
FIGURE XXV-4: Development of different chambers of forestomach and abomasum
From data of Table XXV-3 it is also evident that there are differences
among the individual ruminant species in the rate of maturation, when
rumination can be considered as complete. At birth, reticulo-rumen is hard-
ly visible in calf, and the largest stomach section is the abomasum. Parallel
to the ageing, the reticulo-rumen grows faster, and reaches, even surpass-
es measures of abomasum. The reticulo-rumen/abomasum volume index is
able to express this progression: at the age of 1 month it is only 0.5, at the
age of 2 month 2 and at the age of 4-5 months the value is 4-6. The same
development can be seen at the other ruminant species, but the rate of
growth may be different.
855
What is the key factor of this progression? The development of retic-

ulo-rumen basically depends upon the released volatile fatty acids of the
fermented solid feed. Nevertheless, for the formation of appropriate muscu-
lar wall thickness, the physical stimuli of feed particles are also important.
These statements are verified in model trials, too. Parallel to the volumetric
development of the rumen, it is populated also by micro organisms of the
environment and those from the adult ruminant. Saliva of the cow can get
into the calf mouth by licking, but there are also available commercial,
mostly freeze-dried bacterial preparations.
25.2.1. Digestive characteristics of newborn calf

As one of the most important physiological mechanisms, the OESOPHAGEAL-
GROOVE REFLEX should be mentioned. By means of its function the liquid feed
(milk, milk replacer, water) get directly into the abomasum, bypassing the
developing rumen. Its closure is a reflex process, influenced by the way of
liquid intake (suckling, drinking, in sip or draught), the temperature of liq-
uid (the most appropriate is 38-39oC), the age (in older animal it is more dif-
ficult to bring about) and the presence of certain chemical substances (e.g.
the salts of colostrums). Painting by copper sulphate the palate, the reflex
can be elicited even in older age, which can by useful to get medicine direct-
ly to abomasum. Excitation, increased blood adrenalin level inhibits the
realisation of the reflex. Cold, rarely and irregularly distributed milk replac-
er also fails to stimulate the closing of oesophageal groove: compounds, get-
ting in the developing forestomachs undergo bacterial fermentation, which
may lead to chronic blowing.
The main site of PROTEIN DIGESTION is the abomasum. The here pro-
duced prorennin first transforms into rennin and in the presence of calci-
um ions clots the major milk protein, the casein. Having optimum pH (4-5)
for its action, rennin hydrolyses the denatured casein. Whey of milk passes
along and by the existing low trypsine activity becomes digested in the small
intestine. Measurable activity of pepsin-hydrochloric acid complex develops
by weeks 4-5, when the activity of pancreatic trypsine is already consider-
able. DIGESTION OF LIPIDS partly takes place in the abomasum, by means of
the salivary lipase, which is completed from days 8-12 by the pancreatic
lipase activity in the small intestine. Among the CARBOHYDRATE sources
young calves are able only to the digestion of glucose and lactose. Digestion
of maltose, sucrose (saccharose), and starch becomes possible by the
856
increasing pancreas and brush border enzymes production. Fermentation

of fibre fractions develops parallel to the growth and increasing microbial
activity of forestomachs.
The first nutriment of calf is the COLOSTRUM. Since cows have epithe-
liochorial placenta, the majority of antibodies cannot reach the calf foetus.
The main source of antibodies (containing approx. 6% immunoglobulin) for
calves is the colostrum. The golden rule is, the within two hours after
birth calf should ingest at least 2 litres of colostrum. By passing the
time the concentration of antibodies in colostrum decreases and chances of
absorption worsen, because closure time is achieved at about 24 hours after
birth. Closure means that the intestinal mucosa looses the capacity to
uptake intact protein molecules, included immunoglobulins. Trypsine
inhibitor activity of the colostrum decreases by that time, too. Table XXV-4
compares colostrum to mature milk. The yellowish coloured colostrum can-
not be boiled, because it clots. It contains more fat (in very fine dispersion),
which is the main energy source for the calf. The protein concentration in
colostrum is also higher, than that of mature cow milk: the main cause of
this difference owed to the immunoglobulins: while in the mature milk their
level is 0.09%, in the colostrum is approximately 6%. The casein level is also
higher in colostrum, being the main protein source of calf (remember, the
activity of pepsin-hydrochloric acid complex and pancreatic trypsine is low).
There is less lactose in colostrum, but the magnesium concentration is four
times higher than in mature milk. The function of the latter is to help elim-
inating meconium by causing a slight osmotic dilution of gut content, the
digesta. High magnesium can cause diarrhoea among orphan piglets, fed on
cow colostrum.
857
Table XXV-4: Composition of colostrum and whole, mature cow milk
Vitamin A and carotene content in colostrum are also higher, which

is important in the calf survival. Vitamin levels in colostrum basically
depend on dry cow supply. Generally colostrum contains more vitamin B1,
vitamin B2, vitamin B12, folic acid and cholin, though even concentrations
of the mature cow milk are not negligible, compared to many foods. It
should be emphasized that amino acid and vitamin requirements of calf are
similar to that of monogastric animals, because of the rumen microbial syn-
thesis at that time is low. In practical conditions, some high-quality
colostrum should be frozen for instance when a cow of the herd has low-
quality colostrum at calving. Although the colostrums can be thawed in a
microwave oven without major disruption of antibodies, thawing it in warm
water is preferred.
25.2.2. Drinking of milk replacers

After the days of colostrum and mother milk nursing period (generally 10
days), the use of milk replacers is wide spread. For economic reasons the
858
use of cow milk (mixed milk of herd, appropriately treated milk from cows
in mastitis) is also possible, but caution is essential. Namely, after recent
data, if a calf received milk, infected with bacteria (e. g. Staphylococcus,
Streptococcus), the chance of occurrence of a mastitis in her milk producing
period is higher. Calf feeding on milk replacer is not a new idea, ISTVÁN
LÁNGHY has already proposed in 1831, in his Hungarian book (“The intelli-
gent, skilled and careful cattle breeder and veterinary surgeon “), published
in Pest.
Some specialists, especially if calf has been bought and introduced to
the farm at that time, recommend starting the artificial rearing by black tea
drinking. Tea should be supplemented by glucose (30-40 g/l), sodium chlo-
ride (9 g/l) and vitamin A (1 million UI/l). The appropriate dosage is 3 times
2 litres during the 24 hours. The next day a mixture of milk replacer and
black tea is given and day 3 begins the exclusive drinking of milk replacer.
Most of the good milk replacers contain 18-20% plant oil or animal fat.
Emulsification of lipids is facilitated by 1-2% soybean lecithin. The protein
source is approximately 80% skimmed milk powder. (As a matter of fact, the
precious milk fat is replaced by less valuable plant and animal lipids.)
Supplementation of minerals and vitamins, concentrated in the milk fat
should be done. Among others, magnesium, copper, zinc, cobalt (for the
developing rumen microflora) and iodine should be emphasized. Under the
influence of economic factors, part of skimmed milk powder can be replaced
by milk whey powder and soybean protein isolate. Digestibility of the men-
tioned protein sources depends mostly upon the age of calf (Table XXV-5).
859
Table XXV-5: Digestibility of proteins in milk replacers in calves
Certainly, the digestibility of skimmed milk is the highest, but as time

passes and efficacy of proteolysis in abomasum increases, even gets higher.
Considering the soybean proteins, the higher digestibility are found for pro-
tein isolate, extracted (solvent) soybean protein has a intermediate value
than that of fullfat soybean, especially in young calves is low. Digestibility
of fats and oils in milk replacers, supposed the presence of appropriate
emulsifier, is very good (85-95%). Generally, calf has a very efficient lipid
digestion system, having lipase from two sources (parotis and pancreas),
and as natural emulsifiers, sufficient bile and lysolecithin are present.
By knowing the natural and chemical composition, as well as the
digestion coefficients, the energetic value of milk replacers (DE) can be cal-
culated. Using data of the approximate (Wendee) analysis, the N-free extract
should be considered as lactose. Gross energy content of the individual
compounds (ash= 0, crude protein= 24, ether extract= 39, crude fibre= 17
and lactose=17 MJ/kg) should be multiplied by the relevant digestion coef-
ficient to receive digestible energy (DE) concentration. Digestibility of ash
and fibre is taken as zero that of lactose very good, energetic value of 1 kg
lactose is 16 MJ DE. Protein digestibility varies between 65 and 95%,
according to its origin; milk proteins are digested better. Lipid digestibility
is generally very good. As a conclusion, the energetic value of an average
milk replacer is 15-17 MJ DE/kg.
The air-dry milk replacer powder should be diluted by water (general-
ly one to ten) and offered at a temperature of 39oC for calves at 10% of LW.
Recently, for replacement heifers, an accelerated calf rearing program has
been introduced, in which the milk replacer with a higher nutrient density
and in the amount of the total of 15 to 20% of LW is fed. Calves in this pro-
gram are weaned 10-15 days later than calves in the traditional program,
860
but the growth rate is higher and by the time of weaning, calves are eating
0.7 kg of concentrate daily. However, excessive weight gain in the pubertal
heifer should be avoided. If the milk replacer contains organic acids (e.g.
formic acid), it can be diluted by tap water and distribute at 10-25oC (“pen-
guin-milk”). Low environmental temperatures increase the energy require-
ment of calves, especially those housed outdoors. For this reason, milk
replacer with higher (at least 20% fat or water-soluble oil) (e.g. Berol Nobel
preparation) should be fed during such periods, which help to meet the
extra energy needs.
Parallel to the rumino-reticular development, their cavity populates
with microbes, the capacity to degrade fibre improves and the released
volatile fatty acids, in turn, stimulate the maturation of forestomachs, thus
there is a self-enforcing process. During that time, the newborn “monogas-
tric-like” blood glucose level (5.0-5.5 mmol/l) decreases to the adult rumi-
nant’s characteristic one (2.5-3.0 mmol/l), while the VFA concentration ele-
vates. The predominant VFA is the acetic acid, because the propionic acid
is “filtrated” by the liver for gluconeogenesis.
The cattle should achieve an optimum ratio, when the reticulo-rumen
is 60-70% of the total fermentative capacity. In FIGURE XXV-5 are shown
the nutritional aspects of the calf rearing. The factors, which have to be har-
monized, are the milk or milk replacer, hay, concentrate and drinking
water. The appropriate combination of the above mentioned, the time
schedule (timing of the distribution) depend upon the preparation and the
applied technology. The most reliable indicator of used technique is the
unbroken growth and good ratio of the body parts (legs, head etc.) to each
others and to the live weight.
861
FIGURE XXV-5: Calf rearing using milk replacer
It is well seen in the figure that until week 7-9 this is the milk, milk
replacer and hay, later the concentrate and hay feeding, which are domi-
nating in the daily feed intake. Calf begins to eat small pieces of solid feed
before the age of day 10. The appropriate feedstuff for that is the so-called
“calf hay” (fine, early cut meadow hay or leguminous hay, cut in button
phase), offered ad libitum. At the same time, hay has a low fermentation
potential and provides little energy for the calf – grain concentratie mixture
should be fed too, so that adequate nutrients can be provided to allow the
calf to grow and reduce the nutrients that need to be provided by the milk
replacer. From the age of two weeks, to facilitate habituation, concentrate
should be provided, but considerable concentrate intake occurs only from
week 4 of age. To avoid abrupt dietary changes, technologies have been
developed, which provide only a single pelleted solid feed besides milk
replacer, made by corn and alfalfa meal. Parallel to the advancing age, pro-
portion of corn decreases and that of alfalfa increases. Ad libitum drinking
water supply is indispensable. Calves with additional ad libitum water
besides the starter and milk replacer drink by 1.5 to 2.0 litres of water daily
and their average daily gain improves 100-120 gram. Extra water does not
cause scour.
862
25.2.3. Deficiency syndromes and metabolic troubles of calves

BLOAT. When in the abomasum and developing forestomachs undesirable
bacteria overgrow (E. coli, Clostridium perfringens), gas production may
occur. Predisposing factors are the too dilutes (poor in fat) milk replacer, low
fibre in the concentrate or insufficient amount of “calf hay”. Besides, each
factor, inhibiting the function of oesophageal groove reflex (e. g. plant pro-
tein instead of milk one, subclinical myopathy etc.) facilitates the emergence
of a chronic bloat. Normally gases that are produced in the rumen can be
released, but when stable foams are present then this process is not able to
release the gases and bloat occurs.
RICKETS (or rachitis) may develop in case of calcium, phosphorus
and/or vitamin D deficiency, but in calves, lack of vitamin D alone, still

besides suitable calcium and phosphorus supply can cause rachitis.
Troubles of mineralization can be detected by the bent/twisted legs, hump-
back, swollen joints and the rickety rosary on the ribs.
DEFICIENCY OF MAGNESIUM can be suspected, if the sight is dull and fool-
ish, ears are hanging (Schlappohren”), star-gazer bearing, hypersensitive
calves are observed. (Lack of hair and discharge of saliva cannot be seen.)
Magnesium deficiency develop when fails the necessary Mg-supplementa-
tion (300 mg Mg/kg DM, in form of water-soluble salt) of milk replacer.
Higher amount of magnesium (600 mg Mg/kg DM, especially with 2% NaCl)
increases the incidence of calculi in their kidneys and bladder (MURPHY,
1992). The low digestibility of added lipids is a predisposing factor, because
they can form indigestible soaps with calcium and magnesium, which are
excreted by the faeces. Magnesium deficiency is not typical in adult cattle.
It should be separated from the thiamine deficiency (cerebrocortical necro-
sis, CCN), which is more characteristic in growing cattle and sheep. In bone
samples (get by biopsy or necropsy), the Ca to Mg ratio is higher than 90 to
1, instead of the normal 60 to 1 value. Proposed dietary treatment is 5
grams MgO, or 8 grams magnesium carbonate per os.
ZINC DEFICIENCY may develop primarily and by calcium overdosage sec-
ondarily. On predilection spots of skin parakeratosis develops. There is a fall
of hair around the eyes (ocula), on the nose (muzzle) also signs of paraker-
atosis, pasterns are eczematous and there is an intense discharge of saliva,
caused by the alteration of the mucous membrane of the mouth. To evalu-
ate this status, analysis of hair is sufficiently informative.
COPPER DEFICIENCY is relatively frequent on sandy or previously
863
marshy territories. It can be formed as primary copper deficiency, when

there is no sufficient copper in the feedstuffs, and secondary, when owing
to inhibiting interactions (Mo, S surplus) the absorption is low. Besides
anaemia, osteoporosis is typical. Gait, especially on the hind legs, is stiff.
There is a hair depigmentation around the eyes and on withers, because
melanin production requires a copper-dependent enzyme (poly-phenil-oxi-
dase.
MANGANESE DEFICIENCY is widely spread on slightly alkaline soils. Sex
ratio of newborn calves shifts towards males: more, poor vitality bull-calves
are born, their joints are swollen. Joint alterations can be explained by the
role of manganese in cartilage production (by means of glycosil-transferase
enzyme); the higher mortality of heifer foetuses by the higher manganese
requirement of ovaries.
DEFICIENCY OF VITAMIN A AND/OR BETA CAROTENE. Carotene level of calf
blood and liver depend on the mother’s feeding during dry period. Newborn
calf with poor supply has weak general resistance, incidence of diarrhoea
and pneumonia is higher and the mortality rate also increases.
Development of the skull and backbone periosteum slows down and the
arachnoideal rags (fringes) get infiltrated and closed by fibroblasts, which in
turn, contributes to the increase of cerebrospinal liquid. The latter com-
presses venas of the retina, papilloedema develops. Alteration of the sight
purple causes night blindness (nyctalopy), damage and atrophy of nervus
opticus (while passing through the foramen opticum) leads to a complete
blindness. Epithelium of cervix uteri, salivary glands and bronchi transform
into dequamative cells and these organs cannot fulfil their function. Supply
should be carefully design, because even a slight overdosage may prove to
be toxic: 138 μg/kg LW still decreases ascorbic acid and tocopherol level of
blood plasma; above 200 μg/kg LW increases cerebrospinal pressure and
more than 1327 μg/kg LW may induce clinical signs of hypervitaminosis
(toxicity).
In case of VITAMIN E AND/OR SELENIUM DEFICIENCY the protection of tis-
sues against free radicals is insufficient and the accumulated peroxides
cause cell membrane damages. As clinical and pathological manifestation
heart and skeletal muscle degeneration (cardio- and cardiomyopathy) can
be detected. At the first strong movement heart attack can appear in calves.
Some parts of Scandinavia, South-West of Hungary, some of the Middle
area of the United States and the Keishan territory in China have selenium
864
deficient soils. By treating of the highly pregnant cow and the newborn calf
by vitamin E and selenium injection, development of deficiency syndrome
can be prevented.
Calves in THIAMINE DEFICIENCY (cerebrocortical necrosis) show anorex-
ia, leg weakness, uncoordinated motion, heart function is arrhythmic,
breathing is irregular, intense lacrymation, grinding of the teeth (brygmus)
and opisthotonus can be observed. Thiamine therapy (injection, then per-
oral) in the early phase may be successful. The thiamine deficiency may
develop if no sufficient or only destroyed (heat or chemical treatments) vita-
min B1 is present in the milk replacer.
Calves are extremely sensitive to LEAD TOXICOSIS.
865
CHAPTER XXV: DAIRY HEIFER AND COW
25.3. Feeding and nutrition of dairy replacement heifers
25.3.1. Antecedents
The raising up heifer for high yielding dairy or for suckling beef cow is fun-
damentally different. Herewith the dairy heifer will be discussed. Raising up
is a complex process, comprises among others keeping, technology, disease
prevention etc., right now only the nutritional factors will be reviewed.
By calculating the reticulo-rumen to abomasum ratio (see calf nutri-
tion), one can follow the development, by the end from the newborn, func-
tionally monogastric calf becomes a real ruminant. This process is basical-
ly similar in each ruminant species. This is the point, until now the raising
of the replacement heifer does not differ from those of other production
goals (e.g. slaughtering animal). There is a reservation that extreme feeding
technologies cannot be used. For example either the calf rearing technolo-
gy, using drastically minimized amount of milk replacer, in order to stimu-
late early dry matter intake, or the to intensive concentrate feeding cannot
be applied. Table XXV-6 compares two one-sided technologies, using exclu-
sively concentrate or hay besides milk replacer. Reviewing the most impor-
tant metabolic parameters at the age of week 16, neither of the groups can
be considered as optimal.
Table XXV-6: Comparison of 16-week-old calves, fed on concentrate or hay
* sign of subchronic, latent acidosis!
The lower volatile fatty acid concentration in blood of concentrate fed

calves can be explained by the predominant propionic acid production. This
might also be explained by the fact that the calf is functioning as a mono-
gastric animal and is producing and absorbing sugars and not VFAs. The
later is used for gluconeogenesis in the liver; consequently, they cannot get
867
in the peripheral circulation. In case of hay feeding the predominantly

released acetic acid reaches the blood. The higher blood glucose level of the
concentrate group testifies the above mentioned glucose building from pro-
pionic acid. During the digestive process sugars are being produced and
then absorbed by the calf. Average daily gain and nitrogen retention are sig-
nificantly higher at the concentrate fed animals. However, the intensive con-
centrate feeding caused a subacut lactacidosis, which may be disadvanta-
geous to the development of rumen epithelium. Consequently, the rearing of
replacement heifer calves should be balanced and besides the milk replac-
er both concentrate and good quality hay or a pellet from concentrate and
roughage should be ad libitum offered.
Achieving the state of mature rumen functioning (age 6 month, live
weight approximately 180 kg), the feeding intensity of replacement heifer
and that of the fattening cattle should differ. Concerning the future breed-
ing animals, theoretically five intensity levels could be applied: very low
(from time to times resulting in severe underdevelopment), very high and
between the two extreme a temperate intensive, an intensive and a very
intensive one. The combination of the mentioned intensity levels in the dif-
ferent phases of rearing is also possible. To help the right evaluation and
decision among them, let us outline two relevant trials.
HUSZENICZA et al. (1988), after a wide range of data collection, retro-
spectively formed three groups of heifers, fed on differently during raising.
Holstein-Friesian heifers of Group 1 received low nutrient supply (low
dietary energy); the growth rate was also low. Owing to the continuously
small live weight gain, the puberty delayed and the time of artificial insem-
ination was later than months 18. They did not achieve live weight of 500
kg at calving. Next group was fed according to the NRC of that time (1978)
allowances, i.e. applying continuously a temperate intensive, resulting in
average daily gain of 600-700 grams. Heifers were able to be inseminated
before the age of months 18 and their live weight surpassed 500 kg at calv-
ing. Heifers of the Group 3 were kept in summer on scorched pasture,
obtaining a very low nutrient supply. In autumn, in stall keeping, these
heifers received extra concentrate portions, resulting in a small frame, but
fatty animals, able to be inseminated until the age of 18 months and their
live weight achieved 500 kg at calving.
The first conclusion of the described trial is that the puberty (onset of
the first fertile oestrus) mostly depends upon the live weight than on the
868
age. Any of the heifers could have been inseminated only after achieving the
350 kg live weight limit. This biological principle is true not only for the
dairy cow, but also for females of other breeds, even species. The explana-
tion relies on the function of fat tissue. Each body weight represents an
average body fat portion. If the amount of active adipose tissue does not
reach a minimum threshold concentration in females, then the activation of
steroids into oestrogen, as well as the leptin production proves to be insuf-
ficient. Thus, there is a lower minimum fat concentration in the heifer body
allowing puberty and cyclic sexual activity, which is generally well indicat-
ed by the live weight. While the percentage of adipose tissue does not
achieve a minimal threshold, the puberty will delay. The other extreme, get-
ting fat, also inhibit the sexual maturity.
What is against the system, when in the first phase a very low; later
a high feeding intensity resulted in small, but good condition heifers?
Whereas the insemination time of these animals did not fall behind that of
the moderate intensive fed heifers, their reproductive and health status
after calving proved to be worse. It means that more mortality, sorting out
and worse fertility could be detected in Group 3. The worst situation has
been found in cows of Group 1 after calving. At that time (end of eighties)
conclusion was drawn that the moderate intensive raising (700 g average
daily gain) can be proposed for the practice.
DACCARETT et al. (1993) compared heifer development of 700 gram
average daily gain and that of more by 15% in four different phases of rear-
ing (6-9, 9-12, 12-15 and 15-18 months of age). Each of the collected
parameters (live weight, body length, withers height, heart girth) turned to
be higher in heifers of richer nutrient supply. However, differences were not
proportional to the 15% plus nutrient intake, in some cases the difference
was smaller, in other cases greater than 15%, but every times for the ben-
efit of 115% supplied heifers. Since the disproportionate extra growth of the
individual body parts occurred in different phase, it is advisable to provide
the extra 15% nutrient supply continuously during the whole raising period.
Heifers of higher feeding intensity were successfully inseminated one month
earlier, and their age at calving averaged 23.49 months against the 24.83
months of the moderate intensively fed animals. Based on these data, to
keep an average daily gain of 800 gram can be proposed during the whole
raising period. How to check the intensity level? In farm conditions, the
weighing, at least once a month is sufficient to control growth rate and
869
adjust feed supply, if the monthly weight gain differs from 24 kg.
How can be commented data of the described heifer trial that the
extra growth (gain) of the different body parts was not proportional to the
extra 15% nutrient intake? For explanation should look for the isometric
and allometric phases of growth.
25.3.2. Concept of isometric and allometric growth

Isometric growth means that the growth rate of an organ, body part or tis-
sue is the same as that of the live weight. On the contrary, allometric growth
of a given organ (part, system, tissue) means that their growth rate differs
(disproportionate to) from that of the live weight, i.e. slower or faster.
Accordingly, growth of body length in the age of 15-18 months or the heart
girth in the age of 12-15 months proved to be allometric, because besides 5
and 6% average daily gain, the cited parameters answered by 57 and 37%
plus to the 15% extra nutrient supply. Growth of body length withers height
and heart girth would have been isometric, if it had been equally grown to
the average daily gain.
Generally, development of digestive organs and gonads can be char-
acterised by isometric growth. This statement is also reinforced by the fact
that puberty rather depends on a minimum body weight (at least 250-280
kg) than on achieving a certain age. On the contrary, in udder development
both isometric and allometric phases occur. In this way during the first
three months of life it is isometric, from achieving live weight of 100 kg till
the puberty (plus 2-3 oestrus cycles) allometric, which is followed by a new
isometric period, lasting until calving. During the allometric phase, the
growth of mammary gland is much faster, than that of the body weight.
After that, it is the beginning, repeatedly occurring progesterone effect,
which down-regulates udder development.
In the knowledge of these data, the damaging effect of the too low or
too high feeding intensity during the allometric phase can be interpreted.
The parsimonious feeding inhibits organs to grow according to their genet-
ic potential, they become underdeveloped. This is reflected not only in the
size of mammary gland, but also in body measurements. As consequences,
calving difficulties and lower milk production may be expected. Extremely
high feeding intensity (just fattening) enhances adipocytes functional activ-
ity and alters reproductive functions, fertility. The consequently high leptin
blood concentration inhibits IGF-I and serum induced proliferation of
870
bovine mammary epithel cells. There is also a local tissue production of lep-
tin by bovine mammary epithel cells, stimulated by insulin and IGF-I.
Increased systemic and local secretion of leptin may explain in part the
inhibitory effect of high-energy diet on mammary development in heifers.
The continuously rich nutrient supply is a negative feed-back for the growth
hormone (GH) production. Low GH concentration slows down development
of udder parenchyma, which can be shown by the decreased quantity of
total DNA. Possible aftermaths are the lower milk production, higher inci-
dence of calving difficulties and shorter useful lifespan (i.e. less longevity)
for more frequent culling and replacement.
25.3.3. Practical recommendations

For the practice the following key-numbers are recommended. For replace-
ment heifer should attain a certain live weight, i.e. approximately the 55%
of the adult, mature weight. The generally accepted average adult weight are
as follows: Aryshire 545 kg, Brown Swiss 682 kg, Guernsey 500 kg, Holstein
682 kg, Jersey 454 kg and Milking Shorthorn 568 kg (NRC, 2001).
Testifying the appropriate development of the individual body parts, a min-
imum withers height should be considered. The numerical value of these
two parameters can be given only as a range, i.e. the time of first insemina-
tion falls between 13-16 months of age, according to the actual feeding
intensity (FIGURE XXV-6).
FIGURE XXV-6: Recommended breeding time, live weight curve and withers height for
Holstein-Friesian heifers.
871
The goal is that the first calving should occur no later than 24
months of age (i.e. 2 years, or 730 day) and at that time the live
weight is 525 kg, withers height 133 cm and body condition score
(BCS) 3.5. To achieve this, the dairy (Holstein-Friesian-type) heifer
should be inseminated no later than 15 months of age, having at least
350 kg live weight and 121 cm withers height. FIGURE XXV-8 under-
lines that these corner-numbers should be treated flexibly and according to
the economic environment plus-minus deviations may occur. The optimum
in live weight for first insemination in the USA is considered today 850
ponds (385 kg), 66 inches heart girth (165 cm) and 50 inches height at with-
ers (115 cm) (KEOWN, 2005).
Finally, there are systems, which are more intensive than the physi-
ologically intensive one, recommended mostly in Europe. In the United
States (e.g. Brigham Young University, Cornell University) the even higher
feeding intensity is used. It means an average daily gain of 1000-1200 g,
which in turn make possible insemination (mostly using sex specific sperm
and/or embryo transfer) at the age of 12-13 months. Advantages of this
scheme are the decrease unproductive standing time of heifer, more milk in
the first lactation and faster genetic progress, owing to the shorter genera-
tion interval. As possible disadvantages, the decreased longevity and the
higher incidence of calving difficulties are mentioned. The actual NRC
(2001) recommendations prescribes for Holstein Friesian heifers an
average daily gain of 900 grams during raising, which assures that the
live weight at first calving achieves the 75-79% of the adult, mature
cow.
Which are the tools to control the intensity of rearing? Using the
heifer-relevant tables, the maintenance requirements are given in NEm and
the average daily gain in NEg. More practical approach is, having a given
feedstuff base (e.g. a pasture), to calculate the possible daily weight gain
and decide about the possible supplementation (mostly in form of concen-
trate).
Protein requirements of maintenance and growth can be drawn
together. This number cannot be considered as big one; consequently the
protein supply generally does not mean trouble during rearing of heifer.
However, too much degradable protein may cause reproductive failures.
Thus, the UFP of daily ration should be carefully taken into consideration.
In case of positive UFP (or in other words, in case of negative protein bal-
872
ance in rumen: rare situation!) even NPN compounds can be given. If UFP
is negative (equal to positive protein balance), the rumen available energy is
not enough for the assimilation of released ammonia, consequently it is not
advisable to feed urea and energy supplementation should be given in form
of concentrate or molasses. Excess ammonia and the produced urea are
damaging to the reproductive organs.
Practical nutrition of heifers can be based on forages: silages,
roughages (hays, corn fodder and stalk) and pasture. It is not difficult to
cover protein need, because it is relatively small. Excess rumen ammonia
and high blood urea level should be avoided by continuously monitoring the
UFP value of daily ration. As a rule, only a small amount of concentrate sup-
plementation is necessary. Having very good quality pasture, the grazing
may be supplemented only by good quality meadow hay (see New Zealand),
but more often than not there would be an imbalance between protein and
energy if only forage was fed.
873
25.4. Feeding and nutrition of lactating dairy cow

Dairy cow can be, above all, characterised by its lactation curve. Putting the
curves of milk production and that of voluntary dry matter intake, the key
problem of high-yielding dairy cow nutrition can be seen.
25.4.1. The energy and protein balance
In consequence of the genetic advances, increase in milk production after
calving is striking, has a long persistence and decreases only slowly. On the
contrary, the dry matter intake capacity, even using the best nutritional
strategy, achieves its maximum much slower. It means that at the begin-
ning milk production surpasses feed intake and nutrient supply being con-
sumed. The majority of production, health and reproduction problems of the
high-producing dairy cow can be related to this physiological situation.
FIGURE XXV-7 shows the energy output by milk and changes of the
live weight.
FIGURE XXV-7: Energy intake (MJ ME), calculated requirement and live weight changes
(kg) in the first 18 weeks of lactation (after ROBERTS, 1981, modified)
Difference between milk output and feed intake is counterbalanced by

mobilization of own tissues, consequently live weight drops. Each kilogram
decrease in live weight provides organism with 20.60 MJ NEl and 138
grams metabolizable protein (MP). No more than 550 gram losing weight
daily is considered as physiological during the first 60 days of lactation.
While there is a decline in body weight, the endocrine background (GH,
ACTH and glucagons surplus) is not appropriate for fertile oestrus. For over-
turn hormonal balance towards domination of gonadotropin hormones and
insulin, at least the weight loss should be stopped and the regeneration
874
should start. In turn, it is not necessary to achieve the ideal breeding con-
dition (BCS of 3.0-3.5) for the reproduction; it is enough having an improv-
ing condition.
The situation is similar in case of the protein balance. At the begin-
ning of lactation, the nitrogen balance is negative (dotted field under the
zero line), later slight anabolism is shown (dotted field above zero level),
finally the building of foetus and foetal membranes are visualised by double
striped field. The mostly used indicator of N-balance is the urinary methyl-
histidine excretion. The amount of the latter is proportional to the protein
mass of the body. Protein needs of embryo and foetal membranes practical-
ly are negligible.
25.4.2. Synthesis of lactose, milk protein and fat.

Precursor of milk sugar (lactose) is the blood propionate. In case of energy
deficit, the blood propionate concentration drops and because of the lactose
level of milk is constant, the quantity of produced milk decreases. Milk fat
is composed of fatty acids, deriving almost equally from the blood and the
de novo fatty synthesis in the mammary gland. One of the most important
precursors of milk fat is the blood acetate, which is released from the fibre
components of feeds. Beta-hydroxybutirate contributes to milk fat synthe-
sis, but neither this precursor nor plasma long-chain fatty acids have major
roles as energy substrates except under condition of starvation (see ketosis).
On the other hand, amino acid oxidations constitute the third major source
of energy for milk synthesis, besides glucose and plasma triglycerides. Milk
fat concentration is genetically fixed between physiological ranges. In case
of less acetate availability, milk fat level decreases, but the quantity of pro-
duced milk may remain unchanged. If the fibre content of total mixed ration
is too low (less than 16%), the so-called “LOW FAT SYNDROME”, or “milk fat
depression” develops. Propionic acid from the starch of grain stimulates cir-
culating insulin concentration, which does not allow metabolites to the
mammary tissue. Nevertheless, according recent discovery of BAUMAN and
GRIINARI (2003), a rumen bacterial biohydrogenation fatty acid intermediate,
the trans-10, cis-12 conjugated linoleic acid (CLA) is the most important fac-
tor in milk fat depression. It means that a fatty acid produced by rumen
bacteria affects mammary gene expression. This is a brand new field of
basic sciences, the so-called nutritional genomics or “nutrigenomics” (see
Chapter XVIII.
875
To promote the optimum milk fat production, the 3 to 1 acetic to pro-

pionic acid proportion is desirable in rumen. To achieve this, feed mixture
of 18-23% crude fibre content should be given. To assure the appropriate
energy supply in the first 60 days of lactation, high amounts of concentrate
should be given and fibre level may drop below ideal. Consequently, a slight
decrease in milk fat concentration in the mentioned period can be accepted
Milk fat composition can be influenced by the fatty acid composition of
intake fat, in order to provide healthy food for the consumer. (For details see
practical cow nutrition.) Crude fibre levels are not commonly used in the
USA to formulate rations anymore, since ADF and NDF and other terms bet-
ter predict responses observed.
Milk proteins are derived primarily from plasma amino acids, with
some synthesis of non-essential amino acids from acetate and glucose
through the Krebs-cycle. Although protein concentration in milk has a
genetically determined, close correlation with the milk fat level (milk pro-
tein, % = 1.9+0.4 milk fat, %), but this equation is valid only in average
nutritional circumstances. Nevertheless, milk protein level can be influ-
enced by feeding. Thus, reports are available about the positive effect of
rumen protected methionine (e.g. Smartamine M, Mepron 80) upon milk
protein concentration. Milk composition is influenced by the lactational
stage, too (FIGURE XXV-8).
FIGURE XXV-8:Changes in lactose, fat, protein and milk concentration quality during
the lactation (from top down botton) (GASPARDY et al. 2004)
876
Life of dairy cow consists of production-reproduction cycles and nutrition

should be adjusted to the peculiar needs of the given period. The basic unit,
a cycle, i.e. the time elapsed between two parturitions; in idealized case is
365 days, in case of first-calved cow 400 days (FIGURE XXV-13). If the calv-
ing to calving interval is longer by a day than the described value, it means
approximately half to one EURO economic losses. The 365 days is divided
into the 305-day-long lactation and the 60-day-long dry period. To achieve
these numbers, cow should be successfully inseminated no later than day
85 of lactation. On herd level it means that inseminations should fall
between day 60 and 90. Another practice is to inseminate at 45 days, prior
to minimize the effect of the negative energy balance that occurs in a nor-
mal cow.
The 305-day-long standard lactation can be divided into three, rough-
ly equal parts. The first third can be characterised by tissue mobilization,
live weight losing, stabilisation, beginning regeneration and the fertile
insemination. In the first 60-80 days of this period the cow cannot ingest
sufficient amount of dry matter to assure the energy and protein needs of
the produced milk.
At about the end of the first period (in good keeping and feeding con-
ditions between days 60 and 90) the mobilization of body reserves stops and
the improving condition offers opportunity for the successful insemination.
For the mid lactation characteristic is that cow is able to ingest sufficient
feed to cover milk production. This also means, more or less, the mainte-
nance of live weight. The third part of lactation is appropriate to improve the
condition. Requirement of 1 kg body weight gain is 26.80 MJ NEl and 276
g metabolizable protein, which can be in form of silage and forages, being
milk production already declining.
The ideal body condition score (BCS) of one-day fresh cow is 3.5; the
achieve that, the extra nutrient supply should be given during the third part
of the lactation, and not during dry period. During the dry period the aim is
to maintain the already attained ideal condition. Overwhelming nutrient
supply during dry period is the main predisposing factor of the “fat cow syn-
drome” (fatty liver and ketosis). The explanation for that should be searched
in the special fat metabolism of ruminants and in the regulation of volun-
tary feed intake. Lack of physiological adaptation of thyroid gland makes dry
cow susceptible to get fat (PETHES et al. 1984).
WHY DRY COW DOES BECOME OVERWEIGHT ? First of all, there is a protein
877
and fat accretion, caused by the changed hormonal background (anabolic

effect of gestation). Nutrient requirements of dry cow is the lowest of any of
the phases of the lactation cycle, so it is quite easy to over feed a cow dur-
ing this period of time. Further on, fibre utilisation of dry cow improves,
owing to the slower outflow rate. (However, lactating cows are more efficient
in using consumed energy approximately 80 vs 50%.) The lower is the dry
matter intake (in case of a dry cow 10-12 kg, instead of the fresh cow’s 20-
24 kg DM ingestion); the slower is the rumen exchange rate. The longer stay
of feed particles (in the first line fibre) in rumen increases efficacy of fer-
mentation and degradation of compounds becomes higher. Nutrient
requirement of growing foetus, placenta and udder development should be
taken in account only during the last 2-4 weeks of gestation. Consequently,
during the first 6 weeks of dry period it is enough to provide a daily ration,
equivalent to the maintenance requirement plus the needs of 2-5 kg theo-
retical milk production and preferentially it is given in form of silage and for-
ages. This mean that we do not want the cow to gain during this period of
time, but the fetus require nutrients equivalent to 2-5 kg of milk produc-
tion. In the last two weeks of dry period the concentrate supplementation is
desirable (for details see “Practical feeding” section).
Appetite of fat cows after calving is bad and the curve of voluntary dry
matter intake is depressed. For this reason, these cows cover part of the
milk energy from the mobilization of body reserves (mostly those of fat and
less of protein). A vicious circle develops, because the high blood free fatty
acid concentration worsens appetite further. It worth mentioning that over-
feeding can be made not only by using concentrate, but also by feeding
energy rich silage. The first cases of classical “fat cow syndromes” were
detected in herd, where dry cows were fed on good quality corn silage ad libi-
tum. Recent data shows that obesity, developed in the declining phase of
lactation and only maintained during dry period, is also harmful.
Data of the following interesting trial verify the above stated. Using
feed mixtures of different energy concentration during the third period of
lactation “thin” (BCS=2.0) and “fat” (BCS=4.0) groups of cows have been cre-
ated to the time of drying off. During the whole dry period and first phase
of lactation both groups have been kept on optimum feeding. Despite that,
considerable differences have been detected while compared cows of the two
groups: liver content of “fat” cows proved to be significantly higher: 15-30
vs. 0.8-15%. (Fat level up to 8.0 can be considered as physiological in dairy
878
cow). The voluntary dry matter intake and milk production (30 vs. 27 kg) of
“thin” cows were higher. In spite of the lower milk production, “fat” cows
have lost more BCS than their “thin” counter partner (1.2 vs. 0.5). “Thin”
cows had their first fertile oestrus (35 vs. 39 days) and successful insemi-
nation (89 vs. 99 days after calving) earlier and their sperm index turn to be
lower (1.9 vs. 2.4 dosage per stated pregnancy). Incidence of multifactorial
diseases (mastitis, ketosis and lameness) in “fat” cow increases. Measure
and lipolytic activity of adipocytes practically did not differ in the two
groups.
25.4.4. The body condition scoring system

An excellent practical tool to monitor the changes of body condition is the
CONDITION SCORING SYSTEM. This method is a more accurate indicator of the
body composition than the simple weighing. The significance of its using at
the critical times (on day of calving, at about day 30 and 200 of lactation,
at drying off and once in the middle of dry period) of the dairy cow produc-
tion-reproduction cycle should be emphasized. If body condition is lower
that the desirable, additional concentrate may be given, or, in case of feed-
ing total mixed ration, the cow can be assigned in a higher production
group. When there is an overcondition, the portion of silage and hay can be
increased, or cow is assigned in a lower production group.
When condition scoring is carrying out, the rump, tailhead and the
loin should be observed, touched and cow is classified between BCS 1 and
5 (FIGURE XXV-9). While scoring, stand behind the cow and place the
hands on the pin bone and pelvic bone and feeling for the amount of fat cov-
ering. Score the rump and then the loin area to one-half unit.
879
FIGURE XXV-9: Dairy cow (so-called Scottish) body condition scoring system
If the BCS is 5, the tailhead is buried in fat tissue, skin is distended.

No part of pelvis can be felt even with firm pressure. At the loin area folds
of fatty tissue are over short ribs and the bone structure cannot be felt. On
the contrary, in case of BCS 1, deep cavity is around tailhead and no fatty
tissue felt between pins. The pelvic bone easily felt and the skin is supple.
Overweight cows are good candidate for “fat cow syndrome”. In animals of
BCS 2, shallow cavity lined with fatty tissue at tailhead. Some fatty tissue
felt under pin bone. Pelvis (hook or hip bone and thurl) easily felt. At loin
area the ends of short ribs feel rounded. Their upper surface felt with slight
pressure. Depression is visible in loin. High-yielding cows in peak lactation
may have this state. Ideal BCS is considered 3.5 at calving. In BCS 3 no vis-
ible cavity around tailhead and fatty tissue is easily felt over the whole
rump. The skin appears smooth and pelvis is felt with slight pressure. At
loin the ends of short ribs can be felt with pressure and there is a thick layer
of tissue on top. There is only a slight depression in the loin. Cows of BCS
4 have a rump, where folds of fatty tissue are visible around tailhead. Fat
patches are present around the pin bone and pelvis is felt only with firm
pressure. No visible depression in loin between backbone and hip bone and
short ribs can felt even with firm pressure. Animals of BCS 5 are grossly
obese, The final condition score can be adjusted one-half score if the loin
differs from the rump by more than one point. (Table XXV-7). Recent data
(SCHRODER and STAUFENBIEL, 2005) demonstrated that measuring backfat
thickness by ultrasonographic measurement is of added value to other body
condition scoring methods, being objective and precise. A change in back-
880
fat thickness of 1 mm equals to approx. 5 kg of total body fat content.
Table XXV-7: Calculation of final body condition score (BCS)
25.4.5. Homeorrhetic control

To understand metabolism and its endocrine regulation of milk production
should be familiar with the notion of HOMEOSTASIS and HOMEORRHESIS. In this
way, fundamental role of growth hormone in the first period of lactation can
be explained. Repartitioning of ingested nutrients also depends upon these
two mechanism, i.e. which bodily function (heat production, maintenance,
fat and protein accretion, milk production etc.) will receive priority in a given
physiological state. (For repartitioning agents see “Beef cattle” unit.)
Homeostasis is the short term control of the harmony and steadiness of
interior milieu. Homeostatic control means a minute to minute maintenance
of the physiological balance, like body temperature, blood sodium and glu-
cose concentration etc. Homeorrhesis, introduced by BAUMAN and CURRIE
(1980) as “coordinated changes (or transition) in metabolism of body tissue
to support a physiological state (termed chronic control)”.
Life is a chemical process and the chemistry of life can be considered
as a cycle, in which feed intake and energy utilisation are in the back-
ground. The partitioning of nutrients to various body tissues involves
accordingly two types of regulation. Homeostatic control involves mainte-
nance of physiological equilibrium or constant conditions in the internal
environment. For example regulation to maintain constancy of body core
temperature, feed intake and partitioning nutrients during absorptive and
postabsorptive periods (ratio of insulin to glucagons has an important role
in the latter). Thus, a constant supply of nutrients to peripheral body tis-
sues is maintained by promoting short-term storage of nutrients following a
feed intake and mobilization of these nutrients during the postabsorptive
period. Homeorrhesis is a second type of control in partitioning nutrients.
Homeorrhesis can be defined as the orchestrated or coordinated
changes in metabolism of body tissues necessary to support a physi-
ological state. The Greek derivation of homeorrhesis is “uniform
881
flow”.
Hormones of the anterior pituitary play an important role in these
regulations. There are glycoproteins, like FSH, LH and TSH and proteins,
like prolactin and GH (growth hormone). Prolactin has an ancestral com-
monality with growth hormone in structure, molecular weight and amino
acid number. Protein hormones bind to specific receptors located on the
surface of target cell. On the contrary, steroids bind cytoplasmatic. Median
eminence of hypothalamus releases small peptide hormone (LHRH of
GnRH), which get via portal system of capillaries the hypophysis. Since the
frequency or amplitude of pulse release spontaneously varies, it is called
“pulsatile release”. Sensitivity of the anterior pituitary may be changed by
the modulatory effect of oestrogens and progesterone. There is also a pro-
lactin inhibitory factor (PIF) of dopamine structure production in the medi-
an eminence of the hypothalamus in a pulsatile fashion. Prolactin can be
released by TSH administration and by stress situation, too.
Generally, concerning homeorrhetic control on hormone level, there
are two basic models. 1) Nutrients are removed for anabolic or secretory
processes by tissue, under the influence of a stimulatory hormone. Depot
tissue buffers the central nutrient pool flux, which are regulated homeosta-
tically in response to sensors of the nutrient pool. 2) Homeorrhetic hormone
forcibly mobilises reserves from depot tissue in support of processes being
promoted simultaneously in functioning tissue. For example in high lactat-
ing dairy cow fed normally, GH stimulates through IGF-I (insuline-like
growth factor I) fat mobilisation and mammary milk synthesis. If a cow is
fasted, GH stimulates exclusively adipose tissue mobilisation. In case of a
chronic undernutrition GH inhibits, through IGF-I decreased production,
the milk synthesis in udder and increase at the same time the lipolysis.
Noradrenalin provoked lipolysis (see below) depends both on NEl intake and
milk production.
Lactation of Polar bear and Elephant Seal are the most convincing
example of homeorrhetic control of lactation, but tendencies are valid for
the high-yielding cow, too. Dam has only four weeks for suckling and dur-
ing this period of time has no opportunity to eat. By mobilizing 8 kg of its
body reserves daily and very concentrated milk is produced, allowing babies
to have an average daily gain of 4 kg. Thus, homeorrhesis is functioning
exclusively for the survival of offsprings. In dairy cow there is a similar
metabolic adaptation around parturition. It was shown that surviving
882
adipocytes from pregnant cow cannot be stimulated by noradrenaline treat-

ment. On the contrary, fat cells from lactating cows answered by significant
lipolysis on noradrenalin stimulation. Lipolytic activity of adipocytes gener-
ally depends on feed intake (unlinear, negative correlation) and on the milk
energy output (positive, linear correlation)
The ENDOCRINE CONTROL OF NUTRIENT PARTITIONING supports pregnancy
and lactation. As main factors are the changes in hormone levels, blood flow
to tissue or the number of receptors in target tissue. Placental lactogen coor-
dinates maternal liver and adipose tissue metabolism in manner to supply
nutrients to the developing foetus. Changing the oestrogen to progesterone
ratio alters blood supply to the uterus and, thus, could affect nutrient avail-
ability to the foetus. Oestrogen, progesterone and placental lactogen all play
a positive role in growth of mammary glands. Before the onset of lactation,
the prepartum surge in prolactin and oestrogen and the simultaneous fall
in concentration of progesterone represents one of the key components of
the push necessary for final stages of differentiation of mammary gland
(number of receptors will increase). After calving, prolactin and growth hor-
mone coordinate lipid metabolism in adipose tissue and liver in a manner to
partition nutrients to mammary gland. Oestrogen and prostaglandine F2α
reduce lipoprotein lipase in adipose tissue and increase it in mammary
gland (reciprocal changes).
25.4.6. Principles of mineral and vitamin supply

MINERAL SUPPLY of dairy cow can be understood only using the homeostasis
concept. According to the site of regulation, there are elements, regulated at
the level of absorption, at the level of excretion or both (For more details see
Chapter XI). Accordingly, homeostasis can be achieved by modifying the
absorption: by increasing ingestion less and less calcium, iron, zinc and
manganese will be absorbed. Iodine and cadmium make exception, where
absorption is parallel to the intake, which can lead even to toxicity. The
main tool of getting the homeostasis is the change of urinary excretion in
case of magnesium, fluorine, selenium, sodium and chlorine. Excretion by
milk may be adjusted in case of iodine and molybdenum. Tissue mobilisa-
tion and storage may also be adapted, as for iron in spleen and hemopoiet-
ic tissues, for the calcium the bone and teeth and for molybdenum the soft
tissues. Besides the absorption, the endogenous excretion may also be
altered (Mn). One of the main routes of elimination, which is strongly pro-
883
portional to the intake, is the saliva (Na) and the sweat (Na and Cl).
Absorption and utilisation of minerals are modified also by interac-
tions. It can occur in the soil, feed plant and gut lumen. There are, enhanc-
ing interactions, for example the chelate formation, but there are also
antagonisms. As the most typical ones, the sodium-potassium, the calcium-
phosphorus and the cupper-sulphur+molybdenum inhibiting competition
should be mentioned.
Calcium-phosphorus metabolism in dairy cow. More calcium
ingestion means more calcium absorption. The increased blood calcium
level is a negative feed back for the parathormone (PTH) secretion as well as
for the vitamin D activation mechanism (hydroxylation of 25-OH-cholecal-
ciferol to 1,25-dihydro-cholecalciferol). In lack of active vitamin D (calcitriol)
decreases the synthesis of calcium binding protein (CaBP) in the gut wall
and consequently intestinal calcium absorption worsens. In case of limited
calcium intake the inverse process is realised.
When blood plasma is high in calcium, it causes thyroid gland to pro-
duce more calcitonin, which inhibits Ca absorption from the bone and
increases Ca secretion by the urine. The opposite, if plasma calcium is low,
this causes the parathyroid gland to synthesise more PTH which on the one
hand stimulates the mobilization of Ca from the bones and indirectly caus-
es osteoclasts to start mobilize calcium from bone deposits. On the other
hand, PTH reduces excretion of calcium via the urine and increases absorp-
tion of Ca by stimulating renal vitamin D activation.
On the base of this regulation, the mechanism of MILK FEVER can be
interpreted, when the abrupt reduction of blood calcium is the main cause
of the symptoms. Generally 5-10% of the cows may be affected, typically
occurring 1 to 24 hours after calving. Incidence positively correlates with
age and milk production. The early symptom is a loss of appetite, because
the digestive tract becomes inactive, and muscles of the GI-tract are less
active. Cows are dull and listless, ears and muzzle are cold, and walking is
uncoordinated. In later stages cow will lay down and unable to rise, heads
are turned to side. This situation may be the source of further periparturi-
ent health problems, like ketosis, retained placenta and mastitis. According
to the clinical symptoms, three stages are differentiated: stage 1 (standing,
hypersensitive and wobbly), stage 2 (down on chest, drowsy, muscle flac-
cidity) and stage 3 (cow is on side, comatose and muscle flaccidity is
advanced), while the Ca is very low, Mg unchanged in blood (see György for-
884
mula).
It may occur more frequently, when dry cow received calcium rich
diet. This down-regulates parathyroid gland activity and sets PTH produc-
tion on a low level. With the beginning milk production the calcium output
becomes so high that bone mobilization would be necessary. Lacking suffi-
cient PTH in blood, bone calcium mobilization fails to realise, blood calcium
level drops and the classical form of milk fever develops. There is some ini-
tiative to counterbalance the eventually high Ca level in dry cow ration by
giving surplus phosphorus (to shrunken Ca to P ratio). Although excess of
phosphorus really decreases Ca absorption, but at the same time inhibits
vitamin D activation mechanism in kidneys, too. As a consequence, atypi-
cal milk fever develops. The conclusion is that during dry period the
absolute numbers of Ca and P recommendation should be considered, i. e.
6.1 gram calcium and 4.2 gram phosphorus for 100 kg LW. The dietary form
of Ca and P (biological available of source being fed) will also influence the
amount of Ca and P that needs to be fed. These values cover both the main-
tenance and the gestation needs. Having appropriate vitamin D supply, a
slight Ca overfeeding can be accepted. Feeding management during the dry
feeding period can have a dramatic affect on Ca metabolism after calving.
The cationic and anionic balance of daily ration, i.e. [(Na+K)-
(Cl+S)]/100 g DM, during the dry period has a great influence on the inci-
dence of milk fever. Positively charged (cationic) are calcium, magnesium,
sodium and potassium, negatively charged (anionic) are chlorine and sul-
phate. Feeding a cationic diet, the incidence of milk fever increases, because
they cause bone and kidney to become refractory to PTH stimulation.
Anionic diets decrease incidence of milk fever, because they increase blood
Ca pre- and mostly postpartum. Bone responsiveness to PTH stimulation
grows and as an indicator of bone Ca mobilization the blood hydroxyproline
level elevates. The same, high Ca amount in dry cow, if, supplemented the
feed mixture by chloride or sulphate, cannot induce milk fever. A negative
dietary cation-anion difference is suggested for cows in the last 3 weeks of
gestation (close-up dry cow) to reduce incidence of hypocalcaemia and a
positive value is needed for lactating cows.
Lactating dairy cow requires 4.4 g Ca and 3.4 g P for 100 kg LW and
2.8 g Ca and 1.7 g P for each kg produced milk. Availability of calcium
sources is different; those of organic bond are higher (30-60%) than those
in fodder plants (25-40%). Age of animal may have an influence too, because
885
in older cows the calcium utilisation is lower by 20%. This is one of the
explanations, why the incidence of milk fever increases with the age; the
other cause is that PTH receptor sites in bone decrease, too.
Phosphorus deficiency of dairy cow may cause alimentary infertili-
ty. To monitor status, the blood analysis is less appropriate and key num-
bers are not generally accepted. Therefore, the balance calculation (inges-
tion by the feed compared with recommendations) is satisfactory. According
to the modern approach, phosphorus requirements should be given in DM
concentration, which is 0.42 percent for the first 8 weeks of lactation and
0.32 percent for the remaining time. Magnesium requirement of milk pro-
duction is known, for lactating cow 2.0, for dry cows and heifers 1.6 g/kg
DM are recommended. Occurrence of Mg deficiency (“grass tetany”) can be
expected only in special soil conditions (Northern part of Germany, Poland
and Netherlands).
Sodium, potassium and chlorine. Sodium supply in dairy cow may
present as a limiting factor of milk production. Each litre of milk contains
0.63 gram sodium. Contracted requirement is 0.18% Na (i.e. 0.46% NaCl) in
dry matter. This can hardly be covered by silage and forages; the recom-
mended supplementation is 0.2-0.6% in dry matter, i.e. 30-120 gram com-
mon salt (NaCl). Sodium status of dairy cow can be evaluated by the sodi-
um concentration of urine, faeces and saliva (Table XXV-8).
Table XXV-8: Evaluation ofth dairy cow’s sodium status (KUTAS, 1985)
Potassium to sodium ratio has an influence on ovarian function: if

the amount of potassium is more than tenfold, the incidence of cystic alter-
ations of ovaries increases. It can be expected, while using potassium fertil-
izer or by irrigating pasture using liquid manure. In lack of sufficient zinc
supply in females decrease conception rate, in breeding bulls worsens the
sperm quality (REGIUSNE, 1988). For control the status, the hair analysis is
generally accepted, the lower limiting value is 100 ppm high-coloured,
washed hair. The main functions of manganese are in cartilage formation
as well as ovarian development and functioning. If the supply of pregnant
cows is deficient, the intrauterine mortality of heifer calves increases and
886
the sex ratio at birth are shifted towards bull calves. Joints of newborn and
young animals, having troubles in cartilage formation, are swollen. Mn defi-
ciency of adult cows, similarly to the phosphorus, may cause alimentary
infertility. Control is possible by hair analysis; the critical lower limit is 7
ppm.
Copper deficiency may be expected either on sandy soils or having
inhibiting interactions (Mo and/or S dominance). On the contrary, on soils
poor in Mo and S, dangerously high copper concentration may appear in
some plants. Main signs of copper deficiency are the congenital ataxia of
calves, troubles of oestrus and conception in cows, from time to time
nymphomania. The recommended daily supplementation is either 2 grams
copper sulphate per animal or chelated copper injection. To evaluate the
copper status, both the blood coeruloplasmin level and the hair analysis
(critical lower limit: 6 ppm) are applicable. Iodine deficiency may occur on
soils, poor in iodine, or by feeding large amount of crucifer plant with thyre-
ostatic substances (see Chapter VIII). Main sign is the birth of strumous
calves (goitre), but the general languishing may disturb reproductive
processes, too. The best way to control the supply is the milk analysis for
iodine. Vitamin E and selenium are destined for protecting cow’s organism
against free radicals. If their supply is insufficient, the incidence of mastitis
and of non-infective foetal membranes retention increases. For control use
either the hair analysis (limit being 0.25 ppm) or the gluthation-peroxydase
(GHSx) activity of red blood cells.
Vitamin A and carotene. According recent data, in lack of beta-
carotene, even besides good energy, protein and mineral supply, reproduc-
tion troubles may occur in dairy cow. LOTHAMMER (1976) proved the inde-
pendent vitamin function of beta-carotene in cattle. It means that for assur-
ing the protection against free radicals and the sufficient number of LH
receptor and the subsequent progesterone production, beta-carotene sup-
ply is indispensable and cannot be replaced by vitamin A. Beta carotene pri-
marily is responsible for the fertile oestrus, while vitamin A for the nidation
(or implantation, i.e. the embedding of fertilised egg into the endometrium).
In case of beta-carotene deficiency the level of LH in blood is low, oestrus is
long-lasting, from time to time bloody, ovaries cysts and thin (watery)
oestrus mucus can be observed. If beta-carotene supply is just enough, but
there is a vitamin A deficiency, the heifers show regular oestrus, conception
is easy. However, having no sufficient vitamin A, there is no implantation
887
and animals show another oestrus 30-35 days later.

In the real life practice the situation is more complicated: surplus
carotene may replace vitamin A and in turn, excess of vitamin A may have
a carotene-sparing effect. Mineral supply may also have an influence,
because in lack of phosphorus and zinc (sandy soils!) the β-carotene→vita-
min A transformation in liver is not sufficient and although having good
blood carotene concentration, vitamin A deficiency develops. Carotene sta-
tus can be evaluated on the basis of the blood plasma concentration: the
critical lowest value is 5.6 µmol/l (300 µg%). Minimum requirement is 300
mg beta-carotene per cow; optimum recommendation is 100 mg/cow/day
for maintenance and 25 mg/kg produced milk. Similarly to iron, there is a
carotene blockade, i.e. if blood carotene level attained 22 µmol/l (1.2 mg %),
no more absorption occurs. This regulation prevents eventual vitamin A tox-
icity. During practical nutrition it should be borne in mind that corn silage
has small and biologically low activity carotene isomers content.
Vitamin B-group. For dairy cow only the thiamine (B1), niacin, biotin,
cholin and vitamin B12 supplementation may have some importance. During
the first third of lactation, owing to the excess concentrate feeding and
slight rumen acidosis the thiamine production in rumen may prove insuffi-
cient. High dosage of niacin (6 to 12 g/cow/day) moderates fat mobilisation,
improves milk production. Biotin supplementation at about 20 mg/cow/day
can improve hoof health. Rumen protected cholin may improve the trans-
port of lipids, thereby reducing the incidence of ketosis. Deficiency of vita-
min B12 may occur on soils poor in cobalt.
25.4.7. Biologically active substances of feedstuffs

GOSSYPOL of cottonseed decreases blood tocopherol level and conception
rate. In leguminous and other grazed plant isoflavonoid-type PLANT OESTRO-
GENS (genistein, daidzein, biocanin A, formononentin) may occur (KEGL,
1988; VETTER, 1991). In extreme cases they can cause pseudo-oestrus. On

the contrary, on a pasture of average composition they contribute to getting
physiologically lively reproductive status in spring. CYANOGENETIC GLYCOSIDE
content of green sorghum (especially its frostbitten second cut), linseed
meal and water mannagrass may cause prussic acid (hydrocyanic acid,
HCN poisoning.
STRUMINOGEN (THYREOSTATIC) COMPOUNDS (e.g. vinil-thio-oxasolidon) of
crucifera (Brassica spp. and their crossings, like perko, Emerald, typhoon
888
etc.) inhibit iodine uptake of thyroid gland. Besides a general laziness of

metabolism the vitamin A supply is also altered. Drinking water of high
NITRITE and NITRATE concentration and nitrate-accumulating plants (green
oats, rye, and amaranth) may disturb thyroid function (KAKCSKOVICS, 1987)
and reproductive functions through inhibiting carotene®vitamin A transfor-
mation and the absorption of vitamin A.
Deteriorated feeds. In malfermented/badly fermented silage and hay-
lage the process of decarboxylation leads to the formation of BIOGENIC AMINES
(histamine, spermine, putrescine, cadaverine etc.). They irritate mucous
membranes of the mouth and gastro-intestinal tract and cause damage of
liver and nervous system (BOKORI et al. 1985). Secondary metabolic products
of moulds, MYCOTOXINS, among others, may cause (KÉGL, 1987) pseudo-
oestrus (F–2 toxin), immune-suppression and delayed ovulation (T–2 toxin).
Toxic, inorganic elements (I, Pb and Cd). Notwithstanding essentiali-
ty of IODINE, its excess intake may cause poisoning. LEAD, ingested by waste
gases polluted roughages, can be accumulated in bones, presenting danger
for the cow and even the human consumer, because acidosis will mobilise
it. CADMIUM get in soil by industrial pollution and being a mobile element,
accumulates in plants (KADAR, 1992), and after ingestion, in animal too,
causing primarily reproductive troubles.
25.4.8. Principles of practical dairy cow nutrition

25.4.8.1. I. COMPOSITION OF THE RATION
a) Determination of objective: on the basis of live weight, milk pro-
duction, fat percentage, lactation number, lactation period, expected daily
body weight change and environmental temperature.
b) Determination of requirements
MAINTENANCE
0.3514×W0.75 MJ NEl (where W = live weight, kg);
4.04×W0.75 g crude protein, or 3.6×W0.75 +50; or: 3.41×W0.75 g MP
4.4 g Ca/100 kg LW and 3.4 g P/100 kg LW.
First lactation cows should receive 120 and second lactation cows
110% of the calculated values. If milking parlour is far situated, energy
requirement of one km walking equals to +3% NEl, in case of grazing let’s
give plus 10-20% NEl and below -5oC environmental temperature +8% NEl
extra energy (mud and rain will also increase energy requirements).
889
MILK PRODUCTION
First actual milk production should be converted into 4% fat concen-
tration fat corrected milk (FCM):
FCM, kg = 0.4 × milk, kg + 15 × milk fat, kg.
Requirement of one kg FCM is: 3.10 MJ NEl, 87 g crude protein, 2.8 g Ca
and 1.7 g P. If milk protein concentration is known, the metabolizable pro-
tein need can also be calculated: MP-requirement of 1 kg milk production =
(milk protein, gram)/0.65
CHANGES IN BODY CONDITION. During the first 60-80 days of lac-
tation 1 kg decrease in body weight provides 20.60 MJ NEl and 320 g crude
protein (= 138 g MP). The accepted maximum daily decline in live weight is
500-550 grams. Day 81-305 of lactation: requirement of 1 kg body weight
gain is 26.80 MJ NEl, 500 g crude protein (=276 g MP), 20 g Ca and 10 g P.
The desirable average daily weight gain is 200 grams.
DRY MATTER AND WATER INTAKE
Minimum dry matter requirement is 2% of the live weight. Maximum
dry matter intake (DMI) can be calculated as follows: DMI, kg = 0.025 W +
0.1 FCM. Drinking water intake basically is related to the feed intake, it is
4.2- to 4.7-fold of the dry matter intake. More exactly, milk production, the
weekly minimum temperature and sodium intake should also be taken into
account:
DrWI, kg/day= 15.99+1.58×DMI+0.90×M+0.05×NaI
where DrWI=drinking water intake, kg/day; DMI=dry matter intake,
kg/day; M=daily milk production, kg and NaI=daily sodium intake, grams
(MURPHY et al. 1992).
CRUDE FIBRE REQUIREMENT is 18-23% of DM, but in case of high-
yielding cows during the first 60 days of lactation the allowed minimum can
be of 16%. The average gross SALT need is 0.46% of dry matter. Ranges of
actual supplementation, according the milk production are 0.2-0.6%.
Exactly formulated: 30 grams for maintenance and 2 grams per kg FCM
milk production is recommended. Recommended gross values for minerals
in dry matter are 0.2% Mg, 0.2% S, 40 mg/kg Zn, 40 mg/kg Mn and 10
mg/kg Cu. Minimum vitamin A requirement is 7,800 IU/100 kg LW. (If
added in form of carotene, 1 mg beta-carotene=400 IU vitamin A. Beta-
carotene requirement: minimum 300 mg/cow, but desirable 100 mg for
maintenance plus 25 mg/kg milk produced.
890
DRY COWS. Contracted requirement of maintenance and gestation is

0.436×W0.75 MJ NEl and 7.4×W0.75 +25 grams crude protein (210 g MP
during the first 5-6 weeks of dry period and 272 g MP 2-3 weeks before calv-
ing per cow/day, 6.1 g Ca/100 kg LW, 4.2 g P/100 kg LW and 7,800 IU vita-
min A/100 kg LW. Dry matter needs are 1.75-3.0% of LW, with a 20-34%
crude fibre and 0.20-0.24% salt. For pregnant heifers energy needs are
higher by 15.4% and those of protein by 10.5%.
c) Scheme of ration, optimization. The following natural intake limits
should be taken in consideration: green forages, grass 8-10% of LW, silages
4-5% of LW, haylages: 3-4% of LW, roughages 2% of LW. Minimum daily hay
requirement (no shorter than 2-4 cm) is 4-5 kg/cow. Draft should be opti-
mized according to dry matter, fibre, energy and protein. If “task has no
solution”, the system of conditions should be widened/loosed.
d) Control. Generally deviation equivalent to the requirement of 1 kg
milk production (3.1 MJ NEl and 87 g crude protein » 49 g MP) is accepted
in the practice. If the main components are in order, the mineral supply
should be arranged. In case of a single lack of Ca, let us give limestone (2.5
g/g Ca deficiency). In case of Ca and P deficiency, we use different feed
phosphates. If the Ca need are in order, or in excess and P is deficient, lack
of P should be supplemented and the surplus Ca is ignored.
25.4.8.2. PRACTICAL CLASSIFICATION OF FEEDSTUFFS

PRIMARILY IS IT A FIBRE OR ENERGY SOURCE ?
Classification according to the fibre concentration is useful while planning
the dry period high and the fresh cow low fibre supply. By knowing neutral
detergent fibre (NDF), the expected voluntary dry matter intake can be pre-
dicted. Fibre sources are feedstuffs of more than 50% of NDF content. In the
energy sources the NDF level is lower than 40% (Table XXV-9).
891
Table XXV-9: Classification of cow feedstuffs according to the fibre content*
*“roughage” name is used in this manual for forages, more than 18% crude fibre con-
tent
**lemon, orange, grapefruit pulp
CLASSIFICATION ACCORDING TO THE ENERGY AND PROTEIN CONTENT
Considering possible variants, a matrix can be compiled: rich, medi-

um and poor in energy, as well as rich, medium and poor in protein. In
brackets the rumen degradability of protein is given. For example blood and
fish meal are rich both in energy and protein and their rumen degradabili-
ty is low. Wheat bran is of medium energy and protein concentration and
degradation of its protein is medium, too. On the other hand, wheat straw
hardly has some energy and practically does not provide protein for organ-
ism (Table XXV-10).
892
Table XXV-10: Classification of feedstuffs according their energy and protein content
Rumen degradability (dg): A = 71-90, B = 51-70, C = 31-50,

D < 31%, “-“: practically cannot be considered as a protein source
ACCORDING TO THE PHYSICAL APPEARANCE ruminant feedstuffs are liquid

(molasses, vinas, etc.), high moisture or succulent feeds (green forages,
roots and tubers, silages etc.) and dry (hay, cereals etc.). Using the above
mentioned categories, practically each potential ruminant feedstuff can be
characterised.
Recently, organic foods are increasingly popular. Organic farming
needs specific rules to assure consumer confidence. In Switzerland, a posi-
tive list of feed ingredients has been compiled in order to create trans-
parency and security (HELLER, 2000). The leading Research Institute for
893
Organic Agriculture (FiBL) published a list of products and ingredients that

are allowed in organic farming. Anything not on the list is not permitted
(“limiting list”).
25.4.8.3. Feeding systems

Knowledge of feeding technology at cow extremely important, because ani-
mals have selective feed intake and there is a correlation between fibre
degradation and feeding frequency. If high amount of concentrate is given
all at once, the released organic acids decrease rumen pH up to the critical
5.9-6.0. At this pH protozoa and their symbiotic cellulolytic bacteria die out,
efficacy of fibre degradation and the methane production drops. If concen-
trate is given in several small portions, the continuous saliva production
may compensate acidification in rumen. Fall of pH below 6.2 cannot be
avoided in the fresh period when high amount of concentrate should be
given.
GRAZING. The necessary pasture surface should be determined in site.
In the adjustment period of grazing, as buffering feedstuffs silage and hay
should be given. Extra concentrate supply (about 250 grams/kg of 4% FCM,
or at a constant rate per cow) can be given either in the milking parlour (it
is not recommended in case of high production) or after milking, using indi-
vidual chips and appropriate concentrate distributor automats. The milk fat
level of grazing cow is typically lower, but the concentration of conjugated
linoleic acid (CLA) higher, compared cows on TMR.
BASE FEED PLUS SUPPLEMENTARY MILK CONCENTRATE. Using this system,
each individual cow receives a basal feed, which covers needs of the main-
tenance and 8-12 kg milk production. Basal feed consist of silage and hay.
Cows, producing more milk, receive 400-500 grams concentrate per kg pro-
duced milk. Disadvantage of this method is that it can be used only in tied,
stall keeping and the very low production cows are overfed. Electronic feed-
ing equipment is available, which allows individual cows to be fed a speci-
fied amount of feed and this can be separate out over the day.
FLAT-RATE FEEDING. Each cow receives the same amount of concentrate
and the silage and forage are provided ad libitum. It is a prerequisite that
reproductive status of cows should be more or less synchronized. This
method economises labour but full genetic potential of top animal cannot be
realized.
COMPLETE MONODIET (TMR=TOTAL MIXED RATION). Each feedstuff com-
894
ponent is present in form of a homogenous mixture, which is practically

continuously offered for the cows (optimum five feeding daily, but normally
fed 1 to 3 times per day). In this way the acidifying effect of great concen-
trate intake can be prevented. Rumen microflora and fauna are not obliged
to frequent adaptation. During the first phase of lactation the ratio of con-
centrate to forage is 60 to 40 (with a NEl density of at least 10.4 MJ/kg DM),
in midlactation 50 to 50 and in declining third of lactation 40 to 60 in dry
matter. The starch concentration in TMR should be based on effectiveness
of fibre and on the relative rate of starch fermentation. The maximum non-
fibre carbohydrate (NFC) level for high-yielding cow is 44% and the mini-
mum forage NDF concentration is 19% in DM (NRC, 2001). To assure struc-
tural fibre supply, at least 4 kg hay should give besides the cut hay in TMR.
While optimizing daily ration, the highest value of the feed intake capacity
should be considered. The long-stem hay is an excellent source of effective
fibre, but with a better understanding of the interactions of dietary starch
and fibre, it can recognised that an adequate fibre can be provided in diets
with ensiled forages, too.
If the ideal concentrate to forage ratio shifted, milk production drops.
To high silage and hay portion decreases nutrient intake, too much con-
centrate ingestion alters metabolic status
By using highly digestible silage of small particle size and hay of lower
fulfilment unit (e.g. Italian rye grass) feed intake and milk production can
be improved. In the last two months of lactation protein supplement (extr.
soybean, sunflower) can be replaced by good quality alfalfa.
Improvement of metabolic condition by adding niacin and thiamine,
buffering rumen in fresh period may also enhance milk or milk constituent
production. A new approach of calculating TMR is the CNCPS (Cornell Net
Carbohydrate Protein System). One of the main characteristics of this sys-
tem is the accurate prediction of feed intake on the basis of NDF and fibre
fractions. This model is dynamic and considers not only the protein but also
the carbohydrate degradability in rumen.(see also Chapter X).
25.4.8.4. Preparation of yearly feed balance

On the basis of herd size, lactation number, milk production, available pas-
ture, expected own feedstuff production the quantity of feed to buy can be
predicted.
895
25.4.8.5. Practical veterinary interventions

a) Dry period. The question is not that the use of concentrate is useful, or
prohibited during dry period. Fat cow syndrome can develops on high qual-
ity silage feeding, too. The daily net energy intake should be calculated and
the adaptation of rumen microflora to substrates, (remember advisable con-
centrate feeding during the last 2-3 weeks of dry period). Considering their
high calcium content, the feeding of leguminous feedstuffs (e. g. alfalfa hay)
is not desirable. The occurrence of milk fever can be minimized by giving
vitamin D (25-hydroxy-cholecalciferol) 1-2 weeks before expected calving. At
the same time, in the practice this can often cause more problems, because
if the calf is born early or later then calcium metabolism problems can
occur. High amount of vitamin E (600-700 I.U.) alone or with 50 mg sodi-
um selenite helps in preventing non-infective retention of placenta, mastitis
and cardiomyopathy of newborn calves. Current modern practice feed 1000
IU of supplemental vitamin E/day to dry cows. Another peroral compounds
may given, which help liver function and enhancing glyconeogenesis (propy-
lene-glycol, rumen-protected methionine, niacin, methionine-hydroxy-ana-
logue). Application of some of them should be continued after calving.
b) Lactation. In the first period of lactation the concentrate feeding is
high and that of fibre low. Consequently, rumen pH will be lower than the
optimal. This chronic lactacidosis can be compensated by supplementing
feed mixture or concentrate with buffers. Buffer supplementation increases
rumen pH, improves fibre degradation and acetic acid production.
Recommended concentrations are 1.5% NaHCO3 in concentrate or 0.75% in
total dry matter. MgO, in half dosage is also applicable. By buffering and fre-
quent feeding the so-called “low fat syndrome” may be prevented.
Sulphur to nitrogen ratio in dry matter should be 1 to 10, which in
case of corn silage feeding may require S supplementation, using sodium or
magnesium sulphate, even sulphur powder. To assure good reproductive
performance, the continuous P, Mn, vitamin A and β-carotene supply is
indispensable. Based on health data, current modern practice feed 500 IU
of supplemental vitamin E/day to lactating cows.
To help to achieve a good marketing value of milk (fat and protein con-
centration, somatic cell count) is also part of the tasks of vet. The typical
composition of the modern Holstein Friesian is as follows: 88.32% water,
3.22% protein, 3.25% fat, 0.69% ash, 4.52% carbohydrate, 2.51 MJ/kg
energy, 100 mg/kg cholesterol and 64.9% of total fatty acids are saturated,
896
28.3% monounsaturated and 6.8% polyunsaturated (USDA, 2004). By

using rumen protected methionine (eventually methionine and lysine),
the protein level of milk can be elevated. During the summer months it is
usual to increase the energy density by adding fat. In these cases the
unsaturated fatty acid should not exceed 4% and level of total fatty acid
should not exceed 6% of the dry matter. Higher amount nay interfere with
fibre degradation in the rumen.
The tendency is to reduce total fat content, as well as to decrease its
saturation. The idealistic milk of the future would contain no more than 8%
saturated fatty acids, less than 10% polyunsaturated fatty acids and 82%
monounsaturated fatty acids (Wisconsin Milk Marketing Board, 1998). It is
also a desire to increase the level of unsaturated trans fatty acids, because
of the health concerns. Feeding of rumen-protected linseed or palm oil in
form of calcium salts, the composition of milkfat can be influenced. The
result is a healthier food product.
It is the cis-9, trans-11 CLA isomer (or rumenic acid) represents 75 to
90% of the total CLA in milk fat and it derives from the biohydrogenation of
fatty acids in the rumen and from the endogenous synthesis in the mam-
mary gland. The other bioactive product, the vaccenic acid (trans-11, 18;1)
is also an intermediate of ruminal biohydrogenation of both linolein and
linolenic acid. The key enzyme of the transformation, the ∆9-desaturase can
be found in the mammary gland and other tissues. First the CLA in cooked
beef has been proved antimutagen and later the anticarcinogenic activity of
rumenic acid has been proved. Ingestion of vaccenic and rumenic acid
enriched butter reduced the incidence of mammary tumours in rat model of
breast cancer. Rumenic acid has also significant antiatherogenic effect in
animal model. By feeding plant and marine oil supplement, feeding of fresh
pasture can increase rumenic acid concentration in cow milk 2- to 3-fold.
Besides CLA, the range of functional foods in milk is wide, including omega-
3 fatty acids, oligosaccharides, nucleosides, probiotics, having positive role
in struggling against cancer, cardiovascular diseases, hypertension,
immune deficiency and skeletal diseases.
In some regions of the world (e.g. USA) the use of growth hormone in
order to stimulate milk production is also applicable. The biological prior
obligation of GH use is an excellent environment and feeding as well as a
good body condition score of cows. Exogenous growth hormone primarily
increases milk volume, with relative little effect on milk composition. Since
897
lactose is the major osmotic constituent of milk, increases in milk secretion

can only be achieved by improving glucose supply of the mammary tissue.
In Europe the use of GH is not authorized. In the United State the use of
monensin for lactating cow was approved, mostly for reducing the risk of
ruminal acidosis and ketosis. Results with yeast cultures proved to be vari-
able; however for improving ruminal fibre degradation are widespread
included in diets for lactating cows. For monitoring metabolic state, the
measurement of keton bodies in urine and that of urea level in milk offer
easy to execute methods (for details see “Veterinary nutritional consulta-
tion” subchapter).
898
CHAPTER XXV: BREEDING BEEF
25.5. Beef cow and replacement heifer

In beef cow operation the reproduction is of crucial importance of prof-
itability. The minimum requirement is one calf/cow/year. While describing
principles, the numerical data relate to the European type Hereford breed
and its crossings. The beef cattle is generally kept extensively, in winter
receiving dry bedding, good quality silage and hay. Central role of repro-
duction is reflected by the naming, calling it as „cow-calf-production” or
operation. Efficiency of the latter depend upon the number of calves
weaned, i.e. on their weaning weight (influenced by the dam’s milk produc-
tion and the feeding in the kindergarten), the yearly calving percentage, the
feeding costs of the mother cow and the selling prize of the weaned calf.
Calf mass per cow reflects contracted the number and weight of
calves. It is influenced by the cow’s reproductive performance (mostly
depending on body condition); by calf survival and its genotype (breed). Calf
weight is determined by the dam’s feeding during gestation and lactation.
The role of gestational feeding is more important, because deficient supply
during the suckling period can be compensated by mobilizing the cow body
reserves. However, the worsening condition delays conception and the year-
ly cycle is getting upset.
In details, the feeding during gestation determines the birth weight
and the viability of calf. The ongoing lactational milk production and body
weight gain of calves mostly depend on the cow’s nutrient supply during
pregnancy, as well as her body condition score and the onset of cyclic ovar-
ian activity. In turn, feeding during lactation influences the weaning weight
of calf and the time of conception. Nutrient supply of beef cattle generally is
marginal. Thereby, the main question always is the energy supply. Even a
protein shortage will reflect as an energy deficiency, because of the lower
activity of cellulolytic bacteria and the concomitant less volatile fatty acid
production in the rumen.
25.5.1. Feeding of replacement beef heifers

There are four periods in the raising up of replacement beef heifers.
A) FROM BIRTH TILL WEANING. Individual identification should be
applied; “labelling” at least one third more calves than that of the planed
replacement number. Besides the suckling, the concentrate supplementa-
tion in calf kindergarten is indispensable, except small frame breeds, which
are prone on getting fat. In other words, heifer calves that are to be retained
899
CHAPTER XXV: BEEF CATTLE FEEDING & NUTRITION
for breeding stock replacement may need a liberal feeding program so that
they can be bred at a fairly young age. Calf kindergarten and school is a
roofed and fenced in feeding space, where the adult cow won’t go in.
Concentrate, mineral and salt supplement and calf hay are fed there. Bull
calves for breeding purposes also need a feeding intensity, assuring opti-
mum growth. Calves, intended to fatten and slaughter, in the USA may treat
by steroids. In Europe it is not allowed, but for replacement heifer it should
not be used anywhere.
B) FROM WEANING TO MATING. Taken Hereford as a model, optimum
weight for weaning is 180-200 kg. The goal of rearing is, that the heifers
achieve by mating the 65 to 70% of their mature weight. In case of a
Hereford, it is 300-330 kg and desirable having attained no later than the
age of 14th months. This can be assured by a growth rate of 500-700 grams
average daily gain (it is lower, than in case of dairy replacement heifers, but
the actual weight will depend upon breed. The most important concern in
feeding replacement heifers is to make sure that they are fed an adequate
plane of nutrition so that they are not undernourished or stunted in growth
for an extended period of time. Replacement heifers may be kept both on
pasture and in drylot after weaning; the important is the continuous con-
trol of their nutrient supply. The minimum daily requirement of calcium
and phosphorus ranges from 10 to 20 grams per day. Mating season should
be started with the heifers, which should be kept separately from the adult
cows. Allowing heifers only a short, 35-day-long breeding season, non-con-
ceived animals should be culled. This is a selection for good reproduction
traits, too and this is the reason of extra number at individual identification
after birth.
In a field trial of VARHEGYI and VARHEGYINE (1987) average (180-190 kg
LW) and retarded in growth (150-160 kg LW) in autumn weaned heifers
were designed in two groups (Table XXV-11). They are interested that feed-
ing 770 grams extra corn meal daily for the retarded heifers, whether are
they able to recover their condition. After their data, there is some compen-
sation and heifers of both groups could be mated in May. (Of course, it
depends on where you at in Europe or the World – mating should be
sequenced with the available feed supply for a given location)
900
Table XXV-11: Characteristics of beef heifers (Hungarian Spotted×Hereford F1) receiving

feeding of different intensity during raising up
C) DURING THE FIRST PERGNANCY. Supposed that conception happened

at the age of 15 months, at time of calving cow is 24 months old. At that
time is desirable the achievement of the 85-90% of the mature live weight.
To assure that, above the needs of gestation, nutrient requirements of 400
grams average daily gain should be provided (it depends upon breed too).
D) PHASE OF FIRST LACTATION: further 250 grams average daily gain
should be ensured for the cows. For that reason, it is advisable to keep first
lactation cows separately. Cows attain mature size at the age of 60 months.
Optimum numbers for European Herefords are as follows: body weight at
puberty (first fertile oestrus) 331 kg, at the age of 426 days (14 months).
Consequently, optimum time range for mating is between the ages of 13 to 15
months.
If during winter the nutrient supply is insufficient, reproductive per-
formance of pregnant heifers depress. For example, feeding only grass hay
of poor quality, the onset of the first oestrus delays, conception rate and
weaning weight of calves are lower than animals with concentrate (1 to 3
kg/day/heifer) supplementation, even when lactational nutrition does not
differ.
25.5.2. Nutrient requirement of the beef cow

As it was mentioned, that the key nutritional factor of the beef cattle oper-
ation is the energy supply. Energy requirements of beef cow are calculated
in MJ NEm, both for maintenance and milk production and for pregnancy.
901
In case of beef heifer energy needs of maintenance is also given in MJ NEm,

but for the daily live weight gain the MJ NEg is used for calculation. Energy
requirement for maintenance equal to 0.3347×W0.75 MJ NEm. Efficiency
for building of the foetus and foetal membranes, the transformation of ME
to NE (the “k” value) is low, only 13%. Needs of the developing foetus can be
described by using and exponential equation, which shows that 70-80% of
retention in foetus occurs in the last 50 days of gestation. During this peri-
od, calculating for example with a calf birth weight of 36 kg, the daily net
energy accretion is 9 MJ and that of the protein 55 grams, which can be
realised, if the cow has an average daily gain of 400 grams..
Energy requirement of milk production can be predicted on the basis
of the milk fat concentration: NEm, MJ = 0.418 FAT% +1.46, which gives
on an average 3.14 MJ NEm per kg milk. Milk production achieves 15 kg
during the first phase of suckling, declining close to zero to the end of the
seven-month-long lactation period. In case of growing heifer, pregnant
heifer and first lactation cow there is a need for calculating also the require-
ments of daily weight gain, which is given in MJ NEg.
Prediction of voluntary dry matter intake for beef cow is as follows:
DMI, kg = W0.75 (0.1462 NEm-0.0517 NEm2-0.0074). In the equation the
metabolic body size and the energy concentration of the dry matter (in
Mcal/kg; to receive MJ, multiply by 4.184) figure and it is generally equal to
2.ö-2.1% of live weight.
Protein requirement of beef cow. Protein needs can be calculated
by using the factorial method. All output (incl. retention in conceptus and
also the milk production) stand in the numerator, divided by denominator.
i.e. by the digestion coefficient and the biological value of protein appearing
in the small intestine. Protein accretion into the conceptus is 55 grams daily
in the last third of gestation, to assure that 90-100 grams crude protein
should be provided in the daily ration. The protein amount, excreted by
one litre of milk is on an average 33.5 grams; therefore in the daily
ration 55-60 grams crude protein (52 g MP) should be given.
Influence of environmental temperature: being the lower critical tem-
perature for beef cow approximately zero degree Celsius, each degree below
that increases energy needs by 2%.
25.5.3. Yearly production-reproduction cycle of beef cow

According to the general scheme, calving is timed to the end of winter; how-
902
ever there are some operations with calving periods in autumn, too.
Idealized form of the cycle can be seen on FIGURE XXV-10. Time of calving
is on February 15 (plus minus 1 month), when Period I begins and this is
the so-called fresh period, lasting 80 days. It is finished by the remating,
on May 5. During the 125-day-long subsequent Period II, beef brood cows
are simultaneously suckling and pregnant. In autumn, at about September
8, calves are weaned and the 110-day-long Period III begins, i.e. the middle
of the gestation. Needs of cow during this period are low. The following
Period IV (from the December 26) is the last 50 days of pregnancy, while
70-80% of foetal development takes place and its extra requirement should
be taken into account. With the birth in the middle of February is the begin-
ning of the new cycle. Period III and IV give the dry period.
FIGURE XXV-10 Production-reproduction yearly cycle of beef cow (opposite the clockwise
Nutrient requirements of beef cow are practically to give in compari-

son to the Period III because in this period there is no milk production yet
and needs of pregnancy are very low. Taken the energy and protein require-
ments of this phase as 100%, the others can easily be expressed (Table XXV-
12).
903
Table XXV-12: Nutrient requirement of beef cow in the periods of yearly cycle
**in form of injection
Accordingly, in Period II (cow is still suckling and is already pregnant)

and in Period IV (needs of gestation are already important), energy and pro-
tein requirement are approximately by 25%, and these of Period I (the first
80 days of lactation) by 50-60% higher, than in Period III. Calcium and
phosphorus needs are relatively lower, than in case of the dairy cow. Extra
energy, concentrate supplementation, two weeks before mating, has a cer-
tain flushing effect, conception rate, even the percentage of twin calves
increases. Vitamin A and carotene status should be monitored and by feed-
ing carrot or/and vitamin A injection can be given. Especially animals on
dried mature forage will have limited carotene intake to fulfil the vitamin A
requirement. However, cows usually have approximately three months of
vitamin A reserve stored in their liver. Thus, vitamin A supplementation
became critical only in winter. When it is not possible to feed beta-carotene
sources (carrot) or carotene-vitamin A supplement under grazing condi-
tions, every 90 days 1 million I.U. vitamin A should be injected. In oppo-
site case reproductive performance and calf viability may be altered. In
tables used in practice, energy requirement of maintenance and milk pro-
duction are given in MJ (or Mcal: USA) NEm and that of the daily gain in
NEg. Dry matter intake and protein needs (crude protein or MP) are given
according to the live weight category.
In case of the breeding beef cattle two types of body condition scor-
ing are used. One is the Scottish 5-score-system (see dairy cow); the other
is the 9-score Nebraska System. The latter classifies according the fat cov-
904
erage of ribs, shoulder, loin and tailhead region. According to the Nebraska
System, the ideal condition score of cow at calving is five. In this case the
fat content of her body is approximately 19% (NRC, 2000), has sufficient
reserves for the lactation and the onset of cyclic ovarian activity and the
successful conception can be expected in time. Body condition at calving is
in close correlation with the time of successful conception: the lower is BCS,
the later can the cow conceive. In case of those technologies, when there is
also a calving season in autumn, the ideal BCS at parturition is six.
Body condition scoring is a more accurate tools in reflecting the total
body fat and energy content, than weighing. To increase BCS 4 to BCS 5,
cow needs to retain 36 kg dry matter (mostly fat and some protein) in her
body. Supposed that the live weight of a beef cow is 500 kg, for improving
her condition (36 kg increment) and for the development of foetus and foetal
membranes (45 kg), summed up requires 81 kg weight gain, which can be
distributed to the last hundred days, gives 810 grams average daily gain.
25.5.4. Feedstuffs of breeding beef cattle

The most important are grazing on the pasture and the meadow hay, in
ideal situation 50-50%. (In Middle Europe on an average pasture one
hectare is required for keeping a beef cow and a calf. According to the soil
and weather conditions, this number may vary. Beef cows efficiently utilise
corn and milo fodder, corn stalks and even corn and milo stover (stem).
Fodder greens are also applied, in this way Sudan Grass, cereal forages
(green wheat and oats), corn herbage (young whole corn plant before cob
formation), whole plant corn and sorghum. Most of them can be used by
grazing, too. Soybean and pea hay, straws of oats, rye, barley and wheat are
also fed. Corn cob, rice and cottonseed hulls serve mostly as source of fibre.
In opposite to the fattening beef, the use of NPN compounds is rare,
because the energy supply generally is marginal and there is no surplus for
the assimilation of released ammonia (e.g. when low quality forages are
being fed during winter feeding periods). Protein need can be covered by
feeding alfalfa hay, but it may lead to protein overfeeding. To improve the
BCS, the use of alfalfa hay is not enough. In autumn the grazing of corn
stubble field (and other aftermath forages) is an important feed source, pro-
viding considerable amount of cob and grain, too. Sugar beet pulp or it top
can be given fresh, or in form of silage (fermented together with chops of
corn stalk and molasses). In wintertime, if freezes and the fresh sugar beet
905
pulp get frozen, the cows cannot ingest sufficient amount from it, thus sup-
plementation is necessary.
Concentrate is used only for calves, or for flushing, or improving BCS.
Continuous salt supply (in most cases in form of licking salt) and in the
majority of cases, phosphorus supplementation is necessary. Depending on
the local soil characteristics, copper, selenium, possibly zinc, manganese
and cobalt maybe required. It can be realize by mixing them into the licking
salt, or by the putting long-release bolus in the reticulum. At the end of win-
ter, by feeding carrot and giving vitamin A premix or injection the healthy
reproductive functions can be supported.
,
906
25.6. Fattening beef

25.6.1. Energy requirements
From data of Table XXV-13 and FIGURE IX-3 it can be seen that utilisation
of metabolizable energy (k) depend upon the field of usage (maintenance or
gain) and on the metabolizability (q = ME/BE) of the ration, in other words,
on the concentrate to roughage ratio. It is important from the point of view
of beef cattle, the effect of metabolizability in case of maintenance and gain
is are quite divergent from each other and the difference is not constant.
Consequently, the two types of partial energy should be expressed by dif-
ferent unit, i.e. maintenance is calculated using NEm and live weight gain
by using the NEg. But ME efficiency for growth can be biased by several
other factors; one of the most important is the feeding level. One feeding
level equal to the maintenance energy requirement; above feeding level of
1.2, corrections are necessary.
Table XXV-13: Utilization of ME for maintenance and gain
*Forage of 8.4 MJ ME/kg, concentrate of 13.4 MJ/kg
Component of needs for fattening. Maintenance requirement is as fol-

lows: NEm = 0.322×W0.75 MJ. This is equal to the fasting heat production
in thermoneutral zone. It can be measured by the determination of respira-
tory coefficient. On the other hand, NEg was measured by comparative
slaughter technique. Maintenance requirement depends only on the meta-
bolic body size (i.e. the surface of the animal). On the contrary, NEg-require-
ment is influenced by sex (heifer, bull, steer), by the mature live weight,
frame size (small, medium and large) and by the average daily gain. For the
improvement of adult, lean culling cow the energy need/kg weight gain is
25.94 MJ NEg.
907
25.6.2 Prediction of voluntary dry matter intake

The average dry matter intake is 2.3% of live weight. There are some devia-
tions, because animals of small frame tend to eat relatively more, and those
of large frame relatively more. The daily dry matter intake of the medium
frame size bull can exactly be calculated on the basis of metabolic body size
and net energy concentration of the dry matter (Mcal/kg):
DMI, kg = W0.75 (0.1493 NEm - 0.046 NEm2 - 0.0196). For example, if
a corn silage contains 6.99 MJ NEm (i.e. 1.67 Mcal) per kg dry matter and
a Hereford bull is of 400 kg of live weight, the daily dry matter intake is 9.07
kg
(DMI= 4000.75×(0.1493×1.67-0.046×1.672–0.0196)=89.44×(0.2493
0.1283-0.0196)=89.44×0.10144=9.07),
which is 2.27% (9.07×100/400=2.27) of its body weight. The com-
monly used large frame breed in Europe are the Charolais, Belgian Blue,
Blond d’Aquitaine, Simmenthalian and Holstein Friesian, from the medium
frame the Hereford and its crossings and from the small frame size the
Aberdeen Angus. The above described equation relates to a medium large
bull, for heifer values are 10% lower. No correction is necessary in case of
large frame size heifer, but for large frame bulls the value should be
increased by 5%.
25.6.3. Modifying effect of environmental temperature

Cold environment has a greater influence on the energy needs of fattening
beef, than in case of dairy cow, because of the lack of extra heat of milk pro-
duction. Thermoneutral zone of beef cattle is between 15 and 25oC, where
animals use only the physical heat regulation to maintain core tempera-
ment of their body. Chemical heat regulation (fat mobilization) occurs only
below lower critical temperature.
Cold environmental temperature increases, warm decreases volun-
tary dry matter intake. If the cold weather is combined with other drastic
stressors (e.g. deep mud, snow etc.), the appetite may be blocked. Digestion
of high fibre feeds produces heat and added to the effect of environmental
heat. Consequently, drop in feed intake appear earlier while temperature
increases. Digestibility also is influenced by environmental temperature.
Cold speed up peristalsis therefore decreases digestibility of nutrients.
Correction can be made using .A = B + B[(0.001(T-20)], equation, where A
= corrected (adjusted), B =basal value at 20oC and T = environmental tem-
908
perature, oC. For example at zero oC, the digestibility is lower by 2%. If tem-
perature differs from 20oC, the irradiated heat and the maintenance
requirement will change. Deviation from 20oC by ±1oC causes ±0.003 MJ
changes, therefore the multiplier constant (0.3222) of maintenance require-
ment should be corrected.
25.6.4. Protein (nitrogen) metabolism of beef cattle

Crude protein requirement is calculated by means of the so-called factorial
method. It means that based on data of controlled trial, then characteristic
factors (constants) are accepted to use at the key points of N-metabolism.
For example, biological value of protein was taken as 66%, etc. First of all,
the sources of losses (protein output) are listed. From this point of view the
protein accretion in muscle is also an expense). There will stand in nomi-
nator, which is equal to the net requirement. After that, divided by the fac-
tors of transformation and utilisation losses (denominator), the gross pro-
tein need is received. In formula
CP = (F+U+S+G)/(D × BV× CE), where CP = daily crude protein require-
ment, gram, F = metabolic faecal losses, 3.34% of ingested DM. Its sources
are the desquamated epithelial cells and protein content of bacteria, U =
daily urinary protein loss, equal to 2.75×W0.5, S = scurf protein loss,
0.2×W0.6 and G = tissue gain, gram (268-7.02 X). The “X” in expressing pro-
tein accretion is the energy content of 1 kg weight gain (MJ). Its value
depends on the age. On an average one gram deficit in feed reduces body
weight gain by 10 grams. The D = true digestibility, average value 90%, the
BV = biological value, average value 66% and CE is the conversion efficien-
cy, i.e. percent of intake protein appearing in abomasum, average value
100%.
Summarized value expresses both maintenance and weight gain pro-
tein needs. The Hungarian metabolizable protein system starts out from the
net requirement of growth:
MP need, g/day = (NET REQUIREMENT)/100 × (63-0.0371 W) (SCHMIDT
et al. 2000). NRC (2000) takes into consideration of the body composition of
different genotypes while compiling its tables.
For fattening beef the protein degradability is not generally pre-
scribed, because only in the very low live weight category (100-200 kg) may
be necessary to calculate with the undegradable protein requirement. Above
this weight category, the total protein needs of medium and heavy live
909
weight beef cattle can be covered with RDP protein. Consequently, there is
an opportunity to use of NPN compounds. One-time toxic dose of urea, with-
out adjustment is 0.3-0.8 g/kg LW. Drylot cattle with high concentrate
intake may receive relatively more, because there is enough energy in
rumen for the assimilation of released ammonia, moreover, from the more
acidic rumen the absorption of ammonia[ammonium is slower.
Supplementation of sulphur, potassium and phosphorus is essential to
assure optimal bacterial protein synthesis. Accepted practical allowance is
to feed no more urea than 10 g/100 kg LW. On the basis of urea fermenta-
tion potential, the exact amount of possible urea supplementation can be
calculated
UFP, g/kg DM = 11.78 NEm+6.85-0.0357 CP × dg, or
UFP, g/kg DM = 31.64-3.558 CP + [(945 NEm-887-179 NEm2)]0.5,
where CP = crude protein, %, NEm = Mcal/kg and dg = rumen degradabili-
ty of protein, %. (Details about urea toxicity see below). Other NPN sources
(biuret, vinas, ammonium chloride, poultry deep litter etc.), according to the
local regulation, can also be used.
25.6.5. The compensatory growth

The compensatory growth is a capacity of growing animals to recover lag in
body weight gain, developed during a period of low intensity, when the real-
ization of the full genetic potential was not possible (FIGURE XXV-11). The
notion is generally true, but in case of growing beef cattle it is consciously
applied.
FIGURE XXV-11: Compensatory growth of growing cattle
910
Basically economic considerations can be found in the background,

for example limited feed base, high price of concentrate etc.) In case of beef
cattle, if they will not get very stunted during the low intensity period, it can
be used. Namely, in the subsequent period of rich nutrient supply, animals
are able to recover their delay. It is not desirable to apply at replacement
heifers, because for them the continuous nutrient supply is appropriate,
having every period of time an organ, or tissue of higher growth rate.
How can this explain the compensation capacity ? First of all, the
metabolic rate during the period of undernutrition slows down and remains
lower during the first part of recovery, which reduces maintenance require-
ment (FERREL et al. 1986). Secondly, skeletal and muscle tissue continue to
develop, which allows for a rapid “filling of cells” during recovery and com-
pensatory fat accretion, resulting in a return to a normal growth pattern.
Thirdly, efficiency of energy (ME→NEg transformation) and protein use con-
tinues to be more efficient during the recovery period.

The needs of drinking water is considerable influenced by the environmen-
tal temperature and is given related to the ingested dry matter (Table XXV-
14).
Table XXV-14: Water requirement of beef cattle in function of environmental temperature
Other factors influence water requirements, wind and water mineral con-
tent.
25.6.7. Methods of fattening

FIGURE XXV-12 reviews different fattening systems and technologies.
About the frequency of actual utilisation the economic factors have the
greatest influence. Breeds are both beef, or bull of dairy breeds.
CALF FATTENING
a) up to 80-120 kg LW (the so-called “milky” calf; very expensive)
911
b) slaughter weight is 180-200 kg LW – useful thumb rule that it is as heavy

in kg LW as old is in days. A special version of it is the “white muscle calf”,
which is as a matter of fact an artificial iron deficiency (veal( and consecu-
tive. It is popular in some market, but from the point of view of animal wel-
fare it is less and less practised.
FIGURE XXV-12: Beef fattening methods (after BODO, 1988, modified)
BABY-BEEF FATTENING – up to a slaughtering weight of 350-400 kg. It is

appropriate for small frame size breed, because they reach their maturity at
that weight limit.
FATTENING OF GROWING BULL (in case of large frame size breeds, like
Simenthalian, Hungarian Spotted, Holstein Friesian and their crossings).
The desirable final weight is 550-600 kg. It can be achieved by using one of
the following feeding systems.
a) Fattening based on concentrate, using only 1-2 kg of hay daily. The
expected average daily gain is 1500-1600 grams and animals achieve
slaughter weight at the age of 12-13 months. Incidence of health troubles
may be high, namely rumen acidosis, lameness, liver abscesses, special
form of rickets and urolithiasis (for details see later).
b) Fattening, based on forages, less or more concentrate supplemen-
tation. Expected average daily gain 1100-1400 grams, attaining of slaugh-
ter weight at the age of 16-18 months. Among the forages the corn silage
and the agricultural by-products should be highlighted, but pea straw, stem
of seed alfalfa, humid, crashed barley and poultry deep litter may be men-
tioned as peculiarity.
FATTENING OF GROWING HEIFERS. In principle is the same, than that of
the growing bulls (see above), slaughter weight is at about 400-500 kg of
912
LW, because exponential fat accretion begins earlier at females than at

males.
STEER FATTENING, SLAUGHTERING IN WEIGHT OF 600-650 kg, at the age of
two or three ages. It is applied in regions, where pastures are abundant and
the use of compensatory growth makes this technology economic.
The above described beef fattening methods are realised in North
America in the following different forms. The most economic way is keeping
cattles on the pasture, this is the rangeland, or the animals are fed in
range forage conditions. Nevertheless, when a surplus or large quantities of
grain are available, the cattle may be finished for slaughter to meet the mar-
ket demands for tender, tasty and juicy of steaks and other delicious cuts
of meat, the latter system is commonly referred to as fattening cattle in feed-
lot. Large feedlots are found in areas where feed grain and forages are
grown or readily available. Each pen usually holds 150 to 250 head. Cattle
are fed by mobile mixing trucks with side delivery spouts. In some cases, it
may be necessary to feed cattle (related not only for fattening cattle!) under
confined conditions, generally referred to as drylot. Steers and heifer calves
that are weaned during the fall months are shipped to the feedlots to be fed
on growing rations, or to winter wheat pastures. The calves that are fed
growing rations for four or five months are commonly called as “back-
grounded” cattle.
IMPROVEMENT OF MATURE CULLING COWS takes place parallel to the last
some months of milking, or just after that. The aim of the extra feeding is
to achieve marbleness of muscle, i.e. to enhance deposition of intramuscu-
lar and subcutaneous fat in the carcass.
MEAT QUALITY. Compared the average daily gain of bull, heifer and
steer after the age of 16 months, the bulls showed the best weight gain and
feed conversion efficiency. On the contrary, from the aspect of meat tender-
ness (measured by means of Warner-Bratzler method), taste and water con-
tent did not differ.
25.6.8. Feedstuffs for beef cattle

FORAGES. Beef fattening is economic, if only it is based on pasture, forage
greens, silage, haylage and hays. Grazing of young corn herbage and Sudan
grass or other sorghums is common. Protein quality is an important aspect.
Nitrogen content of green grass and good quality hay is mostly true protein.
In good quality silage 40% of crude protein is as amid compound present,
913
and there are some percent ammonia losses. In poor quality silage the
amount of true protein is even lower and ammonia losses are important.
For combining forages, it is important to know that hay supplemen-
tation of silage decreases fattening performances, while concentrate addi-
tion to hay improves them. The explanation of the described phenomenon
is in the substitution value (see Chapter VI).
AGRICULTURAL BY-PRODUCTS. Cereal straws, corn stalk and corn and
sorghum stover, chaffs, corn cob, leguminous hays (pea, soybean) and
straws, corn, rice and cottonseed husk, cap of sunflower, sugar beet pulp,
its top with the leaves. Grazing on field-crop aftermath (stabble field) is also
popular.
INDUSTRIAL BY-PRODUCTS. Sugar beet pulp, (pressed, dry in loose form,
possibly shredded, as pellets, extruded or ammoniated), malt sprout or
returns, distillers dried grain with and without soluble, distiller dried solu-
ble and condensed distiller soluble (moreover distiller wet molasses, sugar
beet and potato), grape marc (husks and skin of pressed grape), wet and dry
hominy feed, husk and skin of pressed apple and tomato (apple and toma-
to pomace) and other by-products of canned food industry.
An example for the traditional daily ration of a fattening bull of 500
kg of LW: 10 kg corn silage, 10 kg top of sugar beet with leaves, 4 kg pea
hay (straw), 2 kg alfalfa hay of medium quality and 3 kg corn meal or
cracked corn (dry rolled).
25.6.9. Growth promoters in cattle fattening

In the European Union, except rumen buffers, utilisation of the described
compounds below is not permitted. On the contrary, in North America both
anabolic steroids (11 commercial products) and the ionosphere antibiotics
(lasalocide, laidlomycine propionate, monensine and salinomycine) are
allowed and applied in the daily (NRC, 2000). Growth promoters of fatten-
ing cattles can be classified into two main groups: A) compounds, influenc-
ing rumen function and digestion and B) preparations, improving the more
efficient utilisation of the absorbed nutrients.
A) Influencing rumen function
ANTIBIOTICS: monensine, lasalocide, salinomycine, narazine, lizocelline
(ionophores), avoparcine (glycopeptide) transform rumen microflora, name-
ly decrease the number of methane and carbon dioxide producing bacteria,
sparing energy for the host animal. These cocidiostats were designed to dis-
914
turb the cation transport through the cell membranes of cocidia, but they
have a similar effect in case of some rumen bacteria. Owing to the regula-
tory mechanisms, the voluntary feed intake generally drops and the eco-
nomic benefit is rather a better feed utilisation than a higher average daily
gain. In Middle Europe the use of monensine sodium (Rumensin premix)
used to be allowed, up to 250 kg LW in a dosage of 125, above 250 ppm. It
should be borne in mind that it is toxic for horse, rabbit and carnivore and
it is incompatible with the thiamulin.
According to some recent data, ionophore antibiotics act not only by
inhibiting methane production and by enhancing propionic acid production,
but also they decrease basal metabolic rate and increase the undegradable
portion of the intake protein.
DIRECT INHIBITION OF METHANE FORMATION
Each compound, inhibiting methane formation from hydrogen and
formiate, attain the above described effect. Namely the nitrates and sul-
phate, withdrawing directly the hydrogen, as well as unsaturated long-
chain fatty acid, which at the same time pushing down the number of
methane producing bacteria, have the same general influence. Derivates of
chloral hydrate (e.g. hemi-acetal chloralhydrate starch, HCS) kill
meathanobacteria, bound hydrogen and vitamin B12 have similar total
effect.
ANTIBIOTICS OF OTHER MODE OF ACTION: flavophospholipol, tylan and
other compounds contribute to the maintenance of the balance of rumen
microflora, which will result in an improving feed utilisation and decreased
number of liver abscesses. It worth mentioning that application of broad
spectre antibiotics into the rumen is not advisable, because it may kill the
useful members of the microflora and fauna, too.
By supplementing with RUMEN BUFFERS (NaHCO3, MgO), as well as ion-
exchanging clays (zeolite, bentonite) the harmful side effects of high con-
centrate feeding can be compensated and the more efficient utilisation of
rumen ammonia facilitated.
B) Repartition agents
Compounds of this group have postabsorptive effects and modify the
utilisation of absorbed nutrients.
STEROID IMPLANTS
In the beef fattening proved suitable the application of feed additive or
subcutaneous, long-release implants, which will eliminated with safety to
915
the slaughter time. Being steroids (melengestrol acetate, MGA, oestradiol,

Synovex, testosteron and the exception, the trichothecen-structured zear-
alenon, Ralgro), they are anabolic, having a direct positive effect on protein
accretion. Their use is allowed in North America, but in Europe it is strict-
ly prohibited.
GROWTH HORMONE (GH: growth hormone or somatotropin), as well as
GHRH (GH releasing hormone). Main factor of their mode of action is that
they improve the utilisation of absorbed amino acids for protein accretion,
promoting in this manner lean body mass growth. Larger animals have rel-
atively smaller maintenance costs, the need of maintenance get “diluted”.
Fat retention decreases, but the exact mechanism of that is not still known.
The total energy utilisation remains unchanged. Most efficient protein
accretion can be realised only at appropriate protein and amino acid sup-
ply. However, there is no danger of residuum formation; its practical use is
discussed: it is allowed in the USA and prohibited in the European Union.
PHENETHANOLAMINE REPARTITIONING AGENTS
The isoproterenol-structured beta-1- and beta-2-agonists compounds
(e.g. clenbuterol, simaterol, ractopamin, ritodrin, salbutanol and L-
644,869), giving continuously to the feed, increase in carcass the proportion
of protein by 10% and that of fat decrease by 9-12%. Especially retention of
saturated fats decreases. They act by binding to the adrenergic beta-2-
receptors of the organism all over the body. On adipocyte surface they block
lipogenic effect of the insulin, blood supply of fat tissue improves (vessel
effect), which in turn, promotes the integration of free amino acids into the
muscle protein. They stimulate the GH and insulin secretion and at the
same time, inhibit muscle protein mobilization. If their application stops,
the regulation of body replaces the fat deposition during some weeks. Their
application is not allowed in Europe.
25.6.10. Deficiency syndromes, metabolic troubles and poisonings of

beef cattle
Grazed and harvested forages frequently may contain toxicants which
directly harm the ingesting animals. Some of the fattening systems (range
keeping itself, high concentrate intake, use of urea) may also have adverse
effect on health.
“WASHY” PASTURES AND SAND IMPACTION. Plants of early spring pastures
are high in water and protein and often low in dry matter and they may
916
cause watery faeces. Consequently, cattles should not be turned to pasture

too early. Adjustment to lush pastures should be gradual, with the addition
of some hay, straw or corn stalk. If the range has sandy or muddy soil, or
the pasture is on a drainage area, cattle may ingest so much sand or soil
that pathological amounts collect in the reticulo-rumen and in abomasums.
Since ailment develops in latency, frequently it results in death of the ani-
mal.
BLOAT. The ingestion of high moisture, immature growth of alfalfa,
clovers and seldom sweet clover has the potential for frothy bloat.
Formation of a frothy, stable foam in the rumen result in a retention of gas
produced in normal rumen function and in inhibition of belching (eructa-
tion). In acute cases the abdominal cavity becomes severely distended,
breathing is laboured, cattles go down and death is often the end. The prin-
cipal foam-causing agents in legumes are the soluble leaf proteins. Legume
bloat is primarily a problem with grazing cattle, but it can prove equally
serious when they are fed in form of green chop. For treatment and preven-
tion the use of poloxalene is recommended, in acute cases as ruminal
drenching, for prevention can be provided in a liquid molasses-based sup-
plement, mixed or included with the dry supplement. To avoid clinical
cases, gradually increase the time that cattle have access to legume pas-
ture. Once cattles are accustomed to alfalfa or clover pasture, leave the ani-
mals on the pasture constantly, even at night. The legume pastures have to
reach the bloom stage, when cattles are initially turned on. The portion of
leguminous plant in pasture should be kept between 35 and 50%.
RICKETS. Under practical feeding conditions for older cattles, consum-
ing normal forages, no calcium or vitamin D supplementation is necessary.

Very intensively (1600-2000 grams average daily gain) fattened bulls on
high-grain ration, without these additions are prone to develop a special
form of rachitis. The weakness of bones manifest in breaking down the heel
bone (at the hock joints, os calcaneus) together with the Achilles tendon,
causing lameness. Thus, affected animal should be culled for emergency
slaughter.
“GRASS TETANY” is characterised by low serum magnesium concentra-
tions (hypomagnesaemia), which results from a simple magnesium deficit in
the diet or more often from reduced availability and absorption of forage
magnesium. Potassium and nitrogen excess in soil and feedstuffs are the
more important factors decreasing magnesium availability. Grass tetany
917
potential in grazing animals associated with spring grain forage, crested

wheatgrass and other cool-season grasses in lush growth stages. Green
chop from equivalent sources may also cause troubles. Grass tetany occurs
in tetany-prone areas (northern Europe, north-eastern Nevada in the USA
etc.), most often in older, lactating cows, but also in fattening beef, recent-
ly turned onto cool-season pasture in spring. Clinical signs include nerv-
ousness, muscular incoordination, staggering and later paralysis. Death
usually happens within 2 to 6 hours if affected animals are left untreated.
For acute treatment the application of intravenous injection of magnesium
sulphate or calcium-magnesium gluconate is applied. In sensitive territories
the magnesium supplementation of the daily ration, magnesium fertilisation
and introducing legumes into grass meadow are recommended.
HARDWARE DISEASE or traumatic gastritis is used to describe the condi-
tion that results from swallowing foreign materials, usually metal (wire, nail,
screw, pins etc.). Hardware disease is a problem in cattle because of their
eating habit and stomach arrangement. The usual source of sharp metal
objects in the feed and in most cases the object is found in the reticulum.
Sharp objects may injure the lining of the reticulum and cause infection and
inflammation. For the prevention, application of strong magnet in the feed
processing equipment and at the outlets of mechanical silo unloaders is rec-
ommended. Powerful magnets may be permanently placed in the cow’s sec-
ond stomach for the prevention, but the treatment is veterinary surgery.
UREA TOXICITY or more precisely, ammonia poisoning develops, if feed-
ing of high level of NPN-sources without previous adaptation and lack of

available energy for the assimilation of released ammonia (see UFP). This
situation result in a rapid accumulation of ammonia in the rumen fluid,
which in turn, causes a rapid rise in rumen pH and rapid absorption of
ammonia across the rumen wall. When the rate of ammonia absorption
exceeds the capacity of the liver to convert it to urea, ammonia accumulates
in the blood and the toxicity may appear.
Symptoms appear to be progressive as follows: nervousness and
uneasiness, excessive salivation, muscular tremor, incoordination, difficult
breathing, frequent urination and defecation, front legs stiffen and animal
becomes prostrate, violent struggling and bellowing. Bloating is common
and death occurs within half to two and a half hours after initial clinical
signs are noticed. Simple but effective treatment for ammonia poisoning, if
applied before titanic spasms occur, is immediately to administer 20-40
918
litres of cold water orally. The cold water will lower the temperature of
rumen fluid and thereby reduce urea fermentation. It will dilute the con-
centration of ammonia and reduce its rate of absorption from the rumen. Of
course, if available, four litres either of diluted acetic acid or vinegar given
with cold water is more effective than cold water alone. Acetic acid will neu-
tralize the toxic effect of free ammonia (NH3) by transforming it to ammoni-
um cation (NH4).
LACTACIDOSIS AND LIVER ABSCESSES. Subacute, latent lactacidosis devel-
op in fattening beef cattle, fed on high-grain ration. The status is similar to

the ailment of fresh cows and sheep. The increase of lactic acid concentra-
tion in rumen results in a slight, but persistent decrease in pH. This situa-
tion is characterized by poor performance and inconsistent feed ingestion.
In severe cases laminitis (“ski shoe” animals) is created and rumen epithe-
lium is altered, too. This allows Fusobacterium necrophorum penetrates the
covering epithelium through points of injury, discontinuity and necrosis,
causing liver abscesses. The abscesses consist of a central necrotic centre
surrounded by pus of yellowish colour and a wall of connective tissue. In
living animals, presence of abscesses worsens feed utilisation and at
slaughter, livers affected with abscesses are condemned for human food.
UROLITHIASIS. On high-grain ration, intensively fattened cattles mag-
nesium phosphate and magnesium ammonium phosphate (struvite) urolits

develop. The sigmoid flexure of urethra is the most common site in bulls
and steers for calculi to lodge. About the pathomechanism and symptoms
see the similar ailment of lambs. Besides the continuous calcium and
ammonium chloride (or simply: calcium chloride) preventive supplementa-
tion of the daily ration, the 40-80 grams ammonium chloride/animal/day
is the appropriate dosage for the acidification of urine after a possible sur-
gery.
The formation of siliceous calculi is associated with the intake of for-
age high in silica. Range grasses commonly contain 2% silica in dry matter
in spring month, level reaching 7% by dormancy or even more after weath-
ering are common. High levels of silica are found in grasses, in some straws
and especially in reed, sedge and (bul)rush. Low water intake enhances the
formation of siliceous calculi. Prevention includes encouraging high water
intake (even warming water on cold days) and force-feeding high levels of
common salt. Acidification of the urine can also help in reducing the for-
mation of silica stones. Feeding good quality alfalfa hay or other leguminous
919
forages up to as much as one-half of the ration greatly reduces incidence of

the problem.
NITRATE-NITRITE TOXICITY. The problem of nitrate poisoning has been
accentuated in the last decades, with increased nitrogen fertilization of field

crops, irrigation with liquid manure of pasture facilitate the nitrate intake
of plants. When plants accumulate high levels of nitrates (0.5 to 2.0%), poi-
soning may occur. Forage plant species especially adept at accumulating
nitrates include sorghums, brome grass, orchardgrass, oats, wheat, barley
and weeds. Stress in plants from drought, hail or frost increase accumula-
tion of nitrates. Cattles may be poisoned after eating either harvested or
grazed forage. Ensiling forages reduces nitrate concentration, but hay mak-
ing not. Ruminants are more apt to suffer from nitrate toxicity than mono-
gastrics because of the nitrate to nitrite conversion in the rumen; the lethal
dose is approximately 9,000 ppm nitrate in dry matter basis. The nitrite
causes the formation of methaemoglobine in the blood, a form of haemoglo-
bin that cannot carry oxygen. The resulting oxygen starvation causes suffo-
cation and death in advanced stages. The sublethal symptoms of nitrate
poisoning may be dark-coloured urine, a discolouration of the mucous lin-
ings of the mouth, the muzzle and the skin of the ears. Prevention includes
diluting nitrates in the ration by mixing low nitrate forage and avoiding the
addition of NPN sources to silage from sensitive plants.
HYDROCYANIC ACID TOXICITY. Both cultivated and native forages may
contain cyanogenetic glycosides. Prussic acid or hydrocyanic acid (HCN) is

released either by plant enzymes liberated when plant tissue is injured as a
result of cutting, mastication, wilting or freezing, or by the action of rumen
flora. Cassava (in root), corn, flax and sorghum may develop toxic levels of
prussic acid in the new growth that follows either a frost, a period of
drought or a period of heavy trampling or physical damage. Neither green
chopping nor ensiling decrease the prussic acid levels, but field curing or
drying in hay making releases 50 to 70% of the toxin. Poisoned animals
show signs of nervousness, abnormal breathing, trembling or jerking mus-
cles, blue coloration of mucous membranes, spasms or convulsions and
respiratory failure, followed by death. Some animals can be saved by intra-
venous injection of a mixture of sodium nitrate and sodium thiosulphate.
Concerning the other plant toxic effects, like photosensitizing and estro-
genic, see Chapter XXVI.
920
25.7. Veterinary technical advice in cattle herds
The exclusive place of cattle in the veterinary consultation is justified

because the possibility of a direct intervention (change of feedstuff or feed-
ing technology etc.) is greater than in case of monogastric, concentrate feed-
ing pig and poultry. First of all, the actual nutrient requirement of the dif-
ferent groups of animals should be determined. Data of recommendations
should be adjusted to the local circumstances. After having qualified feed-
stuffs at the premises, a balanced ration is formulated. This will be followed
by the clinical observation of the herd. Mineral status should also be eval-
uated. Final technical advice takes always into consideration of the econo-
my.
Most of the diseases and reproductive failures of the transient (peri-
parturient) dairy cow are related to the nutrition. The most frequent trou-
bles are the metabolic troubles, like ketosis, milk fever, partly infectious,
like mastitis, physical (lameness, retention of placenta) and reproduction
disturbances (cystic ovaries, anoestrus). In case of metabolic troubles the
causative factor is nutritional (lack of fibre - obesity, fat liver etc.), but also
in case of the other diseases as predisposing factor in decreasing general
resistance, nutrient supply has an important role: mastitis-vitamin E
and/or selenium deficiency, as well as zinc deficiency, retention of foetal
membranes, overcondition, feet lesions, reproduction troubles: lack of zinc,
manganese and beta carotene. The above described gives the key, why in a
modern structure “veterinarians must integrate consideration of nutrition,
housing and whole farm management systems into recommendations of best
practices. Veterinarians are therefore evolving from task oriented providers of
therapy to advice oriented consultants” (LEBLANC et al. 2006).
25.7.1. Determination of the locally valid requirement data

Requirement data of the available recommendation are valid only on aver-
age animal in a thermoneutral zone. If the actual local circumstances are
not of common, values should be corrected. Two interesting examples will
be mentioned.
In heat stress the dry matter intake of high-producing dairy cows
drastically decreases, which can be compensated by increasing of nutrient
density, RUP or UDP portion and by adding protected lipids to the ration,
too (Table XXV-14). The technical tools (fan, shower) can be also very use-
921
CHAPTER XXV: VET TECHNICAL ADVICE - CATTLE
ful. In North America the needs of animals, treated with GH or beta-2-ago-

nists, are also higher than normally and the expected effect can be realised
only having an improved (high quality) feeding.
Table XXV: Recommended nutrient concentration in dry matter during long-lasting heat
stress or in case of using bGH
Legend: bGH=bovine growth hormone; RDP=rumen degradable protein; RUP=rumen

undegradable protein or UDP; NDF =neutral detergent fibre; NSC =non-structural carbo-
hydrate
25.7.2. Veterinary feed evaluation in spot

Cases of veterinary feed evaluation:
-Disease (diarrhoea, bloat etc.) broke out, potentially related to feeding,
-Drop in production or reproduction, possibly caused by inappropriate feed-
ing;
-Appearance of feed differs from the accustomed one in colour, smell, con-
sistency (precaution or possible warranty request)
-Paid, scheduled visit in the absence of emergency or clinically ill animal, to
prevent diseases and decrease in productivity and to assure product quali-
ty, food safety.
Goals of veterinary feed evaluation at the premises
-Determination of the real feeding value (preference, rough energy, protein,
mineral and vitamin content).
-To take up a position concerning harmlessness, included causative
aspects.
-To select suspicious feeds for sending laboratory analysis and to give “first
aid” advice (withdrawal of a feedstuff etc.)
Important remark: this evaluation is fundamental and determinant,
but does not replace the exact chemical analyses.
922
Human prerequisites of feed evaluation

a) Knowledge of feedstuffs characteristics.
b) Complex understanding of the ruminant digestive physiology and
the impact of feeding on metabolism, reproduction and immune response.
c) Practical experiences (e.g. what is an “average” etc.)
To give technical advice in cattle herds is much more difficult than in
case of monogastrics, because of the variety of feeding technologies, applied
feedstuffs and their quality. Abrupt changes in feeding are also frequent.
Feed qualification for nutritive value (metabolic troubles, like ketosis,
milk fever appeared in herd). The two main sources of qualification are the
data of anamnesis and those of organoleptic examination. Advantage of this
examination, compared that in the lab is that sample is seen together with
its surroundings and antecedents. On the contrary, numerical data of lab-
oratory analysis are much more accurate.
Practical execution of veterinary feed evaluation
Collection of anamnesis data
A) Anamnesis relating to the feed.
-Since when, in which quantity and by which group of animals has been fed
on the feed(s) in question. Searching for cause and effect relationship.
-Circumstances of origin, con- and preservation: for example after-fermen-
tation in the opened silo, silage or wet sugar beet pulp in the bottom of the
stall, waiting to be fed etc.
-Available label, accompanying sheet/leaf, data of previous laboratory
analyses or consulting report.
B) Anamnesis related to the animals.
Time of appearance of failure, number of animals involved, clinical, behav-
ioural and other signs. Did feed intake or/and faeces consistency change?
Possible infectious diseases should be excluded (e.g. fever drops appetite,
too).
C) Others. Did some “unexpected or extraordinary” event(s) happen,
for example buying of new animals, change of animal caretaker, change in
the organisation of labour (e.g. holidays, feast etc.). Did they experience pre-
viously such an event?
ORGANOLEPTIC INVESTIGATION OF FEED
a) Parameters to check: appearance, colour, structure, touch, grab,
smell, taste, pH, dustiness.
_
b) Appliances to use: test strips (pH, NO3, Cl ) nitrate, chloride mag-
923
nifier, series of sieves to fractionate feed particles, cow-side kits (e.g. ketone
bodies, mycotoxins etc.).
c) Characteristics of organoleptic examination using at the different
feedstuffs. The main goal is the approximate determination of the feeding
value, which can be further corrected by feed composition tables. For exam-
ples it can be stated that a given alfalfa hay is of good or medium quality;
for the numerical values, data of tables can be used.
The following parameters could be characterised.
PREFERENCE. Circumstances of fermentation, composition and the
resulted preference of grass silages can be concluded from dry matter con-
centration. Vinegary/acetous smell will be reflected in a drop of feed intake.
The smell may also show the presence of some foreign grains or seeds and
also the signs of beginning deterioration.
ENERGY, PROTEIN AND FIBRE CONTENT. Botanical composition
(leguminous, grass or sedge, reed or rush [bent-grass], phenological phase,
proportion of leaves and stem reflect protein and/or fibre level. Brown
colour of hay or silage may be the sign of previous auto-heat production
(getting stuffy) and a heat damage of proteins by the Maillard reaction.
Based on grabbing and smelling of corn silages the dry matter intake and
the related energy value can be concluded, in case of the hays the fibre level
can be evaluated in this way. Even the fibre fractions can be predicted. Grab
may also show the crude ash and especially the soil, sand and muddy pol-
lution (e.g. beet, hays from river flats. In case of soil contamination, the dan-
ger of listeriosis is high (cheese production, young lamb).
APPROXIMATE MINERAL COMPOSITION. Even the exterior gives
some information, because leguminous plants have much higher calcium
content than the Gramineacea grasses; manganese concentration is the
lower in them. Whether does hay contain some medicinal plants, herbs or
weeds, which may considerably modify mineral composition. Grasses from
alkali soil (lick), or from the environment of salina may contain a lot of sodi-
um.
VITAMIN CONTENT. The most important are the appearance and the
colour. Mostly the carotene and vitamin D content can be estimated in this
way. A washed out, pale hay will contain less carotene; vitamin E and
linolenic acid content of a broken corn grain must be damaged.
d) Feed evaluation from the point of view of harmlessness. (feed
refusal, diarrhoea or other clinical symptoms occurred in the herd). The
924
question is that health troubles are related to the using of the feedstuff in
question, or not.
-Smell gives good information about the circumstances of the storage
before distribution (e. g. stuffed green fodder or grass), silages with butyric
acid smell possibly undergone undesirable fermentation, musty or stale hay
must be mouldy and containing mycotoxins, acetonic; sugary/sickly-sweet
smell (aldehyde!) squeals on the rancidity of concentrate. Peculiar smell may
indicate some special feed component, like citrus pulp, tomato or apple
pomace.
-Appearance may draw the attention of the presence of many harm-
ful substances. Namely, dark, fierce green colour of green forages gives an
inkling of a higher nitrate concentration. Greyish, “dusty” forages generally
are infected with moulds. Degree of “dustiness” almost quantitatively indi-
cates the mould number. Dark brown or black sugar beet pulp may contain
a lot of residual sugar, which can lead to lactacidaemia if fed in large quan-
tity. Inhomogeneity (improper mixing) of concentrate, presence of different
particle size (by using magnifier, too) refer to the presence of a “foreign” feed
mixture (e.g. poultry feed for cattle). Clotting of concentrate shows higher
moisture content possibly originates to an inappropriate or too long storage.
Storage insects (pest) can be found by ocular inspection and fractionating
sieves.
-Grab and touch. Hotness of silage is a sign of unnecessary after-fer-
mentation. Coarse, sandy or muddy touch indicate soil pollution If silages
are of greasy touch, it may be spoiled and having no structural fibre (milk
fat’). Clotting of overstored concentrate can be felt by touching, too.
-Scratchy, bite taste of concentrates is a sign of rancidity. Special
tastes enable the identification of some peculiar component, for example
rapeseed. By tasting, the oversalted milk replacer can be picked/screened
out.
Based on the above described and the level of harmful feedstuff in the
daily ration, decision can be made above the future application.
25.7.3. Evaluation of the real supply

Knowing the herd, the feedstuffs and the composition of daily ration, a com-
parison can be made between the nutrient requirements of animals and the
real supply. Attention let us calculate with the real feed ingestion and not
with the theoretical value.
925
25.7.4. Profile investigations

The available data basis can be enlarged by using analytical results of milk,
blood, urine, rumen fluid, saliva, faeces and coloured hair samples from
cows, representing the whole herd. Rumen energetic status is reflected by
the milk urea level (for details see below).
Ketonuric index (KEGL and GAAL, 1992) is a sensitive indicator of
energetic status of the whole organism. Namely, it takes into account not
only the severity of ketonuria, but also the time, related to the calving. Test
stripes give 1 to 4 “positive” crosses; that very quickly developed four
became the cross 5. Number of crosses should be multiplied by ten.
Correction should be made: if cow is before calving, the double of days till
parturition should be added. If the cow is after calving, the double of
elapsed days should be subtracted. For example, a two-cross result 5 days
before calving gives an index of 35 (10×2+2×5) and 5 days after parturition
a ketonuric index of 10 (10×2-2×5). Above 20-25 the cow should be treated,
for example by eating propylene glycol.
Conclusions can be drawn from saliva to sodium state, from the urine
to the acid-base balance, from the faeces and hair to some mineral status
of the cow. Blood analysis gives useful information concerning the general
metabolism, kidney and the liver functions. Investigation of the rumen fluid
shows both the ingested feed and the rumen fermentation
Blood profile monitoring provides a couple of important data, but it
has also limitations. Selected individual animals cannot really represent the
whole herd, because there is a great deviation of the physiological values in
the blood (FIGURE XXV-22). The different production and reproduction
groups have considerable differences, too. Among the measured blood
parameters and the potential metabolic troubles and reproductive failures
there is a multiple interrelationship). The real evaluation of data requires
sophisticated biomathematical procedures, for example multiple regression
analysis, main component and path analysis (CURTIS et al. 1985; CORREA et
al. 1993).
Summarizing the above elucidated, conclusion, based on blood profile
monitoring, and should be drawn only with precaution.
926
FIGURE XXV-13: Distribution of

physiological values in herds
25.7.5. Special field of the veterinary nutritional consultation
MINERAL SUPPLY (TOLGYESI, 1965-1990).

Opposite to the energy and protein supply, in case of minerals the use of so-
called safety margin is generally not used. The replacement value of the
individual minerals is minimal. Concerning the mineral composition of
plant, the type of soil is determinant. If the soil has an acidic pH, there is
good manganese and bad molybdenum absorption, which will reflect also in
the produced feedstuff plant. Neutral or slightly alkaline soils are favourable
to the alfalfa production, but the manganese to molybdenum ratio will be
low. Calcareous, sandy soils generally are deficient in zinc and the cupper
to molybdenum ratio often cannot reach the appropriate (at least 10 to 1)
value. Thus, during consulting in the field of mineral nutrition, the knowl-
edge of local soil types is indispensable.
A frequent problem is the calcium excess in dry cow ration. Combined
production of different forage plants, for example alfalfa (high in calcium,
low in manganese) with grasses (less calcium, high in manganese) may
mean an economic solution. Unfavourable low copper to molybdenum ratio
can be compensated by feeding of sunflower. On salty soil boron tends to
accumulate, causing diarrhoea. Since only the dicotyledonous plants accu-
mulate boron, the production of monocotyledon plants (grasses, corn)
should prefer in similar conditions.
927
Although corn protein has a high methionine+cystine concentration,

the absolute amount of sulphur in corn (both plant and grain) is small and
may limit bacterial protein synthesis in the rumen. Optimum sulphur to
nitrogen ratio in the feed for lactating dairy cow would be 1 to 10 and to
achieve that corn silage should be combined with plants, rich in sulphur
(rape, oil radish, mustard and perko), or use Na2SO3 supplementation.
Fertilizer should be selected not only according to the main component, but
also its pH-influencing effect. For example (NH4)2SO4 not only supplies
nitrogen, but also acidify soil, enhancing in this way manganese uptake by
plants and decreases undesirable molybdenum absorption. The significance
of farmyard manure and urban sewage-sludge, as micro element source,
should be re-evaluated.
MILK FEVER
From the first description of the milk fever (1819), it continues to appear in
many dairy farms. Its incidence is especially high, if during the dry period
high amount of calcium-rich leguminous forages (alfalfa) or milking
concentrate is fed. The actually absorbed amount of calcium depends upon
the supply and if overfed, the utilisation decreases. There is a complex
regulatory mechanism in the background, for details see unit „Digestive
physiology above). Cow requires some days for the adaptation of a new
calcium concentration in her daily ration.
During the consolidated lactation, if absolute calcium offer is
assured, the described regulatory mechanisms assure the covering of the
requirements. After calving, the abrupt calcium excretion by milk decreas-
es blood calcium level and milk fever develops. This tendency is true also
without pathoclinical consequences, among physiological ranges. Calcium
requirement of pregnant cow is already increasing 1 to 2 days before calv-
ing. After UK recommendations, during the dry period low calcium-low
phosphorus ration should be fed, but 7 days before the expected calving it
is advisable to increase daily calcium supply from 50 to 100, and that of
phosphorus from 30 to 80 grams, by distributing 4 kg milking concentrate.
Limitation of calcium ingestion during the dry period in practical cir-
cumstances sometimes difficult, for example the available forage is only
alfalfa silage, haylage or hay. As it was discussed above, the excess phos-
phorus supplementation is not the measure to choice. According to the
accepted scientific opinions, while given the actual requirement, the opti-
928
mum ratio is given automatically. On the contrary, optimal, close calcium

to phosphorus ratio does not mean obligatory sufficient amounts. Increased
calcium and phosphorus offer before calving get possible of more absorption
of these minerals, just then, when blood calcium begins to drop. After calv-
ing, according to the needs of milk production, cows receive ration of high-
er calcium and phosphorus level. In such a herd, where the limitation of cal-
cium intake in the dry period is not possible, the acid-base balance of cows
should be shifted towards acidic by feeding cationic diet (in the majority of
cases by supplementing daily ration with ammonium chloride). This will
result in such a CAB, which facilitate calcium mobilization from the bones.
As a new tool, giving Ca-containing gel for fresh cow, gave good experiences.
MAGNESIUM DEFICIENCY
Some of the milk fever cows have also hypomagnesaemia, but this is not
typical. There is another link between these elements: too much magnesium
in the feed mixture decreases calcium absorption. The magnesium require-
ment of calves is 0.07% dry matter, for dairy cows 0.2 to 0.25% of dry mat-
ter (NRC, 2001). Feed mixtures of average composition generally cover cow’s
requirement. The recommended magnesium level in milk replacers is 0.7%
of dry matter. Critical serum magnesium concentration for dairy cow is 0.41
mmol/l, the normal value being 0.7 to 1.3 mmol/l. (For comparison, the
normal serum magnesium in human is 0.7 to 1.0 mmol/l.) Calves show
signs of clinical deficiency, when having less than 0.25 mmol/l plasma mag-
nesium level.
In Northern parts of the European continent there are some soils with
low magnesium content. If these territories are intensively fertilized by
potassium and nitrogen, the magnesium concentration of the produced fod-
der green decreases. In the blood of high-producing dairy cows, fed on these
feedstuffs, the magnesium level get lower in blood. Clinical signs (“grass
tetany”) develop, if only the blood magnesium level drops below 0.4 mmol/l.
This a pathognostic sign. Frequently, there is also a slight decrease in
serum calcium concentration. Normal values for inorganic calcium, phos-
phorus and magnesium in healthy cows is 2.1 to 3.0, 1.6 to 2.3 and 0.7 to
1.3 mmol/l, respectively. In fresh cows all data is a little bit lower (in the
same order): 1.75 to 2.1, 1.0 to 1.25 and 0.6 to 1.25 mmol/l, respectively.
Since the magnesium content of milk is considerable (0.012 to 0.015%),
parallel to the increasing milk production the Mg requirement also grows.
929
The availability of magnesium in forages anyway is low, 7 to 33% net

absorption. This may explain that in some herd, as a safety margin, con-
tinuous magnesium supplementation is given.
KETOSIS OF DAIRY COWS

Forced fat mobilization, small insulin to glucagon ratio and low malonyl-
coenzyme A activity are the preconditions of the appearance of ketosis.
Ketosis has two basal types, one of them can be characterised by low blood
sugar and insulin concentrations, and another is accompanied by high glu-
cose and insulin level in blood. The first is called also as “fasting” or “pri-
mary” ketosis (synonyms: Type I ketosis, acetonaemia, ketonaemia) and it
is a metabolic disease of dairy cows in the first six weeks of the lactation,
characterised by anorexia, loss of live weight and milk production and nerv-
ous troubles. Incidence of this type of ketosis may be high among high-
yielding cows, when owing to the inappropriate feeding they are undercon-
ditioned. If cow was overfed during the dry period, the occurring ketosis is
frequently combined with fatty liver. The most frequent it can be diagnos-
tized in weeks 3-4 of lactation.
For an occurrence of high incidence, besides failures of nutrition, the

presence of genetic predisposition, high milk production, calving complica-
tions and unfavourable environmental conditions are necessary. Ketosis
basically is the consequence of a negative energy balance: to compensate
energy losses by the excreted milk, first the blood sugar drops, later a lim-
ited protein and an important fat mobilization follows. As a result, an accu-
mulation of amino acids, free fatty acids and keton bodies (aceto-acetic acid,
aceton, beta-hydroxy-butyric acid) is shown in the blood. Most of the keton
bodies are produced by the liver, but mammary gland (udder) and rumen
wall also contribute. Relative lack of insulin and excess of glucagon stimu-
late fat mobilization in the whole body and keton body production in the
liver.
Thus, insulin has basically an anabolic and glucagon a katabolic
effect. Production of keton bodies from palmitic and butyric acid is the most
intensive between days 30-60 of the lactation. Considerable glyconeogene-
sis (production of glucose from propionate, lactate and amino acids) can be
observed only in case of appropriate carnitine-palmitoiltransferase enzyme
activity in the liver. Rate of keton body production is determined by the
transport of long-chain fatty acids into the liver mitochondria. Ketone utili-
930
sation of peripheral tissues spares glucose. In turn, ketone bodies them-

selves take part in the regulation of metabolism.
Type II ketosis may already emerge some days after calving and often
is connected with other health troubles (metritis, mastitis, laminitis and
other hoof diseases. In the blood of affected cows not only the ketone body
concentration, but also the glucose and insulin level is high. Under stress
influences a fatty liver also easily develops. Outbreak of the ailment is grad-
ual and in the beginning only a decrease in the voluntary feed intake and
milk production, lethargy and defecation of dry, covered with mucous coat
faeces can be observed.
Thus, energy deficit in the postpartum period may lead to clinical

ketosis. The subclinical ketosis (increased ketone bodies concentration in
blood) can be considered as a preliminary phase of the clinically detectable
diseases. To detect energy deficit, the determination of blood sugar, non-
esterified fatty acids (NEFA) level, as well as the ketone bodies (aceton,
aceto-acid acid and beta-hydroxy-butyrate) level in blood, urine and milk. If
avoiding of enhanced production and accumulation of ketone bodies, the
milk production will grow, too. Total energy balance shows a close correla-
tion with the blood sugar and ketone bodies concentration, as well as the
milk production of the following week. Increase in blood sugar and ketone
body concentration can decrease milk production of the clinically healthy
cows, too.
Clinically sick cows may lose even 100 kg of live weight within some
days, which is aggravated by the perverse appetite (pica) of refusing to eat
concentrate. Exhaled air, excreted urea and milk have an acetone smell.
Gait is staggering. Recovery can be expected, if only at the same time, the
predisposing and companion diseases are treated. Dead animals are thin
(even emaciated), they had very small fat depots and the liver, kidneys and
heart are infiltrated by fat. Liver is pale, soft and crumbly, in dragging, long-
lasting cases necrotic. Histological damages can be found in the hypoph-
ysis, pancreas and thyroid gland. Onset of the sexual cyclic activity delays
and sick cows cannot be inseminated.
GRAVERT (1986) in his study measured feed consumption, milk pro-

duction and calculated cows` energy balance. Weekly energy deficit proved
to be 515 MJ NEl in the 4th and 164 MJ NEl in the 16th week of lactation.
For 100 MJ energy deficit has fallen 5.01 kg loss in live weight and 2.64 kg
931
body fat mobilization. The best indicator was the acetone concentration of
milk. Heritability of acetone level cannot be proven in first lactation cows; in
older animals the h2 decreased with elapsed time: 0.17 in the first, 0.09 in
the second and zero in the third month of lactation.
Diagnosis of primary ketosis, based on the clinical observation and

necropsy, should be approved by the Rothera test. The problem is that this
test use urine samples, where the physiological ranges and individual devi-
ations are great, therefore the diagnostic value is small. The calculation of
ketonuric index, introduced by KEGL and GAAL (1992) improve the accuracy
of diagnosis. However, sensitivity of the Rothera probe, made on milk sam-
ple is higher. Diagnosis is confirmed by a low (2.5 mmol/l) blood sugar con-
centration. During the differential diagnosis, milk fever, displaced aboma-
sum, reticulo-peritonitis, some poisoning, kidney damages, listeriosis and
rabies should be excluded.
Ketone bodies components in the milk are the acetone, aceto-acetic
acid and beta-hydroxy-butyrate, which are all transformed into acetone for
determination. For the semiquantitative detection from blood and urine
samples several reagent mixture (e.g. Ross reagent, Legal probe, Lange
probe) and test stripes (e.g. Ketostix) are used. The active ingredient of them
generally is the nitroprussidic sodium. Under the influence of aceton and
aceto-acetic acid, nitroprussidic sodium becomes deep purple; b-hydroxy-
butyric acid does not cause change of colour, therefore special kits are
available for its detection. Quantitative determination can be carried out by
using photometer, gas chromatography, enzymatic breakdown and
diffusion procedure, using automated FIA (flow injection analysis) system.
TREATMENT consists of intravenous infusion of glucose, intramuscu-
lar injection of glucocorticoids and peroral pouring of glucose precursors,
like propylene glycol. The PREVENTION is much more important. The essence
of that is to achieve ideal condition, namely 3.5 at drying off and also 3.5
BCS at calving. Two-three weeks before calving the rumen microflora and
–fauna should be adapted to the concentrate, by giving, with gradual
increase, up to 1 kg/150 kg live weight. After parturition the dry matter
intake should be stimulated by chopping of the forage and giving roughage
of small fulfilment unit and in case of necessity, by application of rumen
buffers (NaHCO3, MgO and sodium sesquicarbonate). Propylene glycol can
also be fed.
Incidence of ketosis is lower, if distribution of the concentrate is more
932
frequent. Therefore frequent (practically ad libitum) feeding is of crucial

importance in prevention of ketosis. Benevolent effect can be explained by
the stabilisation of rumen fermentation and better energy supply. Change
in distribution order (first the forage, after the concentrate) has also good
influence. The best solution, of course, the use of a good total mixed ration.
In herds, seriously affected by ketosis, the use of propylene glycol,
sodium propionate, special yeast, protected methionine (2×10 grams
SMARTAMINE M or MEPRON 80 daily), niacin and cholin is recommended dur-
ing the last two weeks of dry period and the first weeks of lactation. In the
knowledge of the role of L-carnitine in the transport of long-chain fatty
acids, it can be supposed that peroral application of protected carnitine or
in form of injection may be helpful. Since in the Type II ketosis there is an
insulin resistance, the use of trivalent, organic chromium (as part of the
glucose tolerance factor) seems to be useful. Recent findings suggest benev-
olent effect of the application of continuous, long-release monensin capsule
in the last three weeks of pregnancy.
If the ketone level in milk is higher than 0.4 mmol/l, the shape of
lactation curve gets deformed and in the early lactation shows a nadir.
Higher than 100 µmol/l beta-hydroxy butyrate and 0.4 mmol/l aceto-acetic
acid concentrations in milk are grounded suspicion for ketosis. The daily
milk production drops above 0.7 mmol/l milk acetone concentration. In an
epidemiological study, cows of 1.4 mmol/l blood acetone level produced less
FCM milk by 190 kg during the first 100 days of lactation, compared to cows
of blood acetone level below 0.7 mmol/l. Under the influence of subclinical
ketosis, the calving to reconception interval lengthened by five days and the
incidence of ovarian cysts increased (Table XXV-15).
HERD HEALT DIAGNOSTIC VALUE OF MILK ACETON AND UREA CONCENTRATION

Mixed herd milk samples are excellent diagnostic tools for a consulting vet-
erinarian. It is homogenous; sampling does not mean a stress for the ani-
mals, measurement of some of the following constituent is regularly com-
pulsory. The milk is appropriate medium for the measurement of the fol-
lowing parameters: somatic cell count (←subclinical mastitis), fat concen-
tration (←fibre supply), protein concentration (←available energy in rumen),
lactose (←endocrine disorders), progesterone (←discrimination of oestrus
and luteal phase of ovaries), keton bodies, urea, allantoin (←microbial pro-
tein production) and some minerals and vitamins. The ratio of fat to protein
933
is a good indicator of dietary energy balance, because value, higher than 1.4
shows an energy deficit.
Table XXV-15: Possible consequences of milk acetone concentration*
*Critical beta-hydroxy-butyrate concentration: above 0.1 mmol/l

Simultaneous measurement of milk acetone and urea offers a new
opportunity for evaluation of nutritional and reproductive status of a group
of dairy cows. Milk ketone level indicates which group of cow is in danger
of ketosis and by appropriate measures cases of clinical ketosis can be pre-
vented, drops in production and reproduction, caused by subclinical keto-
sis, can be palliated in time. It is advisable to take milk samples parallel to
the monthly obligatory performance tests for measuring milk composition
and somatic cells. Milk urea reflects the blood urea level of 1 to 2 hours ear-
lier, which in turn, shows the rumen ammonia concentration of 1.5 to 2
hours earlier. Consequently, these time shifts to feed ingestion should be
taken into account in timing milk samples. Besides the rumen ammonia
concentration, blood and milk urea is influenced by the gluconeogenetic free
amino acids, but the most decisive factor is the energy to protein ratio of the
feed mixture and the degradability of intake protein. In other words, it is the
UFP value of the ration.
Relation between the length of service period (calving to reconception)
and herd milk urea concentration can be described by a hyperbolic curve:
both low and high urea value increase the incidence of reproductive failures.
The best reproduction state could be experienced at a milk urea concentra-
tion between 4.5 and 5.0 mmol/l. Low milk urea levels means a decrease in
the activity of rumen fermentation. In lack of ammonia nitrogen, bacteria
produce less energy for host cow, too. Too high urea in blood may alter the
composition of uterinal fluid, worsening implantation chances.
For simultaneous evaluation of acetone and urea data, three periods
should be considered. Table XXV-16 shows significance of the different
934
values in the first 50 days of lactation (period of negative energy balance).
Table XXV-16: Milk acetone and urea concentrations and corresponding nutrient supply of
cows in the phase of negative energy balance (first 50 days of lactation).
*according some authors only above 2.0 mmol/l

Legend: RDP= rumen degradable protein; RFC= rumen fermentable carbohydrates
RUP or UDP= rumen undegradable protein
Table XXV-17 gives help for the evaluation between days 51 and 110
and for the last two thirds of the lactation, after day 110. Practically no
hyperacetonemia occurs in these periods. Basically, the highest standard
urea values has been adapted to the given production-reproduction phase.
Table XXV-17: Interpretation of milk urea concentration between day 51 and 110 of lacta-
tion and after day 110 till drying off.
Legend: see Table XXVI
Using key values of the above tables, the nutrient supply of the dif-
ferent groups of herd can be monitored and if necessary, corrected, both at
the level of composition of daily ration and the nutritional technology.
935
FOR FURTHER READING
Aiello, R.J., Kenna, T.M. and Herbein, J.H. (1984): Hepatic gluconeogenetic and ketogenic
interrelationships in the lactating cow. J. Dairy Sci. 67, 1707-1715.
Bauman, D.E. and Currie, W.B. (1980): Partitioning of nutrients during pregnancy and lac-
tation: A review of mechanisms involving homeostasis and homeorhesis. J. Dairy
Sci. 63, 1514-1529.
Correa, M.T., Erb, H. and Scarlett, J. (1993): Path analysis for seven postpartum disorders
of Holstein cows. J. Dairy Sci, 76, 1305-1312.
Daccarett, M.G., Bortone, E.J., Isbell, D.E. and Morrill, J.L. (1993): Performance of Holstein
Heifers fed 100% or more of National Research Council requirements. J. Dairy Sci.
76, 606-614.
Drackley, J.K. (1999): Biology of dairy cows during the transition period: the final frontier?
J. Dairy Sci. 82, 2259-2273.
Duffield, T.F., Sandals, D., Leslie, K.E., Lissemore, K., Mcbride, B.W., Lumsden, J.H., Dick,
P. and Bagg, R. (1998): Efficacy of monensin for the prevention of subclinical keto-
sis in lactating dairy cows. J. Dairy Sci. 81, 2866-2873.
Fekete, S, Huszenicza, G, Kellems, R.O., Szakall, I., Febel, H., Husveth, F., Nagy, P,
Kulcsar, M., Kosa, E., Gaal, T., Rudas, P. and Oppel, K. (1996): Influence of defi-
cient intake of high and low degradable protein on body composition, metabolic
adaptation, production and reproductive performance in early lactation dairy cows.
Acta Vet. Hung. 44, 309-333.
Geishauer, T., Leslie, K., Kelton, D. and Duffield, T. (1997): Evaluation of five cowside tests
for use with milk to detect subclinical ketosis in dairy cows. J. Dairy Sci. 81, 483-
443.
Goff, J.P. (2006): Major advances in our understanding of nutritional influences on bovine
health. J. Dairy Sci. 89, 1292-1301.
Gustafsson, A.H., Anderson, L. and Emanuelson, U.: Effect of hyperketonaemia, feeding
frequency and intake of concentrate and energy on milk yield in dairy cows. Anim.
Prod., 1993. 56. 51-60.
Holtenius, P. and Holtenius, K. (1996): New aspects of ketone bodies in energy metabolism
of dairy cows: a review. J. Vet. Med. A. 43. 579-587.
Huszenicza, Gy.-Fekete, S., Haraszti, J., Molnar, L. and Solti, L. (1988): The ovarian func-
tion during the first postpartum period in dairy cows reared with different intensi-
ty. Acta Vet. Hung.. 36, 153-164
LeBlanc, S.J., Lissemore, K.D., Kelton, D.F., Duffield, T.F. and Leslie, K.E. (2006): Major
advances in disease prevention in dairy cattle. J. Dairy Sci, 89, 1267-1279.
Macri, A., Schollenberger, M., Drochner, W., Tafaj, M. and Morar, M.V. (2005):
Investigation on the “in vitro” degradation of zearalenone in rumen fluid. Mycotoxin
Res. 21, 65-67.
Magdus, M., Fekete, S., Frenyo, V.L., Miskucza, O. and Kotz, L. (1988): Milk production
and certain parameters of energy metabolism in dairy cows fed rations of varying
energy and crude protein contents and fat. Acta Vet. Hung. 36, 43-59.
NRC: Nutrient requirements of beef cattle. National Academy Press. Washington, D.C.
Update 2000
NRC: Nutrient requirements of dairy cattle.7th rev. ed. National Academy Press.
Washington, D.C. 2001.
Paterson, A.B. (1945): The diagnostic value of Rothera’s test on milk. Vet. J. 101. 199-203.
Pethes, G., Bokori, J., Rudas, P., Frenyo, V.L. and Fekete, S. (1985): Thyroxine, tri-
iodothyronine, reverse-triiodothyronine and other physiological characteristics of
periparturient cows fed restricted energy. J. Dairy Sci. 68, 1148-1152.
Schmidt J., Varhegyi J.-ne, Varhegyi J. and Turine Cenkvari E. (2000): Energy and pro-
tein evaluation of ruminant feedstuffs (in Hungarian). Mezogazda Kiado. Budapest
936
Schroder, U.J. and Staufenbiel, R. (2006): Invited review: Methods to determine body fat
reserves in the dairy cow with special regard to ultrasonographic measurement of
backfat thickness. J. Dairy Sci. 89, 1-14.
Tolgyesi Gy. (1969): Miroelement content of plants and its agricultural relevance (in
Hungarian) Mezogazdasagi Kiado. Budapest
Williams, A.G. and Coleman, G.S. (1992): The rumen protozoa. Springer-Verlag. New York-
Berlin-Tokyo-Budapest. pp 361-367.
937
Chapter XXVI
SHEEP FEEDING AND NUTRITION
26.1. Feeding and nutrition of the breeding sheep

26.1.1. Requirements
26.1.2. Maintenance
26.1.3. Pregnancy
26.1.3.1. Early and mid pregnancy
26.1.3.2. Late pregnancy
26.1.4. Voluntary intake and energy balance
26.1.5. Lactation
26.2. Feeding and nutrition of suckling and fattening lambs
26.2.2. Newborn and suckling lamb
26.2.2.1. Natural nursing
26.2.2.2. Artificial lamb rearing
26.2.3. Methods for fattening
26.2.3.2. Intensive express meat lamb’s production
26.2.3.3. Semi-intensive fattening of growing lambs
26.6.2.3. Improvement of old, culling sheep for slaugh-
tering
26.3. Deficiency syndromes and metabolic disorders of
sheep
Chapter XXVI: SHEEP FEEDING AND NUTRITION
26.1. Feeding and nutrition of the breeding sheep

© Jose Antonio GUADA Vallepuga
he world sheep population amount to 1,080 million (FAOSTAT data
I 2005) most of which are settled in Asia (42%) and Africa (24%), while the
rest are mainly concentrated in Australia-New Zealand (14%) and
Europe (11%). Thus, about 66% of the sheep population are settled in
underdeveloped countries where they make a significant contribution in
terms of wool, meat and milk production. In the European Union sheep and
goat meat shares a 3.5% of total meat consumption although the variation
is large and in some countries, like Greece, Ireland, UK and Spain, this con-
tribution ranges from 5 to 15%. Similar figures are recorded in Australia
and some Arab countries. Milk sheep accounts for only 1.4% of world milk
produced by large and small ruminants but 66% of the milk sheep produc-
tion is concentrated in Europe and Near East, in particular in those coun-
tries surrounding the Mediterranean basin, accounting in most of them for
up to 7-35% of the total milk production. Moreover the ability of sheep to
survive and produce in different and sometimes extreme climatic condi-
tions, makes sheep farming a useful tool in the building of sustainable pro-
duction systems focussed on soil conservation and land use, particularly in
semi-arid areas of low productivity.
There is a wide variation of production systems, ranging from contin-

uously housed flocks, mainly of dairy ewes, to extensive grazing systems of
meat breeds, although the most common practise is to combine periods of
grazing and indoors feeding throughout the same day in milking ewes or
during periods of the reproductive cycle in breeding ewes. Indoors feeding
usually corresponds with the periods of higher requirements, late pregnan-
cy and lactation, and depends on availability of pasture and the manage-
ment system. In annual laming systems pastoral resources are high at the
time of lambing in spring, but in husbandry of prolific ewes, with lambing
940
intervals of 8 months or less, housing is required in out of season lambing.

Even under these conditions is usually difficult to fulfil the daily require-
ments of nutrients in late pregnancy and lactation and intervals of under-
feeding are followed by periods of body reserves restoration. Feeding strate-
gies should attempt to avoid detrimental effects of these processes at short
and long term by appropriate management of body condition and feeding
levels compatible with acceptable performances.
26.1.1. Requirements
Energy requirements are given as metabolizable energy (ME) because it is a

convenient unit for a strait forward estimation of feed allowances while the
variation in the efficiency of utilization of ME for maintenance and lactation
is of scarce importance for practical purposes. With regards to pregnancy
although there is some evidence that utilization of ME can be as variable as
for fattening there is still insufficient information to consider this dietary
effect in practical rationing.
Protein requirements are expressed as metabolizable protein (MP) per

MJ of ME since this ratio is appropriate to calculate the microbial contri-
bution to MP due to the fact that microbial synthesis is mainly dependent
on the energy supply. At maintenance feeding level about 8.4 g of microbial
protein are synthesised per MJ of ME providing 5.4 g of MP/MJ if true
microbial protein content and intestinal digestibility are assumed to be 0.75
and 0.85 respectively (ARC, 1982). To meet the degradable N required for
microbial synthesis the diet should contain about 11 of crude protein/MJ
ME when the protein is 80% degradable in the rumen (8.4/0.8) which would
provide 1.8 g of additional MP as by pass protein, given a total supply of 7.2
g MP/MJ ME. This ratio is similar to that estimated from requirements for
all physiological stages except lactation in which an extra supply of
undegradable protein seems to be required although the deficit is lower
than expected due to the higher efficiency of microbial synthesis promoted
by the increased rumen turnover rate at the high levels of feeding in lacta-
tion. In conclusion, providing that energy and degradable N requirements
are met the rumen ecosystem ensures a supply of MP close to requirements
in most of the physiological conditions. Two important practical implication
are derived from this fact. One is the importance of maintaining an ade-
quate rumen environment in order to maximize microbial synthesis and the
second is the decline in MP supply, and the consequent increase in
941
undegradable protein requirements, as a result of energy undernutrition

promoted by food shortage or constraints on voluntary food intake.
26.1.2. Maintenance
Maintenance is of particular relevance in sheep due to the large non

productive interval between weaning and mating in annual lambing sys-
tems. Mature ewes will be in state of maintenance during large part of this
period once they have recovered the body reserves lost in lactation and preg-
nancy. Also maintenance constitutes the basal reference to estimate addi-
tional requirements for production.
Energy allowance for maintenance was estimated assuming an aver-

age efficiency of ME utilization of 0.68 and 256 kJ/kg LW0.75 of net ener-
gy requirements for a 3 years old housed sheep (SCARM, 1990) which is
similar to the fasting metabolism of 0.217 MJ/kg LW0.75 proposed by the
AFRC (1993) plus an 18% of allowance for activity. In harsh pastoral con-
ditions the activity allowance can be twice as higher increasing maintenance
requirements to 0.430 MJ/kg.LW0.75. Protein requirements account for
0.35 g N/ kg LW0.75 of endogenous losses and a variable amount of N
retained in the wool that on average will range from 0.5 to 1.3 g/d for fleece
weights of 2 and 5 kg rendering 0.6 g of clean wool fibre per g fleece. Wool
growth is closely associated to ME intake and the rate of protein deposition
in wool per MJ of ME intake has been estimated to be of 8 g/MJ per kg of
fleece as proportion of body weight (SCARM, 1990). Thus a 50 kg ewe with
2 kg fleece weight as shorn can be estimated to retain each day 0.32 g of
protein wool per MJ of ME (8×2/50). The efficiency of conversion of
absorbed protein to wool is extraordinary low (0.20-0.26) due to its high
content in sulphur amino acids which is three to four times higher than in
milk and meat protein. However endogenous losses are replaced by
absorbed amino acids with much higher efficiency (up to 1.0 according to
AFRC, 1993). Therefore it can be estimated that a supply of 7.1 g of MP/MJ
ME would be adequate to meet maintenance requirements including wool
growth (0.32/0.26 + 0.35×6.25/0.376). It is note worthy that this value is
close to the 7.2 g MP/MJ ME provided by a diet containing enough protein
to satisfy the N requirements of rumen micro organisms as shown above
(11g of 80% degradable crude protein per MJ ME).
Requirements for Ca and P are given in relation to live weight for
942
maintenance and as additional requirements per unit of production for

pregnancy and lactation. Requirements for sulphur are determined pri-
marily by its essentiality for microbial protein synthesis and consequently
are better expressed by reference to N supply; being recommended for sheep
a S:N ratio of 0.08. Sulphur is generally considered to be particularly impor-
tant for sheep, because the sulphur content of wool is about 4%, incorpo-
rated to the cystine molecules. The main sources of sulphur are the methio-
nione and cystine, although rumen microorganisms can also use inorganic
sulphur, usually provided in the form of sodium sulphate. Deficiency of sul-
phate causes depressed wool production and poor fleece characteristics,
including a lack of crimp due to delayed keratin formation. Prolonged and
severe copper deficiency has very similar signs. Additional S is required, if
diet contains urea or other forms of NPN and in such a situation is recom-
mended incorporate sodium sulphate into the diet at a ratio of 1 to 9 by
weight urea.
Signs of sodium deficiency are anorexia and consequent poor per-

formance. Some plants (e.g. alfalfa) are poor in sodium. During silage mak-
ing the sodium concentration decreases, because plant extracellular mate-
rial is lost in the effluent. Forage crops grown in readily drained and vol-
canic soils are also low in sodium. For evaluating the sodium status, the
sodium to potassium ratio in the saliva is of high diagnostic value: the nor-
mal ratio is 20, and the ratio in sodium deficient sheep drops to 0.45.
Supposing continuous water availability, the supplementation is easy and
without danger of toxicity by providing salt blocks or licking salt, because
sheep have a well-regulated appetite for salt. Deficiencies of K and Cl are
unlikely but a Cu deficiency, either in the diet (< 5 mg/kg D) or induced by
high levels of Mo and S, results in wool depigmentation and loss in crimps
and strength. Also, deficiencies of Zn (<20 mg/kg DM) and Se (< <0.05
mg/kf D) result in a marked reduction in wool growth.
Ovulation
Nutrients required for ovulation are quantitatively insignificant but the

number of ova released (ovulation rate) and the length of the annual
anoestrus period are highly dependent on long-term nutrition or body con-
dition score of the ewe at mating. Moreover, changes in the nutritional sta-
943
tus during the 2-3 weeks preceding ovulation also affects the ovulation rate.
An effect described as dynamic in contrast with the static nature of actual
body condition at mating.
The response of the ovulation rate to body condition can be increased

by overfeeding in the previous days (flushing), providing the ewe has an
intermediate body condition, or alternatively decreased by underfeeding. In
ewes with extreme body condition the response declines, indicating that
ewes well fed prior to flushing are already free of constraints to express their
genetic potential for ovulation while below a threshold of body condition the
replenishment of body reserves is a priority for survival. The response may
also differ between breeds, being lower in those of higher potential which are
more consistent lamb producers.
The mechanisms mediating this response are not fully understood

but they probably involve atresia of follicles at several stages of development
as a result of shifts in the composition of follicular fluid in response to an
integrated effect of metabolic hormones and gonadotrophins. In this respect
it is interesting to note that the follicular development from primordial to
ovulatory follicles is a 6 moths process and therefore it is exposed to events
occurring in the preceding breeding cycle, such as weight losses during lac-
tation, even in annual lambing systems, which would explain the impor-
tance of weaning weight as determinant of ovulation rate in absence of
flushing.
Recovery of body reserves to achieve the target body condition is a

slow process, since 1 score point is equivalent to about 13% of live weight
change and depletion levels of 1.5 points during the whole reproductive
cycle are not uncommon. The need of supplementary feeding to achieve this
goal will depend on the intensification of the production systems. In inten-
sive systems characterised by short breeding cycles the reduction of lacta-
tional losses by early weaning ( 4 to 6 weeks) together with the practise of
flushing by prolonging the ad libitum feeding beyond lactation until next
breeding are strategies that allow to maintain a high ovulation rate in spite
of the short recovery period available.
944
26.1.3. Pregnancy
Additional requirements for pregnancy in Table XXVI-1 are expressed per kg

litter birth weight. This is generally known for a particular breed but other-
wise it can be predicted from the maternal weight by the coefficients derived
by DONALD and RUSSEL (1970): 226, 361 and 416 g//kg LW0.75 for single,
twin and triple bearing ewes. Thus it can be estimated that a 50 kg ewe car-
rying single or twin lambs would require during the last month of pregnan-
cy an average of 10.5 and 12.7 MJ of ME/d, equivalent to 1.5 and 1.8 times
its maintenance requirements (0.376×500.75=7.1 MJ). How to provide these
amounts under the constraints imposed by the voluntary food intake will be
later discussed.
26.1.3.1. Early and mid pregnancy

Requirements are only quantitatively significant during late pregnan-
cy when the foetus growths exponentially accumulating most of his final
weight. However during early and mid pregnancy nutrition has also an
important albeit indirect effect on embryo survival and placenta develop-
ment respectively. There is evidence that both under and over feeding after
mating can reduce embryo survival particularly in multiple ovulating ewes.
Underfeeding seems to delay embryonic development causing asynchrony
945
with the hormonal state of the uterus which results detrimental to the
embryo survival. Thus, secretion of cytokines by the embryo (interferon
Tau), which suppress the luteolytic effect of prostaglandins produced by the
uterus, has been shown to be lower in underfed ewes (0.5 times mainte-
nance) while the level of uterine prostaglandins was increased. High feeding
levels during post mating increase the blood flow through the liver and the
hepatic catabolism of progesterone reducing the circulating levels of this
hormone which has a negative effect on embryo survival. Thus overfeeding
during this period should be also avoided and a level of feeding close to
maintenance during the first month of pregnancy rather than prolonging
the flushing after mating will help to reduce embryo mortality. An excess of
degradable protein reflected in high blood urea concentrations as a result of
liver detoxification of ammonia absorbed from the rumen, may also increase
embryonic mortality and foetal gigantism although the effect is attributed to
alteration of uterine environment by ammonia itself rather than to blood
urea. Micronutrient imbalances, such as Se, Co and vitamin A and E have
also adverse effects on embryo survival during implantation.
During mid pregnancy foetal growth can be affected by nutrition

through its effect on placental size. By the 6th week of pregnancy the pla-
centome number is already established and the placenta starts a rapid
growth increasing in weight up to the 90th day of pregnancy followed by a
decline of about 30% over the final 50 days pregnancy when the exponen-
tial rise in foetal weight is most apparent. The response of placental growth
to nutrition of the ewe during this period and the consequent effect on foetal
growth is however variable depending on body condition at mating and age
as shown in
FIGURE XXVI-1 summarizes published results on the response of

placental weight to the level of feeding during mid pregnancy. While sub-
maintenance feeding decrease placental weight and lamb birth weight in
mature ewes with poor body condition at mating, the opposite effect has
been observed in ewes of good body condition and particularly in young ewe
lambs. Therefore in ewes with good body condition at mating moderate loss-
es of weight during mid pregnancy, resulting in the loss of half a unit of con-
dition score, followed by liberal feeding during late pregnancy may be a use-
ful management strategy to match natural fluctuations in pasture resources
946
by taking advantage of the enhanced placental growth which seem to occur

during feed restriction. In this conditions it is likely more advantageous to
improve nutrition at some early stages to ensure that the animals are in
good body condition at mating favouring a high ovulation rate.
FIGURE XXVI-1. Response of placental weight to the live weight change (% of LW) durin-
mid pregnancy.
26.1.3.2. Late pregnancy
About 70% of foetal weight is laid down during the last 8 weeks of
pregnancy which makes foetal growth vulnerable to undernutrition at this
sage. The evolution of energy and protein accretion has been quantify by
serial slaughtering of pregnant ewes with different foetal burden and data
can be related to birth weight with enough accuracy. Requirements in Table
XXVI-1 has been assessed assuming an efficiency of ME utilization of 0.13
which is slightly lower than the coefficient found in pregnant sows (0.2-0.3)
because in the ewe the total heat increment during pregnancy is attributed
to the gravid uterus while in the sow is partly explained by the assessed
increase in maternal maintenance requirements.
Protein requirements are also based on measured rates of protein

accretion in the gravid uterus (Robinson —-) and an estimated retention in
947
the udder (Rattay —-), assuming efficiencies of retention of 0.85 and 0.68 in
each site respectively. Since protein accretion accounts for a constant pro-
portion of the energy retained in the gravid uterus (c. 78%), the require-
ments of MP are also relatively constant when expressed in relation to ME
for pregnancy. As previously observed in maintenance, protein require-
ments in pregnancy are also met by microbial and undegradable protein
when sufficient energy is provided and the degradable N requirements are
satisfied by a conventional protein source. However, in late pregnancy vol-
untary intake frequently constraints the consumption of the required ME
and body fat is consequently mobilised as a source of energy. This mobi-
lization reflected in high circulating levels of free fatty acids and also beta-
hydroxibutyrate as a result of reduced fat liver oxidation due to the exten-
sive utilisation of glucose by the foetus, can lead to pregnancy toxaemia
when the energy deficit is large, particularly in fat ewes more prone to mobi-
lization of body reserves due to their lower appetite. Avoiding over-feeding
in mid pregnancy and improving the energy and protein content of the diet
offered before parturition helps to prevent the disorder by increasing the
intake of glycogenic substrates. At the onset of symptoms the oral adminis-
tration of Na propionate and propylene glycol as glycogenic substrates can
be used as therapy.
Since a low intake of fermentable energy inevitably leads to low yields

of microbial protein, the resultant deficit has to be balanced by proteolysis
of maternal tissues or by an increased supply of undegradable protein
which explains the positive response of birth weight to protein supplemen-
tation observed during energy under-nutrition. This is particularly relevant
in many extensive systems where the negative effect of low quality
roughages on energy intake and lamb birth weight can be alleviated by
undegradable protein supplemen-tation, taking advantage of the reduced
protein degradation in late pregnancy as a result of the increased rumen
outflow rate. Among the essential amino acids, cysteine is probably the first
limiting due to the high rate of accretion in the lamb’s coat exceeding the
microbial supply. Although cysteine can be synthesised from methionine a
depressed microbial yield as results of low energy intake can promote a defi-
ciency reflected in the wool growth of the ewe and the impaired development
of secondary wool follicles of the foetus. Therefore a protein source of low
degradability rich in sulphur amino acids is an advisable supplement in late
pregnancy to sustain foetal growth during energy undernutrition.
948
While protein restriction has an important impact on foetal growth,

deficiencies of Ca and P have little effect if any on skeleton development due
to the foetal control on mineralization even at expenses of maternal deple-
tion. It is not surprising therefore that ewes can be affected by hypocal-
caemia during the last weeks of pregnancy with flaccid paralysis as the first
symptom even though the tetanic variety is also frequent. Differential diag-
nosis of this disorder from pregnancy toxaemia is often difficult since both
arise during the same period but urine analyses to detect ketone bodies in
the case of toxaemia may be a useful aid. A rapid recovery of hypocalcaemia
can be achieved by intravenous administration of Ca gluconate.
Inadequate intake of other essential mineral elements and vitamins

invariable lead to lower viability of the new born lamb.
26.1.4. Voluntary intake and energy balance
Voluntary intake tends to decrease in late pregnancy due to compression of

the rumen by the growing uterus although endocrine changes before partu-
rition might also be a contributory factor. The decline is higher in ewes over-
fed in mid pregnancy and increases with litter size and poor quality
roughages. A likely estimation of dry matter intake can be made from pre-
diction equations developed from diets based on hays and silages (ORR and
TREACHER, 1984, 1989, AFRC, 1993) although adjustment is necessary when
extrapolated to breeds of different live weight.
The evolution of ME intake estimated from the AFRC (1993) equation

is compared in FIGURE XXVI-2 with energy requirements of single and twin
bearing ewes during the last weeks of pregnancy. Because voluntary intake
is restricted by ruminal distension, poor quality roughages fail to provide
sufficient energy, mainly in twin bearing ewes. Meeting the needs by means
of concentrate supplementation is not always practicable due to the high
levels of supplement required as a result of the substitution effect of con-
centrates on roughage intake which has been shown to increase with stage
of pregnancy and foetal burden. Therefore some degree of energy undernu-
trition is inevitable at this stage if concentrate feeding has to be restricted
to avoid the risk of acidosis.
949
FIGURE XXVI-2. Voluntary intake and energy requirements (x maintenance) of single and
twin bearing ewes fed on roughages of different energy content.
Mobilization of body reserves can attenuate the energy deficit and

apparently a mild degree of undernutrition (approx. 15% of daily energy
requirements) can be tolerated without a detrimental effect on lambs birth
weight. Translation of this energy deficit to changes in body condition or net
maternal weight (live weight change minus foetus, placenta and annexes) is
difficult due to the variable composition of body weight changes. The fol-
lowing relationship between net body weight change (x, g/d) during the last
90 days of pregnancy and maternal energy balance (y, kJ/d) estimated in
vivo from the dilution of deuterium oxide in body water (DAPOZA et al. 1999):
y = 11.2 x – 636; RSD=734
shows that the energy content of net maternal weight ranges from 43
to 18 MJ/kg in ewes loosing from 20 to 100 g/d respectively as a result of
the variable proportions of body fat and protein mobilised. In this regards it
is interesting to note that energy balance (y=0) is achieved at a positive
weight change (57 g/d) due to protein and water retention concomitant with
fat mobilization for an equivalent amount of energy. From the above rela-
tionship it can be estimated, as detailed in Table XXVI-2, that a 15% of ener-
gy deficit during the last month of pregnancy is reflected in a maternal
weight loss of 1.4-2.0 kg for single and twin bearing ewes of 50 kg live
weight, equivalent to 0.2-0.3 points of body condition score, ( see Chapter
III).
950
Combining predictions on voluntary intake with these estimates it is

possible to calculate levels of supplementation to achieve defined levels of
energy balance. FIGURE XXVI-3 illustrates the level of concentrated feed
supplementation required to meet the energy requirements of single bearing
ewes or to induce a loss in body condition of about 0.5 points in twin bear-
ing ewes ranging from 40 to 60 kg live weight when roughages of different
energy value are offered ad libitum. This level of energy deficit, equivalent to
0.20 of daily requirements of twin bearing ewes is probably the threshold for
ewes in good body condition below which a significant reduction in the birth
weight of lambs can be expected, particularly in their fat content and abili-
ty to survive in view of the role of brown fat in heat production. This implies
that a medium-high quality forage needs to be offered. to avoid feeding large
amount of concentrates or to incur in higher deficits that may affect lamb
birth weight and survival.
951
FIGURE-XXVI-3: Levels of concentrate supplement required to meet the energy require-

ments of single bearing ewes during the last month of pregnancy or to avoid a loss in
body condition higher than 0.5 score points in twin bearing ewes ranging in weight from
40 to 60 kg and fed on roughages of different energy content.
Regarding dietary protein a concentration of 120-130 g/kg DM of a

75% degradable source could be sufficient when the energy requirements
are met but in conditions of underfeeding the supply of escape protein
should be increased to account for the decrease in microbial synthesis
linked to the lower energy supply. This can be achieved by raising the crude
protein concentration or preferentially by lowering its degradability as
shown in FIGURE XXVI-4. Moreover, an extra supply of glycogenic amino
acids may favour the supply of glucose decreasing the risk of pregnancy tox-
aemia.
952
FIGURE XXVI-4: Crude protein concentration required in the diet of twin bearing ewes
(50 kg live weight) receiving a 8.5 MJ ME/kg DM roughage and different amounts of a
concentrate supplement (12.5 MJ ME/kg DM).
26.1.5. Lactation
The length of the lactation period is widely variable depending on the aims
of the production system. Most sheep are reared for meat and wool produc-
tion but in some countries of Asia and South Europe milking the ewes dur-
ing 5 or 6 moths for cheese manufacturing is a widely spread. practise.
Milking used to start after weaning of lambs at 4 weeks although more
recent breeds of dairy ewes are milked just after the calostral phase and the
lambs are reared on milk replacers to avoid the abrupt reduction in milk
yield resulting from the change of suckling to machine milking that persists
for the rest of lactation,. In meat breeds early weaning at 4-6 weeks is a
common practise in intensively managed systems although the suckling
period can be prolonged until 3 moths in annual lambing systems when the
availability of forages makes suckling more economically advantageous
than feeding the early weaned lambs on diets rich in concentrate feeds.
In lactation nutrient requirements are proportional to the level of milk

production which in dairy ewes depends on the genetic potential of the
breed and the lactation stage as milk yield declines after the lactation peaks
at increasing rate. The shape of the lactation curve can be described by
mathematical models and milk yield predicted at any stage from the model
953
and the genetic potential. Although different models has been proposed the
most common are those based on the equation described by WOOD (1967)
for dairy cows as the following developed POLLOT and GOOTWINE (2000) for the
Awasi breed:
Yt = a×t0.32×e(-0.0106*t)
where Yt is the daily production at the time t (days) and “a” is a lev-
elling factor of the curve that can be estimated by dividing the potential pro-
duction in a 200 days lactation by 293. The constants 0.32 and -0.0106
describe the increasing and decreasing rates of the lactation curve and its
ratio (0.32/0.106 = 30) gives an estimate of days at the peak. For example
a ewe producing 500 kg milk in the whole lactation with an “a” value of
1.706 (500/293) can be predicted to produce 3.7 kg at the peak (30 days)
decreasing to 2.6 kg after 100 days in lactation.
In meat breeds milk production is related to the growth rate of the lit-
ter. The conversion rate of milk dry matter to live weight gain by the suck-
ling lamb is close to 1.1:1 which allows to estimate milk production with
reasonable accuracy from the lambs growth rate during the first 4 weeks
when the solid food intake is negligible. In terms of liquid milk this conver-
sion rate is equivalent to 6 kg of liquid milk per kg of live weight gain
(1.1/0.18) which results in estimated average milk yields of 1.3 and 2.0
kg/d for single and twin suckling lambs gaining 210 and 170 g/d per lamb
in the first 4 weeks. If 7.9 MJ of ME are required to retain the net energy of
one kg milk (4.8 MJ for 7% fat) with an average efficiency of 0.61, these lev-
els of milk production raise the daily ME requirements to 10.0 and 16.1 MJ
in addition to maintenance. These are equivalent to feeding levels of 1.4 and
2.3 times above maintenance for a 50 kg ewe, which makes of early lacta-
tion the period of highest nutrient requirements in the productive cycle. To
fulfil these requirements is not always feasible even with high quality diets
because the evolution in milk yield is not fully encompassed by voluntary
intake. While milk yield peaks at 2-3 weeks, voluntary intake needs about
5 weeks to reach its maximum which makes inevitable some degree of mobi-
lization to meet the energy deficit as illustrated in FIGURE XXVI-5 based on
the evolution of milk yield in meat breeds and voluntary intake predicted by
the equation developed by BOCQUIER et al. (1987).
954
FIGURE XXVI-5: Evolution of energy requirements ( ) and intake (…) expressed as times
maintenance and consequent changes in body condition score (.-..-) during the lactation
of a 50 kg ewe rearing single (●) and twin lambs (▲) and fed on mixed diets with 9.4 and
10.0 MJ/kg DM of ME.
In this figure empty body weight changes (live weight minus gastroin-
testinal content) were estimated assuming energy equivalents of 35.8 and
44.8 MJ ME /kg weight lost or gained respectively based on an energy con-
tent of 26 MJ/kg empty-body weight (AFRC 1993) and efficiencies of 0.84
and 0.58 for milk synthesis from body energy and for weight gain from
dietary ME, respectively. The assumed constancy of this values is clearly a
simplification as the composition of body weight change is more variable
than in pregnancy (COWAN, 1979) and the energetic efficiency of milk syn-
thesis seems to decline as the contribution of body fat increases.
The extent of mobilization is related to body condition (COWAN, 1980)

which would explain the positive relationship observed in dairy ewes
between milk production and body condition at lambing. In meat breeds the
955
level of milk production is lower and a decrease in yield can be partially

compensated by the increased intake of solid food by the lamb from the
third week if a palatable starter is available. Moreover an early intake of
solid food is desirable in early weaning as it ensures a more smooth transi-
tion from milk to solid food. Therefore the extent of body reserves mobiliza-
tion has lower effects on performance in meat than in dairy ewes.
According to the INRA (1988) an acceptable daily negative energy bal-

ance should not exceed during the first 3 weeks the equivalent to 1.0 and
0.6 times the fasting metabolism for ewes scoring 3.5 and 2.5 at lambing,
which in the above example results in energy concentrations of 10.0 and 9.4
MJ/kg DM for ewes in good and medium body conditions rearing twin and
single lambs. These are equivalent to daily intakes of concentrate DM
increasing from 0.6 to 0.9 and 0.3 to 0.5 kg of during the 6 weeks lactation
when a hay of 8.5 MJ ME/kg DM is offered ad libitum. Flat rates of supple-
mentation required for a similar threshold of energy depletion during the
first 3 weeks are given in FIGURE XXVI-6 for ewes of body weight ranging
from 40 to 60 kg consuming forages of different quality. It illustrates the
importance of the basal forage quality to avoid high levels of supplementa-
tion that may lead to lactic acidosis and inappetance. If shortage of good for-
age obliges to high levels of concentrate feeding a better choice is to limit the
level of supplementation during the first weeks in order to allow the rumen
adaptation even at expenses of higher levels of body fat mobilization. A pro-
gressive increase in the level of supplementation from 3 weeks onwards
would help to promote an increasing recovery of body condition at the time
of weaning that will act as flushing favouring the early cycling and the ovu-
lation rate if the feeding level is maintained after weaning.
956
FIGURE XXVI-6 Flat levels of concentrate supplementation required to maintain an

acceptable pattern of mobilization of energy reserves in the lactation of single and twin
bearing ewes ranging in live weight from 40 to 60 kg and fed on roughages of different
energy content.
While the ME equivalent of 1 kg de body reserves accounts for 4.5 kg

of milk (35.8/7.9 = 4.5) its protein content (119 g/kg) only provides the MP
required to produce 1.6 of milk (119/7.9×9.5)= 1.6). Therefore milk yield
can be only maintained if dietary undegradable protein is provided to bal-
ance the protein deficit arising from body reserves mobilisation as conse-
quence of reduced feed intake. This justifies the response in milk yield to
undegradable protein supplements commonly observed in underfed lactat-
ing sheep. Assuming a 0.7 degradability of it ca be estimated that protein
contents ranging from 150-160 g/kg is
Daily energy, protein, vitamin A and E, as well as mineral require-
ments are summerized in Table XXVI-3 and Table XXVI-4.
957
958
959
26.2. Feeding and nutrition of suckling and fattening lambs

26.2.1. Nutrients requirements
The recent NRC (2006) recommendation calculates the energy requirements

of maintenance using NEm and NEg values is given according to the aver-
age daily gain. The protein needs are given in metabolizable protein, but the
other forms of expressions are given, too. Table XXVI-5 gives the nutrient
requirements of growing lambs, using the compilation of NRC (2006).
CANNAS et al. (2004) developed a version of Net Carbohydrate and Protein
System for Sheep (CNCPS-S).
26.2.2. Newborn and suckling lamb

26.2.2.1. Natural nursing
Newborn lambs have small nutrient depots, therefore they need concen-
trated feed immediately from the birth. Supposed that the ingested amount
is sufficient, the requirements are covered by the colostrum and sheep milk.
Growth rate of the suckling lambs is high; in appropriate circumstances the
birth weight is doubled during 2 weeks. The sheep milk is more concen-
trated than the cow’s one, containing aapproximately 20% dry matter.
Generally the daily gain of lambs’ corresponds to the dry matter content of
the suckled milk, because the digestibility and utilization of the milk com-
ponent is close to 100% (NAGY and FEKETE, 2003).
Unlike calves, lambs have relatively small abomasums, this is why
the suckling frequency is higher. At the same time, the daily dry matter
intake capacity is bigger, which can be satisfied only by 15 to 20 sucklings
per day. For this reason lambs are kept together with the ewes.
First the lambs prefer roughages to the concentrate. The young har-
vested, good quality grass hay has the highest preference (“lamb hay”). The
intake of the solid feed promotes the rumen development and the estab-
lishment of the ruminal flora and fauna. Lambs become functionally rumi-
nant earlier than calves, i.e. at the age of 4 to 6 weeks of age. From this on
most of the vitamin supply is assured by the production of rumen microbes.
If the green forage intake is low, vitamin E deficiency may occur.
961
CHAPTER XXVI: SHEEP FEEDING AND NUTRITION
962
Through the suckled milk lambs consume high amounts of fat, which
requires antioxidants, namely vitamin E and selenium. In endemic seleni-
um-deficient areas skeletal and cardiac myodegeneration can develop, espe-
cially if the supply of pregnant ewes were inadequate. Selenium can be sup-
pled by sodium selenite containing subcutaneous injection or special con-
centrate feeding.
26.2.2.2. Artificial lamb rearing

The artificial rearing is not common in Merino-type flocks, because the
dam’s milk production, althougj assures only a medium growth rate, it is
just sufficient to raise up twin lambs. At the same tyme, the spread of pro-
lific breeds and strains (e.g. Romanov, Booroola) the birth of 2-3 newborn
lambs makes necessary the artificial raising.
Commercial milk replacers are available, when the way of application
is given. Generally, the following principles should be considered.
a) Lambs are allowed to suckle only the colostrum during the first 24 to 48
hours after birth, otherwise it will be difficult to get them accustomed to the
artificial drinking device.
b) Owing to the small stomach capacity, the milk replacer should be offered
ad libitum.
c) Acidified, preserved milk replacer (“penguin milk”) can be given without
previous warming.
d) Keeping of a strict hygiene during preparation and distribution of milk
replacer is essential.
e) Good quality hay and concentrate should be offered ad libitum from the
the end of the second week
963
FIGURE XXVI-7compares the natural nursing, the early weaning and the artificial rearing
of newborn lambs.
964
26.2.3. Methods for fattening
26.2.3.1. Feeding of suckling (“milky”) lambs

Using dominantly sheep milk lambs can achieve 14 to 18 kg of live weight
till the age of 7 to 9 week-old. Besides suckling pelleted concentrate is fed,
too. Milky lamb has good slaughter quality, the meat is soft, due to the con-
siderable lactose intake. This form of slaughter animal is generally produced
seasonally, for Easter.
26.2.3.2. Intensive express meat lamb’s production

The aim of this type of production is a slaugher lamb of 28 to 35 kg of LW,
at the age of 120 to 130 days of life. Lambs are fed exclusively on pelleted
concentrate. From the point of view of digestive physiology, this type of feed-
ing cannot be considered as optimum, but economically advantageous. The
transport to far distance requires special care (e.g. regular drinking), other-
wise it may raise animal welfare concers.
The growth rate is especially high till the age of 4 months. Until that
this is the protein accretion which is dominant instead of the fat deposition.
After the age of 4 months the feed intake drops and the feed conversion effi-
ciency declines. The occurrence of health problems (e.g. parakeratosis of the
rumen epithelium, chronic bloat etc.) becomes more frequent and as a
result the fattening turn to be uneconomical.
One of the prerequisite of the successful express fattening is the early
weaning: instead of the traditional 3 months, lambs should be separated
from their mothers at the age of 35-42 days. By that time they have to
achieve a live weight of twice, 3-times that of the birth weight (it means
approx. 10 to 13 kg of LW). Lambs have to get accustomed to the solid feed,
the daily concentrate intake should be at least 400 grams.
Supply of the appropriate pelleted concentrate should already be
given from day 14 as a creep feed; drinking water and licking salt blocks
should be provided continuously. Because the capacity of rumen and abo-
masum is small, the pelleted concentrate should be offered ad libitum after
the weaning, too. In the practice one-phase or two phase feeding technolo-
gies are used. The concentrate of the one-phase technology as well as the
Phase I concentrate (“starter”) of the two-phase technology should not con-
tain NPN-substances. On the contrary, Phase II concentrate (“finisher”: feed
of the last month of fattening) may compromise urea, too. Concentrates con-
965
sist of cereals, (solvent) oilseed meals, alfalfa meals, occasionally NPN sub-
stances, minerals and vitamins. The expected average daily gain amounts
200 grams. The role of dehydrated alfalfa meal is to provide rumen unde-
graded protein, fibre and calcium. Using feed flavours the voluntary dry
matter intake may be enhanced and added synthetic lysine cen balance
amino acid profile.
The technology of the intensive express meat lamb production uses
exclusively pelleted concentrate to the animals in groups of 40 to 60. This
predisposes lambs to health troubles, because the feed is low in fibre (10
to12% crude fibre), and this form of fibre, owing to its small particle size,
cannot considered as a functional one. This is the reason, why intensively
raised lambs should not be used as breeding animals in the future. Fibre
deficient lambs incline to eat bedding material, like straw, which, in turn,
decreases the concentrate intake. The above described facts queries the jus-
tification of this fattening practice from the proint of view of animal protec-
tion.
The high growth rate requires large amount of calcium and phospho-
rus. The latter is assured by the cereals, but the adequate calcium supply
generally needs limestone supplementation. Phosphorus excess makes
lambs, especially rams, prone to the formation of urolith. By decreasing the
urine pH by ammonium chloride supplementation the danger of the devel-
opment of urinary calculi can be prevented or diminished. Table XXVI-6
gives the daily mineral requirement of some model growing lambs, using the
compilation of NRC (2006).
966
967
26.2.3.3. Semi-intensive fattening of growing lambs

The aim of this type of production is a slaugher lamb of 40 to 50 kg of LW,
at the age of approximately 6 to 8 months. The animals are fattened using
mostly bulky feeds, predominantly meadow grass. The carcass contains
more fat (suet) but it is preferred on the Middle-Est markets.
Lambs are generally weaned at the age of 3 months, but they already
go to graze during the suckling period together with their mothers. If the
pasture /and hay can also be fed. During this type of fattening the feed uti-
lization is bad, because parallel to the age the fat deposition increases and
the protein accretion decreases. Nevertheless, considering the relatively low
price of forages and rougheges this type of fattening may be proved eco-
nomical.
26.6.2.3. Improvement of old, culling sheep for slaughtering

Breeding ewes are generally culled at the age of 4 to 6 years, considering
their production level and the state of the teeth. Their feed conversion effi-
ciency is bad, the meat production is low and the meat quality is poor.
Therefore only an improvement of the condition can be justified. Cull ewes
may receive only juicy feed like fodder beet. Owing to the bad teeth quality,
the chewing of roughages is incomplete and consequently the feed utiliza-
tion bad. Cereals should be given in ground form; continouos water and
common salt supply is essential. The duration of this condition improve-
ment lasts generally 2 or 3 months. Considering the expected muscle
enlargement, the protein requirement slightly exceeds that of the mainte-
nance.
968
26.3. Deficiency syndromes and metabolic disorders of

sheep
ACUTE LACTACIDEMIA.
Ailment develops after the ingestion of large amount of readily fermentable
carbohydrate (e.g. grazing of corn fodder field, or corn aftermaths) without
previous adjustment. The concentration of lactic acid in rumen abruptly
increases from the normal 50 mg/100 ml to 1 to 2 grams/100 ml and the
pH decreases from the normal 6 to 7 up to 4.0 to 4.2. Lactic acid is pro-
ducing bacteria (like lactobacilli, Streptococcus bovis etc) are normal con-
stituents of the ruminal flora, but without available substrata they are not
harmful. In the very acidic milieu the rumen protozoa will die. The high
osmotic concentration of the rumen content causes saliva and blood plas-
ma diffusion into the lumen through the ruminal wall. This repartitioning of
the body fluids results in collapse and death (per acute form). Acute and
subacut forms of disease are dominated by metabolic acidosis and necrosis
of the rumen epithelium; total germ count of rumen increases, primarily by
the lactate-producing bacteria, secondary the E. coli and Proteus strains,
causing a consecutive putrefaction of the rumen content. Affected sheep
show prostration, nervous signs, anuria and defecation of small amount of
yellowish green. Part of the animals becomes distended/blows up; breath
(halitus) is penetrating sourish. To treat acute cases, the intraruminal injec-
tion of 20 to 50 ml 5% of sodium hydroxide is the first measure, after the
inflammation should be alleviated and the lacking thiamine production
replaced. To avoid this ailment, the feeding of readily fermentable carbohy-
drates should be gradually, in order to make sheep rumen microflora accus-
tomed. Before leaving animals to graze on a stable field, early morning in the
stall meadow hay, corn stalk or straw should be fed.
ALGAE TOXICITY
Algae poisoning is rare but the danger from poisoning is always present if
sheep drink water from dams or lakes on which algae grow forming a scum
on the surface (see Non-Infectious abortion unit in Chapter XX). Freshwater
algae like strains of Microcystis, Anabena and Aphanizomenon) are usually
blue, green or yellowish-green in colour. Sheep are particularly susceptible
to poisoning. Algae contain toxic substances which damage liver. The algae
969
get dangerous only, when they grow very profusely in warm (dense bloom),
still water and are then disturbed by windy weather which disperses them
through the water or concentrates them along the edges where stock are
liable to swallow them as they drink. Clinical signs take a very acute form
in which mortality occurs rapidly and a less acute form in which there is
jaundice and photosensitisation, ending in deaths within one or two weeks
or a chronic unthriftness. Haemorrhages and free blood in various internal
organs is the main sign at necropsy. There is no antidote known. Treatment
should be symptomatic. Namely, affected animals should be placed in a pro-
tected area out of direct sunlight. Large quantities of water and good quali-
ty hay should be made available. Administration of activated charcoal slur-
ry and laxative dose of heavy mineral oil has proved useful in removing the
toxin from the organism. However, for the prevention, chemical control of
algae is possible. This should be applied as a spray to ensure even distri-
bution and avoid the danger of killing fish.
BLOAT
Most frequently it results from grazing on protein-rich legumes (lush green
alfalfa or clover), without previous adjustment. The characteristic sighs are
swelling on the left side, marked uneasiness leading to breathlessness and
rapid death. In a freshly dead sheep the rumen is full of froth. As a first aid,
drenching with about 1 dl of paraffin, sunflower or rapeseed oil may relieve
affected sheep. The rest of the flock should be removed from the. For details
see Beef cattle (Chapter XXV).
COPPER DEFICIENCY and POISONING.

Sheep (and cattle) is especially susceptible to the shortage in copper supply,
because the absorption is very poor. Inhibiting interactions with sulphur
and molybdenum further lower copper availability. Copper being part of
important metalloenzymes, the clinical signs can be realized: loss of wool
crimp (“steely wool”, in breeds with coloured fleeces decolouration, too),
congenital or delayed swayback (enzootic ataxia), anaemia, susceptibility to
infection and fragile bones. Necropsy reveals cavitations of the cerebral
white matter and internal hydrocephalus. In the brain stem chromatolysis
and in the spinal cord bilateral symmetrical demyelination is shown. Form
of anaemia hypochromic, microcytic or macrocytic) depends upon the stage
of deficiency. Diagnosis can be verified by the low copper concentration of
970
brain and spinal cord (below 79 μmol copper/kg DM) and the blood plasma
(below 9.0 μmol copper/l. Liver analysis is not sensitive enough. The super-
oxyde dismutase (SOD) activity in the red blood cells refers to a prolonged
deficiency. Treatment has no reason; for the prevention the application of
copper fertilizers (copper sulphate), oral application of CuO, supplementa-
tion of feed and also injections are available.
Copper toxicity.
Sheep are extremely susceptible to copper poisoning. Acute form develops
following the intakes of 20 to 100 mg/kg LW copper, mostly in form of fer-
tilizer or feed additive. In typical cases, caused by prolonged intake of low
amounts of copper (e.g. eating growing pig feed), toxicity remains subclini-
cal, until the limits of liver storage capacity. If the liver copper concentra-
tion exceeds 750 mg/kg DM, copper is abruptly released into peripheral
blood circulation, causing severe intravascular haemolysis and icterus (tox-
aemic jaundice). Intake of plants, containing hepatotoxic alkaloids (wild
heliotrope, lupines and ragwort) can facilitate copper poisoning. For treat-
ment of acute cases ammonium tetrathiomolybdate is recommended, 3.4
mg/kg LW s.c. daily, three-four times. Daily oral administration of 100 mg
ammonium molybdate with 1 gram sodium sulphate per animal or drinking
of water with 20 mg/l ammonium molybdate helps prevent further clinical
outbreaks. By applying 50 mg/kg LW penicillamine enhances urinary
excretion of the copper. For the prevention, 7 mg/kg molybdenum should
be added to the feed or top-dressing the pasture with 70 g Mo/ha, in form
of molybdenized superphosphate.
ENTEROTOXAEMIA OF FEEDER LAMBS.

Enterotoxaemia („overeating disease“; „pulpy kidney disease“) an acute,
mortal disease of finishing lambs on high-grain rations. This is the most
common of all ovine toxic renal disease. Ailment is caused by the epsilon
toxin, produced by an anaerobic bacterium, Clostridium perfringens Type D.
This bacterium is widely spread in the nature, for example it is found in the
soil and in the digestive tract of the healthy sheep. Affected lambs in a high
state of nutrition (e.g. lush feed of grain, milk or grass), ingesting large
amount of starch and/or sugar, while .bacterium is able to grow and pro-
duce toxin. Accumulating toxin damages the gut lining and enters the cir-
culation. Therefore, it may occur not only in finishing lambs, but also in
suckling and milk replacer consuming lambs, as well as in high-pregnant
971
ewes. Influences, modifying peristalsis (e.g. gastrointestinal parasites) are

predisposing factors. The toxin is so lethal to the renal tubules that they are
completely destroyed in some hours, resulting in rapid death of the animal.
Being the pulpy kidneys, the pathological finding during necropsy is char-
acteristic and toxin can be detected in the rumen content. Prevention con-
sists in applying Type D toxoid injection. Vaccinated ewes assure sufficient
passive immunity for the suckling lambs in the first 4-6 weeks of their life.
After that the use of an antiserum is possible. Incidence of the enterotox-
aemia in the feedlot may be reduced by lessening the grain ration, but in
this case the growth rate will slow down.
IODINE DEFICIENCY is common in many regions, where the soil and water is
poor in iodine. Secondary iodine deficiency develops, when animals ingest
feedstuffs, containing thyreostatic substances (for details see Chapter VIII),
or drink water high in nitrates and nitrites. Cyanogenic glycosides of some
forage are also mentioned (e.g. white clover). These compounds inhibit the
iodine uptake of thyroid glands. Such plants are generally the Crucifer, in
most of cases rape, rapeseed, cabbage, kale, turnip, Schwedish turnip.
Since selenium takes part in deiodinase enzyme activity, selenium deficien-
cy indirectly makes animal susceptible to goitre. In most of the cases, the
insufficient iodine supply of pregnant ewe results in the goitre of newborn
lambs. Most of the affected lambs are born dead or will die within a few
hours. Characteristic is and obvious enlargement of the thyroid gland on
the neck and swelling of neck and tongue. Another occasionally appearing
symptom is the birth of lambs without wool. For the prevention, iodized salt
should given (2.2 g potassium iodate/kg common salt) for ewes during the
last half of pregnancy.
NEWBORN LAMB STARVATION occurs, if colostrum intake of lamb is insufficient,

owing either owing to its weakness or the inadequate milk quantity. The
colostrum requirement of newborn lamb during the first day is strikingly
high, according to the environmental conditions, 210 to 280 ml/kg LW. The
consequences of deficient colostrum intake are the hypoglycaemia of lamb,
alteration of the normal gut maturation and growth and immune incompe-
tence. To prevent this situation, maternal underfeeding should be avoided
and especial attention should be turned to the newborns` colostrum supply.
972
PHOTOSENSITIZATION.
Potentially hazardous plants may be present in native meadows, seeded for-
age crop fields and intermittent croplands. Among others, it worth men-
tioning the Saint Johnswort (Hypericum perforatum), summer grass, yellow
vine (Tribulus), heliotrope, algae, alsike or Schwedish clover (Trifolium
hybridum), crowfoot and buckwheat (Phagopyrum esculentum) with photo-
sensitizing properties. Photosensitization is the result of the lightly pig-
mented parts of the skin sunlight. This is brought about by the presence in
the skin of photosensitizing agents which reacts to light. This agent is usu-
ally a pigment of plant origin, or sometimes, chemical origin, such as phe-
nothiazine used to be. “Yellow big head” is also a form of photosensitization
and it is caused by grazing on pasture including summer grass, yellow vine
or catheat (Tribulus terrestris) and green millet (Panicum effusum). It is par-
ticularly common in young sheep. The ears and face swell and may exude a
yellowish fluid. The whites of the eyes take on a muddy-yellow colour. The
swelling becomes hard and mummified. Losses are often quite heavy though
many sheep recover. Treatment is impracticable on large scale. Affected
sheep should be removed from sunlight and allow them to graze at night.
For more details see Chapter VIII.
PLANT OESTROGENS.
After the intake of oestogenous compound found in leguminous pasture from

ladino clover, alfalfa and subterranean clover (Trifolium subterranum) repro-
ductive troubles occur both in sheep causing infertility. (Feeding these for-
ages as hay or haylage is not known to cause a major problem.) An exces-
sive intake of plant estrogens causes interference with the normal functions
of the reproductive cycle. The effect is progressive and cumulative, causing
either permanent or temporary infertility. Dystocia becomes frequent. One
of the most characteristic signs is the “high tail” which causes ewes to carry
their tail high, causing by the relaxation of certain pelvic ligaments. Wethers
shows false lactation and formation of a large sac situated between the
bladder and pizzle. It is filled with urine and bulged externally below the tail.
As prevention, avoid clover-dominant pastures during the mating season
and lambing.
PREGNANCY TOXICOSIS (SHEEP KETOSIS or “TWIN-LAMB DISEASE”).

During the last quarter of the pregnancy, the glucose requirements of
973
foetus(es) are considerable. From the point of view of nutrient supply,

foetus has a priority over against the maternal tissues. If high-preg-
nant ewes have not a satisfactory carbohydrate (i.e. concentrate) pro-
vision, the temporary energy deficit is covered from the fat content of
the mother’s body. If it long-lasting, pregnancy toxaemia (or sheep
ketosis) develop in the pregnant ewe. Undernutrition is by far the most
important factor in practice while stress is frequently responsible for
triggering it off. It is usually facilitated, even besides normal condition,
by reduced feed intake, such as hypocalcaemia, change in diet or
movement.
Mobilized maternal triglycerides are decomposed into glycerine
and free fatty acids, which in turn, transform into keton bodies.
Parallel to the already existing low blood glucose, the mentioned
decomposition products accumulate in the organism. Long-lasting
hypoglycaemia (around 1.5 mmol/l) alters the susceptible brain tissue.
Thereby the clinical signs are characterized by dullness and blindness.
An affected ewe stands on her own and can be approached without
showing any obvious response. She makes no attempt to eat, have
uncertain gait, then extreme apathy and may stand sleepily for hours
in the same position, then after a comatose phase animals die. During
necropsy in the uterus of died ewes well developed, mature, but died
embryos can be found. One of the most remarkable finding is the liver
of dams, which are enlarged, friable and ochre (GAAL and FEKETE,
1982).Kidney are also pale yellow and friable. Lesions can be found in
the brains; urine gives positive keton answer (e.g. Ketostix, or sodium
nitroprussic powder) and besides low blood sugar concentration the
beta-hydroxy-butyrate level is higher than 3.0 mmol/l. The ewe most
possibly will have a body condition score of 2 or less and having two or
more foetuses. Treatment includes replacement of glucose and elec-
trolytes parenteral, and giving propylene glycol orally. For avoiding the
pregnancy toxaemia, ewes should be grouped according their condi-
tion, pregnancy stage and foetal number, and if necessary, additional
concentrate should be given.
THIAMINE DEFICIENCY.
The ailment is dominated by opisthotonus, having polyencephalomalatia

(PEM) and cerebrocorticalis necrosis (CCN) in the background. The diseases
974
generally occurs in older (3-4 months of age) fattening lambs, in case of

abrupt change of feed. Drastic change of pasture, from a good to a very poor
one may also a cause. In such a case the thiaminase-producing bacteria
grow in the rumen. In wintertime appears only sporadically, if thiaminase-
containing silage, hay with high bracken fern (Pteridium aquilinum) or rock
fern (Cheilantes sieberi) contamination or sulphate of sulphite anion-con-
taining industrial by-product (sugar beet pulp, wet corn gluten feed) is fed.
Lack of appetite, sudden depression, medial dorsal strabismus (“star gaz-
ing”), opisthotonus, and disturbed gait are followed by tonic-clonic convul-
sions in untreated cases. Lack of the thiamine pyrophosphate disturbs the
energy supply of brain cells (mostly in the cortex), causes neuronal necro-
sis and swelling of astrocytes. Secondary ischemic lesions occur due to
brain swelling. During necropsy in the swollen, oedematic and haemor-
rhagic brain cortex greyish yellow necrotic foci can be found, which are aut-
ofluorescence. Vessels of the meninx are swollen and the mucous mem-
brane of abomasum and intestine are in inflammation.
Ingestion of large amount of green bracken fern plant produces signs
of acute poisoning related to thiamine deficiency in monogastric animals
and preruminant lamb. Bracken fern contains not only thiaminase, but also
other toxins; therefore symptoms are more complex, than in case of a sin-
gle thiamine deficiency. Signs in sheep include bright blindness as a sign of
a progressive retinal atrophy that derives its name from the hyperreflectivi-
ty of the tapetum in affected sheep. Bladder tumour and bone marrow alter-
ations are naturally occurring, too.
Therapy should begin immediately after onset of signs, because neu-
rons are dying by millions. Treatment of thiamine deficiency is highly effec-
tive if diagnosis is made early enough. Injection of a solution of thiamine
hydrochloride at 10-15 mg/kg LW is suggested, given initially intravenous-
ly every two hours, then intramuscular for the second day. Peroral supple-
mentation of 0.1-0.5 grams thiamine should last 1-2 weeks. According
another protocol, the introductory therapeutic dosage of thiamine is 0.5-2
grams per animals, half of that should be given intravenously, another half
subcutaneously. From the point of view of differential diagnosis, the liste-
riosis (taken up by muddy silage), copper-deficient (congenital) ataxia, pes-
ticides poisoning and scrapie should be excluded.
975
UREA POISONING.
The most common cause of the poisoning in fattening lambs is the giv-
ing of feed supplements containing urea without previous adjustment
period. It can occur by ingestion (licking) of nitrogen fertilisers. The sin-
gle toxic dose is 0.4 g/kg LW, showing higher tolerance than in case of
cattle and lower that in goat. Urea fermentation in the rumen releases
ammonia. The absorbed ammonia causes elevation in blood pH and
nitrogen concentration. Affected animals show general weakness, stag-
gering, muscle twitching and other nervous signs. Profuse salivation,
convulsion ending in comatose state and death is typical. The
drenched vinegar drops pH in rumen and transforming ammonia into
the less toxic ammonium. Assimilation of the absorbed ammonia in the
ornitine cycle can be supported by 0.2 g/kg LW glutamate peroral or 5
ml/kg LW of 5% and of pH 6 malic acid with glucose intravenously. For
further details see Beef Cattle (Chapter XXV)..
UROLITHIASIS.
Mineral deposition within the urinary tract in the form of small stones
(calculi) can occur in both males and females, but the problem great-
est in ram and especially in wethers. This hard deposition collect in the
sigmoid flexure and urethral process, thereby interfering with urine
flow and kidney damage may also occur. Kidney damage is more fre-
quent in preruminant lambs, because they are unable to degrade
oxalates, which will crystallize out in the kidney tubules and the cen-
tral nervous system to cause chronic renal failure and nervous signs.
Symptoms of the typical urolithiasis include tail twitching, uneasiness,
kicking at the abdomen, dribbling urine and straining in an attempt to
urinate. In advanced stages the urinary bladder may rupture and urine
spills into the abdominal cavity, giving rise to an extended abdomen
referred to as „water belly“, and death follows.
The most common calculi in fattening lambs on high-grain
rations with unbalanced mineral composition (high in phosphorus and
low in calcium) is the struvite type, which contains magnesium, mag-
nesium phosphate and small amount of calcium. Mucoproteins, cell
debris and epithelial cell may enhance stone formation. The formation
of crystals from compounds normally in urine is facilitated by changes
in urine concentration and pH. The mucoproteins and mycopolysac-
976
charides have also an important role in the formation of magnesium

ammonium phosphate, magnesium phosphate, and silica and calcium
carbonate stones. Surgery of a simply snipping of the urethral process
may be helpful. Struvite crystals are soluble under pH 6.8, therefore
withholding of feed for 24 hours in conjunction with oral dosing of
ammonium chloride or (7 to 10 grams/day for a lambs of 23-30 kg LW)
can acidify the urine. Acidification should be maintained for a week fol-
lowing surgery. Since the most frequent struvite type urolits are asso-
ciated with high-concentrate rations high in phosphorus, limestone
should be given to set 2 to 1 calcium to phosphorus ration. If the inci-
dence of urolits is high, as prevention, 0.5% ammonium chloride or
sulphate should be mixed in the concentrate. Vitamin A addition to the
diet or injection help regenerations of the urethral epithelium.
In grazing sheep the ingestion of plants rich in silica (e.g. sedge,
rush, cane) may promote the formation of silica calculi. The addition of
1-4% of salt in the ration in conjunction with ad libitum drinking pos-
sibility proved to be benevolent by diluting urine. For grazing animals,
before allowing them to the pasture, small amount of concentrate with
9% of salt or 2% of ammonium chloride may be offered.
WHITE LIVER DISEASES OF LAMBS

The outbreak of this ailment is rather seasonal, having the highest inci-
dence in spring. Morbidity may achieve 40%, besides a 15% of mortality. In
the acute form (7-10 days) is dominated by eyelid and labial oedema and
inflammation (“weepy eyes”), caused by a secondary photosensitization of
hepatic origin. Chronic form (4-6 weeks) shows normocytic normochromic
anaemia, anorexia and loss of weight and crusting lesions on the ears.
Ailment is caused by cobalt and later a subsequent vitamin B12 deficiency.
Forages containing less than 0.07 ppm are considered to be deficient for
ruminants. Cobalt-deficient soils are not uncommon all over the world.
During necropsy an extreme emaciation, bone marrow hyperplasia and
haemosiderin deposits are found in the spleen. Livers are grossly enlarged,
pale, fatty and friable, showing the fatty infiltration.
In the background of clinical and pathological picture stands the
decrease of two coenzymes in the tissues, caused by inadequate vitamin B12
absorption. First of them (methylcobalamin) contributes in the methionine
synthesis, the other (deoxyadenosylcobalamin) does in methylmalonyl coen-
977
zyme A mutase, a key enzyme of ruminant energy metabolism. The function

of the later is to facilitate the “processing” of propionate. In lack of this
enzyme, the methylmalonyl coenzyme A accumulates and inhibits the oxi-
dation of fatty acids in liver, elucidating the development of ovine white liver
syndrome (OWLS). The prevention may start in the field, by supplementing
the superphosphate fertiliser with cobalt sulphate. Cobalt salt can be
administered orally, at monthly intervals with an anthelmintic, in a dosage
of 1 mg Co/kg LW. A good oral method is the administration of Co-contain-
ing bolus (possible with Cu and Se). In a dosage of 120 mg cobalt sulphate
per kg licking salt will also protect sheep. For the treatment of acute cases,
intramuscular injection form is also available.
WHITE MUSCLE DISEASE

or “stiff lamb disease”; nutritional muscular dystrophy”. This syndrome is
associated with selenium and/or vitamin E deficiency. In can be occurring
in three forms: congenital, delayed and in hog(get)s. In the first case the
pregnant ewe received insufficient supply from selenium and vitamin E and
among newborn and suckling lambs the incidence of white muscle disease
is high. Affected either born dead or die within some days. In the delayed
cases (from birth up to 3 months of age) lambs are emaciated, their heart-
beat is arrhythmic, their EKG is irregular. Affected lambs show stiffness,
unwillingness to move and losses may reaches 15%. The similar disease of
older animals (9-12 months of age) usually occurs, if the selenium and vita-
min E supply during wintertime was poor; Symptoms generally comes sud-
denly, during or after driving. Acute signs of stiffness, listlessness, inability
to stand and death within a day. Many times they excrete reddish brown
urine.
Lesions are seen both in hearth muscle and body muscle. Affected
muscle is light in colour, resembles fish muscle, usually with white streaks
or white spots through it. Lesions develop through a hyaline degeneration
followed by coagulative necrosis and calcification (Zenker-type necrosis).
Individual muscle fibres may be oedematous and swollen, with loss of cross
striations and sarcolemnal proliferation. A bilaterally symmetric distribu-
tion of muscle lesions in the thigh and shoulder regions, are characteristics,
deep muscles overlying the cervical vertebrae also show white striations.
Acute cases should be treated by sodium selenite and vitamin E injection:
for prevention the selenium and vitamin E provision of pregnant ewes
978
should be increased in touched territories.

Se-poisoning is caused of plants grown on soils containing selenium,
or by overdosage of feed additive. Leg deformities, loss of hair, hoof troubles
are characteristics. Selenium toxicity in lamb, showing congenital deformi-
ty (especially in the hind legs) is traceable to selenium injury during foetal
development.
FOR FURTHER READING
AFRC (1993) Energy and protein requirements of ruminants. CAB International.

Wallingford, UK
Bocquier, F., Thériez, M. and Brelurut, A. (1987 : The voluntary intake by ewes during
the first weeks of lactation. Anim. Prod. 44, 387-394.
Cannas, A., Tedeschi, I.O., Fox, D.G., Pell, A.N. and Van Soest, P.J. (2004): A mechanis-
tic model for predicting the nutrient requirements and feed biological value for
sheep. J. Anim. Sci. 82, 149-169.
Cowan, R.T., Robinson, J.J., Greenhalgh, J.F.D. and Pennie, K. (1979): Body composition
changes in lactating ewes estimated by serial slaughter and deuterium dilution.
Animal Production 29: 81-90.
Cowan, R.T., Robinson, J.J., McDonald, I. and Smart, R. (1980): Effects of body fatness
at lambing and diet in lactation on body tissue loss, feed intake and milk yield of
ewes in early lactation. J. Agric. Sci. Cambridge, 92, 123-132.
DLG- Futterwerttabellen- Wiederkauer (7., rev. ed.): 81997): DLG Verlag. Frankfurt am
Main
INRA (1988) : Alimentation des bovins, ovins & Caprins. (Ed. Jarrige, R.). Institute
National de la Recherche Agronomique. Paris.
Nagy, B. and Fekete, S. (2000): Sheep feeding and nutrition (in Hungarian. In: Fekete, S.
(Ed.) (2003): Veterinary Nutrition and Dietetics (in Hungarian). University Textbook.
Budapest – Zamardi –Zebegeny, pp 581-612.
NRC (2006): Nutrient requirements of small ruminants. Sheep, goats, cervids, and new
world camelids. National Research Council, National Academy of Science.
Washington, DC
Orr, R. J. and Treacher, T.T. (1984: The effect of concentrate level on the intake of hays
by ewes in late pregnancy. Anim. Prod. 39, 89-98.
Orr, R. J. and Treacher, T.T. (1989: The effect of concentrate level on the intake of grass
silages by ewes in late pregnancy. Anim. Prod. 48, 109-120.
979
Chapter XXVII
FEEDING OF GOATS
© Sandor Gy. Fekete and Eva Cenkvari
27.1. The „standard” goat

27.2. Biological and nutritional physiological specialities in
goats
27.4.1 Dry matter intake
27.4.2. Energy and proteinrequirements
27.4.3. Mineral requirements
27.4.3.1. Macroelement requirements
27.4.3.2. Microelement requirements
27.5.Feeding the kids
27.8. Nutrition and feeding of breeding bucks
27.9 .Feeding strategies in goats
27.9.1. Grazing
27.9.2. Alternative choice of feeds
27.10. Frequently used feeding systems of dairy goats
981
Chapter XXVII: FEEDING OF GOATS
proximately 90% of the world’s total 500 million goats are raised in
A Asia and Africa. In Europe, France has the largest stock with about
2 million goats kept on farms. People who breed goats for living keep
an average of 300 dairy stocks. The best animals now are capable of pro-
ducing 1500 to 2000 kg of milk per year. Small stocks of 50 to 60 goats are
kept as an additional income source. Beside the by-products such as kid
meat and kid skin the main products are milk and cheese produced of that,
respectively. On average, 50 types of cheese are produced in France.
Variability in body size and in geographic distribution among goat breeds
exceeds that of any other farm animal. Although goats can consume and
utilise a wide variety of low cost forages, feed costs can account for at least
55% of the production.
The most widespread goat varieties are Saanen (of 65 kg adult live
weight), and Alpine (of approx. 60 kg adult live weight), respectively. Further
worldwide spread breeds are the French Alpine, the Nubian, the Saanen,
the Toggenburg, the South-African, the Dutch pied, the East-African dwarf
and the Angora goats. In Angora goats, also wool is utilized. There are
numerous goats breeds in India where a large proportion of the world’s
goats are found. Goat milk has a similar gross composition to that of cows’,
although there are differences within the fat fraction with a higher propor-
tion of small fat globules in goat milk. Cashmere is much finer than either
wool or mohair and weight for weight, has a substantially higher insulating
value. Goats are the most commonly applied model animals in metabolic
feeding trials. Especially dairy cows are well to simulate with those, espe-
cially with the Cameroonian dwarf goats. In spite of the above facts, their
biological characteristics in many aspects, are different, from sheep and
from cattle as well.
982
27.1. The „standard” goat
A good considered „standard” goat when it is of 60 kg live weight and pro-

duces 4 kg of milk of 3.5% milk fat (approx. 1000 kg milk per lactation).
That animal takes up 2.65 kg dry matter of the INRA reference grass (123
g×W0.75) which is the value most similar to the one of dairy cows, conse-
quently we use UEL as the unit of bulkiness in goats (INRA, 1985). Daily dry
matter intake is close to 1.5 to 5.0% dry matter of the live weight (mature
average= 2.25% of LW while in sheep only 1.54% of LW) and the daily vol-
untary dry matter intake exactly equals with 2.65 units of bulkiness in
dairy goat.
27.2. Biological and nutritional physiological specialities in

goats
Goats are of a very adaptive nature (they can be kept even in the most arid
areas and in very intense production systems). Goats can be maintained
well on poor pastures because they have the ability to choose the high-qual-
ity parts of plants. They are especially selective about feedstuffs (voluntary
feed intake), but it does not mean that they are too „choosy”, but contro-
versially, they search and find the most valuable plant parts, whereas sheep
does not (e.g. it grazes tree leaves, even from among stings). Goats are effi-
cient browsers and selectively consume a variety of shrubs, woody plants,
weeds and briars.
Their nutritive needs may be higher than in other ruminants if an
acceptable level of production is to be attained. They will eat anywhere from
1.5 to 5 (mature average= 2.25) percent of their live weight (on dry matter
basis), while cattle and sheep normally eat only 1.5 to 3 percent (mature
average=1.54 of % LW at sheep, NRC, 2006). Similarly to cow the goats’
rumen is the largest compartment of their stomach representing about 80%
of total stomach area. It appears that goats and sheep have similar capaci-
ties to digest forages of medium to high digestibility (OM digestibility>60%).
When fed on low-quality roughages without nitrogen supplementation goats
are likely to have an advantage over sheep in being able to maintain
digestibility due both the selection and the higher rumen to LW ratio.
Conventional feeds are likely to have the same digestible energy value (DE)
983
and presumably metabolisable energy value (ME) value for both sheep and
goats. Their rumen has a relatively larger volume compared to their live
weight (considering that of dairy cows) and it also includes more cellulolyt-
ic microbes, therefore their fibre digestion is more effective, and solid feed
particles have longer retention time than in other ruminants. Urea content
in saliva is of the highest content compared to all the other ruminants, so
that is less demanding towards feed protein. Their rumino-hepatic circula-
tion is more effective and nitrogen that get into the circle is less wasted. In
goats, one third of protein requirement can be practically covered by NPN-
sources.
Similarly to dairy cows, production and reproduction cycle takes a whole

year in goats, too. We calculate with 3.5 months of dry period and 8.5
months of lactation period. In their production cycle, there is a strong sea-
sonality yearly. Goats in Europe are inseminated in autumn (September to
October) and they lactate for 1.5 months after, then they are dried off.
Pregnancy takes 153 days and kidding takes place in February to March.
(Therefore if insemination was on 1st Oct., then they are dried off on 15
November. Then they are dried off for 3.5 months and they kid ca. on 1st
March and then thye start to lactate for 8.5 months.)
If there is no synchronization, insemination would take from
September to December. In specialized farms ovulation is synchronized (e.g.
with vagina tampons). Despite in sheep, flushing has no stimulating effect
in goats. Its productivity ranges from of 170 to 200%. Total weight of off-
springs reaches 10 to 12% of live weight of the mother. FIGURE XXVII-1
shows the changes of live weight of goats during pregnancy, drying-off and
lactation.
984
FIGURE XXVII-1: Production and reproduction cycle of dairy goats
In the last two months of pregnancy the increase in live weight, because of
the increase of the foetuses, is quite fast, but dry matter intake shows a
slight decrease. That hides the opportunity of pregnancy toxicity. After kid-
ding, similarly to dairy cows, feed intake can not cover nutrient requirement
because of high level of milk production and ketosis can come up because
of the mobilization of nutrients stored in the body.
27.4.1 Dry matter intake.

Dry matter intake capacity of female goats (40 to 80 kg weight) at mainte-
nance and of male goats for lie weights of 60 to 120 kg can be described by
the equation below:
DMI, kg/day = 0.522 + 0.0135×W
The feeding regimes below are based on the daily nutrient requirements of
a dairy goat of 60 kg live weight and 4 kg milk production with 3.5% milk
fat. The “standard goat” is supposed to have an ingestive capacity of 2.65
UEL with an adjustment of 0.23 UEL for each 1.0 kg difference in 3.5%milk
yield. Based on a diet containing maize silage and lucerne hay with some
concentrate supplementation, feed dry matter intake of does can be accu-
985
rately fitted by the following equation:

DMI, kg/day = 0.062×W0.75 + 0.305×Y
where W = live weight in kg
Y = milk yield in kg/day at 3.5% milk fat
27.4.2. Energy and proteinrequirements

Unfortunately, there is no adapted feed evaluation normative in Hungary for
goats. NRC (1981) gives energy requirements in ME and protein require-
ments in crude protein. INRA (1988) calculates energy requirements in UFL
(unité fourragère lait = net energy for lactation). Protein requirements are
given in PDI (protein digestible intestine), refers to the quantity of protein
absorbed in the small intestine). AFRC (Agricultural and Food Research
Council) (1998) calculates energy requirements similarly to cattle in the sys-
tem of ME and NE.
It is recommended for kids up to 30 days of age and averaging 6.5 kg
live weight hand a daily gain of 165 g, an allowance of 0.8 MJ NE/W0.75.
Dairy goats, such as dairy cows, experience a period of negative energy and
nitrogen balance during their early weeks of the lactation cycle. The efficient
rationing in the dairy goat at this stage of lactation therefore requires a thor-
ough knowledge of the extent and composition of tissue mobilisation.
In the feed evaluation system in the United Kingdom the following informa-
tion are needed to calculate the daily energy and protein requirements of
dairy goat:
-requirements in terms of net energy (NE) and net protein (NP;
-efficiency factors to convert NE and NP into ME and MP,
information above is required for maintenance, milk production, live-weight
change=LWC) and pregnancy (SUTTON and ALDERMAN, 2000).
In the calculations and examples below, the assumptions are those
adopted in AFRC (1998): 65 kg Saanen-type multiparous goat; milk con-
tains 37 g fat and 29 g total protein (26 g true protein)/kg; pregnant with
twins; later in the lactation cycle the diet has a metabolisability, “q” value
(ME/GE) of 0.6 equivalent to 11.3 MJ ME/kg DM. It was decided that the
efficiency factors calculated for sheep and cattle for converting ME and MP
to NE and NP, respectively in ARC (1980) and AFRC (1992).could be applied
to goats. These conversion factors (“k” value) are given in terms of energy
and protein as follows respectively:
-for maintenance 0.71 and 1.0.
986
-for milk production 0.63 and 0.68

-for live weight gain 0.6 and 0.59
-for pregnancy 0.13 and 0.85.
The INRA (UFL system uses almost identical efficiency factors to con-
vert dietary ME values to NE lactation values (NEl) and then to UFL as kg
of barley. MAINTENANCE ENERGY REQUIREMENTS (for the ‘standard goat’
described above) can be calculated to be 9.9 MJ ME/day from INRA (1988)
and 10.2 MJ ME/day from AFRC (1998). Both INRA and AFRC give similar
estimates of the maintenance requirements as 53 g PDI or MP per day.
Estimates of maintenance requirements in AFRC (1998) and INRA 1988)
include a factor for activity estimated for housed goats (Table XXVII-1).
Maintenance requirements. Table XXVII-2 includes the requirements of

nutrient of goats for maintenance of different live weights.
987
In the last two months of pregnancy the increase in nutrient require-

ments can be neglected. During the last two months of drying off period
total live weight increases significantly. In this period nutrient requirement
of the foetus should be considered. Because of the steady growth of the foe-
tus a decrease in dry matter intake can be experienced in the last 2 to 6
weeks of the dry period because the foetus increases and the volume of
digestive tract decreases. For that reason, pregnancy toxicity can occur. To
avoid that, we have to consider the increased requirements of energy and
protein, and in the last month of pregnancy, proportion of UDP also has to
be increased. It can be explained by the relative energy deficiency which
decreases the rate of microbial protein synthesis. When UDP requirement is
met, it is to be considered that goats have a highly developed taste, there-
fore it is advised to give extra feeds (e.g. gluten feeds) of the best quality
mixed with taste and aroma additives (Table XXVII-3).
To summarize the specialities of nutrient supply in pregnant goats, it

can be stated that during drying-off taking up surplus feedstuff involves no
danger, as e.g. in dairy cows. For foetus development, 25 to 50% of extra
energy can be delivered without danger of getting fat. Since in the last 2 to
6 weeks of pregnancy, restricted energy supply decreases the efficiency of
microbial protein synthesis, a part of daily crude protein supply should be
of bypass (undegradable) character. It was assumed that goats lose 1.0
kg/week for the first 4 weeks of lactation but none in weeks 5-8 compared
with losses of 0.5 kg/week adopted by INRA (1988). A summary is set out
in Table XXVII-4 giving the estimates for the minimum requirements with no
safety margins added by AFRC (1998), whereas INRA (1988) define their
988
estimates as allowances, not minimum requirements.
Both types of ketosis (pregnancy toxicity before kidding, and ketosis

of freshly kidding goats of high milk yield, respectively) can occur. At the
same time, opportunity of their occurrence is lower compared to mother
ewes and to dairy cows as well.
REQUIREMENTS DURING LACTATION: a dairy goat of 60 kg live weight pro-
duces 4 kg milk with 3.5% milk fat, which means 700 to 1000 kg of milk
production in the whole lactation period of 8.5 months. To meet its require-
ments, a nutrient supply is needed which is 3 to 4 times the daily require-
ment of the maintenance level. (It can be considered the same as the daily
nutrient requirement of a dairy cow producing 30 to 40 kg milk daily!) Table
XXVII-5 includes daily nutrient requirement data of does and dairy cows
compared.
989
Besides the above figures, either 25% or 50% or 75% of the daily nutrient
requirement is to be calculated as an extra for the grazing activity depend-
ing on the pasture quality and on the geographical characteristics (an extra
70% is considered for hill-side pastures).
By compilation SAHLU et al. (2004) developed a metabilizable energy-
metabolizable protein feed evaluation system for goats. For values a 15%
difference in maintenance energy requirement between intact males com-
pared with females and males castrates or wethers was assumed (NRC,
2006).
-Growing intact males (dairy)
MEm= 0.624 × LW0.75 MJ ME
-Growing females and wethers (dairy):
MEm= 0.537 × LW0.75 MJ ME
-Mature intact male (dairy):
MEm= 0.576 × LW0.75 MJ ME
-Mature females and wethers (dairy)
MEm= 0.501 × LW0.75 MJ ME
Table XXVII-6 and Table XXVII-7 comprise the nutrient requirements of a
modell doe, growing kids and buck, respectively, using the compilation of
NRC (2006).
990
991
992
27.4.3. Mineral requirements

27.4.3.1. Macroelement requirements. In most respects, Ca and P metabo-
lism in goats appear to be similar to that in cattle and sheep. Goats can
utilise inorganic sulphur, possibly through the microbial synthesis in the
rumen. It was reported that calcium sulphate supplementation not
increased only mohair production, but milk production in Alpine goats as
well, when dietary sulphate concentration was increased from 0.16% to
0.26%. The calcium to phosphorus ratio in the feed should be 2:1. Trace
mineralised salt may be fed for trace mineral supplementation at 0.5% of
grain mix. The estimated Ca and P requirements of goats with different ages
and production periods are given in Table XXVII-8.
27.4.3.2. Microelement requirements. Magnesium deficiency (hypomagne-

saemia) in goat can cause tetany and hyper-irritability, while excess Mg may
depress Ca absorption. ARC (1980) suggested that the value of 3.0 mg of
Mg/kg live weight is recommended daily. Using a value for Saanen-type
goats, Mg requirements for milk production are 0.43 g Mg/kg milk.
The metabolism of Na, K and Cl is closely related. Deficiency symp-
toms include depraved appetite, depressed fed intake, emaciation and
reduced milk and mohair production. Goats appear able to adapt to wide
variations of salt (NaCl) providing that drinking water is freely available. It
has been suggested that diets for goats should supply 5 g NaCl/kg DM and
5 g K/kg DM for growing and for lactating goats. It is well-established that
complex interactions occur between copper and a number of other ele-
ments. The maximum dietary copper level of 20 mg Cu/kg DM should be
taken into account. A maximum dietary Cu level of 20 mg Cu/kg DM is to
993
be followed as a toxicity guideline. At least 50 mg Zn kg/DM is advised

which can be possibly be increased up to 80 mg Zn/kg DN for breeding
females or in the presence of dietary Zn-antagonists. It is suggested that a
maximum of 8-10 mg I/kg DM can be regarded as a safe dietary level in cat-
tle and sheep, which should be also be adopted for goats. Recommended
manganese requirement will vary considerably (60 to 120 mg Mn/kg DM)
according to dietary interactions. A guideline recommendation for copper
and some trace elements are given in Table XXVII-9.
27.4.4. Vitamin requirements.

The apparently low incidence of the observed vitamin deficiencies may arise
because in many areas goats are not kept intensively, they have a relative-
ly low level of production and their choice of feeds is wide. Rumen microbes
can synthesise all the water soluble vitamins required by goats. They
include the B-vitamins and vitamin C and also the fat-soluble vitamin K,
but they require dietary sources of vitamins A, D3 and E. Young kids
require the vitamin B complex in their diet until their rumen develops.
Therefore goat feed formulators are only concerned with adding vitamins A,
D and E to goat feeds. Table XXVII-10 gives the mineral requirement of a
model, growing kids.does and bucks, respectively, using the compilation of
NRC (2006).
994
995
Water requirement. The goat’s water houshold is very economic and

their heat economic is better than that of sheep (because of less tallow and
hair production. In goats water turnover per unit of body weight approach-
es that of camels. Lactating goats should consume 2.8-3.8 litres of water per
each kilogram of milk produced.
Nubian goats can produce 1 kg milk from 1.5 kg water, while dairy
cows needs 5 kg water to produce 1 kg milk.
27.5. Feeding the kids
A variety of rearing systems is used for goats including natural rearing, par-
ticularly for fibre-producing breeds, artificial rearing of herd replacements
and various systems for meat kids. Naturally reared kids are normally left
with their dams until about 1 month before the start of the normal breed-
ing season, which generally means that the kids are 12 to 16 weeks of age
(AFRC, 1998).
A variety of regimes can be used as it is demonstrated in Table XXVII-
11. The first three days after birth are the most critical days in the life of a
newborn kid, so it is essential to allow them to suckle in that period to
receive colostrum. However, in some herds caprine-arthritis encephalitis
(CAE; see Metabolic disorders and nutritional diseases section) is a concern,
and kids from those herds must be bottle-fed heat treated colostrum instead
of nursing their mothers. Feeding milk or milk replacer could continue until
the kids are of 8 to 12 weeks. Some milk substitutes are produced specifi-
cally for kids, but milk replacers made for calves or for lambs can also be
996
fed. For dairy production, it may be more economical to separate the kids
from their mothers and to feed them with milk replacer. 1.5 kg per day fed
two times per day.
These mainly comprise skim-milk powder with some added fat,
although some contain whey powder with added soya protein. In general,
crude protein levels are within the range of 220 to 360 g/kg and ether
extract content varies from 120 to 260 g/kg. The recommended concentra-
tion of milk powder in the liquid also varies from 125 to 200 g/kg for dif-
ferent products, and the recommended feeding temperature ranging from
ambient temperature to 42°C. Rearing kids on automatic machines may
result in excessive drinking, which causes diarrhoea. Once kids reach 3 to
4 weeks of age, they seem to be able to cope with this challenge. If milk sub-
stitutes are offered ad libitum, drinking from teats fixed directly to the milk
container seems to give the best results.
In all rearing regime kids should be encouraged to eat solid feed, such
as calf or lamb rearing concentrates and good hay from 2 to 3 weeks of age.
Weaning is a critical phase in kid management. Kids can be weaned by age
and/or weight provided an adequate amount of solid feed is being con-
sumed. Although goat kids can be weaned as early as 4 weeks of age, it was
found that weaning at 8 weeks was optimal (PINKERTON et al., 1993).
Following weaning, goat kids should be fed and managed to achieve daily
gains of 0.10 to 0.15 kg. Such gains would allow the preferred breeding
weight of 30 to 32 kg to be reached by 28 to 32 weeks of age. A typical post-
weaning feeding program continues the kid starter offered prior to weaning
changing to kid grower at about 16 weeks, and then to a gestation diet at
about 3 weeks. When the feeding of milk replacer ends, the young goats
should be able to consume 0.25 to 0.5 kg of a kid starter grain mix daily.
Typical kid starters contain 16% crude protein and 11% crude fibre. Good
quality hay can be fed to kids beginning at one week of age to develop rumen
function. Most cashmere and Angora kids kept in naturally rearing man-
agement systems akin to those pertaining in sheep flocks. Weaning nor-
mally takes place at about 14 weeks of age. Although most male kids are
generally weaned no later than 12 weeks, by which age they can be sexual-
ly mature.
Bocks are slaughtered at a very early age (at 1 to 2 month of age).
During this period of time feeding efficiency is 1.2 to 1.4. Very seldom they
are slaughtered at the age of 3 to 4 months with a live weight of ca. 20 kg.
997
Female kids are not slaughtered at an early age. Female breeding animals
are inseminated already in the first year. The goal is that they reach 55 to
55% of their adult live weight (30 to 35 kg) by that time (until autumn).
The aim is to mate kids at 7 months of age to kid at 12 months. To achieve

this, the weight of kids at mating should be at least 60% of theadult weight,
i.e. 40 to 45 kg for Saanen-type goats require a relatively high pre-mating
live weight gain of about 150 g/day. Yearling goats require nutrients for tis-
sue maintenance and growth. Kid starter can be fed to goats at 3-6 months
of age at a rate of 0.25-0.5 kg per day along with good quality forage and
pasture. Silage and NPN-sources should not be fed to goats at this age.
When goats are 6-9 months old, they require 0.5-0.75 kg of grain mix with
good quality hay.
Feeding after mating must allow for continued growth as well as for
development of the foetus. It is preferable that these aims are achieved
mainly by using high-quality forages rather than by excessively relying on
starchy concentrates which can cause over-fattening.
The actual daily quantities of protein and total digestible nutrients
(TDN) needed per goat daily in a range of body weight categories are shown
in Table XXVII-12 as practical feeding guidelines.
MANAGING BODY CONDITION. As the breeding season approaches producers

should be concerned with the body condition of their breeding does. (FIG-
URE XXVII-2) Failure in reproduction, low twinning rates and low weaning
998
rates will result if does are too thin. Overly fat does can suffer pregnancy
toxaemia, but fat does are rarely a problem. The term body condition refers
to the fleshing or fatness of an animal. Since goats are as far as possible
grazed, the 1 (extremely thin and weak) to 5 (extremely obese) point scale
can be advised to be applied as in beef cattle breeding practice. The body
condition score (BCS) just before the breeding season is between moderate
condition (3) and good condition (4) to maximize the number of kids born.
Pregnant does should not have a BCS of 3.5 or above towards the end of
pregnancy because of the risk of pregnancy toxaemia. In addition, a body
condition score of 3 to 4 at kidding should not drop too quickly during lac-
tation. Body condition score is also used to determine whether flushing will
be of benefit to breeding does. Flushing means increasing the level of feed
offered to breeding does, mostly energy, starting about one month prior to
the introduction of the bocks to increase body weight, ovulation rate ad
hopefully litter size.
FIGURE XXVII.2: BCS at goats

PREGNANCY. In most cases, it is a goodpractice to dry off dairy goats at about
6 weeks prior to kidding. During this period the foetus gains 70 percent of
its birth weight. Good pasture or quality hay will maintain the pregnant doe
during this period. If the doe is thin, supplementation of 0.2-0.5 kg of grain
mix is warranted. Four to five days before kidding, the grain allowance
999
should be decreased by substituting 50% of it with bran. By eliminating

much of the bulk in the digestive tract kidding is facilitated. Goats should
kid with adequate fat reserves to sustain high yields in the early stages of
lactation. It is recommended therefore that a suitable feeding regime in the
last third of pregnancy should be based on high-quality forages available ad
libitum with concentrates increasing to about 0.5 kg/day until parturition.
MILK PRODUCTION
Lactation cycle of goats is very similar to that of cows (INRA, 1988). Milk
yield reaches a maximum at 1-2 months after parturition and then declines.
Live weight falls rapidly to a minimum about 6 weeks after parturition and
then slowly recovers until a rapid increase in the last 3 months of pregnan-
cy. Feed intake increases more slowly than milk production and declines
slowly until the third month of pregnancy when it stabilizes. Dairy goats
experience a period of negative energy and nitrogen balance during the early
weeks of lactation cycle. After parturition, milk yield increases rapidly to a
peak (6 to 8 weeks after parturition in most breeds) and then declines slow-
ly. Goats are normally dried off after about 10 months of lactation, in prepa-
ration for kidding at 12-month intervals.
INRA (1988) suggested that goats lose about 1 kg adipose tissue per
week during the first month post-partum and 0,5 kg during the following
month. From about the fourth month of lactation, dairy goats begin to
regain their live weight. INRA (1988) recommended a target of 1.2 kg live-
weight gain per month for multiparous goats and 2.2 kg per month for prim-
iparous. Since peak feed intake normally occurs later than peak milk yield,
as in dairy cows, it is important that the feed offered during this period
should be of high quality, though not too rich in concentrates.
Milk yield is highly correlated with ME intake with the correlation
increasing as lactation progresses. Milk yield responses to extra concen-
trates in goats consuming various forages ad libitum vary from 0.4 to 1.6 kg
milk/kg concentrate dry matter. In some dairy breeds, intake of hay
increased when the CP in the concentrates was increased to about 180g/kg
DM in early to mid lactation, but they did not respond to higher CP con-
centrations.
1000
27.8. Nutrition and feeding of breeding bucks
Except one month before and after insemination period, and during the lat-
est period (from September till December in Europe) of pregnancy, it is suf-
ficient to deliver a maintenance level for breeding bocks. In the above men-
tioned period an extra 50 % nutrient of maintenance level should be deliv-
ered.
Bucks require more forage than does. Additionally, they should
receive 0.5 to1.0 kg grain daily. Overweight bucks make inefficient breed-
ers, so it is important to have the bucks in good physical conditions, but
they should not be fat or have excessive fleshing.
27.9. Feeding strategies in goats
27.10.1. Grazing. Saanens producing 1 to 3 litres milk per day increase

their apparent grass intake and milk production with pasture allowances up
to 8-9 kg DM per head per day in early lactation and up to 6-7 kg DM per
head per day in late lactation. Herbage intake is estimated to be 3.0 to 3.3
and 1.8 to 2.0 kg DM/head per day, respectively.
27.10.2. Alternative choice of feeds

Forage. Generally the level of forage intake is higher when goats have plen-
ty of time for choosing the most nutritive parts of forage. The nutritive value
of really ingested forage is improved and always higher than that of given
forage. Consequently, one should pay attention to the variation in the qual-
ity of the given green or conserved forages and to separate the different cuts
according to their quality. If the hay is of poor quality, it should be distrib-
uted in a large excess to encourage intake.
Green fodders. Generally the level of grass intake is higher on zero-grazing
than on pasture provided that forage is fed ad libitum and feeding rack is
open all the time. The level of intake of legumes is higher than that of grass-
es. Preserved forages. The voluntary intake of silage is generally lower than
that of hay from the same grass. Wilted silage (haylage) can also be more
efficient than unwilted silage for lactating goats. On the other hand, the risk
of listeriosis is very much increased when the diet contains silage, particu-
larly maize silage because goats are every sensitive to this disease. Cereal
1001
straws (wheat, barley or oat) are accepted by goats in limited quantity (15-
30 g/kg W0.75), and they can not meet maintenance requirement. Roots and
tubers. The proportions of roots and tubers must be limited in goat diets
amounting to about 30-50% of the total dry matter and should be fed
together with forages.
Concentrates. Goats can be given a wide range of cereal grains, oil
seeds, legume seeds or even industrial by-products. Rolled seeds or pellet-
ed meals are more accepted by goats than whole seeds or finely ground
meal. Does in late pregnancy and in early lactation are especially demand-
ing towards the quality of concentrates.
27.10. Frequently used feeding systems of dairy goats
Feeding based on hay

-1.3 kg grass hay
-5.5 kg wet sugar beet slices
-0.2 kg oat or barley
-0.3 kg soybean or sunflower solvents
It is a well established feeding strategy in goats, but not in dairy cows, that
roughages are given in two rations and the concentrates are fed at the milk-
ing stand.
Monodiatetic feeding based on silage

-5.0 kg maize silage
-0.5 kg alfalfa hay
-0.6 kg dried sugar beet slices
-0.3 kg soybean or sunflower extract
NDF (neutral detergent fibre) of dry sugar beet slices are especially well uti-
lized – because of long retention time- in goats.
Feeding based on cut grass

-13.0 kg cut pasture grass
-0.45 kg maize meal
-0.15 kg field peas or sweet lupines
In all the above cases feed salt and minerals and vitamins are to be
1002
supplemented separately or mixed in the concentrate. The suggested gross

quantities include ca. 20% extra nutrients compared to net requirements.
That is required because goats need to choose, as ca. 20% of daily ration
remains.
Silage or haylage can also be fed to goats. However, silage should not
be fed to goats younger than six months of age. In addition to forages and
concentrates, goats have a preference for hay or for pasture grass.
Rangeland pasture can be made complete with mixture of corn, cottonseed
meal, fish meal and molasses. The supplementation significantly increased
body weight gain and production of clean fleece. Higher rumen undegrad-
able protein content of corn, cottonseed or fish meal supplement may pro-
duce significantly longer mohair of goats.
Table XXVII-13 includes the ingredients of different concentrates
according to the various daily protein concentrations of the daily diets.
A wide range of diseases can affect goats, and dairy goats in particular, and
herd health programmes should be implemented to prevent these. However,
goats kept extensively in communal grazing areas may be remarkably free
of internal parasites and diseases. Management and husbandry are partic-
ularly important during kid-rearing in intensive systems. Some diseases will
have little effect on herd productivity, such as isolated cases of carcinoma
or a more general effect, such as with pneumonia, or a specific effect on fer-
tility, such as toxoplasmosis, or an effect both on goats and humans, such
as brucellosis.
1003
ABORTION
Goats are animals „dependent on luteum”, and therefore they react easily to
external influences with abortion.
Stress abortion: in case of energy deficiency, glucose level of does declines,
which results in abortion due to stress. It is typical in the 90-110 days of
pregnancy, and newborn kids are alive in this case. Direct reason for this is
that the steroid production of the adrenal cortex in kids are still oestrogenic
and induces abortion.
Habitual abortion: reason for this is that the mother suffers from hormonal
imbalance and the aborted foetus is usually dead, macerated, and its occur-
rence can not be connected to an exact period as in the previous case.
Deficiency of vitamin A, iodine, or copper can also cause abortions. Parasites,
certain drugs, poisonous plants, and stress can also cause a doe to abort.
If abortion is widespread in the herd, there is most likely to be an infectious
cause. Goats will suffer from iodine deficiency if they are not supplemented
in iodine-deficient areas. The iodine needs of pregnant does are the highest
(0.8 ppm DM of feed) and the most common manifestation of iodine defi-
ciency is abortion or stillborn kids. Weak kids will have enlarged, goietered
thyroid glands. The kids may have a reduced hair coat, but the most obvi-
ous signs are the large bilateral swellings on the neck. The thyroid gland
may be several times the normal size. If iodinized salt is already fed, the
needs may be increased if the does are grazed on brassica plants (turnips,
cabbage, forage rape) while they are pregnant.
For the differential diagnosis: Chlamydia psittaci is the most common
cause of infectious abortions. Toxoplasmosis is another major cause of abor-
tion in goats. It is contracted by goats ingesting cat faeces. Certain additives
(monensin) can help prevent abortions due to toxoplasmosis.
ACUTE ENTEROTOXAEMIA
If too much concentrate is fed after kidding, high level of lactate is produced,
and the pH of rumen drops to 4.8, and acute lactate toxicity occurs. With
the aid of Clostridium perfringens C or D enterotoxaemia develops. To avoid
this, daily ration has to be fed in some smaller portions.
ARTHRITIS
Diets containing excessive calcium have been associated with calcification
1004
of periarticular tissues, ankylosis, and a stiff gait. The pathogenesis is con-

sidered to be hypercalcitonism induced in non-lactating animals by dietary
calcium levels appropriate for lactating animals (SMITH and SHERMAN, 1994).
To avoid both metabolic bone disease in bucks and non-lactating pet goats,
dietary calcium should be limited to 4 to 6 grams daily for mature animals
weighing 70 kg.
COCCIDIOSIS
Coccidiosis is often considered to be a disease of intensification, affecting
goat kid particularly; however it may also occur under more extensive con-
ditions. It is a disease resulting from infection of the intestinal tract by par-
asitic protozoa called coccidia, causes scours (diarrhoea) in goats, particu-
larly in kids. Coccidiosis occurs in damp, crowded areas. Keeping kids away
from those areas prevents serious problems. Animals gain immunity against
this organism by nine months of age, and clinical disease rarely occurs in
adult animals (HALE and WELLS, 2004).
KETOSIS OR PREGNANCY TOXAEMIA

This is a condition of late pregnancy and early lactation most commonly
occurring in the last six weeks of gestation in does with multiple foetuses
and in the first 4 weeks in heavily lactating does (MENZIES, 1998). It is typi-
cal that not so much keton bodies as levels of non-esterified free fatty acids
in particular (NEFA) increase. It occurs both before and after parturition.
Ketosis sometimes occurs in late pregnancy in goats carrying multiple foe-
tuses. The disorder is the result of a discrepancy between glucose demand
of the foetuses or that of milk production and glucose availability from the
diet resulting in body lipid mobilisation to increase glucose supply. The pri-
mary cause is a deficit in energy intake as in cattle and sheep.
Factors that predispose does to develop pregnancy toxaemia can be
divided into two types: inadequate nutrition and adequate nutritional sup-
ply while external or animal factors (e.g. disease) affecting intake. To coun-
teract insufficient feed intake the ration must be of high energy and protein
content (e.g. 35% grain to 65% forage). Does that are very thin (BCS<2.5)
are at increase risk despite a good ration. Very fat does (BCS>4.0) will use
their body fat reserves in late gestation having decreased voluntary intakes.
All this will predispose them to formation of keton bodies that further sup-
presses appetite. Does decrease grain intake, which is followed by decreased
1005
intake of silage and forage. If these signs go unnoticed, the does may go on
to exhibit neural disorders, which include abnormal gait and stance, appar-
ent blindness, stargazing and necrosis of the brain cortex secondary to pro-
longed hypoglycaemia (low blood glucose).
Prevention of ketosis is normally a matter of reformulating the diet to
increase daily energy intake by increasing the proportion of concentration
in the diet. For prevention and therapy, Na-propionate and propylene glycol
(600 mg/ml) at a rate of 60 ml/bid per os for a minimum of 3 days and chlo-
ral-hydrate (NRC, 1981) are appropriate additives. Improved nutrition and
feeding management are needed. Correction of ketoacidosis using bicar-
bonate or bicarbonate precursors might be necessary.
Most changes are attributable to primary hypoglycaemia resulting
from the failure of nutrient intake to meet the combined needs of the doe
and the foetuses or of the doe’s milk production. Since hypocalcaemia is
often a secondary disease associated with pregnancy toxaemia, clinical
signs of hypocalcaemia should be evaluated. This is a disease that needs to
be prevented rather than treated. If one doe is clinically ill many more in the
herd are likely to be at risk.
LACTIC ACIDOSIS
If the proportion, absolute amount or type of grain changes too quickly,
then lactic acidosis will develop. Animals fed on diets with little fibre or
chopped too finely are more at risk of lactic acidosis. Chronic feeding prob-
lems will manifest as variable appetite, depressed milk fat and chronic
laminitis. Milk fat is depressed because of the lack of adequate quantity of
fibre in the ration. With more severe lactic acidosis, the protozoa die, the
rumen becomes static and the goat becomes depressed and dehydrated.
Diarrhoea smells acidic and is yellow in colour. In very severe cases, there
is no diarrhoea because of total gut status. The goat may appear “drunk”
and ataxic
Supportive therapy includes iv. fluids, rumen transfaunation (rumen
juice from a healthy animal), alkalizing solutions for the rumen, antibiotics
and nursing care. Forage should be fed before grain and the daily amount
divided into at least 3 separate feedings. A total mixed ration (TMR) helps to
keep the rumen flora supplied with carbohydrates constantly.
1006
PARTURIENT PARESIS
Mechanism of development is similar to that in dairy cows, but rate of its
occurrence is lower of the PERIPARTURIENT HYPOCALCAEMIA (milk fever). Goats
are less susceptible to hypocalcaemia or „milk fever” than cattle at compa-
rable level of production. Both Ca intake and Ca/P ratio should be con-
trolled in late pregnancy to reduce this risk. Hypocalcaemia is usually seen
in high producing dairy goats one to three weeks post-kidding and is much
rarer than pregnancy toxaemia. Initially the doe is ataxic, nervous and
hyperactive but quickly becomes sternally recumbent. The doe stops eating
and the ears are cold. The course of disease can be as short as a few hours
and occasionally may occur as “sudden death”. Serum calcium levels are
deceased, usually less than 1.7 mmol/litre (normal 2.1-2.8 mmol/litre).
Clinical cases of hypocalcaemia are usually treated with calcium boroglu-
conate solution (20 mg Ca++/ml) iv. or sc.).
Long-term undernutrition is required for PRIMARY HYPOCALCAEMIA to
develop. Goats require calcium rich diets after kidding. Alfalfa hay can pro-
vide this. Cereal crop forages such as wheat or oat hay/straw should be
avoided in unless the ration is balanced with other calcium sources. The
ration in late gestation and in early lactation should have calcium:phos-
phorus ratio greater than 1.5 to 1. Prevention of pregnancy toxaemia will
also help to prevent hypocalcaemia as well.
POLIENCEPHALOMALACIA
This a neurological disease caused by real or relative thiamine (vitamin B1)
deficiency. Kids or does on high carbohydrate diets may have their normal
rumen flora upset. A change in bacterial types may cause either deficiency
of thiamine or production of an enzyme which inhibits thiamine activity.
The end result is the disease poliencephalomalacia (softening and necrosis
of the grey matter of the bran).
POSTHITIS (PIZZLE ROT)

In case of overfeeding of protein (nitrogen), urea content of urine increases.
Urea of urine staying in praeputium is depleted by Corynebacterium renale,
and it causes local inflammation after its proliferation. The prepuce
becomes ulcerated, swollen and very painful. Treatment consists of diet
change to lower protein rations (e.g. 12 to 14%).
1007
UROLITHIASIS
Bucks and wethers are prone to urinary tract blockage due to urinary cal-
culi (stones). The most common types are calcium phosphate and struvite
(magnesium phosphate) from high grain diets rich in phosphorus but defi-
cient in calcium and potassium. Sand may becomes blocked anywhere but
most frequently is at the urethral process. If not noticed and blockage is
total and the bladder ruptures in 24 to 36 hours. After rupture, the
abdomen swells with urine and the goat appears more depressed. This con-
dition is called “water belly”. Male kids are highly susceptible to this disease
if they are fed high cereal diets, requiring the addition of buffers to prevent
them going off feed with acidosis.
The addition of Ca to maintain a Ca/P ratio of 1.5 to 2:1 in the diet
and the avoidance of any Mg compounds is effective in preventing the occur-
rence of this disorder. Common salt (NaCl) should be included at 1% of total
dry matter intake. Plenty of fresh, palatable water should be available. Diets
high in potassium should be avoided. Vitamin A requirements should be
met (good quality green hay and pasture will do this). Acidification of urine
(by mixing of ammonium-chloride and potassium-chloride in the ration at
½% of dry matter intake) assists recovery and therapy since it dissolves the
remaining stones.
WHITE MUSCLE DISEASE

Selenium deficiency may cause acute muscle necrosis known as white mus-
cle disease. Usually young fast growing kids are affected anywhere from
birth to full maturity. Kids are acutely painful, reluctant to move, but may
still eat. It has been shown that selenium supplementation helps with cell
mediated immunity.
Feed supplementation should be done with caution. Selenium can be
added to feed supplements and premixes to a maximum of 0.3 ppm.
MISCELLENOUS
Against toxic plants, goats are usually resistant. For example, grazing of
Hymenoxys odorata causes severe mortality among sheep, but goats toler-
ate this well. Goats can also tolerate tannin in plants well, probably because
of its efficient degradation in the rumen.
1008
FOR FURTHER READING
Agricultural Research Council, ARC (1980): The nutrient requirements of farm livestock. No
2. Ruminants. London
Agricultural and Food Research Council, AFRC (1998): The Nutrition of Goats, CAB
International, Wallingford
Donkin, E.F. (1997): Productivity and diseases of Saanen, indigenous and crossbred goats
on zero grazing. Ph.D. Thesis. Medical University of Southern Africa. p. 18
Drochner, W., Flachowsky, G. Pallanf, J., Pfeffer, E., Rodehutscond, M. and Staudacher,
W. (2003): Recomendations for the supply of energy and nutrients to goats. DLG
Verlag. Frankfurt am Main
Hale, M. and Wells, A. (2004): Goats: Sustainable production overview. ATTRA
Publications, No. IP248. 13.
INRA, (1988): Alimentation des Bovin, Ovins et Caprins, INRA, Paris.
INRA (1998): Alimentation des Bovin, Ovins et Caprins. INRA, Paris
Menzies, P.I. (1998): Metabolic and nutritional diseases. Ed.: Ministry of Agriculture, Food
and Rural Affairs, Ontario, Canada, p.1.
Morand-Fehr, P., Sauvant, D. and Brun-Bellut, J. (1987): Recommendations alimentaires
pour les caprin. Bulletin Technique, C.R.Z.V. Theix, INRA, 70. p. 213-222.
NRC (1981): Nutrient requirements of domestic animals, Nutrient requirements of goats.
National Research Council, National Academy of Science. Washington, DC
NRC (1989): Nutrient requirement of domestic animals. Nutrient requirement of dairy cat-
tle. National Research Council. National Academy of Science Washington DC
NRC (2006): Nutrient requirements of small ruminants. Sheep, goats, cervids, and new
world camelids. National Research Council, National Academy of Science.
Washington, DC
Pinkerton, F., Escobar, N. and Pinkerton, B. (1993): Feeding and management of goat kids.
Fact Sheet, Kika de la Garza Institute for Goat Research, Oklahoma, pp 1-10.
Sahlu, T., Goetsch, A. l., Luo, J., Nsahlai, I.V., Moore, J.E., Galyean, M.L., Owen, F.N.,
Ferrell, C.E. and Johnson, Z.B. (2004): Nutrient requirements of goats: developed
equations, other considerations and future research to improve them. Small
Ruminant Res. 53, 191-220.
Smith, M.C. and Sherman, D.M. (1994): Goat medicine. A Waverly Company. p. 555.
Solaiman, S.G. and Castaldo, D.J. (1994): Feeding programmes for goats, Feed Intern. May,
p. 28.
Sutton, J. D. and Alderman, G. (2000): The energy and protein requirements of pregnant
and lactating dairy goats. The Agricultural and Food Research Council Report,
Livest. Prod. Sci. 64, 3-8.
1009
Chapter XXVIII
PRODUCTION FISH NUTRITION AND FEEDING
© Åshild Krogdahl
28.1. Physiological characteristics of importance for nutrient require

ment and feeding
28.1.1. Species diversity
28.1.2. Digestive organs and digestion
28.1.3. Excretion of water soluble compounds and water replace
ment
28.1.4. Energy and protein requirements for maintenance and
growth
28.1.5. Specific requirements of vitamin C, long chain fatty acids
and astaxanthin
28.1.6. Nutrition and disease defence systems in fish
28.2.1. Minerals
28.2.2. Oxygen
28.2.3. Temperature, growth potential and feed intake
28.2.4. Other important environmental interactions
28.2.5. Fish, at least coldwater species, show increased growth in t
28.3.1. Feedstuffs
28.3.2. Technical requirements of fish feed
28.3.3. Feeding regime
28.5.1. General
28.5.2. Deformities
28.5.3. Cataract
28.5.4. Thiamine deficiency
28.5.5. Nutrient to energy imbalance and fat accumulation
28.5.6. Carbohydrate excess
28.5.7. Iron excess
28.5.8. Mycotoxins
1011
CHAPTER XXVIII: PRODUCTION FISH NUTRITION & FEEDING
ish cultivation has long traditions in several countries and dates back
F many centuries in Asia. Cultivated fish supply essential nutrients for

many human populations. During the last decades fish cultivation
has become a world wide industry representing important export income
also in countries without long traditions in fish cultivation. The number of
species under cultivation is countless and great research efforts are made
to bring new species under cultivation. The present chapter regards fish
species of importance for intensive cultivation and for which there is a body
of scientific literature to build on. As intensive fish cultivation has become
of significant economical importance quite recently, fish nutrition research
is also fairly young compared to nutrition of other domestic animals. Our
knowledge regarding fish nutrition and feeding is far from complete and for
many cultivated species it is quite insufficient. The situation is best for
salmonids. Also for catfish, basses, carp and some other species knowledge
on nutrition is sufficient enough to form a basis for development of efficient
diets and feeding strategies. The information below reflects the fact that
research on salmonids has supplied a larger share of our knowledge in fish
nutrition.
In fish cultivation the animals are in most cases kept in large groups.
The animals are relatively small and the individual fish is difficult to observe
without great disturbances of the whole group. Feeding and disease treat-
ment on an individual basis is therefore not feasible. Preventive strategies
are imperative in aquaculture medicine. Nutritional disorders may be pre-
vented through careful consideration of diet composition and conductance
of appropriate feeding routines according to the changing nutrient and ener-
gy needs of the fish throughout their life stages. Optima regarding feeding
frequency, timing, pellet size, etc., change depending on species and devel-
opmental stage. If a nutrient deficiency develops, the likelihood of correct-
ing the situation depends on the nutrient in question and stage of develop-
ment. However, in most cases, the situation becomes apparent only after
severe symptoms are visible, such as deformities, cataracts, and reduced
disease resistance. At this time the situation may be difficult to correct in
all fish.
The diet in many disease situations, e. g. when contagious diseases
strike, it is the only possible vehicle for medication. However, as diseased
1012
CHAPTER XXVIII:PRODUCTION FISH NUTRITION & FEEDING
fish often eat less or not at all, the healthy fish get more medicine. Moreover,
diets with medicine are less palatable. The result is inefficient medical treat-
ment and low growth in all fish.
28.1. Physiological characteristics of importance for nutri-

ent requirement and feeding
Cultivated fish differ from traditional domestic animals in several ways that
affect nutrition and feeding. The most relevant of these characteristics are
presented below.
28.1.1. Species diversity

Fish are classified as herbivore (carp and milkfish), omnivore (channel cat-
fish and tilapia) and carnivore (salmonids basses, breams, flounders and
groupers) according to the ingredients of their natural diet. Some species
feed on dead, others on living materials, some feed solely on microorgan-
isms, others on larger plants and animals, and some are scavengers eating
whatever they can find. Interestingly, species that have a similar dietary
selection may show great variation in intestinal anatomy. Even within
species we find differences between developmental stages. The dietary selec-
tion of fish larvae is less complex than that of adults.
28.1.2. Digestive organs and digestion

The physiological diversity of the digestive tract among cultivated fish
exceeds that of domesticated homoeothermic animals. Most species start
out as carnivores and have straight simple gastrointestinal tracts at hatch-
ing. Through the larval and juvenile stages the GI tract develops into more
complicated structures. Some fish continue to have short, relatively simple
gastrointestinal tract, others long, some are stomach-less, some have sev-
eral hundred pyloric appendages, and some have digestive organs that may
ferment dietary fibre components (FIGURE XXVIII-1). Fish do not have a
colon similar that of homeothermics. The distal intestinal structures usual-
ly differ from the more proximal compartments and may be very complex.
Their functional morphology and biochemistry indicate that, not only water
and minerals, but also protein, lipids and carbohydrates may be hydrolysed
and absorbed in this region. Absorption of macromolecules also takes place
in this region, which seems to be of great importance in antigen presenta-
tion and immune function.
1013
FIGURE XXVIII-1: Diagrammatic representation of typical digestive configurations. (a)

Euryphagous carnivore with a y-shaped stomach (salmon, trout, lingcod, sablefish, and
halibut). (b) Euryphagous omnivore empha sizing animal sources of food: pouched stom-
ach or intestinal sac (catfish and tilapia). (c) Euryphagous omnivore emphasizing plant
sources of food: stomach absent (carp and goldfish). (d) Stenophagous planktivore; tubular
stomach with muscular gizzard (milkfish). (From SMITH, 1989; presented in HALVER and
HARDY, 2002).
Present knowledge indicates that, qualitatively, digestive processes in

fish are very similar to processes in homoeothermic monogastrics. However,
quantitatively, differences exist between fish species as well as compared to
homoeothermic animals. Fish have teeth with which they catch feed, but
they do not chew the feed. Neither do they salivate. Accordingly, salivary
amylase is absent. The stomach secretes pepsinogen and HCl from oxynti-
copeptic cells. No gastric lipase production has been observed. Fish pan-
creas seems to produce the same range of digestive enzymes as in
homeothermics, but their molecular as well as functional characteristics
may differ. Adaptation to cold environments seems to have resulted in high-
er specific activity for several of the fish enzymes. The pancreas itself is
organized as a distinct organ in some fish such as sturgeons, sharks, and
eels. However, most fish species have a pancreas diffusum scattered in the
1014
mesenteric tissue as in Atlantic salmon, and/or along the intestine, on the

gall bladder, and in the liver (hepatopancreas, as in sea bream). Pancreatic
ducts collects into one or a few larger ducts in some fish. In others, pan-
creatic juice enters the intestine via many small ducts (salmonids and cod).
Carnivorous fish digest and absorb starch less efficiently than other
fish. Atlantic salmon, in particular, show very low efficiency and no feed-
back response upon starch feeding. Accordingly, starch digestibility
decreases with increasing inclusion in the diet. Amylase activity in intestin-
al chyme from Atlantic salmon is about 1/20 of the activity found in rain-
bow trout. Most raw starch is almost indigestible in fish and need to be
heat-treated under moist conditions to be made utilizable.
The highest intestinal microbial activity is found in the distal com-
partments, but the total number seems to be far lower than in mammals.
No information exists on the significance of microbes in fish nutrition, but
its importance is considered to be low.
28.1.3. Excretion of water soluble compounds and water replacement

The gills function as important excretory organs for water-soluble com-
pounds, relieving the kidneys of several excretory tasks, for example NH3
released during amino acid oxidation.
Fish in seawater loose water rapidly through the skin due to the high
osmotic pressure of the surrounding water. Hence, very little water is avail-
able for excretion through the kidneys, and the urine is highly concentrat-
ed. To replace lost water, marine fish have to drink seawater. Water is
absorbed along the entire intestinal tract secondary to Na+ absorption.
Sodium is efficiently transported across the intestinal wall by several trans-
port mechanisms, creating the necessary water drag. The body is supplied
with excessive amounts of Na+ which is efficiently excreted by the gills. In
fresh water the situation is the opposite. The low osmotic pressure of the
environmental water compared to the fish, forces water into the fish e. g.
with food ingestion, through the skin, and other surfaces exposed to the
environment. The water excess is excreted in urine, which is dilute and
voluminous under these conditions.
28.1.4. Energy and protein requirements for maintenance and growth

As cells of other poikilothermic (ectothermic) animals, fish cells are less per-
meable to ions than cells of homeothermics. Poikilothermics therefore need
less maintenance energy for ion transport across cell membranes. Also loco-
motion, in general, requires less energy in fish than in other domestic ani-
mals. Maintenance energy and protein requirements of fish have been esti-
mated to levels in the range 1/5 to 1/20 of the requirements in domestic
homoeothermic animals. The values vary with environmental temperature
and differ between species. Requirements around 40 kJ/kg0.7 and 0.4 g
digestible protein/kg0.7 have been estimated. Accordingly, required
1015
digestible protein to digestible energy ratio, DP/DE, for maintenance shows

values in the range of 5–10 g/MJ, which is similar to the requirements of
domestic, homoeothermic animals.
Fish require digestible protein and energy for growth and reproduc-
tion at the same level as homoeothermic monogastrics, e.g. chickens and
pigs. They need about 1.3–1.5 kg digestible protein of good quality per kilo
of deposited protein and 1.3–1.5 MJ digestible energy per MJ deposited
energy. Deposition of muscle protein requires a DP/DE of about 40 g/MJ,
whereas deposition of adipose tissue only requires 1 g/MJ. As maintenance
has a low cost in fish, dietary nutrients are mainly used for growth when
fish grow at optimal rate. Depending on the fat content of the gain and life
stage, required dietary DP/DE for the production, maintenance, and
growth, varies between 15 and 35 g/MJ. Overall utilization of nutrients is,
in general, much higher in fish than in homoeothermic animals for which
maintenance is much more costly. Retention of digestible energy and pro-
tein well above 60 % is not uncommon in salmonids under experimental
conditions.
Another factor adding to high nutrient utilisation in fish production is
low cost of reproduction. The number of offspring is so high that cost of
reproduction may be considered negligible.
28.1.5. Specific requirements of vitamin C, long chain fatty acids and

astaxanthin
Absolute requirement for ascorbic acid, vitamin C, is a characteristic of all
fish. Fish, as humans, lack the enzyme L-gulonolacton oxidase, which cat-
alyzes the final step in ascorbic acid synthesis from glucose. As vitamin C
is among the least stable vitamins and fish feed processing is a critical step
for any nutrient of low stability, fulfilment of the vitamin C requirement has
been difficult in the past. However, utilizable, stable vitamin C derivatives
have been developed. Vitamin C deficiencies are no longer a serious prob-
lem in modern fish production.
In contrast to the situation in anadrome fish such as salmonids,
many marine fish species seem unable to convert linolenic acid (C18:3 ?-3)
to eicosa-pentaenoic (C20:5 ?-3, EPA) and docosa-hexaenoic (C22:6 ?-3,
DHA) acid. Accordingly, they have an absolute requirement for the long ?-3
fatty acids. Whether salmonids, which have the necessary enzyme machin-
ery, are able to carry out the conversion efficiently enough under all condi-
tions is not clear.
Astaxanthin is an essential, fat soluble nutrient for Atlantic salmon
and gives the natural flesh colour of salmonids and shrimp. However, the
nutritional requirement is much lower than the level needed for good flesh
colouration. It is not known whether astaxanthin is an essential nutrient for
other fish species.
1016
28.1.6. Nutrition and disease defence systems in fish

The immune system of fish has in principle the same components as
domestic homoeothermic animals, but the non-specific immune responses
appear to play a relatively more important role in fish. Thymus, spleen,
head kidney, liver, and intestine are important immune-competent organs.
Mucus secreted by the skin and intestines contains immunoglobulins,
lysozyme and complement. However, the organisation of the immune mech-
anisms within these organs and tissues has not yet been revealed in detail.
Different fish species seem to differ regarding their defence mechanisms, at
least quantitatively.
As all biological processes, the function of physiological and immuno-
logical mechanisms involved in disease prevention depends on a well-bal-
anced nutrient supply to the organism. Estimates of nutrient requirements
define minimum levels needed to prevent rather severe deficiency symp-
toms, but they may not be sufficient to prevent marginal deficiencies that
can reduce resistance to infections. Nutritional deficiencies may be a pri-
mary, but often overlooked, cause in outbreaks of infectious disease. FIG-
URE XXVIII-2 and XXVIII-3 summarises interactions between feed, envi-
ronment, nutritional status, and health. Nutrient deficiencies may cause
diseases or reduce the organism’s ability to resist attacks from microorgan-
isms and parasites. Also imbalances, either between essential nutrients
and/or relative to energy supply, may cause suboptimal disease resistance.
FIGURE XXVIII-2: Factors affecting fish health. Nutrient imbalances en altered

nutrient requirements may reduce disease resistance.
1017
As for other animals, diseases and stress may alter nutrient require-
ments and utilisation and hence induce nutrient deficiencies and imbal-
ances. Transport and handling, and changes in temperature, salinity, O2-
level, population density, day length, feed quality, and tank colour are chal-
lenging conditions that have been found to increase stress parameters such
as blood glucose and cortisol in fish. Some dietary components, on the other
hand, may reduce stress responses when included at surplus levels, e. g.
vitamin C and tryptophan.
FIGURE XXVIII-3: A schematic illustration of nutritional conditions that may affect char
acteristics of the immune system.
A number of components, some commonly found in fish feed ingredi-

ents, others foreign to fish, have been investigated for their potential ability
to enhance fish immune function and thereby to prevent disease outbreaks.
Among these potential immune-stimulants are: various nutrients supplied
at over-fortification levels, bacteria, yeast and spirulina, chitin, peptides,
non-starch polysaccharides from plants, synthetic compounds such as lev-
amisol, and others. The compounds have been evaluated under conditions
of vaccination, handling, changes in environmental conditions, and disease
outbreaks. Although some experiments indicate that some of these compo-
nents may be beneficial for the fish health, the financial return is question-
able.
1018
28.2.1. Minerals
Fish, living in water, are in contact with their environment to such an extent
that some mineral requirements, e. g. Ca, Mg, Na, K, Fe, Zn, and Cu, may
be fulfilled by absorption directly from the water. On the other hand, excess-
es in the water of any nutrient or other compound may be detrimental to
the fish. Fish are in closer contact with any harmful factors and organisms
present in the environment than homoeothermic animals. Examples of envi-
ronmental factors that have caused fish death are: algae, plankton, bacte-
ria, jelly fish, virus, acids, pesticides, detergents, etc. Great care must there-
fore be taken to secure fish for clean water and otherwise good environ-
mental condition.
28.2.2. Oxygen
Oxygen is also an essential compound delivered to the fish via the water and
absorbed by the gills. Oxygen saturation level in water decreases with
increasing temperature and salinity. At high temperatures, O2-level may
become too low for efficient nutrient oxidation in the fish. If so, feed intake
and subsequently growth will decline. Each species has its own optimal O2
range outside of which growth will be compromised. It is advisable to
decrease feeding rate when the water temperature approaches upper
threshold limits for the fish. For Atlantic salmon the optimal O2 range is
about 5–20 ppm. The lower values correspond with the O2-saturation level
around upper temperature limits for the species, 15–18ºC. Oxygen supply
is much more critical in tank and pond cultivation than ocean cultivation.
At high fish densities it may be difficult to maintain optimal oxygen levels in
the water. Aeration and oxygenation are means frequently used to secure
sufficient oxygen supply to the fish.
28.2.3. Temperature, growth potential and feed intake

Changes in environmental temperature have direct effects on body process-
es in fish. Maintenance and growth processes therefore increase with
increasing temperature within the species natural limits. Nutrient require-
ments vary accordingly. Hence, fish must be fed increasing amounts of feed
with increasing temperature for efficient utilization of feed and growth
potential. Growth potential of different fish species vary greatly. Tables and
formulas have been developed as tools for the farmers in prediction of
growth and feed requirements at different fish sizes and environmental tem-
peratures. The formula below is a good alternative to feed and growth
tables.
Predicted growth=(Initial weight1/3 + (TGC×Sum of day-degrees/1000)3
1019
TGC=(Final weight⅓-Initial weight⅓)×1000/Sum of day degrees;

Unit of weight: gram
Since conditions and efficiencies differ between farms, the factor of the for-
mula, called the Thermal Growth Coefficient (TGC), needs to be estimat-
ed for each farm to obtain the best prediction of growth. Daily feed require-
ment is calculated from the predicted growth, knowledge on energy require-
ment, and information on available energy concentration of the diet to be
fed.
Good estimates of initial weight, fish biomass, are also essential for
optimal feeding. To find good tools for biomass estimation still represents a
great challenge to fish farmers. For juvenile fish, the task is easier than for
larger fish. Samples of small fish are easy to take and average weights can
be recorded. In larger units with large fish, for example in the ocean, it may
not be possible to obtain representative samples. Different alternative
approaches have been developed. One is based on the placement of a metal
frame, equipped with instruments for recording fish length, height and
speed in the net, through which a random sample of the fish will swim dur-
ing a day. This equipment has been found useful in estimations of average
fish weights. Good records of the number of live fish in the tank, cage, or
net are necessary for the calculation of total biomass. For large net cages,
diving to count and remove dead fish is often used.
28.2.4. Other important environmental interactions

Fish, at least coldwater species, show increased growth in times of increas-
ing day lengths. Moreover, for salmonids, growth is greater in sea water
than in fresh water. Water pH is also of utmost importance for fish per-
formance. Different species have different tolerance levels for changes in pH.
Values between 5.5 and 4.2 are most toxic to fish, possibly because release
of aluminium from the environment is at its greatest at this pH range.
Aluminium is very toxic to fish.
28.3.1. Feedstuffs
Ample supply of marine ingredients, i. e. fish meal and oil from a variety of
fish species caught in distant waters, has been a key factor in modern devel-
opment of intensive fish cultivation. Demands from fish feed producers have
forced general improvements in the quality of these products from rather
low qualities, extracted and dried at very high temperature and harsh con-
ditions, to gently dried, high quality oils and meals. Demand for marine
products for aquaculture as well as for traditional animal production has
increased greatly during the last decades, whereas volumes of fish catches
1020
have largely been unchanged with annual variations in catches related to

environmental phenomenon, such as El Niño. The result is a trend of
increasing prices with great variations from year to year. Great efforts are
therefore applied into the search for alternative protein and lipid sources for
aquaculture. Some marine resources are still underexploited, particularly at
lower levels of the nutritional hierarchy, such as krill. However, techniques
for their efficient harvesting and utilization are lacking.
Dried by-products from the meat industry, such as meat and feather
meal and blood meal, are also used in modern fish feeds. For cold water
species, these products have limited use because the high melting point of
the fat fraction reduce nutrient digestibility in general. However, for warm
water species they are commonly used. Severe restrictions on the use of
meat by-products in animal production, due to the fear of transmissible
spongiform encephalopathy (TSE), prevent extensive use of these products
also in aquaculture.
Plant feedstuffs, the main feed ingredients for animal production, are
available also for aquaculture at low prices and in large amounts on the
world market. However, as fish in general have low needs for carbohydrates,
high content of carbohydrates, in particular non-starch polysaccharides
(NSP), often limits their use. Moreover, the presence in plant feedstuff of
several bioactive components, some with distinct antinutritive effects, rep-
resents a major obstacle for their use in fish feed. The compounds are for-
eign to most cultivated fish, and they may interfere with metabolism or
cause allergic, inflammatory, or intolerance reactions reducing production
and health status. Among these foreign compounds are a variety of indi-
gestible carbohydrates, protease and amylase inhibitors, lectins, saponins,
tannins, phytoestrogens, phytic acid, and goitrogens. Feeds with high con-
tent of plant feedstuffs generally cause lower nutrient digestibility and feed
efficiencies than diets based on marine products. Moreover, salmonids react
with severe intestinal inflammation of the distal intestine upon ingestion of
diets with soybean products at levels above about 10 %. The symptoms, are
described as follows: 1) shortening of the heights of the mucosal foldings as
well as of microvilli, 2) loss of the normal supranuclear vacuolization of the
absorptive cells in the intestinal epithelium, 3) widening of the central stro-
ma within the mucosal foldings with increased amounts of connective tis-
sue and 4) profound infiltration of inflammatory cells in the lamina propria.
The mechanism behind the reactions involves increases in crypt prolifera-
tion, apoptosis, and heat shock protein level. Reduced nutrient digestibili-
ty, particularly regarding lipids, is also prominent in these fish. FIGURE
XXVIII-4 illustrates morphological alterations in soybean-fed Atlantic
salmon. Indications exist that also other fish species may be affected by soy-
bean antinutrients, but cod and halibut seem to be resistant. The causative
component of soya has not been identified.
FIGURE XXVIII-4: Histological detail of the distal intestinal villous folds of Atlantic salmon
1021
fed the 0% (A), 10% (B), or 35% soya (C) diets (H&E, ´ 280) (KROGDAHL et al, 2004).
28.3.2. Technical requirements of fish feed

Three types of fish feed are used in fish production:
-Wet feed (30% dry matter)
-Semi -moist feed (60–70% dry matter)
-Dry feed:
Compression-pelleted, temperature below 100ºC (88-92% dry matter)
Extruded and pelleted, temperature above 100 ºC (> 95% dry matter).
Wet feeds are still in use in certain areas of the world and were also used in
the early days of modern salmon cultivation. At that time, frozen blocks of
fish were thrown into the net cages and eaten by the salmon as the block
thawed and dissolved. In some areas of the world, farmed fish are fed wet
mixes of ground fish and dietary supplements, thrown to the fish by pow-
erful pumps. However, as wet feed cause environmental pollution and have
both nutritional and hygienic challenges, they should be replaced by more
technically improved diets.
Semi-moist feeds are suitable for the utilization of by-catches of fish
and by-products of the fish industry in areas distant from fish meal facto-
ries. Moreover, some fish species may show preference for and grow better
on moist feed under certain conditions, for example under challenging con-
ditions of low temperatures and high salinities. Wet and moist feeds will
supply fresh water and reduce bodily costs of water replacement. Dry feed
dominates in intensive fish cultivation. It is easy to transport, store, and
1022
handle, and is easily kept under good hygienic condition.

Besides containing nutrients in the right concentrations and propor-
tions, good fish feed must fulfil several technical criteria. High stability to
mechanical stress, low nutrient leakage to the water, correct pellet size, and
suitable sinking rate are necessary. Fish eat while the feed is in the water.
Some fish may prefer to eat floating feed, some prefer while it is sinking,
while others willingly eat from the bottom. Diets should be tailored and
processed to meet the needs and priorities of the various fish species.
Extrusion cooking, i. e. short time exposure to moist heat at high
pressure and temperature is at present the most efficient processing tech-
nology to reach all technical criteria. Binders, most often a grain with high
starch content or a modified starch of certain sources are used in pellet pro-
duction. Gelatinization takes place during moist heat treatment in the
extruder and the starch turns into an amorphous, continuous structure
throughout the pellet, adhering the particle elements. Proteins may also
serve as texurizers in fish feed. Depending on the temperature and pressure
used in the process, the pellets expand more or less when released from the
extruder through the dye. Tiny holes are formed throughout the pellet as
the pressure is released and the water suddenly boils and evaporates. These
holes will easily take up fat or other liquids. For high fat diets, vacuum may
be used to fill the holes more efficiently. The fat applied after extrusion will
prevent leakage of nutrients from the pellet to the water.
The size of the pellet should also be tailored according to the needs of
the fish. Mouth and oesophagus size set limits for pellet size, i. e. small fish
need small pellets.
28.3.3. Feeding regime

In a practical feeding situation, fish size, stage of development and environ-
mental conditions dictate feeding regime. Small fish have high relative
growth rates but their gastrointestinal volume limits meal size. They there-
fore generally need to be fed smaller meals and more often than larger fish.
There are both species and sex differences in stomach volume. Moreover, as
fish grow faster at higher water temperatures optimal meal sizes and fre-
quencies vary with temperature. Individual fish may show great daily vari-
ation in feed intake. Some fish such as halibut may eat only every other day
or even more seldom. Hierarchy develops rapidly among fish in a tank or net
pen, and the dominant manage to get most feed and grow fastest. Large
body weight differences develop rapidly if measures to counteract hierarchy
development are not taken. There should always be enough pellets in one
meal for all the fish in a group to eat at the same time. Relatively small pel-
lets and larger meals may help limit size differences. However, low frequen-
cy of feeding may increase aggression among fish between meals.
Feeding technology is under continuous development. In some fish
farms, feed is given solely by hand or by manually-controlled feeding
1023
machines. Hand feeding gives the farmer information on appetite which may
be a good indicator of the general condition of the fish. Although of decreas-
ing importance, hand feeding is still a supplement to automatic feeding. In
many farms, feeding is fully automated. For fish in large net pens, equip-
ment employing two different principles is used to adjust feeding to appetite
and thereby reduce feed waste to a minimum. By mounting sonar systems
under the nets, position of the fish in the net can be monitored and adjust-
ed according to eating activities. Low fish density near the feeding area indi-
cates that the fish are satisfied and feeding is stopped. With the other type
of equipment, uneaten feed that sinks all the way to the bottom of the net
pen is recorded. Feed delivery is cut automatically when the number of pel-
lets increases.
Fish can learn to push buttons to release feed. So-called “demand
feeders” have been developed and successfully put to use for rainbow trout
and yellow tail. The system secures that fish get feed when they are hungry.
Such systems seem to lower variation in fish size.
Fish species may be divided into two groups according to larvae morpholo-
gy at hatching: those that have a very simple digestive tract which changes
to the adult complexity after first feeding; and those that hatch with a fully
differentiated digestive tract. Most fish in cultivation are of the first group.
At hatching, the yolk sac still contains nutrients which secure a gradual
transition from endogenous to exogenous nutrient supply.
Most cultivated fish species hatch without the ability to utilize the
present day’s formulated, dry feeds. Salmonids and monkfish are excep-
tions, and formulated feeds that support efficient and healthy growth from
the very beginning of start feeding have been developed for these species.
For most fish, however, live feed is still necessary to bring the larvae
through the initial developmental stages. The natural diet of most marine
fish larvae is zooplankton, mainly various species of copepods. However,
copepods reproduce slowly and cannot be grown at high densities. Huge vol-
umes of water are therefore needed for the production which often is carried
out in lagoons and large pool systems. Such cultivation systems supply feed
varying in quality and quantity depending on the season. Two zooplankton
species, the rotatorium Brachonus plicatilis and the brine shrimp Artemia
spp., are available for larvae feeding and are good alternatives. These organ-
isms tolerate high densities. They grow well on simple nutrient sources such
as yeast and marine oil. The artemia is a larger organism than the rotatori-
um and both grow in size during development from two organisms. Young
rotatoria are used for start feeding of the smallest fish larvae, whereas,
adult artemia are suitable for the final stages before start feeding of the fish
1024
CHAPTER XXVIII: Production Fish Nutrition & Feeding
larvae.
Cultivation of the plankton must be carried out under controlled con-
ditions, and loading of the plankton with nutrients concentrates just before
feeding the fish, is necessary to meet all requirements of the larvae and for
success in many species. Emulsion particles that can hold the necessary
nutrients have been developed. A main challenge has been to supply that
larvae with sufficient amounts of long chain ?-3 fatty acids, in particular
C22:6 DHA. The lipid fraction of artemia, which does not normally contain
DHA, may contain 20 % DHA after 24 hours of supplementation with a
DHA-containing emulsion.
In the fish larvae, pinocytotic absorption of nutrients may compensate
for low pancreatic function in the earlier stages. Endogenous digestive com-
ponents of the live feed, such as enzymes, may also add importantly to the
digestive processes in the fish larvae. For fish species showing slow devel-
opment of the digestive tract, inclusion of digestive enzymes, partly hydrol-
ysed protein and fatty acids in phospholipids form may be beneficial.
28.5.1. General
Classical experiments have been conducted to estimate requirements of
vitamins and minerals in several fish species. In addition to indications of
minimum-required inclusion levels, the experiments have also supplied
information on deficiency symptoms. Anorexia, low growth, fatigue, low dis-
ease resistance and increased mortality are deficiency symptoms observed
for most nutrients in fish, as well as in other animals. Many of the more spe-
cific symptoms of nutrient deficiencies observed in fish show similarities
with those observed in birds and mammals; neurological disorders, skin
lesions, deformities etc.. The vitamins and minerals that most likely would
be involved in a possible nutrient deficiency are: vitamin C, vitamin E, pan-
tothenic acid, pyridoxine, niacin, thiamine, phosphorus, zinc, iodine, cop-
per and selenium. Table XXVIII-1 gives a list of common symptoms and their
possible causes regarding nutrient deficiencies in fish. The list of diet-relat-
ed factors that may challenge fish in cultivation is long. Some selected
nutritional disorders typical in fish cultivation are presented below.
1025
&CHAPTER XXVIII: Production Fish Nutrition & Feeding
Table XXX-1. Nutrient disorder symptoms that may occur in some species, the likely nutri-
tional status and possible nutrient involved
28.5.2. Deformities
Our lack of knowledge on nutrient requirements and interactions in fish has
1026
become apparent as a consequence of the recent major shifts in sources of

nutrients in fish feed. Major changes have taken place for many fish
species, from high protein, low lipid and low carbohydrate, to lower protein,
higher fat and higher carbohydrate. The change from fish meal and oil to
higher levels of plant-based ingredients may represent even greater chal-
lenges for the physiology and health of the fish. The changes have shifted
nutrient balances, in particular relative to energy density.
The recent increases observed in intensive aquaculture in frequency
of skeletal and other deformities and in digestive disturbances, may partly
be the result of such deficiencies and imbalances – imbalances that could
not be foreseen due to incomplete information regarding requirements and
nutrient utilization. FIGURE XXVIII-5 illustrates the result of spinal defor-
mities in salmon that may or may not be a nutritional disorder. Similar dis-
orders have been observed as the result of a too high incubation tempera-
ture at larval stages.
FIGURE XXVIII-5: Deformities reflecting severe fusion and compression of spine elements
(Photo: TRYGVE POPPE).
28.5.3. Cataract
Cataract has been a large-scale feed-related problem in salmon production
in at least two periods (FIGURE XXVIII-6). The first time the situation was
brought about by elevated levels of calcium in fish meal. Certain fish species
used for fish meal production have higher proportions of bones in the body
1027
than others resulting in fish meal with high calcium levels. Inclusion of by-
products from fish filleting plants in fish meal production also gives prod-
ucts with high levels of calcium. The use of high-calcium fish meal as an
ingredient in salmon diets reduces zinc absorption from the intestine and
may cause zinc deficiency with cataract formation and blindness as a typi-
cal symptom in fish. The other situation with nutritionally dependent
cataract developed after the general ban of blood meal from animal feeds
due to fear of transmitting TSE prions from meat by-products. Histidine
deficiency turned out to be the likely major cause of the cataract. Blood
meal is particularly rich in histidine. It was not easy, in either of the cases,
to find the cause of the symptom. Zinc level appeared sufficient in the first
case, whereas in the second, histidine requirement had not been studied in
detail and hence no reference levels were available. The two cataract out-
breaks illustrate that any change in feed ingredients may change nutrient
supply and utilization deficiencies may develop.
FIGURE XXVIII-6: Cataract in Atlantic salmon. Note the cloudiness of the lens. (Photo: 0
TRYGVE POPPE).
28.5.4. Thiamine deficiency

Some fish species, such as herring and capelin, show high so-called thi-
aminase activity concentrated in internal organs. The compound(s) respon-
sible for the activity is (are) not well known. Heat treatment inactivates the
compound. Storage of moist or semi-moist diets based on such ingredients
for some hours without heat treatment results in total destruction of the
thiamine. The phenomenon occasionally caused thiamine deficiency in fish
farms in the early stages of intensive aquaculture.
28.5.5. Nutrient to energy imbalance and fat accumulation

Fish, as other animals, will over-eat energy to meet requirements of
several essential nutrients when they are present in the diet at somewhat
1028
less than optimal levels. A nutrient to energy balance on the low side of the
requirement may therefore be the reason for excessive lipid accumulation in
fat deposits. For lean fish the liver is the most important lipid storage organ
and lipid accumulation is a common symptom of nutrient deficiency (see
Table XXVIII-1). Swollen, pale livers with high fatty infiltration are a common
finding and a great problem in the production of many fish species, e. g.
Atlantic cod. Not only may the liver function be compromised, the fat in the
liver is also an unwanted and wasteful deposition of feed energy. Fatty liv-
ers may develop also in fatty fish such as salmonids, e.g. when fed ingredi-
ents with oxidised (rancid) fat. Trash fish is a critical ingredient in this
respect. The affected livers show extreme fat infiltration. Even renal and
spleen haemopoietic tissues may be affected with anaemia as a conse-
quence.
28.5.6. Carbohydrate excess

Starch is a common ingredient in fish diets as it has favourable technolog-
ical characteristics giving texture and stability to the pellet. Levels around
20 % are not uncommon. However, most fish have limited capacity for uti-
lization of carbohydrates. Surplus carbohydrates cause accumulation of
glycogen in the liver and enlargement of the organ in many species.
Glycogen accumulation may impair liver function. In rainbow trout recovery
time after exposure to anaesthetics has been observed to correlate with liver
glycogen accumulation; the higher the glycogen level, the longer it takes the
fish to recover.
28.5.7. Iron excess

In the late 1980s, the outbreak of the “Hitra disease”, which was caused by
Vibrio salmonicida, affected Norwegian salmon farmers with dramatic con-
sequences. There are indications that the infection was exacerbated
because the fish were overloaded with iron from fish meal. At that time, the
meal was dried on hot steel drums that gave off rust. Iron sequestration is
one important strategy for animals to prevent growth of invading bacteria.
However, the iron-binding capacity of the fish was exceeded, and the free
iron allowed increased bacterial growth. A disease challenge trial showed
that mortality caused by Vibrio salmonicida strongly depended on dietary
iron levels.
28.5.8. Mycotoxins
Some of the most potent animal toxins are produced by moulds. Aflatoxin,
produced by the blue-green mould Aspergillus flavus, common in soil, is a
powerful carcinogen for humans and similar effects have been observed in
rainbow trout. Among cultivated fish species, rainbow trout appears to be
very sensitive. In both humans and rainbow trout, clinical levels of hepato-
carcinoma are reached 4–6 months after exposure. For rainbow trout,
1029
effects have been shown at dietary levels as low as 0.01 ppb (=pars pro bil-
lion, 10-9). At 80 ppb acute toxic effects are induced with massive hepatic
necrosis and haemorrhage. The mould may contaminate feedstuffs such as
peanuts and cottonseeds.
FOR FURTHER READING
Halver, J.E: and Hardy, R.W (2002): Fish nutrition. Academic Press. San Diego
Lim, C. and Webster, C.D (2001): Nutrition and fish health. Food Products Press.
Binghamton
Wilson, R.P. (200): Handbook of nutrient requirement of finfish. CRC PRess. Boca Raton
1030
Chapter XXIX
KEEPING AND FEEDING EXOTIC ANIMALS

29.1. Environment
1031
Chapter XXIX: KEEPING AND FEEDING EXOTIC ANIMALS
Introduction
The row of unconventional pet extends from monkeys to myriapodous.

Although it can be discussed whether a private man, without the appro-
priate knowledge and skill has the ethical right to keep them, the veteri-
narian should be prepared to their treatment and to give technical advices
concerning handling, housing and feeding. The most frequent source of
problems is that pets are bought, guided by an unexpected/sudden idea,
without the relating knowledge and preliminary studies. Inexperienced
owner will ask for help from the veterinarian, not only concerning diseases,
but also housing, care, feeding, hygiene and other basic tasks around the
animal, too. To be able to answer this challenge, vets have to know the biol-
ogy of these species in the nature, and the derived from the latter, require-
ments in the captivity. The two most important factors of the disease pre-
vention are the appropriate housing and feeding, covering nutrient needs.
The present overview is destined for helping the general orientation and
does not replace the thorough study of the relevant literature.
29.1. Environment
The range of required environmental temperature and relative humidity for
the majority of exotic species is narrow and a significant deviation from that
itself may cause disease. This is especially true for the poikiloterm (“cold-
blooded“) species (reptiles, amphibian, fish and invertebrate), which are not
able to assure their stable body temperature and thus they are in a total
dependency of the environmental conditions. The homoioterm (“warm-
blooded”) animals (birds and mammals) are much more resistant to envi-
ronment, but they also hardly tolerate increased air motion (>0.2 m/s); rab-
bit and ferret being especially sensitive to that. Although both extreme heat
and cold are harmful, opposite to the majority of production animals, for the
exotics the extreme cold is more dangerous, causing typical clinical signs
(Table XXIX-1). While in case of birds and mammals the requirement to the
environment is known, that of cold-blooded animals only to a less degree. If
there are no literature data concerning the requirements of a given species,
inside the cage or terrarium/vivarium, “nooks” of different microclimates
1032
should be formed by placing separate heater or/and vaporizing device. In

this way, animal itself can choose its optimal environment. Attention: this
will work well only in case of completely healthy individuals, because sick
animals may be apathetic! For reptiles and amphibian the skin respiration
is highly important and dirty materials, floating in the water bath may stop
pores of skin, resulting in drowning/asphyxia. For some species (e.g.
Mediterranean tortoise) environmental temperature of winter sleep (hiber-
nation), i.e. 6 to 8 oC should also be provided in a given period of the year.

Data, concerning the nutrient requirements of exotic animals are sparse;
consequently those of wild living or production animals are used. To make
things more complicated, preference does not mean obligatory essentiality.
For example, Grey African parrot has been fed on grains and seed (often
only on sunflower seed), or Greek tortoise has been kept exclusively on veg-
etable feedstuffs. Later turn out that both species require animal protein,
too. For exotic animals of hardly known requirements, a broad variety of
feedstuffs should be offered, independently from the fact that the animal is
basically an herbivore, omnivore or carnivore. If needs of a species is
absolutely unknown, observation and recording are necessary during some
weeks, concerning the preference and refusal of several individual feedstuffs
and the faeces consistency, produced after ingestion one of them. In these
cases the owner’s compliance is indispensable. Table XXIX-2 summarizes
the most important types of rations. It has to be borne in mind that feeding
of vegetables and fruits rather frequently involve pesticide or heavy metal
poisoning.
The two most frequent nutritional problems of the exotic animals in
captivity are the starvation and the obesity. Starvation develops mostly in
some cold-blooded species, when snakes and frogs refuse eating in captivi-
ty. The reason for that are either the inappropriate feed or the colder than
optimal environmental temperature. A fatal vicious circle may start: loss of
body condition, secondary ailments, for example skin lesions, which finally
result in death of animal. Some birds and mammals refuse eating for the
sake of stress, caused by confinement. Veterinarian, besides the curative
activity should always be ready for giving advices about feeds and feeding
techniques, as well as the keeping conditions. The other extreme is the obe-
sity, together with all its damaging consequences (like fatty liver etc.) is also
common among exotic animals in captivity. In most cases obesity develops
as a consequence of insufficient movement, accompanied by overfeeding,
but it may be caused by endocrine-metabolic troubles, too. Prevention can-
not be restricted to a decrease of rations, but at the same time, feedstuffs,
high in fibre (grass, greens and vegetables, acacia foliage etc.) should be
offered. Latter not only contributes to the obesity prevention and/or treat-
ment, but also helps in overcoming behavioural stress.
1033
In the amphibian and reptiles commonly found constipation

(coprostasis) should be cured by giving warm water bath, enema, using
paraffin oil or colonic injection of anti-mastitis cream. Diarrhoea, besides
eliminating causes, can be treated by activated charcoal and by feeding
banana and/or apple.
Lack of minerals and vitamins causes specific symptoms. The most
common are the skeletal diseases. While feeding pure meat or cereals, phos-
phorus excess and calcium deficit will develop. Affected animals show gen-
eral weakness; their bones are painful and fragile. Eyelids and nostrils of
tortoises get swollen, the animal will lose its capability to taste and smell
and stops eating. Secondary bacterial infections may cause suppuration.
The concomitant vitamin A deficiency facilitates the development of pneu-
monia. In lizards, iguana the swollen mandible indicates the emerging nutri-
tional osteodystrophy. Moderate cases can be successfully treated by feed-
ing ground limestone, calcium hypophosphoricum, water-soluble calcium
pill (“sprinkling” pills) and grinded egg-shell.
Thiamine (vitamin B1) deficiency also occurs, because in many fish
species and in one-day chicken body thiaminase enzyme is present. Table
XXIX-3 classifies fish species, according the presence of thiaminase enzyme.
By giving such a feed for carnivore mammals or birds of prey and reptiles,
deficiency signs may develop. Heat treatment inactivates this enzyme, but
at the same time, decreases preference of feed for the animal. Clinical
appearance of thiamine deficiency may differ according to the individual
species, but neurological troubles, muscle spasms and convulsions, as well
as paralysis are generally characteristic. Both parenteral and peroral thi-
amine applications are effective, if started in time. Parallel to the treatment,
diet should be either removed or supplemented by yeast or vitamin prepa-
ration.
Total omission of green feeding results in vitamin C deficiency in the
fishes. Typical signs are the scoliosis, lordosis and haemorrhage in the skin.
In these cases scalded or frozen lettuce, spinach or green pea can be put in
the water of aquarium.
1034
Table XXIX-1: Clinical consequences of cold environmental temperature
1035
Table XXIX-2: Some characteristic diets for exotic animals
1036
Table XXIX-3: Presence of thiaminase in different fish species (NRC, 1968)
1037
Chapter XXX
FEEDING OF ORNAMENTAL FISH, AMPHIBIANS

AND REPTILES
© Sandor Gy. Fekete and James Sales

30.1.1. Biological characteristics
30.1.3. Practical feeding of ornamental fish
30.1.4. Important nutritional diseases of ornamental fish

30.2.2. Nutrition
30.2.3. Nutritional diseases of amphibians
30.3. Feeding of reptiles

30.3.2. Tortoise and turtle nutrition
30.3.3. Feeding of herbivorous lizards
30.3.4. Feeding of carnivorous snakes and lizards
30.3.5. Nutritional diseases of reptiles
1039
CHAPTER XXX: ORNAMENTAL FISH - AMPHIBIAN, REPTILES
F ish nutrition as a science started in the second half of the 19th cen-
tury, when culturists evaluated various feeds, including horse meat,
pig spleen and marine fish, to increase performance of salmonids.
On the contrary, ornamental fish nutrition as a science is still in its infan-
cy, and most recommendations are extrapolated from results obtained with
food fishes under intensive farming conditions.

Due to differences in digestion among the numerous species of fish kept as
ornamentals, a general overview about biology and digestive functions is
given. For more detailed descriptions, see Chapter XXVIII of this manual
(KROGDAHL, 2005), and „Fish medicine” of STOKOPF (1993).

General nutritional classification is much more complicated in fish than in
birds or mammals. Combinations of herbivorous, omnivorous and carnivo-
rous as the most important biotic nutritional classifications, and tempera-
ture, salinity and water hardness as prominent abiotic factors, result in
more than 18 distinguisable nutritional groups of ornamental fish. Without
doubt, the digestive tract differs among groups, and from those of terrestri-
al animals.
Fish have no tongue, and amylase production is absent in the buccal
cavity. Whereas carp (Cyprinus carpio) have pharyngeal teeth and no stom-
ach (agastric), fully developed or brush (grabbing) teeth, and a well devel-
oped stomach with pyloric caecae, are found in carnivorous species. In gen-
eral gut walls contain substantially amounts of cells, which secrete diges-
1040
tive fluids, and villi are absent. The liver and pancreas are combined in the
hepatopancreas. Intestinal flora are limited, with Aeromonas, Pseudomonas
and Flavobacterium as the main representatives. The number of intestinal
microbes might decreases under natural keeping conditions in winter to
such an extent that the gut lumen becomes sterile. Faeces and urine are
excreted together through the cloaca, complicating the determination of
nutrient digestibility. With some exceptions, fish usually do not digest
dietary fibre.
The main route of ammonia excretion in fish is not the kidneys, but
the gills. Rate and extent of excretion depend on, among others, nitrogen
intake, body weight and environmental temperature. The homeostatic
organs of electrolyte balance in fish are the kidneys, gills, urinary bladder
and intestine.
With maintenance of life processes taking priority over growth and other
functions, energy concentration should be the most important nutritional
consideration in diet formulation for fish. Energy is not a nutrient, but
released during the metabolic oxidation of carbohydrates, fats, and amino
acids. Protein, in contrast to carbohydrates in most terrestrial animals, are
the main energy-yielding compounds in fish. The net energy derived from
protein is higher in fish than either mammals or birds (Table XXX-1).
Table XXX-1: Energy yields (%) from protein
The energy needs of fish are lower than those of terrestrial animals.
In mammals and birds 20 to 30% of the heat increment of feeding (HiE) is
required to deaminate ingested amino nitrogen and excrete the end prod-
ucts. Because they can eliminate the end products of protein metabolism
(ammonia, bicarbonate, carbon dioxide) without the need to synthesize
urea, uric acid, or other similar compounds, this figure is only 5 to 15% in
fish. Also, maintenance energy requirement (HEm) are lower in fish than
terrestrial animals because they do not regulate body temperature, and use
less energy in maintaining position in the water.
1041
Energy requirements of fish are influenced by several external factors.

An increase in water temperature increases the intensity of energy metabo-
lism. Because of this phenomenon maintenance energy requirements are
specified at „Standard Environmental Temperature (SET)”. The latter differs
among species, with values of 12-15, 21-22, and 23-26oC for coldwater,
coolwater, and warmwater species, respectively. Other water quality char-
acteristics which have an influence on energy requirements include pH,
oxygen concentration, and salinity.
Protein in body tissues contain about 23 amino acids, of which 10
cannot be synthesise by fish and have to be supplied by the diet. Relatively
to terrestrial animals, protein requirements in fish are high in terms of
dietary concentration. However, this is due to a a lower absolute energy
requirement. A diet deficient in energy in relation to protein will cause pro-
tein to be used for energy to satisfy maintenance before growth. In contrast,
excessive dietary intake may restrict protein consumption and subsequent-
ly growth. The above illustrates the importance of a balance between ener-
gy and protein in the diet. The optimum dietary protein concentration for
growth of some ornamental fish are presented in Table XXX-2.
Table XXX-2: Optimum dietary protein concentration (%) for growth of some ornamental
fish species
GE: gross energy, ME: metabolisable energy, DE: digestible energy.
Despite the importance of dietary lipids as sources of energy and fatty

acids that are essential for normal growth and survival of fish, and to
achieve optimal larvae quality in broodstock, information on fatty acid
requirements of ornamental fish is lacking. Research with freshwater and
marine food fishes indicated that fish in general require fatty acids of longer
chain length and a higher degree of unsaturation than mammals.
Although no dietary carbohydrate requirement has been established
1042
for fish, carbohydrates present a cheap energy source, that would “spare”
the catabolism of components such as protein and lipids to energy.
Herbivorous fish are able to digest complex carbohydrates due to the
microflora in their hind gut. Carbohydrate digestibility varies from 50% in
moonlight gouramis (Trichogaster microlepis) to 70% in goldfish.
With fish absorbing some water-soluble minerals from the water
across the gills and skin, studies on dietary mineral requirements are com-
plicated. Phosphorus, essential in growth, bone mineralization and lipid
and carbohydrate metabolism, is one of the most important minerals need-
ed in the diet, because of low contents in natural water. Mineral require-
ments during growth established for ornamental fish are shown in Table
Table XXX-3.
Table XXX-3: Dietary mineral requirements of growing ornamental fish
Vitamins, classified as water-soluble and fat-soluble, are organic

compounds required in trace amounts, usually from the diet, which play a
catalytic role for normal growth, reproduction, and health. Many symptoms
of vitamin deficiency are non-specific, and nutritional deficiencies signs
usually develop gradually, not spontaneously. Identification of characteris-
tic deficiency diseases due to the absence of vitamins is poorly investigated
in fish. Requirements might depend on the intake of other nutrients, size of
the fish, and environmental stress. Recommended requirements and dietary
levels of some vitamins for fish are summarised in Table XXX-4. Water-sol-
uble vitamins are most vulnerable to nutrient leaching, with a large per-
centage of vitamin C, vitamin B12, choline, and panthothenic acid in some
commercial flake diets lost in water within 30 seconds after been fed to
ornamental fish.
1043
Table XXX-4: Vitamin requirement of freshwater fishes
* IU; ** μg
One of the most important quality characteristics determing market

value of ornamental species is skin pigmentation. Despite factors such as
genetics, gender, general health status, and even social hierarchy,
carotenoids, one of the most important groups of natural pigments, are
dominant in determination of skin colour. Natural ocurring carotenoids
include beta-carotene, lutein, taraxanthin, astaxanthin, tunaxanthin,
alpha-and beta-doradexanthins, and zeaxanthin. Astaxanthin, a carotenoid
which is readily metabolised from the yellow pigment zeaxanthin, and found
in high concentrations in shrimp, krill, and crab meal, and certain algae,
imparts red colouration in goldfish and Koi carp. Deficiencies of folic acid,
essential fatty acids, or phenylalanin and tyrosine, might cause colour fad-
ing.
30.1.3. Practical feeding of ornamental fish
The diversity of species kept in home and public aquaria, predominantly in
a multi-species environment, complicates the provision of an adequate diet
which is suitable for all tank inhabitants. The latter might be a combination
of fish differing in nutrient needs, ability to digest different feeds, feed pref-
erences (herbivores, omnivores, carnivores) and habits (surface, middle,
bottom feeders). Most aquarium fish are omnivorous. DEVOSJOLI (1999) pre-
sented the the dietary preference of the most important freshwater goups as
described in Table XXX-5.
1044
Table XXX-5: Feeding classification of the most important freshwater aquarium fish
Due to nutritional deficiencies and the possible transmission of dis-

eases if not stored properly, the traditional method of feeding ornamental
fish with live feed has lost popularity. However, the importance of live feed
as a supplement to improve growth in aquarium fish is accentuate by sup-
pliers in Singapore. Furthermore, in fish nutrition live feeds are used for
feeding of larva fish, with rotifer (Brachionus plicatilis) and brine shrimp
(Artemia) the zooplankters produced in mass quantities. Nematodes of the
Panegrellus spp., and Daphnia, among others, are suitable organisms for
feeding of postlarval ornamental fish. The chemical composition of some
popular live feeds is presented in Table XXX-6,
Vitamins, drugs and other compounds to be supplemented can be
incorporated into a gelatin-based diet, of which a recipe is presented by
GRATZEK and MATTHES (1992): Thoroughly mix 35 g canned sardines or tuna,
30 g fine grounded vegetables, 30 g cereal meal, 5 ml cod liver oil, 2-3 g

brewer’s yeast, 250 mg ascorbic acid, 50 mg a-tocopherol, and 5-10 g gela-
tin. Add warm water to make a slightly pourable slurry. Add supplement,
which should be predissolved in warm water or vegetable oil. Dissolve gela-
tin in 100 ml boiling water, cool down, and mix into slurry. Refrigerate in
plastic bags, and distribute by grating a portion from the the frozen block
in the aquarium.
1045
Table XXX-6: Size and chemical composition of some live fish feeds
(after HOFF and SNELL, 1987)
30.1.4. Important nutritional diseases of ornamental fish

Energy deficiency causes undernutrition in ornamental fish, with an influ-
ence on body width and weight, without affecting the length of the fish
(„lanky fish-syndrome”). Whereas a lack of protein and/or arginine causes
proportional and reversibile growth depression, tryptophan deficiency leads
to reversible scoliosis and lordosis. Inadequate intake of w-3 fatty acids
results in fatty liver, depigmentation of the skin, oedema, shock syndrome,
cardyomyopathy and fin necrosis.
Clinical signs related to vitamin deficiencies are presented in Table XXX-7.
Table XXX-7: Clinal signs related to vitamin deficiencies
Iodine deficiency results in goitre, with histological evaluation show-

ing the presence of excessive amounts of colloid material in the follicles of
the hyperplastic thyroid gland. Insufficient intake of carotinoids leads to
1046
decolouration, with otherwise colourful fishes become greyish. This condi-

tion can be prevented or cured by feeding carotinoid-containing plant
extracts, shrimp or crabmeal. A lack of tocopherols results in ascites and
pericardial oedema, accompanied by microcytic anaemia and increased
fragility of red blood cells. Excessive ceroid pigmentation is found in the
body.
Fish are extremely susceptible to natural antinutritional substances,
mycotoxins, heavy metals and environmental pollutants. Due to this the
significance and use of fish in toxicological studies are increasing. Even a
slight starch overfeeding, especially in carnivore fish, provokes severe kid-
ney damage and lethal glycogen-storage diseases in the liver. Cyclo-
propanoid-structured fatty acids of cottonseed oil, especially together with
aflatoxin B1 ingestion, may induce severe liver cancer (“trout hepatoma”).

More than 4000 species of amphibians are documented, all of which are
members of one of three main orders: (1) Anura (frogs and toads), (2)
Caudata (salamanders and newts), and (3) Gymnophiona (caecilians).
Although most amphibians are protected, some species are authorized in
the pet trade, and even used as laboratory animals. The African clawed toad
(Xenopus laevis) is commonly kept as oocyte donors for molecular biological
researches.

Most adult amphibians are obligate carnivores, with a digestive system sim-
ilar to warm-blooded carnivores. However, some differences exist among the
three orders of amphibians, and even inside the same order. Furthermore,
most tadpoles (larvae) are herbivorous, and have a long, coiled intestine
which permits the digestion of plant material. On the contrary, caecilian
and salamander larvae are carnivorous, with cannibalism occurring under
crowded conditions, especially as metamorphoses approaches.
In some salamanders and several frog species the hyoid bones are
modified to eject the tongue for capture of prey. The oral cavity of salaman-
ders and frogs is spacious, and usually serves only for grabbing and swal-
lowing whole prey. They have „jointed” pedicel (base) teeth, which are
replaced througout life. Whereas ranid frogs lack mandibular teeth, bufid
toads do not have any teeth. Aquatic frogs (Pipidae) lack a tongue, with a
1047
fishlike tongue found in aquatic salamanders. The oesophagus, stomach,

small and large intestines, and cloaca, are similar to those of reptiles, with
pancreas and a gall bladder present. The main function of the liver is trans-
formation of ammonia into urea, which is stored in the bladder and excret-
ed when water is available. The liver, with its pigmented melano-
macrophages and non-pigmented Kupfer cells, also plays an important role
in immune function.
30.2.2. Nutrition
Scientific sounded information on nutrition of adult amphibians is scanty,
although some results have been reported on American bullfrogs (Rana
catesbeiana), a species that is commercially exploited in many Asian and
Latin American countries for the food market and as live laboratory materi-
al. The optimum dietary protein level for growth of juveniles (8 g body
weight) of this species was estimated as 39.2% at a protein:energy ratio of
19.4 mg/kJ. The proposed calcium to phosphorus ratio in diets for amphib-
ians is 1:1 to 2:1. Diets of neonatal or young growing amphibians, which are
fed daily, should be supplemented with calcium and vitamin D twice a week.
Adult amphibians fed whole prey (frozen fish, rats or mice) do not require
calcium and phosphorus supplementation.
In nature amphibians consume a variety of feed items, with differ-
ences in preference during the various stages of metamorphoses. Most
important in practical feeding of amphibians is the relation between the size
of the animal and the feed item. For small frogs small insects such as
springtails (order Collembola) are appropriate, whereas fruit flies
(Drosophyla spp.) could be offered to medium size animals, and crickets
(family Gryllinae), which could up to a fifth of the animal’s body size, to
adults. Eating habits should be considered. Capture of prey could be
through ambush and capture, attracting and capture, foraging and capture,
or scavaging. After capturing the prey, frogs swallow it whole with little or
no chewing. In contrast, large salamanders and aquatic amphibians
(Amphiuma, Cryptobranchus, Necturus and Siren spp.) exhibit some chewing
activity. Time of feeding is important. Most terrestrial salamanders and
many tree frogs are nocturnal, consequently they have to be fed at night.
Nearly all terrestrial amphibians are „sight feeders”, meaning that they pre-
fer moving over immobile prey.
Tadpoles, especially the Dendrobatis and Mantella spp., can be fed
1048
with spirulina powder. Overfeeding should be avoided. The larvae of sever-

al salamanders species require live feed during the early developmental
phase. Daphnia, gammarus (fairy shrimp), and other tiny crustacean
species should be cultured for the above, as species collected in the wild
might be infected by a parasite called body fluke”.
The standard prey for most larger salamanders and frogs is crickets,
whereas fruit flies are usually fed to small amphibians. Domestic crickets
(Acheta domestica) can be cultured on fresh vegetables and fruit. However,
their ratio of calcium to phosphorus is 1 to 13, consequently calcium sup-
plementation is needed. Because of the hard exoskeleton of the mealworm
(larva of the Tenebrio molitor beetle), newly „molted” larvae (5-10 mg) are
preferred. Larvae of this species before pupa formation (approx. 30 mg)
should be avoided, because it may cause gut perforation and peritoneal
abscessation, and subsequently death. Calcium to phosphorus ratio in this
source is 1 to 12. Larvae of a relative of the mealworm, the smaller flour bee-
tle (Trilobium spp.), can safely be fed to small frogs. Two species of flightless
fruit flies (Drosophyla melanogaster and D. hydei) are commercially avail-
able and are easy to culture. Springtails or leafhoppers, with a size of less
than 1 mm, can be cultured on detritus and decaying plant material. Larvae
and imago of this insect are excellent as a feed for neonatal or very small
frogs and salamanders.
Some larger frogs, toads and salamanders can be conditioned to feed
on freshly killed or thawed fish, and mouse and rat pups („pink mouse or
rat”). Wild-caught prey may transmit infective agents, including the lethal
fungus Batrachochytrium dendrobatidis, which causes a disease known as
„chytridiomycosis” in amphibians. Some aquatic amphibians, especially
salamanders and newts, readily accept pelleted diets, and floating pellets
are commonly used for commercially reared frogs. However, terrestrial
amphibians practically never eat this nonmoving prepared form of feed.
Longterm maintenance of most amphibians neccessiates offering of live
feed. Furthermore, I t should be remembered that certain amphibians, espe-
cially some frog species, are specialists in their feed choices, and are very
difficult to feed in captivity.
30.2.3. Nutritional diseases of amphibians

Amphibians are extremely sensitive to environmental contaminants such as
ammonia, nitrites, nitrates, chlorine, salt, organophosphates, and
1049
pyrethroids, which are harmless to warm-blooded animals.

Gastrointestinal diseases, usually due to obstruction by a foreign
object, such as gravel, soil particles or invertebrate exoskeletons, frequent-
ly occur in amphibians. The first signs are anorexia and occasional regur-
gitation. Treatment depends on the nature of the foreign body, and could be
from oral administration of a laxative (e.g. psyllium or mineral oil), to sur-
gery in larger animals.
„Metabolic Bone Disease (MBD)” frequently occurs in amphibians.
The development of this syndrome, which includes gastrointestinal bloating,
soft, pliable bones and decreased bone density, is related to vitamin D3 defi-
ciency. Without Vitamin D3 amphibians are unable to use dietary calcium,
even if offered daily calcium supplements. Except for dietary supplementa-
tion, oral treatment with calcium borogluconate and vitamins are recom-
mended. However, „overdosing” of vitamin D leads to hypervitaminosis D,
and secondary renal failure. Ultraviolet light absorbed through the skin
helps amphibians, and all animals, to convert inactive vitamin D to active
vitamin D3 (cholecalciferol).
Thiamine deficiency is seen in aquatic amphibians fed exclusively
on fish, because of its high thiaminase contents. This condition, that is
characterised by tremors, seizures, and opisthotonos, can be prevented by
routinely supplementing diets with 250 mg thiamine/kg of fish fed.
Since many amphibian species will continue to consume prey as long
as it is available without concerning their energy needs, overfeeding will
result in obesity, with all the consequences related to this disease.
30.3. Feeding of reptiles

Reptiles, belonging to the class Sauropsida, and represented by the orders
Crocodilia (crocodiles, gharials, caimans and alligators), Sphenodontia
(tuataras), Squamata (lizards, snakes, amphisbaenids), and Testudines (tor-
toises and turtles), include over 8000 species, and are found on every con-
tinent except Antarctica. As a result of captive breeding projects and easy
availability, popularity of reptiles as pets are fast becoming as common as
fish or birds in households throughout the world. Crocodilia are farmed for
production of exotic leather and meat. Nutrition guidelines for feeding of
reptiles are critical, not only because of numerous nutritionally related dis-
orders experienced in reptiles kept as pets, but also due to the importance
of nutrition in reptile conservation.
1050

In general, reptiles are grouped in four feeding types, as presented in Table
XXX-8. Some species consume diverse diets, including feeds from two or
more categories. For example, skinks (family Scincidae) and girdle-tailed
lizards (family Cordylidae) are insectivorous-carnivorous, with the older
individuals eating flowers and fruits during marginal food supply.
Table XXX-8: Feeding classification of reptiles
The digestive tract of reptiles is much shorter than those of birds or

mammals. Reptiles are polyphyodont animals, i.e. they develop several sets
of teeth, consisting of enamel, dentine and cement, but without a periodon-
tal membrane, successively throughout life at a rapid rate. Depending on
their feeding habits, three types of dentition exist: (1) acrodont (water drag-
ons, chameleons), with the summit of the alveolar ridge of the jaw without
sockets, (2) pleurodont (snakes, iguanas), with teeth that are fused by their
sides to the inner surface of the jaw bones, and (3) thecodont (crocodiles),
with teeth embedded in a deep bony socket without a periodontal mem-
brane. Unlike snakes and chelonians, lizards do chew their feed, and even
tear off pieces if the prey is too large to swallow. The venom gland of some
snake species is a modified oral secretory gland. The tongue has diverse
functions. In lizards it is used for swallowing, and to pick up scent particles
that are carried to the Jacobson’s organs where they are detected by sen-
sory cells, whereas it is used in the chameleon to catch prey.
The stomach is small and produces hydrochloric acid. Crocodilia
swallow stones, known as gastroliths („stomach-stones”), which, except for
acting as a ballast, assist in post-digestion processing of prey. The passage
of digesta is slow, and can take up to four weeks in some herbivorous tor-
toises. This allows for maximum utilization of nutrients. Peristalsis are
slower in herbivorous than in carnivorous or insectivorous species. Since an
1051
impaired immune system will slow down peristalsis, peroral application of

minerals, vitamins and other drugs is not promising in debilitated animals.
Although absent in most snakes, caeca are prominent in herbivorous rep-
tiles such as tortoises. Similar to birds, reptiles have a cloaca, consisting of
an anterior chamber for faeces collection, a middle chamber for urine and
reproductive products, and a posterior chamber to accumulate all waste
products before excretion. Faeces contain only indigestible plant fibre and
animal materials such as fur, hair, beaks, claws, chitin remnants and
eggshells. Insectivorous reptiles have chitinolytic enzymes, secreted by the
stomach and pancreas.
Reptiles are poikilothermic (ectothermic), with their body temperature
depending on the external environment rather than the internal metabo-
lism. Furthermore, they are heretothermic, revealing a range in body tem-
perature parallel to environmental conditions. Thus, environmental tem-
perature will have an influence on activities, including peristalsis, digestion,
and energy needs. The digestive processes slow down between 10 and 15oC,
and stop at temperatures below 7oC. Maintenance energy requirements are
approximately one tenth (0.05-0.06 MJ ME/W0.75) of that for warm-blood-
ed animals. The above caused a lower feeding frequency in reptiles than in
mammals.
Each reptile species has a „Preferred Optimum Temperature Zone
(POTZ)”, which corresponds to the temperature range of the natural habi-
tat, and is usually between 20-38oC. At lower environmental temperatures
digesta will decompose. Tube-fed reptiles must be kept at their POTZ.
30.3.2. Tortoise and turtle nutrition

Tortoises and turtles are commonly kept as pets. The red- eared terrapin
(Pseudemus scripta elegans) and Spur-thighed tortoise (Tescudo graeca). .
Most terrestrial tortoises are herbivorous, in contrast to certain aquatic tur-
tles, which are carnivorous. Box turtles are omnivorous, consuming both
prey and vegetation.
Tortoises and turtles have no teeth, and use their beak-like, sharp
horny mandible to grap feed and swallow without chewing. The tongue is
undivided, short and muscular. The oesophagus and stomach are folded in
herbivorous species, and able to considerable expansion. Whereas aquatic
turtles excrete more ammonia and urea than uric acid, terrestrial tortoises
mostly excrete uric acid.
1052
Carnivores could be fed on earthworm, crabs, whole fish, mouse

pups, worms and maggots. When fed in water, remains should be removed
30 minutes after feeding. Any meat product containing preservatives (e.g.
sausage, salami) is strictly prohibited. For herbivores vegetarian fare, such
as fruit with a low sugar content, water plants, and vegetables, are suitable.
Herbivorous tortoises are not able to produce lactase, consequently feeding
of milk and lactose-containing milk products will result in severe diarrhoea.
As with all animals, the correct dietary ratio of calcium to phosphorus, and
vitamin D to utilise dietary calcium, is important. Tortoises and turtles are
able to use vitamin D3, but not vitamin D2 (ergocalciferol).
30.3.3. Feeding of herbivorous lizards

One of the most popular reptiles kept as a pet is the herbivorous green igua-
na (Iguana iguana), native to Central and South America. It can grow to 1.5
m in length from head to tail. Because they consume primarily leaves in
their natural environment, they are called folivores. Green iguanas are
hindgut fermenters, with large transverse folds in the proximal colon, sub-
dividing it into five pockets which retain feed for up to 85 hours. Newly
hatched iguanas have no microbial flora in the digestive system, and the
eating of faeces from adults might be an adaptation for colonizing. There is
dispute if hatcling and juvenile green iguanas tend to be omnivorous, or
partial insectivorous. Relatively to nutritional research on other reptiles, a
considerable amount of scientific based work has been conducted with
green iguanas. Despite this, the nutrient requirements of this animal are
still unknown.
Captive diets should be adapted to the development of the animal
(BRAULT, 2005). During the period of intensive growth from hatchling to 30
months of age (puberty), 85 to 90% plant material, 10 to 15% animal pro-
tein, and a mineral-vitamin supplement, should be provided on a daily
basis. From puberty onwards till 5 years old (period of moderate growth),
95% plant material and 5% animal protein, and a mineral-vitamin supple-
ment, could be fed every second day. After 5 years of age growth slows down
drastically, and the amount of feed, dietary protein level, and feeding fre-
quency should be decreased. Mature females will need more dietary calci-
um and protein during egg production.
Despite been herbivorous, green iguanas will take a wide variety of
feed items, including forages, vegetables, fruits, insects, meat, cheese, and
1053
hard-boiled eggs. Care should be practised when feeding cabbage, broccoli,

cauliflower, kale, brussel sprout and other members of the Brassica family.
They contain thyreostatic compounds (see Chapter VIII), which might dis-
turb thyroid functioning. Mainly due to „self-selecting” of nutritionally
incomplete items when offered a „salad-type” diet, but also because of a pro-
liferation of iguana ownership following the release of the movie Jurassic
Park, commercial diets for green iguanas in the frozen, canned, pelleted,
extruded, meal and powder forms have been developed. However, these
diets are extremely variable in nutrient composition. If any nutrients are
omitted or included in inappropiate levels it may affect the health and per-
formance of the animal.
30.3.4. Feeding of carnivorous snakes and lizards

Carnivorous snakes and lizards are almost exclusively fed on whole prey
items (rodents, fish and poultry). The ability of snakes to swallow large prey
is made possible by the flexible (kinetic) skull that can create an enormous
gap. Feeding of live prey is frequently criticized by animal welfare.
Furthermore, live prey may cause bite wounds, trauma, secondary infec-
tions, and even death, to snakes. In addition to whole prey items, inverte-
brae, fish slices, eggs and fruit could be offered.
Feeding frequency varies with size and age of the animal, in some
large snakes it could be every second to third month. However, newborn
and juvenile animals should be fed daily. Feed intake and weight gain have
to be monitored on a regular basis.
30.3.5. Nutritional diseases of reptiles

Growing herbivorous tortoises fed on excessive amount of protein, such as
leguminous seeds, are extremely susceptible to „protein overfeeding syn-
drome”. This secondary osteodystrophy is characterised by abnormal shell
growth, soft shells, irregular shell growth („pyramiding”), and lost of shell
pigmentation. It develops if calcium supplementation is not adapted accord-
ing to protein intake. The terminal phase is renal failure, indicated by
edema (accumulation of fluid between tissue cells) under the skin of the
sublumbar fossa, neck and shoulders. Survival is low.
Vitamin A deficiency, commonly found in carnivorous terrapins,
leads to degeneration and metaplasia of epithelial surfaces such as the con-
junctiva, gingiva, pancreatic ducts, renal tubules, skin and lung alveoli.
1054
Clinical signs are swollen eyes, conjunctivitis, sloughing of skin, „runny

nose syndrome”, necrotic stomatitis, respiratory problems, jaundice (yel-
lowish discoloration of the skin), renal failure, and finally death. Except for
vitamin A supplementation (injection, feeding of liver), secondary symptoms
should be treated. On the contrary, „overdosing” of vitamin A”, as also
found with vitamin D, causes hypervitaminosis (vitamin toxicity), noticeable
by edema under the neck, weakness, listlessness, poor appetite, and dehy-
dration (from kidney damage). Also, because of an interrelationship between
vitamin A, vitamin D3 and calcium, it appears that excessive vitamin A
intake might deplete calcium reserves, resulting in symptoms of „Metabolic
Bone Disease”.
Steatitis („Yellow fat disease)” may develop in growing herviborous
tortoises receiving diets containing an excess of polyunsatured fatty acids
and lack of vitamin E, e.g. commercial dog and cat feed, fish, cheese and
other dairy products. Symptoms include obesity, nodular fat deposition,
fatty livers and jaundice. Oral or injectable vitamin E treatment, with thy-
roxine feeding and anabolic steroid vaccination to break down fat, is rec-
ommended.
Thiamine deficiency due to excessive intake of fish containing thi-
aminase, iodine deficiency resulting in lethary or debility, and „Metabolic
Bone Disease”, as described above for fish and amphibians, are often
encountered in reptiles. Clinical signs related to the latter condition, as well
as for „osteodystrophia fibrososa”, which is caused by feeding of a low-cal-
cium high-phosphorus feed, are intensified during the period of fast growth,
or by feeding of high-protein diets, such as meat or commerical dog and cat
feeds.
Visceral and articular forms of gout have been reported in reptiles,
especially in Red-eared terrapins. Primary visceral gout, usually caused by
excessive dietary protein intake, is the accumulation of urate microcrystals
in organs secondary to a chronic hyperuricemia. Secondary visceral gout is
due to chronic hyperuricemia, and caused by, among others, dehydration,
starvation, and renal damage. For treatment proper hydration is necessary,
and fluids could be be administered. Medications such as an urease
inhibitor (allopurinol) may be used. However the exact dosage and safety of
these drugs in reptiles have not been determined yet.
Hair and feathers on some prey may result in hairball (trichobezoar)
formation. Feeding of fruit contaminated with yeast, especially during
1055
extended storage, and milk, could cause diarrhoea. However, diarrhoea

might also be due to viral, bacterial, parasitic and fungal infections, gener-
al chronic debility, antibiotic treatment („sterile gut syndrome”), inappropri-
ate diets, and vitamin deficiencies. After treatment fluid therapy should be
provided, and, if applicable, dietary fibre levels be increased and sugars
decreased.
FOR FURTHER READINGS
Cogger, H.G. (2001): Reptiles and Amphibian. Federal Street Press. Springfield,
Massachusetts, USA.
Donoghue, S (1998): Nutrition of pet amphibians and reptiles. Seminars in Avian and
Exotic Pet Medicine 7, 148-153.
Fekete, S.: The nutritional value of some protein sources, studied by means of the grubs of
the beetles of family Tenebroidae (in Hungarian). Allategeszsegugyi es takarma-
nyozasi kozlemenyek, 1980. No.2. 125-128.
Frye, F.L. (1995): Iguana, Iguana. Guide for Successful Captive Care. Krieger Publ. Co.
Malabar, Florida, USA.
Gatzek, J.B. and Matthes, J.R. (1992): Aquariology. The Science of Fish Health
Management. Morris
Sales, J. and Janssen, G.P.J. (2003): Nutrient requirement of ornamental fish. Aquat.
Living Ressour. 16, 533-540.
1056
Chapter XXXI
FEEDING AND HOUSING OF SMALL MAMMALS

© Christine Iben
31.1 Biological characteristics

31.2. Diet design and feed composition
31.2.1. Herbivores
31.2.2. Omni-/granivores
31.2.3. Carnivores
31.2.4. Water supply
31.3. Special needs of individual species
31.3.1. Guinea pigs (Caviidae)
31.3.2. Chinchilla
31.3.3. Degu
31.3.4. Rats and mice
31.3.5. Hamster
31.3.6. Gerbils
31.3.7. Ground squirrels (syn. burunduk, chipmunk)
31.3.8. Ferrets
31.4. Feeding neonates
1057
CHAPTER XXXI: SMALL PET MAMMALS
he way of living of wild animals live teaches us the way how they
T should be housed and fed in captivity. Wild animals spend most of

their time foraging, ingesting feed and depending from on the species,
hiding feed. Due to these habits the environment of animal in captivity
should be as big large as possible and proper facilities should offer a struc-
tured surrounding. Those animals which are living in colonies and there-
fore must not be kept separated. Ferrets and rodents are popular pets that
are often presented to veterinarians for advice about concerning their care
including diet and feeding management.
31.1 Biological characteristics

The single species differ due to the differentces in their digestive system
and their nutritional requirements. On the one hand there are the more
grain consuming species like mice, gerbils, hamsters, rats, chipmunks
(Tamias striatus) and squirrels, which have a restricted capability to digest
high fibre diets. On the other hand there are those species that are defini-
tively herbivores such as rabbits, chinchillas, guinea pigs, and degus and
able to digest high fibre diets. On the other hand, ferrets are carnivores
(Table XXXI-1).
Table XXXI-1: Classification of small mammals due toregarding their feed/nutrient require
ments
Rodents (from the Latin verb “rodere”, to gnaw) are identified by their
four prominent, continuously erupting incisors. They do not have canine
teeth and the presence or the absence of premolar teeth is species depend-
ent. The premolar and molar teeth can be open or closed rooted, depending
from on the species. The growth rates of incisors in different species are
shown in Table XXXI-2.
1058
Table XXXI-2: Rate of incisor tooth growth (mm per week) of different species
Like rabbits, guinea pigs, chinchillas and degus are hindgut fer-
menters and use show the habit of coprophagy/caecotrophy. Caecotrophes
are special faecal pellets produced in the caecum) are excreted during the
night and early morning as clusters of grapelike material and are consumed
directly (caecotrophy) from the anus. Caecotrophes contain twice the pro-
tein (25 to 30% of dry matter) of usual faecal pellets, more B-vitamins and
much less fibre. Most other rodents are also coprophagous, too. Young
rodents ingest maternal faeces, thereby inoculating their own intestinal
tracts with autochthonous flora (see Chapter XIX).
The calcium metabolism of guinea pigs, chinchillas and hamsters is
similar to that of rabbits. They absorb calcium very efficiently and the
excess is excreted in the urine, rather than in the bile as it typically occurs
in other species. Prolonged intake of high-calcium feeds in combination with
high vitamin D intake may cause calcification in the soft tissues such as
aorta and kidneys.
31.2. Diet design and feed composition
31.2.1. Herbivores
Hay and green fodder, vegetables and fruits, with or without the supply of
cereals and/or minerals
Risks: These compositions can cause affections due to sudden changes of
feed, deterioration of moist feed or improper minerals. Nevertheless, hay in
combination with green fodder is the most natural and appropriate feed.
The high water content causes higher total fluid intake. Due to low energy
content, animals are considerably longer busy with feed intake and the dis-
position to become overweight is lower.
Compositions of hay and cereals
Risks: Calcium supply may be deficient. Due to the high energy content of
cereals, animals can become overweight.
Hay and commercial food
Risks: Selection of special seeds can cause overweight/obesity or nutrient
imbalance.
31.2.2. Omni-/granivores
Composition of cereals, seeds, nuts, with or without fruits or vegetables
and, depending from on the species, egg, curd cheese , and/or cooked meat.
Risks: When self-made compositions are given, a deficient supply of
1059
minerals and vitamins may occur. Furthermore, high energy intake because
of selective intake of highly palatable fatty seeds or nuts may result in high
energy intake.
Commercial concentrates
Risks: If the concentrates are not well prepared, excessive growth of teeth
can be caused. Hay should be given in a way that the animals have the
chance to chew. Concentrates do not provide enough activity through feed
intake.
31.2.3. Carnivores
Home made mixtures of meat and entrails, cereals, depending on the
species also vegetables or fruits and minerals.
Risks: Lack of feed hygiene can cause diarrhea. The preparation of a bal-
anced diet requires knowledge about of the need of the animals and the
composition of the different feedstuffs. Therefore animals are at risk of
unbalanced diets can occur.
Dry or canned dog and cat foods.
Risks: High energy density and possibly no exercise opportunities can lead
to obesity.
Insects (mealworm, house cricket, flies, maggots and others)
Sufficient mineral supply has to be ensured, as the mineral and vitamin
content of these insects is often often too low.
31.2.4. Water supply

Water has to be offered to all animals all the time, even when if juicy feed is
given (fruit, vegetables, green fodder) and the animals hardly drink.
Anyhow, factors that cause higher water requirement like are diseases, high
environmental temperature or little water content in the feed, can occur.
Sipper bottles are most comfortable. Using sipper bottles water as well as
cage contamination can be prevented. Because not all of the animals are
used to sipping bottles it is necessary to make sure that the animals get
enough water from the bottle.
31.3. Special needs of individual species
31.3.1. Guinea pigs (Caviidae)

Housing. Guinea pigs are gregarious (social) animals living in groups of
about 20 animals or more and this species, therefore they must never be
kept singly. They must not be kept together with rabbits. The cage for one
or two animals must be at least 100×60×50 cm (length×width× height). They
need a small house for to sleep or to hide in and an elevated mooring area.
Because guinea pigs are easily startled, the cage should be placed in a quiet
area. Ideally, a relatively constant temperature of 18–24°C should be main-
tained. Elevated temperatures may cause heat stress. Guinea pigs tend to
1060
contaminate feed and water dishes with their excreta. Proper sipper bottles
and hay racks should be used.Guinea pigs are active during the
daylight.during the day.
Physiology. The teeth of guinea guinea pigs are open-rooted and erupt
continuously. Guinea pigs have a large stomach and a long digestive tract.
The caecum is a thin walled sac divided in numerous lateral pouches by
smooth muscle bands (taenia coli). The caecum is normally found on the
central and left side of the abdomen and may contain as much as 65% of
the gastrointestinal contents. Any changes in access to feed or drinking
water as well as feed composition should be made gradually, over a period
of 5 to 10 days. Changes in husbandry or feeding can cause anorexia and
the animals have to be monitored for their feed and water intake.
Guinea pigs are unable to synthesise vitamin C, because they lack the
enzyme L-gluconolactone oxidase. To prevent hypovitaminosis C, 10 to 20
mg vitamin C per day are necessary. Vitamin C containing fruits and veg-
etables as well as greens can provide adequate amounts of vitamin C.
Quality commercial guinea pig foods are food is formulated with adequate
amount of vitamin C.
Vitamin C -content of some feedstuffs (mg/100g fresh substance): let-
tuce 10, carrots 8, spinach 51, potatoes (cooked) 14, apples 12, bananas 11,
kiwis 100, dandelion 30.
Feeding. Hay has to be available all the time. Guinea pigs consume
about 30 g hay/day. Additionally they need about 150 g green fodder or
about 80 g vegetables or fruits. When hay and concentrates or cereals are
given, they need about 20 g green feedstuffs.
Isoleucine requirement for the 3 weeks old male guinea pigs was
studied by using of crystalline amino acid diets containing 3.65% nitrogen.
Crystalline amino acid diets ranging from 0.2 to 1.2 % isoleucine were fed
for 22 days. A 0.5% level of dietary isoleucine (2.2% of total N×6.25) was the
lowest level given that did not have a response significantly lower than that
of the higher levels, and that generally promoted a thrifty and well-groomed
appearance of the guinea pigs. High cholesterol level in feed leads to
decreased feed intake.
Energy requirement for maintenance is approximately 0.5 MJ DE/kg
LW/day.
Protein requirement for (maintenance is approximately 3 g pro-
tein/kg LW/day (practically 10% protein in dry commercial food)
Feed intake: 40 to 60 g dry matter/kg LW/day (adult animals)
50 to 75 g dry matter/kg LW/day (growing animals)
Water intake: 2 to 3 (or more) ml/g dry matter intake
Feed and feeding related diseases. Hypovitaminosis C (scurvy) can be
caused by nutrient´snutrient’s lability during storage. Also the feeding of
rabbit food without additional vitamin C may cause scurvy. Anorexia,
weight loss, gingivitis and the inability to move or abnormal moving of the
1061
lower limbs were observed in vitamin C deficient guinea pigs. Animals are
reluctant to move because of joint and muscle pain. Young animals and
pregnant sows are affected most severely. When hypovitaminosis C is diag-
nosed or when the suspicion exists, 50–100 mg vitamin C/kg LW should be
given until clinical signs resolve.
Pregnancy toxaemia occurs primarily in obese, anorectic, stressed
guinea pigs. Clinical signs occur within 5 days before and after parturition
and include lethargy, prostration, muscle spasms and death.
Hyperlipidemia and ketonemia can be diagnosed, as well as proteinuria and
an imbalanced electrolyte household. Obesity is often seen in animals kept
singly.
31.3.2. Chinchilla (Chinchilla chinchilla, Chinchilla vevelligera)

Housing. Chinchillas live in Peru, Bolivia and Chile on the rocky mountain-
sides of the Andes up to a sea level of 4000 m. The crevices and caves of
the rock faces provide hiding places..Chinchillas are active during theby
dawn and during the night. They have skilfully madel climbers; thus the
opportunity to climb and to jump should be provided also in captivity by
offering of adequate cages. Chinchillas are sociable and live in colonies.
Males and females can be kept together if there are enough hiding places.
The wild animals are endangered.
Chinchillas are fastidious groomers and must be provided with a dust bath.
Physiology. Chinchillas possess a big stomach, the caecum is big and
equipped with taenias and posches. The caecum is on the left side of the
abdomen and the ingesta is thin fluid. The first part of the colon is wide and
also voluminous and equipped with taenias and posches. The second, nar-
rower part forms the characteristic faeces pellets.
Like guinea pigs and degus, chinchillas lose their milk teeth before
parturition. Chinchillas are nocturnal animals. In an investigation of WOLF
and KAMPHUES (2003), chinchillas ingested more than 70% of their total feed
intake during the dark phase (highest level of activity between 9 PM and 7
AM).
Feeding. Chinchillas are strict herbivores and wild animals consume
grass and shrubs. All recommendations reflect a high overall dietary fibre
requirement. Nevertheless, chinchillas need concentrates or cereals, espe-
cially when there are in times of increased requirement like pregnancy or
stress. Hay has to be the basic feed. In spring, sprouting twigs of fruit trees
can be given, also dandelion, grass or salad. Pipfruit as well as carrots and
celery complete the ration. Furthermore, they need about 15 to 20 g con-
centrate (oats or commercial feed for guinea pigs or chinchillas). Chinchillas
kept at the institute of nutrition consumed approximately 15 g hay and 15-
20 g oats or concentrates per day. Additionally they ate small amounts of
fruits or vegetables. WOLF et al. (2003) found the daily amounts of feed
intake between 2.5 (fresh grass) or 2.6 (hay) and 5.5 (pelleted complete diet)
1062
g of dry matter per 100 g LW. An offered mixed feed based on native com-
ponents led to a selection of individual ingredients (high palatability: carots,
beet pulp, sunflower seeds). The average water intake amounts 1.5-3 ml/g
of dry matter
Like all herbivores, chinchillas do not tolerate sudden changes in
drinking or feeding behaviour. Mouldy feed can be lethal, as well as crow-
foot, poppy seed, meadow saffron and other poisonous plants. The daily
energy requirement for maintenance is approximately 0.5 MJ DE/kg LW.
Feed and feeding related diseases. Any disruption of the digestive
processes can result in diarrhoea, constipation, mucoid enteritis, bloat,
intussusception and rectal prolaps. Inapropriate feed or sudden feed
changes are common causes of these problems, especially when the food
does not contain enough fibre.
31.3.3. Degu (brush tail rat; Octodon degu)

Housing. Degu is found on the west slope of the Andes at elevations of up
to 1500 meters in fairly open areas near thickets, rocks, or stone walls.
Colonies build extensive burrow systems with a main area usually located
beneath shrubs or rocks, and coplex complex tunnels and surface paths
radiating from it.
Degus have a strong social organisation, which is determined by
group territoriality. The territory primarily defended is the central burrow,
where females of the same social group may rear their young together.
Therefore, also in captivity, degus may not be kept alone; more than 2 two
animals should be housed together. Degus are diurnal, with most of their
activity occurring in the morning and late afternoon. As they are very active,
they need a lot of space. They like climbimg up and sitting on perches like
birds. They enjoy tightrope walking, bathing in chinchilla dust and engag-
ing in boxing matches. Like gerbils they are avid nest builders. Apparently
in the wild they spend a lot of time making piles of twig. In the absence of
twigs, they collect paper tissues, which can be offered for bedding. Like ger-
bils, degus can shed part of their tail as a defence mechanism. Therefore
these animals should never be fixed by their tails!
Physiology. Degus are unable to metabolise sugars. If they are fed
anything containing sugar, - and that includes the natural sugars that are
found in fruit, they often become diabetic which shortens their life time.
Additionally, diabetes can cause cataracts and thus degus become blind.
The digestive tract is similar to that of chinchillas and changes in feed or
water supply have to be done carefully as well.
Feeding. In the wild the diet consists of grasses, leaves, bark and
seeds. Hay and commercial food for chinchillas or guinea pigs is the basic
food, in addition to dandelion, grass and twigs can be given.
1063
31.3.4. Rats and mice

Housing. Mice (Mus musculus) and rats (Rattus norvegicus) belong to the
Muridae family and originate from Asia. Both species are in principle, com-
patible animals in principle, if as long as there is no mix with of more than
one male and with females. The cage for rats or mice has to be escape-proof,
easy to clean and equipped with at least a sipper bottle, houses and possi-
bilities opportunities to climb. Mice and especially rats are intelligent ani-
mals and the environment should be enriched with hiding places , evenfor
themselves and for feed. The animals should be able to be as busy with
looking for feed as they are in the wilderness.
At the institute of nutrition, forty eight-month-old male rats were put
together in a big large cage which that was equipped with 10 sleeping hous-
es. Sawdust was used as litter. No fights were observed between the animals
and the life span of these rats was about 3.5 years. The animals were fed
commercial food for rats and selected table-scraps.
Physiology. Mice and rats are omnivores. Incisor teeth are open root-
ed, but molar teeth are permanent rooted. The gastrointestinal transit time
of mice is 8 to 14 hours in mice and 12 to 24 hours in rats. Rats have a
divided stomach, a large caecum and no gall bladder. Although various
nutrient deficiencies have been produced in experimental rats; theese syn-
dromes are uncommon in pet animals fed commercial dry rodent food.
Protein deficiencies can be observed and can result in anaemia, cataracts,
poor growth and impaired reproduction.
Feeding. The energy and nutrients requirements of mice and rats is
well known, because both species are have been used as laboratory animals
for a long time (see Chapter XXXIV). Mice and rats should be fed a clean,
fresh, commercial dry rodent food. For mice, the food (90% dry matter)
should contain 12 to 18% protein, 5% or less fat and 2.5% fibre. For rats,
dietary fibre content should be at least 5% to minimize problems with diar-
rhoea. On a dry matter basis, crude protein requirements are approximate-
ly 10 % for maintenance and 25% for growth and reproduction.
Rats can be fed on commercial dry rodent food, offered free choice.
Rats and mice both like cCommercial food for hamsters, cereals, vegetables
and fruits, and cooked eggs or ,meat or milk products are liked by both
species. Both species generally eat everything; any kind of food for human
humans is taken up. Both species are primarily nocturnal eater.
Rats: Energy requirement for maintenance amounts approximately 0.46 MJ
DE/kg LW0,75/day, the daily protein requirement for maintenance is
approximately 1.25 g CP/LW0,75, according to the protein quality.
For mice the daily energy requirement for maintenance is evaluated as 0.74
MJ DE/kg LW0,75.
1064
31.3.5. Hamster
Housing. There are many species of hamsters. The most commonly seen are
the golden or Syrian hamster (Mesocricetus auratus), the Chinese hamster
(Cricetulus griseus) and the Dwarf hamster (Phodopus sungorus). Hamsters
are nocturnal animals. In the wild, hamsters are solitary animals so that in
captivity these animals may also be kept on their own. Hamsters live in bur-
rows which can reach 2 m below the earth and are equipped with different
chambers. In captivity, these animals may be kept solely, too.
Hamsters are very active in gnawing and they even can escape by
chewing through cages or by pushing open cage lids. Therefore, proper
caging is a critical factor to for the overall well-being of these animals. They
need a place to hide and an enriched environment. Hamsters are territorial
animals and mark their housing area with urine and faeces and the secre-
tion of the sebaceous glands, which are black and located on both sides of
the flanks.
Physiology. Hamsters possess large cheek pouches that are used to
transport and to store feed. When alarmed, hamsters will also temporarily
store their young in these pouches. These cheek pouches are protuberances
of the cheek mucosa which range to the shoulders. Young are born with
erupted incisor teeth, which grow throughout the life, but the molar teeth
are closed rooted. The stomach is divided into glandular and nonglandular
portions. The nonglandular forestomach is lined with keratinized epitheli-
um and is the site of pregastric fermentation. The dwarf hamster has no gall
bladder. The caecum is located on the right side of the abdomen and looks
like a tubination.
Hamsters are coprophagous and nocturnal animals. The Golden ham-
ster hibernates when temperature falls below 10°C; the Dwarf hamster is
active at each every temperatures.
Feeding. Specific nutrient requirements for hamsters have not been
well established yet. In the wild, hamsters are omnivorous, ingesting a vari-
ety of plants, seeds, fruits and meats (insects, snails). They need relatively
high quantities of protein, fat, and the vitamins A and E. Hamsters ingest
different sorts of cereals. They especially like sunflower seeds, pumpkin
seeds and nuts they like very much and, but these seeds should only be
given fed only in limited amounts. The Dwarf hamsters prefer smaller seeds,
e.g. food for canaries or budgerigars. Moist feeds, such as fruits, carrots,
dandelion, and salad as well as cooked eggs, meat or curd complete the
ration. For gnawing, they like sprouting twigs of fruit trees and hay, which
can also be used as bedding material too.
Commercial rodent food that provides 16 to 20% protein appears to
allow good growth. Hamsters preferentially ingest fruits, nuts, cereals and
“rodent treats” rather than nutritionally balanced commercial dry rodent
food. Therefore, these items should be provided in limited quantities when
commercial food is given.
1065
Fed and feeding related diseases. Deterioration of the stored feed as

well as sudden changes of feed can cause indigestion and anorexia.
Lack of protein, vitamin A and E can lead to skin diseases.
Cannibalism can be occur due to a lack of protein,protein; more often, how-
ever, this it is caused by failures in housing and keeping.
Daily food intake (90% dry matter):
- Golden hamster 8-12 g
- Dwarf hamster ~ 2.5 g
Daily protein requirement:
- Golden hamster 3.5-4 g
- Dwarf hamster 1 g
31.3.6. Gerbils (Meriones unguiculatus)

Housing. From the Cricetidae family the Mongolian gerbil is a frequently
kept pet species. Color patterns are agouti or brown, also other variations
such as black, white or cinnamon exist. Gerbils are burrowing animals
native to the desert regions of Asia. They are social animals and are living
in small family groups. As pets, - they are generally friendly and can be eas-
ily handled. Gerbils form monogamous pairs which that stay together all of
their live span.
Gerbils can be housed as described for hamsters; additionally they
need a dust bath. Temperatures should be maintained between 18 to 29°C.
Gerbils do not tolerate high temperatures and high humidity. They are peri-
odically active by day and night. Big sebaceous glands are located close to
the belly button and their secretion as well as urine and faeces are used to
mark the housing area. Some of the pet gerbils exhibit spontaneous, con-
vulsive seizures that are induced by strange environments or excitement.
Physiology. The digestive tract is similar to that of mice or rats. Their
caecum is remarkably big. Because of their special water conservation
mechanisms, they are adapted to marginal water consumption and they
produce only a few drops of urine daily. They can shed part of their tail
when fixed on it. Gerbils are coprophagous.
Feeding. In the wild, gerbils feed on plants, seeds and insects.
In captivity, different seeds as described for hamsters plus oat flakes and
hay, additionally vegetables and fruits can be fed. Like all rodents, they ger-
bils like twigs. Once a week insects, meat or eggs (cooked) should be given.
They can also be fed a commercial dry rodent food.
Gerbils generally eat about 8 meals per day, with a total feed con-
sumption of 5 to 8 g. Because they frequently eat frequent small meals,
rapid weight loss occurs if available feed quantities are limited. Clean water
in easily accessible sipper bottles has to be provided. Components that are
rich in fat, e.g. nuts have to be limited , as they can cause hyperlipidaemia.
can be caused.
Feed and feeding related diseases. A special disease, intestinal
1066
lipodystrophia, appears predominantly in females. and the reason is an

oversupply of saturated fatty acids, especially lauric acid (coconut oil).
Wet tail or enteritis affects suckling animals aged 10 days until weaning.
Reasons are failures in feeding and/or viruses. Older animals do not sicken-
not fall ill. Reasons are failures in feeding and/or viruses.
31.3.7. Ground squirrels (syn. burunduk, chipmunk) (Tamias striatus)

Housing. Asian chipmunks (Tamias sibiricus) and the chipmunks(Tamias
alpinus, Tamias baller, Tamias striatus) are living in on the West of north-
ern America. Chipmunks are the species mainly kept as pets. Their habi-
tats are mixed forests and coniferous woodland with brushwood and suffi-
cient water. Depending from on the climate they make a superficial winter
sleep from October to April, but they interrupt the their sleep for feed
intake. Chipmunks are solitary living, being active during daytime, espe-
cially during early morning .
Physiology. They have a simple stomach and the length of the diges-
tive tract is about 3.5times the body length.
Feeding. In the wild, chipmunks feed seeds, berries, fruits, vegetables,
sunflower seeds, nuts, line seed, insects, also amphibians, reptiles, even
young birds. They are able to transport their winter stock for kilometres
long distances and they store the fed in different chambers of their burrow.
In principal, the nutrient requirement of chipmunks is similar to that of
hamsters. Commercial food for hamsters supplied with hay, sorghum, sun-
flower seeds and other seeds, maize, nuts, fruits, salad, mealworms or meat
can be fed.
Feed and feeding related diseases. Feeding solely cereals causes frac-
tures because of calcium deficiency and P-phosphor oversupply.
Sudden feed changes as well as spoiled food and ingestion of cabbage
cause indigestions and exhalations.
31.3.8. Ferrets (Mustela putorius furo)

Housing. The domestic or European ferret is a member of the family
Mustelidae. Aristotle already has described the ferret in the 4th century B.
C., which was even yet in that time domesticated as early as that because
of its hunting skills. Ferrets are social animals.
Physiology. Ferret possesses with developed anal glands for marking
the housing area. and theThe secretion of these glands smells intensively
nasty. The digestive tract is short (4times the body length) and similar to
those of other carnivores. No caecum is developed. A subcutaneous fat dep-
osition leads to heavy seasonal differences in body weight (January 2800 g,
July 1700 g).
Feeding. Ferrets are carnivores, although also feeds of plant origin up
to 20 % should be offered. In the wild, the intake frequency is high and in
captivity they should be fed at least five times a day. They thrive on highly
1067
digestible foods food containing large amounts of protein and fat, with min-
imal soluble carbohydrate and fibre. Thus, pet ferrets may be fed commer-
cial foods manufactured for cats. Cat food appears to be adequate for all life
stages of ferrets. It should be mixed with vegetables and cereal flakes.
Especially for ferrets marketed ommercial foods are known to be successful
in the feeding of minks and cats. Guidelines for cat food may be used when
assessing the complete and balanced composition of food intended for fer-
rets. Commercial foods should generally contain more animal-based than
plant-based ingredients to ensure high digestibility, palatability and protein
quality.
Energy requirement (maintenance): 0.5 MJ ME/LW0.75
Protein requirement: at least 30% of food dry matter
Amount of food/day:
90-130 g canned food mixed with vegetables and/or cereal or
25-35 g dry cat food
Feed and feeding related diseases
Urolithiasis (mainly struvit-uroliths); pathogenesis and prophylaxis are alike
the same as for cats (see Chapter XXI).
Hypocalcemia during lactation (parturient paresis) caused by lack of calcium
in meat rations.
Rickets and osteodystrophia fibrosa caused by unbalanced and deficient cal-
cium and phosphor contents in the feed or lack of vitamin D3.
Thiamine deficiency can occur when raw fish are fed.
Dilatation cardiomyopathy can possibly be caused by a taurine deficiency.
1068
31.4. Feeding neonates
The energy and nutrient requirement of all species is increased 2 to 3-fold

during late pregnancy and during lactation. At least during lactation, her-
bivores also (guinea pig, chinchilla, degu) should be offered grains to meet
their energy requirement as well as green fodder and hay to meet protein
requirement. Omnivores need higher amounts of high quality protein like
milk products, eggs, meat or insects. The feed should be offered by free
choice access to minimize the risk of diseases. A calcium containing miner-
al lick may help to prevent calcium deficiency.
For rearing orphaned neonates of these species milk replacers for
dogs or cats can be used. Guinea pigs, degus and chinchillas are precocious
and the rotation of teeth proceeds already before birth. They have a full coat
and open eyes. Neonates usually begin eating to eat solid feed at 4 to 5 days
of age. Nevertheless, to receive sufficient energy and protein, they need milk
during the first 3 weeks of life.
The other species are altricial, eyes and ears are closed. All neonates
need a high environmental temperature (30-32°C), which normally is pro-
vided by the mother and has to be made availableprovided for orphaned
pubs as well. Orphaned pubs need to be massaged gently on their belly to
activate urination and defecation. The temperature of the milk replacer hast
has to be 37°C.
1069
Table XXXI-3: Biological and nutritional database of small mammals
1)depending on gender and season: female 600-1200; male 1200-2400; 2)depending on the
race; 3)concentrate (88 % DM); 4)females heavier; 5)1st week of life; LW = body weight; DM
= dry matter
1070
1071
FOR FURTHER READING

Ayers, L.S., Typpo, J.T. and Krause, G.F. (1987): Isoleucine requirement of young growing
male guinea pigs. J. Nutr. 117, 1098-1101.
Carpenter, J.E. and Kolmstetter; C.M. (2000): Feeding small exotic mammals. In (Ed.
Hand, M. S., Thatcher, C. D., Remillard, R. L. and Roudebush, P.): Small animal clin-
ical nutrition. 4th ed. Walsworth, Marceline, Missouri.
Cheeke, P.R. (1987): Rabbit feeding and nutrition. Animal feeding and nutrition. A series
of monographs. Academic Press, New York-London
Fekete, S.Gy., Fodor, K., Prohaczik, A. and Andrasofszky, E. (2005): Comparison of feed
preference and digestion of three different commercial diets for cats and ferrets. J.
Anim. Physiol. a. Anim. Nutr. 89, 199-202
Fox, J.G. (1998): Biology and diseases of the ferret. 2nd ed., Williams & Wilkins, Baltimore.
Jolankai, R.; Iben, Ch and Fekete, S.(2006): Relation of urinary cristal formation and feed
ing in the guinea pig (in Hungarian with English summary). Magyar Allatorvosok
Lapja. 128, 232-238
Isenbugel, E. and Frank, W. (1985): Heimtierkrankheiten. Ulmer, Stuttgart.
Matin, C.M. and Ostwald, R. (1975): Food intake and growth of guinea pigs fed a choles-
terol-containing diet. J. Nutr. 1975; 105, 525-533.
Wolf, P. and Kamphues, J. (2003): A critical assessment of commercial supplementary feed-
stuffs for rabbits, guinea pigs, chinchilla as companion pets. Praktische Tierarzt.
84(9): 674-678.
Wolf, P., Schroder, A., Wenger, A. and Kamphues, J. (2003): The nutrition of the chinchilla
as a companion animal - basic data, influences and dependences. J. Anim. Physiol.
a. Anim. Nutr. 87, 129-133.
1072
Chapter XXXII
CAGE BIRD AND PIGEON FEEDING AND NUTRITION
© Geert P.J. Janssens and James Sales
32.1. Cage birds

32.1.1. Digestive physiology
32.1.3. Diets for cage birds
32.1.4. Nutrient deficiencies
32.1.5. Nutritional status
32.1.6. Correcting nutritional imbalances
32.2. Pigeons
32.2.1. Crop milk and hand rearing
32.2.2. Nutrient requirements and diets for adult pigeons
1073
CHAPTER XXXII: CAGE BIRD FEEDING AND NUTRITION
32.1. Cage birds

Introduction
Despite the popularity of psittacine and passerine birds as pets, knowledge

on their nutritional needs is scarce. Diets for cage birds are based on
extrapolations of the nutritional requirements for commercial poultry and
game birds, food habits of wild birds, and some information derived through
trial and error feeding. However, cage birds show different nutrient require-
ments, substantial differences in their development stage at hatch, growth
patterns, gastrointestinal physiology, and different clinical symptoms asso-
ciated with specific differences in nutrient deficiency compared to domestic
poultry. The aim of commercial poultry production is to minimize cost while
maximizing production of meat and eggs over a short period of time, where-
as longevity and good health are critical in cage birds.
In the wild, birds eat a mixture of feeds, often containing plant and
animal material, creating complete and balanced diets due to the variety
and seasonal changes in natural foods. In captivity such variety is seldom
offered, and opportunity for natural feeding behaviours, such as foraging
and browsing, are denied. Despite the problems of species diversity, variety
in feeding behaviours, sociability and activity, and lack of information from
controlled trials, we nevertheless expect cage birds to live in constrained
habitats, and to consume diets of questionable nutritional value. There is
an urge for dietary guidelines to help aviculturists and cage bird owners to
feed cage birds properly, to guide the commercial food manufacturers in
producing diets that can assure excellent health and longevity, and to help
veterinarians assess a patient’s diet and educate the client in proper feed-
ing methods.
1074
32.1.1. Digestive physiology
The gastrointestinal morphology differentiates the feeding strategies of cage

bird species into granivory, frugivory, florivory, nectarivory or omnivory
(Table XXXII-1). The digestive tract of the bird starts at the beak, followed by
a toothless mouth, tongue, pharynx, oesophagus, crop, proventriculus, giz-
zard, intestine, rectum, cloaca, and vent. Each part of the tract presents a
specific role in acquiring, digesting, and absorbing nutrients, or for
immunocompetence.
Table XXXII-1: Eating habits of some captive kept cage birds.
Dietary nutrient concentrations have to be balanced according to energy

density to ensure adequate nutrient intake. Birds will generally eat an
amount to satisfy their daily energy expenditure when food is available ad
libitum and provided all other nutrients in the diet are balanced. The
amount of dietary metabolizable energy (ME) needed to support basal
metabolism, together with additional energy to fuel activity and thermoreg-
ulation, is described as the maintenance energy requirement. Growing
birds also need energy to support the accretion of new tissues, reproducing
birds need additional energy for accretion of gametes and egg production,
and moulting birds need energy to support feather growth. Two different
equations have been recommended to calculate the maintenance energy
requirements (kJ/d) of cage birds from live weight (LW):
1075
(1) Birds less than 100 g = 2.59×LW1.1
(2) Birds between 100 and 1500 g = 18.95×LW0.55
The nitrogen requirements of the bird have to be met by an essential

level of protein included in the diet. Too much protein is associated with
articular and visceral gout, probably due to inadequate removal of excessive
dietary nitrogen by uric acid. Too low dietary protein levels resulted in more
body fat and greater mortality. A higher level of protein is required with seed
protein sources if the amino acid found in the lowest proportion compared
to dietary requirements (first limiting amino acid) is not supplemented. The
physiological state of the bird will determine the quantitative requirements
for amino acids. Requirements are being lowest in adults at maintenance
and highest in hatchlings and females laying large clutches of eggs, where-
as requirements for amino acids are further increased during the period of
feather development. Different dietary protein concentrations found as opti-
mal through experimental work in some cage birds are presented in Table
XXXII-2. It has been demonstrated that considerable differences exist
between canaries, budgerigars and lovebirds.
Table XXXII-2: Level of protein shown to be adequate for cage birds at a given physiologcal
1076
Calcium, necessary for bone mineralization, metabolic homeostasis,

and eggshell calcification, is needed in greater quantities than any other
mineral. Vitamin A is needed, among others, for vision, cellular differentia-
tion, and immune function. However, requirements for minerals and vita-
mins in cage birds are unknown, and recommendations are done according
to values derived for commercial poultry.
Unfortunately the norm is to judge the nutrient adequacy of a diet

strictly on the total amount of a nutrient in the food. To optimize health,
longevity and reproduction of cage bird species, it will be necessary to eval-
uate both the intake and bioavailability of the nutrient to the bird. As sev-
eral cage bird species husk most seeds before swallowing, calculation of
nutrient intake is a complicated task. Protein quality varies on the basis of
amino acid balance and digestibility. Protein digestibility of different seeds
commonly used for cage birds is shown in Table XXXII-3. Rather than
attempting to supplement with one or two amino acids, improving the over-
all amino acid profile and balance through addition of high quality protein
appears to be safer.
Table XXXII-3. Digestibility of protein in seeds determined with adult cage birds.
1077
32.1.3. Diets for cage birds
The general believe is that an all-nut and -seed diet is a complete diet for
most species of psittacine birds kept as cage birds. Most commercially avail-
able seeds are deficient in certain limiting nutrients (amino acids, vitamins,
minerals), and are not the primary natural diet of these birds. Nuts and
some seeds (sunflower, sesame, pumpkin) commonly fed to cage birds pre-
sented a very high fat and moderate protein content when taking energy
content in consideration (Table XXXII-4). Furthermore, seeds are charac-
terised by a low ratio of calcium to phosphorus, which might interfere in
skeletal development in growing birds and egg production. Seeds and nuts
also lack iodine and carotene. Most fruits and vegetables are safe for cage
birds, however, avocado has reported to be toxic. Excess fruit and vegeta-
bles lower the overall energy density of a diet and risk insufficient energy
intake. Commercial diets, often pelleted or extruded, attempt to provide
optimum nutrient levels for a wide range of species. Mineral supplements
and grit, that aids in the crushing and grinding of food in the wild, are not
considered as essential for cage birds that receive adequate nutrition. Hand-
feeding formulas, either commercial or homemade, for rearing of baby birds,
is purely based on trial and error feeding, and is outside the scope of this
chapter.
Table XXXII-4. Energy and nutrient content of nuts and seeds (hulls removed) commonly
fed to cage birds
1078
32.1.4. Nutrient deficiencies
One reason for cage birds to develop nutritional deficiencies is the tenden-
cy of individual birds to select specific food items from a variety of offerings.
This has resulted in the misconception that cage birds are able to preferen-
tially balance their diets. However, cage birds often select food items based
on texture and colour rather than nutritional content. As a result, individ-
uals may become habituated or fixated on a specific food item that is defi-
cient in a number of essential nutrients, such as sunflower or safflower
seeds.
Inadequate diets that result in nutritional deficiencies can lead to

immune dysfunctions, leaving cage birds susceptible to infectious diseases.
Furthermore, specific nutrient deficiencies may lead to metabolic or bio-
chemical dysfunctions that can produce noninfectious diseases such as
nutritional secondary hyperparathyroidism and thyroid hyperplasia or dys-
plasia. The most commonly recognized deficiencies of dietary nutrients in
cage birds are those of vitamins A and D3, and calcium, whereas protein
deficiency and intake of poor quality protein are seen less frequently.
Vitamin D3, the active form of Vitamin D for avian species, plays an essen-
tial role in calcium absorption and regulation, and clinical signs associated
with Vitamin D3 deficiency are characteristic of that of calcium deficiency.
As well as mineral deficiencies, cage birds are susceptible to mineral excess-
es. The most common disease that is believed to be related to excess min-
eral intake, is probably hemosiderosis (iron storage disease), of which the
cause (genetic and nutritional) is still unknown.
32.1.5. Nutritional status
A combination of physical examination and laboratory findings, dietary his-

tory (including frequency at which a feed is offered) and diet evaluation,
together with clinical experience and judgement, is used to assess nutri-
tional status. Signs of malnutrition might include irregular beak hyperker-
atosis, excessive beak length, thinning, swelling, peeling, and ulcerations of
the skin, and thick, keratinised epithelium on the skin, beak, and nails of
birds. Passeriformes often have large accumulations of exfoliated hyperker-
atotic scales on their legs and feet when malnourished. The feathers of mal-
nourished birds lose elasticity and may be bent or broken, malformed, or
1079
frayed. Accumulations of subcutaneous fat over the crop, sternum, or

abdomen indicate obesity. After a thorough physical examination, weigh the
bird and compare the weight to that estimated from palpating the pectoral
muscles. Birds should be scored for body condition according to fat: lean
ratio, degree of fat loss, and muscle depletion. Some clinical signs due to
nutrient imbalances are presented in Table XXXII-5.
Table XXXIII-5. Clinical signs attributed to some nutrient imbalances.
1080
32.1.6. Correcting nutritional imbalances
Gradually add the new food to the current diet, increasing the amount of
the new food over days to weeks, and use texture and colour to ease the
conversion. Whereas treatment with systemic corticosteroids has been rec-
ommended for patients suffering from vitamin D3 toxicosis, steroids wors-
en the symptoms of vitamin A toxicosis by prolonging high concentrations
of serum retinol and retinyl ester. This illustrates the importance of a cor-
rect differential diagnosis. Patients fed nutrient-deficient diets are often
best managed by switching to a commercial diet originating from companies
with nutritional expertise and established quality assurance. However,
some owners that are feeding imbalanced homemade diets, prefer to
improve the recipe rather than change to commercial products. The biologi-
cal value of protein can be improved by adding egg. Whereas fat content may
be increased by adding egg, cheese, meat (for saturated fatty acids), or veg-
etable oils (for polyunsaturated fatty acids), it will be decreased by adding
carbohydrates or low-fat, high-fibre foods such as vegetables, greens, or fruit.
The diet of an obese patient should be changed to one containing less than
10% (DM), with a gradual weight loss of not more than 1% of body weight
lost per week. Calcium deficiencies are corrected by adding limestone, cal-
cium salts, lactate, or gluconate. Vitamins A and D3 can be provided in cod
liver oil and in cooked liver. Vitamin E, often given to cage birds with feath-
er problems, may be added to diets with corn oil.
CONCLSIONS
Understanding cage bird nutrition is still in its infancy, and there is an

urgent need to establish to what extent available diets meet nutrient
requirements. Due to diversity in species kept, individual variation within
species, and individual preferences by owners, basic nutritional knowledge,
practical experience and common sense are critical factors in advice on
feeding of cage birds. A thorough evaluation of the avian patient is vital in
order to set nutritional goals and to design diets. This is not only valid to
cure nutritional imbalances, but also to prevent them.
1081
32.2. Pigeons
Introduction
Pigeons (Columba livia domestica) have been domesticated since 2500 BC by

man for a variety of reasons including sports (racing pigeons), aesthetics
(fancy pigeons), communication (messenger pigeons) and food (squab).
Furthermore, pigeons are used in modern times as experimental animals in
behavioural analyses and studies for the improvement of human health.
However, information on nutrient requirements and metabolism of nutri-
ents are lacking in pigeons. Nutritional research with pigeons is complicat-
ed by the fact that young growing pigeons (squabs) continuously stay in the
nest and are fed by the parents, initially with a special feed, the so-called
crop milk.
32.2.1. Crop milk and hand rearing
The Columbidae (pigeons and doves) regurgitate crop-milk from the crop by
both parents, a highly nutritive substance being mainly composed of pro-
teins and lipids (Table XXXII-6). A polypeptide with a molecular weight of
6000, the so-called pigeon-milk growth factor, has been identified as the
factor responsible for the relatively high growth rate of squabs.A few
attempts have been made to hand feed squabs on formulas containing 13
MJ/kg metabolizable energy and a crude protein content of 20 to 22%.
32.2.2. Nutrient requirements and diets for adult pigeons
The crude protein requirements of adults pigeons have been established as

between 12 and 18 % at an energy content of around 12 MJ/kg metaboliz-
able energy. Of the feeds normally fed to pigeons (maize, wheat, barley, red
and white millet, sorghum, canary seed, peas, lentils, sunflower and hemp)
intake was highest for peas, and lowest for sorghum and wheat (Table
XXXII-6). However, individual patterns of seed selection behaviour which
persist over prolonged periods, have been identified in pigeons. Digestibility
of grains and seeds (Table XXXII-7) are in agreement with values derived for
poultry where appropriated.
1082
Table XXXIII-6. Chemical composition of crop-milk.
1083
Table XXXIII-7: Intake, apparent metabolizable energy (AME) and digestibility values of
different feed ingredients for adult pigeons
Pigeon feeds mainly consist of whole grains and seeds whereas min-
erals, vitamins and some other nutrients are provided as separate supple-
ments. Pigeons on grain mixtures need grit to facilitate gizzard function. The
use of pelleted and extruded compound diets in pigeon nutrition is still lim-
ited. Drinking water after eating seems to help move grains and pellets from
the crop, and failure of access to water after feeding might result in crop sta-
sis.
Growth improvement of squabs was found when drinking water of

parents was supplemented with L-carnitine, probably due to a quantitative
impact on crop milk production. Pigeons are more sensitive to carbohydrase
enzymes than poultry when looking at nutrient digestibility, but unlike in
chickens, lactose does not seem to have a prebiotic activity against
Salmonella in pigeons.Most likely, the absence of functional caeca might
attribute to this difference.
CONCLUSIONS
The traditional feeding of adult pigeons on combinations of grains and

seeds, that are low in some minerals and vitamins are still valid in modern
times. However, minerals and vitamins are often offered as supplements.
1084
Knowledge on nutrient requirements and nutritional imbalances in pigeons

are extremely scarce, and based on trial and error feeding by owners.
FOR FURTHER READING
Donoghue, S. and Stahl, S. (1997). Clinical nutrition of companion birds. J. Avian Med.
Surg. 11, 228-246.
Fekete, S., Meleg, I., Hullár, I. and Zoldag, L. (1999): Studies on the Energy Content of
Pigeon Feeds II. Determination of the incorporated energy. Poult. Sci. 78, 1763-1767.
Hullar, I., Fekete S.Gy., Mezes, M., Gaspardy, A. and Febel,. H, (2008): Effect of oral L- car-
nitine, L lysine administration, and exercise on body composition and histological and
biochemical parameters in pigeons. J. Anim. Physiol. Anim. Nutr. 92, 411-418.
Janssens, G.P.J., Millet, S., Van Immerseel, F., De Buck, J. and Hesta, M. (2004): The
impact of prebiotics and salmonellosis on apparent nutrient digestibility and
Salmonella typhimurium var. Copenhagen, excretion in adult pigeons (Columba livia
domestica). Poult. Sci. 83, 1884-1890.
Klasing K.C. (1998). Comparative Avian Nutrition. CAB International, New York, NY, USA.
Kollias, G.V. (1995). Diets, feeding practices, and nutritional problems in psittacine birds.
Vet. Med. 90, 29-39.
Koutsos E.A., Matson K.D. and Klasing K.C. (2001). Nutrition of birds in the order
Psittaciformes: A review. J. Avian Med. Surg. 15, 257-275.
Nott, H.M.R. and Taylor, E.J. (1994). Advances in our understanding of the nutrition of pet
birds. Wiener Tierärzt. Monatsschrift 81, 135-140.
Sales, J. and Janssens, G.P.J. (2003). Energy and protein nutrition of companion birds.
Flemish Vet. J. 72, 51-58.
Sales, J. and Janssens, G.P.J. (2003). Nutrition of the domestic pigeon (Columba livia
domestica). World’s Poult. Sci. J. 59, 221-232.
Ullrey, D.E., Allen, M.E. and Baer, D.J. (1991). Formulated diets versus seed mixtures for
psittacines. J. Nutr. 121, 193-S205.
Wolf, P., Rabehl, N. and Kamphues, J. (2003). Investigations on feathering, feather growth
and potential influences on nutrient supply on feather’s regrowth in small pet birds
(canaries, budgerigars and lovebirds). J. Anim. Physiol. a. Anim. Nutr. 87, 134-141.
1085
Chapter XXXIII
FEEDING AND NUTRITION

OF LABORATORY RODENTS
33.1. Zoological features, nutrient requirements

33.1.1. Introduction
33.1.2. Feeding and nutrition of mouse and rat
33.1.3. Feeding and nutrition of guinea pig
33.2. Actual topics in on the feeding and nutrition of laboratory

rodents and rabbits
33.2.2. Diet and experimental results
33.2.3. Diet and well-being
33.2.4. Diet and animal models
33.2.5. The impact of a regular feeding schedule on circadian
rhythms of physiological and behavioural functions
1087
CHAPTER XXXIII: LABORATORY RODENTS’ FEEDING
33.1. Zoological features, nutrient requirements

Particulars, concerning the feeding and nutrition of rabbit, the small pet
mammals, fishes, amphibian and reptiles are given in Chapter XXX.
Herewith the zoological features, digestive characteristics and the nutrient
requirements of the most important laboratory rodents are reviewed. (It
worth mentioning that most of these species are pets, too.) One of the most
important criteria of the classification of the small mammals is the zoologi-
cal characteristics whether their offsprings are altricial (=having the young
hatched or born in a very immature and helpless condition so as to require
care for some time; altrix=nurse, alere=to nurish in Latin, WEBSTER, 1993.
Syn.: nest-living) (e.g. mouse, rat, hamsters, rabbit) or precocial (=capable
of a high degree of independent activity from birth; syn.: nest-leaving or nid-
ifugous) (e.g. guinea pig, gerbil). In the first case the nutrient requirements
of lactating dam are extremely high; in the second case the mother’s gesta-
tion feeds and the creep feed of the suckling are of paramount importance.
Basic reproductive particularities are summarized in Table XXXIII-1.
Labelling of their feed mix may be closed, when only the nutrient com-
position and energy concentration are given or open while even the level of
the natural components are displayed. The open labelling is more appropri-
ate to the feed mixtures of the laboratory animals. Evaluation of the follow-
ing nutrient requirements was made on the basis of the biological answers,
but instead of growth rate, production and nutrient utilization (see produc-
tion animals), the reproductive performance and the longevity were empha-
tically considered.
1088
Table XXXIII-1: Basic data of the rodents’ reproductive physiology
The energy concentration of the mixed feed (13-18 MJ DE/kg) is of

small importance, because most of the rodents eat on energy, i.e. their vol-
untary dry matter intake is adjusted to the actual energy needs.
Consequently, they can generally be fed ad libitum. The most important
exception for this regulation is the guinea pig. In some types of trials,
restricted feeding is recommended from a scientific point of view (BEYNEN,
1992). The overall protein requirement is 12 to 14% ideal protein in the feed
dry matter, which is equivalent to approximately 18-22% crude protein.
Possible discrepancies or small deficiencies of the protein supply can be
alleviated by the phenomenon of coprophagy (mouse, rat) or by the cae-
cotrophy (guinea pig, rabbit). Rabbit, chinchilla, degu, guinea pig have high
fibre requirements (10 to 15%), hamsters do very low (2 to 4%) and the oth-
ers (mouse, rat etc.) are in the middle. The nutrient requirements are
extremely influenced by the genotype (e.g. ob/ob mice, fa/fa rats, nu/nu
mice etc.), as well as by the microbiological status of the animals. Feeds for
SPF or germ-free colonies should be sterilized, which unequally destructs
vitamins. Therefore the gross vitamin levels before the treatment should be
proportionally higher. The use of synthetic antioxidants is not allowed,
because of the possible interaction of other feed components. Above the
items, prescribed for the production animals, the inositol and molybdenum
requirements are given, too.
Feeds for laboratory animals can be based on natural ingredients
(cereal grains, oilseeds, fish meal etc.), on the derivates of natural feed-
stuffs, like starch from potatoes, maize, sugars, casein, animal fats and veg-
etable oils, cellulose from cotton (these are the semisynthetic or purified
diets) and finally on chemically exactly defined components, for example
amino acids, glucose, sucrose, triglycerides, essential fatty acids, industrial
1089
minerals and vitamins (synthetic diets). In case of toxicological test the use
of natural ingredient diet may profoundly alters gene expression and con-
founds genomic analysis, therefore the use of purified diet is indispensable
(KOZUL et al. 2008). The physical form (“presentation”) of the laboratory ani-
mal diets may be different, according to the experimental purposes: floury,
meal, pelleted, crumbled, extruded, cooked („biscuit”, “cake”), flaked (most-
ly for fishes), “popped”, semi-moisture or gel (“jelly”) and liquid.
The digestibility of nutrients and the urinary energy losses in rodents
are similar, or better than in swine. Table XXXIII-2 summarizes the DE and
ME values of the most important raw materials for rats. On an average, the
DE value is 90% of the GE and the ME value is 94% of the DE (NELSON et
al. 1974). The Djungarian hamster has a very effective digestion: the
digestibility of dry matter is 91%, that of the crude protein 80% and 92% for
GE (SCHIERWATER and KLINGEL, 1984). According to the ad libitum or restrict-
ed feeding, the coprophagy or prevention of the coprophagy/cacotrophy may
influence the digestibility of feed but has no impact on passage rate of diges-
ta (WILLIAMS and SENIOR, 1985).
The most frequent feed-related syndromes are the vitamin C deficien-
cy (fishes, guinea pig), pregnancy toxicosis (guinea pig), increased suscepti-
bility to the mycotoxins (many species) and hair ball (zootrichobezoar) for-
mation in the stomach.
Table XXXIII-2: Digestible and metabolizable energy content of feed ingredient for growing
rat (NELSON et al. 1974)
1090
33.1.2. Feeding and nutrition of mouse and rat
MOUSE
The mouse is the most frequently used laboratory animal, but it is kept as
a pet, too. Growth rate, adult live weight of the different strains are signifi-
cantly different (Table XXXIII-3), which should be taken into account during
the feeding. The NRC (1993) published the nutrient requirement of the
“common” mouse (Table XXXIII-4).
Table XXXIII-3: Growth rate of some well-known mouse strains (♀/♂), after POILEY, 1972
*Wild and outbred mouse have an average live weight of 27 grams.
1091
Table XXXIII-3: Recommended minimum energy and nutrient concentration for growing
mice in air dry feed mixture (after NRC, 1995): supposed a feed mixture of 10% moisture
and 16-17 MJ ME/kg
The average daily feed intake of the weaned mouse during the subse-
quent 2 to 4 weeks is 3 to 4 grams, which assures 0.5 to 1.0 gram average
daily gain. The relatively low protein and amino acid levels are valid only, if
protein of high biological value is fed.
1092
RAT
To the digestive physiology of rat is highly a characteristic that animals eats
according to their actual energy need and generally are capable of adjusting
their voluntary feed intake to the energy concentration of the feed mixture.
They are not too demanding to the taste or physical form of the diet, conse-
quently their needs can be covered both using natural ingredient and syn-
thetic compounds. Table XXXIII-5 summarizes the NRC (1995) recommen-
dations for the different rat categories.
Table XXXIII-5: Energy, nutrients and mineral requirements of rats on air-dry matter basis,
16-17 MJ ME/kg energy concentration
“?” the maintenance needs are not known;* in case of feeding on ideal protein (e.g. lactal-
bumin); using natural ingredients 18 to 25% ** coprophagy allowed
1093
33.1.3. Feeding and nutrition of guinea pig

The feed mixture for the guinea pig should basically be composed using
ingredients of plant origin. To assure the healthy functioning of the diges-
tive tract, the crude fibre supply is of high importance. The minimum rec-
ommendation is 10% fibre of low digestibility in the feed dry matter. The
dietary allowances are given in Table XXXIII-6. One of the widely-accepted
recipe of the praxis recommends 35% of alfalfa meal, 12% of soybean meal
(extracted, solvent), 25.25% oats, 23.6% wheat, 1.5% soya oil, 0.55% dical-
cium phosphate, 1.0% calcium carbonate, 0.75% common salt and 0.45%
species-specific micromineral and vitamin premix.
Guinea pigs produce two types of faeces: the normal faeces is light-
brown coloured, thick crescent-shaped and of 10 to 12 mm length, the cae-
cotrophy, on the contrary is approximately 8 mm long, black and its shape
is like the seed of the cucumber. The caecotroph is not directly ingested
from the anus, but after the excretion it is eaten from the bottom of the
cage. Opportunity of the caecotrophy may be considered as part of the envi-
ronmental enrichment (see 3 “R”). Unlike to the majority of rodent the ad
libitum feeding predisposes guinea pig to obesity.
Table XXXIII-6: Dietary recommendation for growing guinea pigs on air-dry matter basis
(NRC, 1995)
Guinea pigs are well-known not to be capable of synthesizing ascor-

bic acid (vitamin C), therefore the continuous external (generally: dietary)
supply is indispensable. Signs of the vitamin C deficiency in adult animals
are haemorrhagic diathesis and scurvy. In addition, the development of
long, tubular bones (mostly the femur and tibia) retards in the growing ani-
1094
mals, with the consequence of the atrophy of the associating muscles

resulting in prone position. The daily vitamin C requirement is 4 to 10
mg/animal or 200 mg/kg air-dry feed. A regular green feeding would also
cover the needs, but it is difficult to use in a laboratory animal facility. The
average ascorbic acid content of some feeds is given in Table-XXXIII-7.
Table XXXI-7: Ascorbic acid content of feeds, freshly harvested (HOFFMANN et al. 1995)
*samples from the feedmills
FOR FURTHER READING
AIN Committee (1977): Report of the American Institute of Nutrition Ad Hoc Committee on
standards for Nutritional studies. J. Nutr. 107, 1340-1348.
Beynen, A.C. (1992): Ad libitum versus restricted feeding: Pros and cons. Scand. J. Lab.
Sci. 19, 19-21.
Hedrich, H.J. and Bullock, G. (2004):The laboratory mouse. Elsevier Academic Press. Pp
463-482.
Hoffmann, P. Schai, E. and Canfantaris, B. (1995): Vitamin C content in feedstuffs. Poultry
Intern. October, 46-47.
Kozul, C.D., Nomikos, A.P., Hampton, T.H., Warnke, L.A., Gosse, J.A., Davey, J.C., Thorpe,
J.E., Jackson, B.P., Ihnat, M.A. and Hamilton, J.W. (2008): Laboratory diet pro-
foundly alters gene expression and confounds genomic analysis in mouse liver and
lung. Chemico-Biol. Interact. 173, 129-140.
Nelson, T.S., May, M.A. and Miles, R.D.Jr. (1974): Digestible and metabolizable energy con-
tent of feed ingredients for rats. J. Nutr. 38, 554-558.
NRC (1978; 1995): The nutrient requirements of laboratory animals. 3rd and 4th Ed.
National Academy Press. Washington, D.C.
Poiley, S.M. (1972): Growth tables for 66 strains and stocks of laboratory animals. Lab.
Anim. Sci. 22, 759-799.
Reeves, P.G., Nielsen, F.H. and Fahey, G.C.Jr. (1993): AIN-93 purified diets for laboratory
rodents: final report of the American Institute of Nutrition Ad Hoc committee on the
Reformulation of the AIN-76A rodent diet. J. Nutr. 123, 1939-1951.
Schierwater, B. and Klingel, H. (1985): Food digestibility and water requirements in the
Djungarian hamster Phodopus sungorus. Z. Säugertierkunde 50, 35-39.
Seki, M., Yamaguchi, K., Marumo, H. and K. Imai (1997): Effect of food restriction on repro-
ductive and toxicological parameters in rats – in search of suitable feeding regimen
in long-term tests. J. Toxicol. Sci. 22, 427-437.
Williams, V.J. and Senior, W. (1985): The effect of coprophagy in the adult rat on rate of
passage of digesta and on digestibility of food fed ad libitum and restricted amount.
J. Nutr. 115, 1147-1153.
1095
33.2. Actual topics in on the feeding and nutrition of labora-

tory rodents and rabbits*
©Merel Ritskes-Hoitinga, Bart Savenije and Burghart Jilge
Practical experience from teaching in laboratory animal science courses has

shown that students (and their supervisors) are often not conscious
(enough) about the influence of diet and dietary composition on the health
of the animals and experimental results. This sometimes leads to the exe-
cution of experiments in which the diets used are such, that the results do
not have any meaning and cannot be published. This is inappropriate use
of laboratory animals.
This overview will hopefully add to the understanding of the impor-
tance of laboratory animal nutrition, avoiding doing experiments using
inappropriate diets and thus unnecessary use of laboratory animals.
Thereby this short overview is expected to contribute to the refinement of
animal experiments, one of the important goals of FELASA.
33.2.2. Diet and experimental results
a. Nutrient requirements
In order to provide each species with the proper nutrient levels of essential
nutrients, nutrient requirements must be fulfilled (NRC documents describe
nutrient requirements for each species). By providing each animal with their
species-specific essential nutrients in the proper amounts, diseases can be
prevented as well as unwanted interference with experimental results. In
case essential nutrient requirements are not fulfilled, unreliable conclu-
sions may be obtained (RITSKES-HOITINGA et al. 1996, RITSKES-HOITINGA 2000).
*based on the FELASA Quick Reference Paper (Text compiled: June 2000 / updates
November 2000, February 2001, and July 2007)
1096
Commercial pelleted diets provide well over the minimum nutrient

requirement levels. This avoids the risk of nutrient deficiencies, but too high
levels of ingredients may cause problems as well. MARTUS et al. (2005)
reported that some chows may contain up to seven times the amount of
sodium that rats would naturally consume, with elevated blood pressure as
a result.
b. Nutrient requirements in different (transgenic) strains
Different species, strains, stocks and individuals can have different nutri-
ent requirements (NRC, 1995). Regarding the enormous development of
many new transgenic strains, it must be taken into consideration that
depending on the nature of a transgenic strain, nutrient requirements may
vary as well.
c. Standardisation
Standard commercial diets usually fulfil nutrient requirements more than
sufficiently, at least when transported and stored under the proper envi-
ronmental conditions. However, there can occur a large variation in compo-
sition in natural-ingredient diets (between and within brand variation) due
to raw material variation, which will differentially influence experimental
results (BEYNEN et al. 1993, RITSKES-HOITINGA et al. 1991). As between-batch
variation can occur, it is advised to buy diets with a batch-analysis certifi-
cate so that one is informed about the actual composition of each batch of
diet that is being used. For GLP-studies this is a necessity.
Purified diets (BEYNEN et al. 1993), formulated with a combination of
natural ingredients, pure chemicals and ingredients of varying degrees of
refinement, have a more standardised composition and give therefore more
reproducible results than the use of natural-ingredient diets. However,
there is a higher risk of creating shortages of unknown essential nutrients,
which are present as “natural contaminants” in natural-ingredient chow
diets (e.g. Chromium and Vanadium, NRC, 1995). Moreover, certain refined
ingredients can cause problems (e.g. short-type cellulose fibre can cause
intestinal obstruction in rats, SPEIJERS 1987).
For rodents a “cook-book recipe” is available for composing a purified
diet, the so-called American Institute of Nutrition diet (AIN-93 diet) (REEVES
et al. 1993). The AIN-93 diet fulfils the nutrient requirements for rodents as
published in 1995 (NRC), except for the vitamin B12 level. The AIN-93 vita-
min B12 level must be doubled to live up to the minimum requirements as
described by the NRC (1995). In general, rats fed the AIN-93 diet perform
1097
equally well as rats on a chow diet, although serum cholesterol and triglyc-
eride levels may be elevated (Lien et al. 2001). Survival rate and body weight
were similar in rats fed chow or AIN-93 diets, respectively, in a two-year
study when fed ad libitum. Maintaining sufficient vitamin levels is important
when feeding the AIN-93 diet restrictedly for two years (Duffy et al. 2002).
Ad libitum food intake is in principle determined by the energy need.
This energy need changes according to the stage of life the animal is in
(growth, maintenance, pregnancy, lactation). When changing the energy
content of the diet (e.g. by adding fat to the test diet), one changes the
dietary intake in grams. In order to make sure that only the dietary fat (and
carbohydrate) intake will differ between the control and test group, one
needs to apply the isocaloric exchange method (BEYNEN and MEIJER, 1993).
d. Feeding level
Ad libitum feeding is considered normal practice for rodents, however it is
considered bad veterinary practice for e.g. pigs, monkeys, rabbits and dogs,
as they become obese (HART et al. 1995). When feeding restrictedly, it must
be secured that the restricted feeding level provides enough essential nutri-
ents. Although ad libitum feeding of rodents is considered “normal” practice,
this must be questioned intensely. Ad libitum feeding as opposed to restrict-
ed feeding has a clear negative impact on rodent health, as it shortens sur-
vival time, increases cancer incidence, shortens cancer latency period and
increases the incidence of degenerative diseases in kidney and heart (HART
et al. 1995). Obesity in rats impairs the animal’s immune function, which is
not seen in normal and food restricted rats (LAMAS et al 2004). These effects
are very reproducible! Moreover, it increases the number of animals needed
if sufficient animals are to survive a 2-year period in long-term toxicological
studies. Moderately restricted feeding avoids or minimizes these problems.
Severe food restriction results in even higher two-year survival rates that
moderate restriction, but this will create deficiencies in the animal (HUBERT
et al, 2000). A meta-analysis by DIRX et al (2003) revealed that caloric
restriction reduces the risk of spontaneous mammary tumours in mice with
55%. This reduction is achieved irrespective of the energy-providing nutri-
ent (fat, carbohydrate, or protein). The physiological mechanisms that are
responsible for these benefits with restricted feeding are as yet unknown.
Research on the mechanisms involved focus on the glucose metabolism and
oxidative stress. A thorough review of past and current theories on the
mechanisms involved with caloric restriction has been published by MASORO
1098
(2005).
KEENAN et al. (1999) state that ad libitum overfeeding of rodents is at
present one of the most poorly controlled variables affecting the current
rodent bioassay. Moderate dietary restriction (70-75% of adult ad libitum
food intake) is advised as a method that will improve uniformity, increase
exposure time and increase statistical sensitivity of chronic bioassays to
detect true treatment effects (KEENAN et al. 1999; LEAKEY et al. 2003a,b,
RITSKES-HOITINGA, 2006). MORIYAMA et al (2006) showed that moderate food
restriction improved uniformity in body weight and some serum parameters
already after four weeks. However, moderate dietary restriction will only
improve uniformity in individually housed animals, where there is control of
individual food intake. A restricted amount of food in group-housed animals
is expected to increase variation due to differences in individual food
intakes, based on the hierarchy in the group. It will be the challenge to find
restricted feeding schedules in group-housed animals, in order to fulfil the
animal’s social needs as well.
e. Contaminants
There are several documents stating maximum allowed concentrations of
contaminants (GV-Solas 1980, BARQA 1992). One of the guidelines that give
maximum limits, to which all toxicologists all over the world are referring to,
are issued by the Environmental Protection Agency (1979). As different
guidelines state different levels, what to choose as the “correct” maximum
tolerated levels? Firstly, one has to decide which guidelines are most appro-
priate in the experimental setting one is working in. One might even have to
develop specific institutional guidelines. Secondly, for each experiment one
can do a literature search to figure out whether contaminants, and if yes,
which will interfere with the specific purpose of that study. That way con-
crete maximum levels of specific contaminants can be established. Even so,
natural-ingredient diets may contain contaminants that are unknown or
normally not accounted for, such as phytoestrogens from soy (NYGAARD
JENSEN and RITSKES-HOITINGA, 2007). Purified diets have lower contaminant
levels than natural-ingredient diets.
33.2.3. Diet and well-being

a. Welfare and enrichment
From preference testing it is known that rats prefer to work for food instead
1099
of obtaining it just like that. For each species there are certain species-spe-
cific essential needs connected to searching and finding food. If these essen-
tial needs are not fulfilled, abnormal behaviour like stereotypies can occur.
Enrichment of the environment is possible by letting the animals
work and or search for food. Knowledge of the natural feeding time and
behaviour are important factors to consider. The time of day at which a
restricted amount of food is given can be an important tool in providing a
better welfare (e.g. in rabbits, KROHN et al. 1999). Giving food rewards is an
important tool to learn and train animals. Which food rewards are chosen
and in what amounts need careful consideration: is there interference with
the experimental results and or health of the animal?
Certain dietary schedules require individual housing. As individual
housing opposes the wellbeing of social living species, alternative ways of
feeding need to be considered. E.g. the animals can be individually fed for a
certain period each day and then socially housed for the remaining part of
the 24-hour period.
b. Transport and acclimatisation
Knowledge of the species is important when transporting animals. Before
transport, getting specialist advice for each particular species is needed.
Rats and mice will acclimatise faster after transport, when food and water
has been provided during the transport (VAN RUIVEN, 1996).
33.2.4. Diet and animal models

a. Choice of model and experimental conditions
Knowledge and choice of species and experimental (including dietary) con-
ditions will have a major impact on results. The effect of linoleic acid on
mammary tumour development in animal models depended on the model
system used and type of parameters measured (RITSKES-HOITINGA et al.
1996). Feeding fish oil to rabbits to examine the possible positive influence
of fish oil on atherosclerosis resulted in liver pathology and more athero-
sclerosis on higher doses of fish oil. This was thought to be the result from
the inability of the herbivorous rabbit liver to cope with the long-chained
unsaturated fatty acids from fish oil (RITSKES-HOITINGA et al. 1998b).
Feeding by gavage is expected to cause stress, influences metabolism
and will therefore lead to other results than voluntary intake (VACHON et al.
1988). VACHON et al. proved that voluntary intake of a certain meal gave
results similar to the human, whereas giving the same meal by gavage did
1100
not (VACHON et al. 1988)! Food restriction may lead to decreased body tem-
peratures in mice due to a shift in the animal’s metabolism and behaviour.
Reduced energy intake leads to an increase in activity in mice. These
changes are strain dependent (RIKKE et al 2003; GELEGEN et al 2006).
b. Diet and pharmacological studies
The effect and pharmacokinetics of pharmacological substances (e.g. oral
antibiotics) are largely dependent on the time of administration in relation
to the time of feeding. How long animals need to be fasted before the “bare”
effect of pharmacological substances tested can be judged, is an important
animal welfare issue (CLAASSEN, 1994). A rat will have an empty stomach
already after 6 hours (VERMEULEN et al. 1997). Fasting for longer periods led
to increased locomotor and grooming behaviour (VERMEULEN et al. 1997).
33.2.5. The impact of a regular feeding schedule on circadian

rhythms(T) of physiological and behavioural functions [ (T)
(Some chronobiological terms are explained at the end of the text]
When individuals of several strains or species of rodents and rabbits are fed
ad libitum they tend to accumulate adipose tissue at various parts of the
body, predominantly in the peritoneal cavity. This is especially so with
increasing age and limited space for physical workout (e. g. NZW rabbits
kept in cages during longtime maintainance). It is well known that in
humans and animals a greater amount of peritoneal adipose tissue not sole-
ly is representing an energy store but in addition has to be regarded as an
endocrine organ secreting a couple of different adipocytokines (leptin,
adiponectin, resistin etc: HALAAS et al. 1995, FRIEDMANN, 1998, ELMQUIST and
FLIER, 2004, YU and GINSBERG, 2005) and causing the metabolic syndrome
(VETTOR et al. 2005, ROBINSON and GRAHAM, 2004). In addition anesthesia and
surgery of fat individuals is rather more complicated than compared to lean
ones. Thus, an excess of adipose tissue does not appear to be advantageous
in men nor in animals. In order to prevent excessive fattening, most often
the quantity of food is restricted: usually a limited amount of food is replen-
ished at various times during the working hours of the caretaker.
As a rule after several hours of fasting the animals start to eat eager-
ly immediately when food is replenished. In consequence, many biochemi-
cal and physiological functions of the gastrointestinal tract and other
organs are phase-shifted and can even be inverted, and nocturnally active
1101
animals may thus become equivalent to light active animals. Since the
impact of shifted or inverted circadian rhythms on experiments usually is
underestimated this paragraph intends to compile some few basic informa-
tion on the impact of periodic food restriction on circadian functions.
Supplied with feed ad libitum, nocturnally active animal species like
the mouse, rat, hamster and rabbit are consuming most of their food dur-
ing the hours of darkness. Correspondingly many follow-up parameters are
on a significantly higher level during the hours of darkness. The left column
of figure 1a demonstrates that 60-80 % of the functions monitored are
expressed during the 12 h dark period. When feed access is restricted to 4
h during the hours of light, almost all of these functions are shifted to the
hours around food access during light time (FIGURE XXXIII-1a, b). One
direct follow-up parameter would be the secretion of mucosal digestive
enzymes: it was shown by SAITO et al 1975 that they, too, are shifted to the
hours of light when food access is restricted to some hours during the light
time (FIGURE XXXIII-2). In the same way carbohydrate absorption (HARA
and SAITO, 1989), bile flow and composition (HO and DRUMMOND, 1975),
serum gastrin and cholecystokinin (PASLEY et al. 1987) and serum insulin
(RUBIN et al. 1988) is shifted to food access. During ad libitum feed access
the differences between the regular peak (highest) and trough (lowest) val-
ues can amount to several hundred percent, and the day-to-day sequence
of peaks and troughs of circadian functions is quite regular. When a regu-
lar schedule of restricted food access is controled by means of a program-
mable computerized feeder, the feeding-entrainable rhythms are regular as
well, the 24-h amplitudes as a rule are conspicuously higher. In either case
the rhythms are quite significant and predictable and it would be an arte-
fact to ignore them. Quite unpredictable ups and downs of functions would
be obtained, if the animals are fed by hand with a large day-to-day differ-
ence of timing. In this case one would expect a large amount of variability
and, thus, unprecise and non-valid results at least of those functions dis-
cussed here.
The phase of periodically restricted food access (RF) in most of physio-
logical functions evidently does override the light:dark regimen, which dur-
ing ad libitum feed access is the main ´zeitgeber´ (T) for circadian rhythms of
animals and men (PHILIPPENS et al. 1977, 1988, SAITO et al 1976 a,b, SAITO et
al. 1980, STEVENSON et al. 1975, STEVENSON and FIERSTEIN 1976). Even many
of those functions that are not so obviously coupled to feed intake, e. g. the
1102
24 h rhythm of corticosteroids (KRIEGER, 1974 (FIGURE XXXIII- 3), locomo-

tor activity (BOULOS and TERMAN, 1980, BOULOS et al. 1989, HONMA et al.
1983, JILGE et al. 1987, JILGE 1992, JILGE and STAEHLE, 1994, JILGE and
HUDSON, 2001), core body temperature (JILGE et al. 2000, 2001 a,b, 2007;
MORIMOTO et al. 1979; TAKAHASHI, 1979), heart rate and blood pressure (VAN
DEN BUUSE 1999) are phase-shifted by RF. Recent experiments with rabbits
demonstrated that not only in rabbit pups but also in adults core body tem-
perature is set by RF: body temperature exceeds the 24-h average starting
about 3 h prior to food access. The anticipatory component was shown to
be of endogenous origin: it persisted even during 72-h of fasting. In the
same way endocrine functions are controlled by RF: in rabbit, the time of
birth and the time of nursing are tightly coupled to the time of food access
(FIGURES XXXIII-4a,b,c; 5a,b). In contrast, during constant dim light the
time of parturition is distributed almost evenly around the 24 h days (JILGE,
1995). All these facts show that time-restricted feed availability has a most
significant effect on phase and amplitude of 24-h rhythms of many physio-
logical functions.
There are two ways how restricted feed access affects circadian
rhythms: masking and entrainment(T) (ASCHOFF, 1986, ASCHOFF et al. 1982,
ASCHOFF and VON GOETZ, 1986, MROSOVSKY, 1996, 1999, PITTENDRIGH and
DAAN, 1976).
MASKING
Masking means that a periodic environmental factor directly controls
the overt rhythm without affecting the circadian oscillator(T) that drives it
(ABE et al. 1989; ASCHOFF and von GOETZ 1986). As a result the rhythm is
synchronized immediately, without transients. When e.g. the circadian
rhythm of locomotor activity, which is free-running in constant conditions,
is exposed to scheduled feed access, the activity rhythm immediately stops
to free-run(T) and re-assembles around the phase of feed access within 1–3
cycles. When after some days feed is offered ad libitum again, the circadian
rhythm continues to free-run at the phase which it had without an inter-
spersed food regimen, i.e. at the phase which can be predicted by linearly
extrapolating the free-running rhythm before RF.
ENTRAINMENT
Entrainment means that an external variable has in fact zeitgeber
properties (for the definition of zeitgeber see: ASCHOFF 1958, 1960;
PITTENDRIGH, 1960). Therefore, if scheduled feeding is a zeitgeber, it acts on
1103
the oscillator system itself which controls the timing of overt rhythms. In
general, the time necessary for entrainment can last up to 30–60 days. The
time of consolidation depends on the time difference between corresponding
phases of zeitgeber and circadian rhythm (PITTENDRIGH and DAAN, 1976;
JILGE et al. 1987, JILGE and STAEHLE, 1993, JILGE, 2000). When returning to
ad libitum feed access again, a free-running rhythm starts out from the
phase of the preceding food regimen. While the honey bee was the first ani-
mal in which entrainment of an oscillator with scheduled feeding had been
proven (BELING, 1929) a ´feeding-entrainable oscillator´ (FEO) was shown to
exist in some strains of mice, the hamster, rat, rabbit, pigeon, house spar-
row and some marsupial species, with the parameter recorded most fre-
quently being the activity rhythm (STEPHAN, 1986, JILGE et al. 1987, JILGE
and STÄHLE, 1993; JILGE and HUDSON, 2001, COLEMAN et al. 1989; KENNEDY et
al. 1991; HAU and GWINNER, 1992; JILGE, 1992, MISTLBERGER, 1993; PHILIPPS
et al. 1993; RASHOTTE and STEPHAN, 1996; MARCHANT and MISTLBERGER, 1997;
CHALLET et al. 1998; STEPHAN and DAVIDSON, 1998; MISTLBERGER and
MARCHANT, 1999; LAX et al. 1999). The FEO is a circadian oscillator in addi-
tion to and separate from the ´light-entrainable oscillator´ (LEO): even when
the LEO had been destroyed, hamsters were entrained by periodic food
access (reviewed by MISTLBERGER, 1994). While the LEO in mammalian
species is known to be located in the suprachiasmatic nuclei of the hypo-
thalamus, lying above the chiasma opticum and bilaterally symmetric to the
third ventricle, recent evidence suggests that the dorsomedial hypothalam-
ic nucleus may be site of residence of the FEO (GOOLEY et al. 2006; MIEDA et
al. 2006). One other FEO location is discussed, which resides in the gas-
trointestinal tract and the liver (DAVIDSON et al. 2003).
There are some functions, however, which are exclusively synchronized
by the light-dark zeitgeber: the enzymes N-acetyltransferase and hydrox-
yindol-o-methyltransferase and the end product catalyzed by them in the
pineal organ, melatonin (REITER, 1993, TAMARKIN et al. 1985), the disc shed-
ding rhythm of photoreceptors (LAVAIL, 1976) and the mitotic index of the
cornea (BURNS et al. 1976).
Consequences of masking and entrainment
The implementation of periodic food access as a rule is answered by the ani-
mal in two steps: an immediate one resulting in a quick overt rearrangment
(“masking”) and a second step, requiring a much longer time, resulting in
entrainment and restitution of homeostasis. The latter one is taking place
1104
almost unnoticed ´under the mask´. The masking of physiological functions

thus appears to be necessary for maintaining vital functions. Thereafter, the
rather time-consuming process of achieving homeostasis implying complete
circadian reorganization is taking place.
When food access is restricted to some hours during the day, one
should keep in mind that many digestive, metabolic, endocrine and behav-
ioral functions are shifted to the phase where food access is taking place.
Especially when nocturnal animals are fed during some hours of the light
period this means that circadian functions in fact can be reversed. The
process of re-entrainment around the phase of food access can require
30–60 days depending on the time difference between the old and the new
phase RF. Physiological functions like locomotor activity, digestive functions
and urine excretion will be affected significantly (JILGE and STÄHLE, 1993).
So far we do not know whether a transient internal dissociation of functions
is perceived by the animal as a stressor, comparable to the jet lag of many
humans following transmeridional flights. Neither do we know at this time,
whether the permanent dissociation of physiological rhythms, in part syn-
chronized by light and by RF each, is perceived as stress: the DNA synthe-
sis of the thymocyte for example is coupled to restricted food access where-
as the mitotic index in the cornea is not altered by restricted feeding (PAULY
et al. 1976). It does not appear to be unreasonable that reproductive,
immunologic, intermediary-metabolic or behavioural parameters could be
affected and that (certain) restricted feeding schedules are threatening
homeostasis.
There are different ways to avoid the accumulation of excessive fat
deposits. One way would be to offer feed with a lower level of calories and a
higher amount of crude fibre. Another way would be to use automatic time-
able feeders and to adjust feeding to a constant time, in nocturnal animal
species preferably in the mid of the dark period. If this is not possible and
restricted feeding is done by hand it is recommended to provide food at the
end of light time. In any case it should be avoided to feed animals restrict-
edly at various times during the light period.
Chronobiological terms explained:

CIRCADIAN RHYTHM (CR): periodic biological function with a frequency of one
cycle per 24+4 h. CR´s are generated endogenously in the suprachiasmatic
nuclei (SCN) of the hypothalamus. The SCN consist of about 20.000 neu-
1105
rons. They are situated at either side of the third ventricle dorsally of the
optic chiasm. They receive their light:dark input from the monosynaptic
retinohypothalamic tract (RHT) which consists of the axons of melanopsin
containing ganglion cells of the retina. This means that even completely
blind non-rod and non-cone transgenic mice are entrainable by the
light:dark zeitgeber.
CIRCADIAN OSCILLATOR(SYSTEM): Neurons generating a CR with a period of
about but significantly different from exactly 24 h. The SCN are considered
to be the ´masterclock´ of mammals which is entrained by an external zeit-
geber. Since the light:dark cycle, entering the SCN via the retinohypothala-
mic tract (RHT) is the main zeitgeber for mammals, the SCN are referred to
as light-entrainable-oscillator (LEO). As delineated above, in some species
an additional oscillator system has been described so far, which is entrain-
able by periodic food access. Hence, the name feeding entrainable oscillator
(FEO) has been suggested.
ENTRAINMENT: synchronization of a CR by an external (or internal) periodic
variable within a limit of 24+4 h.
FREE-RUNNING RHYTHM: circadian rhythm (e. g. of locomotor activity) in the
absence of any external zeitgeber.
ZEITGEBER: external, periodic variable entraining a circadian oscillator.
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CAPTIONS TO THE FIGURES
FIGURE XXXIII-1a:
24-h rhythms of five behavioural functions of the rabbit living in a persistent 12 h
light:dark alternation (LD 12:12). In the left panel the animal was fed ad libitum, in the
right one it was fed during 4 h of L. The 24 h mean values of 39 (left) and 30 days (right)
are plotted as 100 %. The respective average 60 min columns are hanging when below and
standing when above the total average.
During ad libitum access 65 % of locomotor activity, 68 % hard faeces excretion, 69 % of
food intake, 64 % of water intake and 80 % of urine excretion was taking place during D.
In contrast the right panel demonstrates, that practically all events are assembled around
the 4 h of restricted food access (2 vertical lines) during light time. In addition the
amplitudes during a regular RF regimen in all parameters are significantly higher.
FIGURE XXXIII-1b:
Plot of original activity data of one typical animal during 4 h of feed access during L. The
black dots indicate the calculated centre of gravity of the histogram of each 24-h day. It
regularly lies at the end of the 4 h of feed availability. It is clearly seen that almost all of
the activity is expressed during L and that
~ 20 % of activity regularly is anticipating the begin of feed availability.
FIGURE XXXIII-2:
Shift of 24-h rhythms of duodenal mucosal enzyme secretion. The secretory peak values
are shifted from the end of the dark period during ad libitum feed availability now to the
end of L (redrawn from SAITO et al. 1975).
FIGURE XXXIII-3:
Plasma corticosteroid in female SD rats kept in LD 12:12 and fed ad libitum (top) and for
2/24 h (9:30-11:30 - bottom). During RF the corticosteroid peak was shifted by RF for ~ 12
h. In addition, as seen in fig.1 the amplitude during RF was significantly higher as
compared to ad libitum feed access (redrawn from KRIEGER, 1974).
FIGURE XXXIII-4a:
Rhythm of core body temperature of one adult rabbit during a 4-h food restriction
schedule. Note the anticipatory increase of body temperature starting about 3–4 h prior to
1111
food access. Vertical lines indicate the 4 h of food availability; the horizontal line marks the
7-day average of core body temperature.
FIGURE XXXIII-4b:
24-h - temperature rhythm during a 4-h schedule of food access
(2 vertical lines): as seen in fig. 4a temperature rises above 24-h average (horizontal line)
about 3 hours prior to food access. In addition to this anticipatory component a second,
thermogenic peak is seen during food ingestion. After the end of food access a significant
drop of temperature for 0.7 – 1.0 °C is seen.
FIGURE XXXIII-4c:
Circadian rhythm of body temperature during RF and an interspersed 72-h fast. The
temperature rhythm persists during the fast at a slightly reduced amplitude. Detailed
analysis demonstrated, that it was the anticipatory component which persisted while the
thermogenic component (logically) was completely absent. According to chronobiological
definition it is a necessary and sufficient requirement that a rhythm persists in the absence
of a zeitgeber for some cycles in order to be designated an endogenously generated rhythm.
Thus, the anticipatory component apparently is generated endogenously while the second
peak is due to thermogenesis.
Please realize the anticipation on day three of fasting: the animal did not ´know´ that food
would be available.
FIGURE XXXIII-5a (top):

Time of parturition of 15 NZW rabbit does during restricted feed access (stippled sector).
13/15 does gave birth to their litter within 8 h after the beginning of food access. In
contrast during continuous light conditions and ad libitum feed access the time of
parturition was scattered all over the 24-h of the day (JILGE, 1995).
FIGURE XXXIII-5b (bottom):

Time of parturition (*) and of nursing the litter during the first 16 days. The doe nurses her
pups during the first 90 min after the end of feeding.
FIGURES XXXIII-5a and XXXIII-5b demonstrate clearly that the phases of parturition
and suckling time unequivocally are set by the RF-zeitgeber.
1112
FIGURE XXXIII-1a and 1b (below left and right)
FIGURE XXXIII-2 and 3 (above left and right)
1113
FIGURE XXXIII-4a, 4b and 4c

(from top to bottom)
1114
FIGURE XXXIII-5a and 5b (top and bottom)
1115
Chapter XXXIV
BASICS OF FUR ANIMAL NUTRITION

AND DIETETICS
34.1. Nutrient and energy utilization of fur animals

34.1.1. Digestion
34.1.2. Energy
34.1.2.1 Energy requirements for maintenance
34.1.2.2 Energy requirements for growth
34.1.2.3 Energy requirements for fur production
34.1.2.4. Energy requirements for pregnancy
34.1.2.5. Energy requirements for lactation
34.1.3. Protein and amino acids
34.1.3.1 Protein and amino acid requirements for mainte-
nance
34.1.3.2 Protein and amino acid requirements for growth
34.41.3.3 Protein and amino acid requirements for fur pro
duction
34.1.3.4 Protein requirements for pregnancy
34.1.3.5 Protein requirements for lactation
34.1.4. Lipids
34.1.5. Carbohydrates
34.2. Nutritionaland Metabolic Disordersand Diseases in Fur-Bearing
Animals
34.2.1. Dietary imbalances in carnivorous fur animals (ie. ferrets,
minks and foxes)
34.2.1.1. Generally about nutrient deficiencies
34.2.1.2. Reproductive Organs and Performance Effected by
Nutrition
34.2.1.3. Skin Problems (ie. dry skin, itchy skin) Influenced by
the Nutrition
34.2.1.4. Nutritional Anaemia
34.2.1.5. Biotin deficiency.
34.2.1.6. Urinary Calculi (Bladder stones or Urolithiasis)
34.2.1.7. Wet-belly Disease (Urinary incontinence)
34.2.1.8. Chastek Paralysis
34.2.1.9. Yellow Fat Disease (Nutritional steatitis)
34.2.1.10. Taurin deficiency
34.2.1.11. Nursing Sickness
34.2.1.12. Sore Paws
34.2.1.14. Nutritional Muscular Dystrophy
34.2.1.15. Overfeeding of nutrients (briefly
34.2.2. Dietary imbalances in herbivorous fur animals
(ie. chinchilla)
1117
CHAPTER XXXIV: FERRET - FOX - MINK FEEDING
34.1. Nutrient and energy utilization of fur animals

© Anne-Helene Tauson
uring the last century the mink (Mustela vison), the blue fox (Alopex
D lagopus) and the silver fox (Vulpes vulpes) were kept in farms for fur
production. In the recent years the importance of ferret as pet and
laboratory animal continuously increases. The process of domestication has
gradually progressed during the period these species have been kept in cap-
tivity. Being a recently domesticated group of animals, the nutritional
requirements have been less thoroughly investigated than for other farm
animal species, therefore information on nutrient and energy utilization for
the various life processes is scanty. Farming of animals for fur production
is carried out in the holarctic and temperate zones, their main fur produc-
ing areas are found among the Nordic countries: North America and Russia.
The animals are monooestrous seasonal breeders with an annual repro-
ductive cycle regulated by the photoperiod. For a comprehensive treatise on
the reproductive processes in the mink and foxes see TAUSON and VALTONEN
(1992).
Due to the animals’ carnivorous nature, fur bearing animal diets are
mainly based on by-products of animal origin, e.g. from the fishing indus-
try and slaughter houses. Because these products have less processing
value to other domestic animals, fur animal production is complementary to
conventional farm animal production. Knowledge on furbearer animal nutri-
tion is essential not only for production, but also for pet and zoo animals,
too. The main research efforts in recent years have concentrated on the
nutritional demands of the mink, but results found on this species are usu-
ally applied also when formulating feeding recommendations for the two fox
species, too.
1118
34.1.1. Digestion
As in other carnivores the digestive tract is simple with a non-compartmen-
talized stomach and a relatively short intestine. The mink lacks caecum,
and in foxes it is without functional significance. Both the mink and the two
fox species have a short, non-sacculated colon, and the total length of the
intestine is about 4 times of the body length. Feed passage rate is rapid and
in the mink averaging 3.5 h (SZYMECZKO and SKREDE, 1990) and about 95%
of the feed passing the tract in 4 to 5 h (HANSEN, 1978). In the fox digestion
is considered to be completed within 24-30 h (PERELDIK, 1975). Hence, the
time for enzymatic digestion is relatively short and the digestive system is
not adapted to feedstuffs with high fibre content. Owing to the need for
highly digestible nutrients and for a high dietary fat content, fur bearing
animal feed usually has a far higher energy concentration than diets for
other domestic species.
In mink kits, digestibility of nutrients reaches the level of the adult
animal at an age of 10-12 weeks (ELNIF and HANSEN, 1987; TAUSON, 1988a).
This reflects an age-related increase in activity of the proteolytic enzymes,
activities levelling with those of adult animals at the same age as digestibil-
ity has reached the adult level (ELNIF et al. 1988). In the adult animal, the
pancreatic release of proteolytic enzymes is substantial as indicated by
highly negative amino acid digestibility in the proximal part of the small
intestine (SZYMECZKO and SKREDE, 1990). The activity does not seem to be
regulated by the dietary protein level (SIMOES-NUNES et al. 1984), but enzy-
matic secretion may possibly be affected by the protein source (SKREDE and
KROGDAHL, 1985).
There are some systematic differences in apparent nutrient digestibil-
ity between the species. The fox generally being more efficient than the
mink, which probably reflects slower feed passage rate in them. The supe-
riority of the fox has been demonstrated for protein in compounded diets by
AHLSTRØM and SKREDE (1995a). For protein feedstuffs the difference may
amount to 15 units (SKREDE et al. 1980). Fish products with a low bone con-
tent are usually highly digestible for the mink, and a linear decrease in pro-
tein digestibility with increasing ash content was demonstrated by SKREDE
(1978a). Amino acid digestibility of highly digestible feedstuffs do not
diverge much from N-digestibility, whereas in poorly digestible feedstuffs
amino acid digestibilities may be widely dispersed, and on average very low,
1119
and the poorest digestibilities generally found for cystine (SKREDE,1979a, b).
Similar to overall protein digestibility, the separate amino acids are also bet-
ter utilized by the blue fox than the mink in poorly digestible diets (SKREDE
et al. 1980).
Fat digestibility is dependent on chain length of the predominating
fatty acids and on the degree of saturation; long-chained and unsaturated
fatty acids have higher digestibility than short-chained and saturated ones
(ØHMAN, 1976; AUSTRENG et al. 1979). Also, fat is more efficiently digested by
the fox, mainly because of the higher digestibility of saturated fatty acids
(ROUIVINENE et al. 1988; ROUVINEN, 1991; AHLSTRØM and SKREDE, 1995a).
Digestibility may be impaired by a high dietary level of some calcium salts
owing to saponification. ROUVINEN and KIISKINEN (1991) found that the
digestibility of tallow and rapeseed oil in mink was considerably decreased
when the ash content of the diet was increased from 4-14% of dry matter
(DM) when Ca sources were CaCO3 and bone meal, but not when fish offal
meal was used
Carbohydrate is a very diverse group of nutrients, of which the fur
bearing species have very limited ability to digest fibre. The major source of
carbohydrate energy is starch which, depending on origin, may be almost
100% digestible for the mink if gelatinized, but only poorly digested when
untreated. The fox, on the other hand, is far more efficient at digesting
untreated starch at least from 8-10 weeks of age. Carbohydrate digestibili-
ty may be improved by about 5 (oats) to 30 (wheat) units by cooking
(JØRGENSEN and GLEM-HANSEN, 1973), but also fineness of grinding influences
digestibility, and thus GLEM-HANSEN and SØRENSEN (1981) demonstrated that
the finer the grinding of the ceraeals, the higher became the digestibility.
A comparative study with mink, blue fox and silver fox, carried out
with a diet having a high content of fish offal has confirmed the main fea-
tures among the different species for nutrient digestibility as outlined above
(HANSEN, unpublished results; Table XXXIV-1).
1120
Table XXXIV-1: Chemical composition of the experimental diet and nutrient digestibility
(%), mean and standard deviation, in mink, blue fox and silver fox fed diets containing more
than 60% fish filleting scrap (HANSEN, unpublished results)
Data on chemical composition, nutrient digestibility and metaboliz-

able energy (ME) content for mink for the most commonly used feedstuffs in
fur bearing animal nutrition have been published by the Scandinavian
Association of Agricultural Scientists (NJF, 1985, revised 2004). Data on
digestibility and ME content for the most important groups of feedstuffs
have been quoted from the table by TAUSON (1988b). As indicated in Table
XXXIV-1, calculation of dietary ME based on NJF (1985, 2004) will under-
estimate the ME content of fox diets. Ferret’d data must be similar to those
of the mink.
34.1.2. Energy
Studies on energy and nutrient requirements of fur bearing animals are to
some extent complicated by the fact that these animals, compared with
other domestic species, have a high locomotor activity which makes up a
considerable item and depending on individual, varying part of the mainte-
nance energy requirement. Furthermore, animals are kept in non-climatized
houses, and therefore the requirement for thermoregulation is dependent
on ambient temperature. Moreover, the definite requirement for a specific
production process is not easily accessible, since for instance, shedding of
the winter fur starts during the gestation period, and the summer pelt
1121
primes during the lactation period. In kits the growth of the winter fur starts
in the late part of the growing period.
Only few attempts have been made to determine the basal metabolism
in fur bearing animals. In a study on female mink FARRELL and WOOD
(1968a) concluded that basal metabolic rate (BMR) in mink could only be
measured during short intervals when the experimental animals were
asleep. Their measurements, carried out within the zone of thermoneutral-
ity, indicated a BMR of 0.324 MJ/W0.75, but only few animals were meas-
ured, owing to the difficulties in fulfilling the criteria for BMR. Data from
FARRELL and WOOD (1968a), however, are in fair agreement with 0.354
MJ/W0.78 ,which was reported for some mustelid species, including mink,
in the weight range from 1.0 to 15.0 kg (IVERSEN, 1972). The maintenance
energy requirement is of greater practical interest than BMR, but also, for
such data the definition of the experimental conditions must be taken into
consideration in order to give reliable and comparable estimates of the ener-
gy requirement.
Ambient temperature is a factor of great importance for the energy
requirement. FARRELL and WOOD (1968a) found little variation in respiratory
rate between 16-29 oC, and concluded that the thermoneutral zone for
mink is fairly broad. Later results indicate that the lower critical tempera-
ture is 21-22 oC, and that the energy requirement increases by 9.6-15.5
kJ/W0.75 and day per oC reduction in temperature below this (KORHONEN et
al. 1983, 1985). These data support the estimates of CHWALIBOG et al. (1980)
who found that heat production (HE) increased linearly at the rate of 12.1
kJ/W0.75 per oC reduction in temperature within the range of +22 to -3oC.
The lower critical temperature of the blue fox has been estimated to -6 oC
(KORHONEN et al., 1985), but there are no data on the increase in energy
requirements below this temperature. Also effects of wind may influence the
critical temperature and hence, the energy requirement. In mink the insu-
lating capacity of the fur is reduced by 15-20% at 1-5 m/s (SEGAL and
IGNATOV, 1975), and even very little wind influences the critical temperature
(HARRI and KORHONEN, 1985).
Locomotor activity makes up to a considerable part of the total main-
tenance energy requirement in the fur bearing animal species. There are,
however, no experimental data on the specific energy utilization and
requirements for activity. Indications of the importance of locomotor activi-
ty may be gained from the work of FARREL and WOOD (1968b), where daily
1122
intake of digestible energy (DE) increased by more than 25% when the cage
bottom area was approximately doubled. Also, CHWALIBOG et al. (1980, 1982)
have discussed the importance of registrations of activity when evaluating
the energy requirement for maintenance.
Seasonal effects on energy metabolism in the mink have been claimed
by PERELDIK and TITOVA (1950) and CHARLET-LERY et al. (1984) who found that
HE (heat expenditure) in adult animals was lowest during the November-
February period (0.500-0.585 MJ/kg) ranging from 0.585 to 0.710 MJ/kg
during the rest of the year.
34.1.2.1 Energy requirements for maintenance
Estimations of the ME requirement for maintenance (MEm) in mink have
included the balance and slaughter technique (HARPER et al. 1978; HANSEN
et al. 1981, 1984), balance experiments (PERELDIK and TITOVA, 1950), indi-
rect calorimetry (CHWALIBOG et al. 1980, 1982; CHARLES-LERY et al. 1984;
BURLACU et al. 1984), and, in a single study with direct calorimetry, heat loss
at maintenance level has been studied (WAMBERG, 1994). Furthermore, a
vast material based on data obtained from production experiments in the
Scandinavian countries has been used to state recommendations for daily
ME supply for mink and foxes (HANSEN et al. 1991). In Table XXXIV-2 some
experimental results regarding MEm in mink are summarized, and the data
indicate a considerable variation between sources, which to some extent
may be explained by different experimental conditions and varying levels of
animal activity. According to CHWALIBOG et al. (1980) the estimate of MEm in
adult male mink of 0.527 MJ/W0.75 gave an overall efficiency of utilization
of ME for production of 0.67 which seems to be reasonable. Results from a
later study on growing mink kits (CHWALIBOG et al., 1982), however, gave a
MEm, which was considered too high, and an unacceptable overall efficien-
cy of ME utilization of 125%. These results point out the difficulties in
achieving appropriate estimates of MEm even under controlled experimental
conditions. In some of the quoted studies on mink, MEm has been stated
per kg body weight instead of per kg metabolic weight. The relevance of the
above has been discussed by FARRELL and WOOD (1968b) and HANSEN et al.
(1981) who claimed that since locomotor activity, which makes up to a con-
siderable part of MEm, is not dependent on surface area of the animals,
MEm is not more accurately expressed in terms of metabolic live weight.
Recent estimates of MEm in suckling mink kits have ranged from
0.356 MJ/W0.75 (FINK et al. 2001) with an efficiency of ME utilization in milk
1123
for body gain of 0.53 for protein (kp) and 0.71 for fat (kf) to 0.448 MJ/W0.75
(TAUSON et al. 2004a) with an average utilization efficiency of milk ME for
growth (kg) of 0.67. Data on MEm in the fox species mainly derive from pro-
duction experiments. Recalculation of data on recommended energy supply
and live weights of adult male blue and silver foxes from HANSEN et al. (1991)
and on adult male and non-reproductive female blue foxes and male silver
foxes from PERELDIK (1975) to ME supply per kg metabolic body size are
given in Table XXXIV-3. The agreement between the Scandinavian and the
Russian data is good for both species, and MEm falls within the same range
as the mink data in Table XXXIV-2.
34.1.2.2 Energy requirements for growth
Determinations of energy requirements in growing mink kits have indicated
MEm values in growing animals ranging from slightly above 0.600
MJ/W0.75 to about 0.730 MJ/kg of LW (Table XXXIV-2). Attempts to esti-
mate ME utilization for growth (kg) have generally been unsuccessful.
Hence, CHWALIBOG et al. (1982) estimated MEm to 0.680 MJ/W0.75, which
resulted in an unacceptable kg value of 125%. Also, the investigations of
BURLACU et al. (1984) gave a high MEm and an estimate of kg above 1. When
recalculating CHWALIBOG‘s et al. (1982) data to MEm 0.527 MJ/W0.75
(CHWALIBOG et al. 1980), kg was estimated to 0.83, which possibly may be too
high because the experiment was carried out in the period of intensive
growth when protein retention still is high. HANSEN et al. (1984) assumed
fixed values for ME utilization for protein (kp) and fat (kf) deposition, and
found that MEm was little affected by using kp of 0.45 and 0.5 and kf of 0.8
and 0.9, respectively. Based on these assumptions, HANSEN et al. (1984)
concluded that the ME requirement for growth (MEg) was made up to 30-
35% of the total ME intake during the period of intensive growth at an
approximate kit age of 8-10 weeks. Over the total 4.5 month experimental
period MEg was estimated to 22-24% of ME. The results quoted above
emphasize the difficulties in achieving reliable estimates of the energy
requirements for specific production processes, and point out the substan-
tial influence of locomotor activity on the total ME requirement.
Recommendations on energy supply for growing mink, blue fox and
silver fox kits kept under normal farm conditions have been given by HANSEN
et al. (1991). These data are based on production experiments carried out
in the Nordic countries, in which the animals generally have been fed ad libi-
tum. Recalculation of HANSEN’s et al. (1991) data to ME supply per kg meta-
1124
bolic body size has given the results presented in Table XXXIV-4. In mink
the true body growth is almost completed by the age of 16 weeks. Weight
gain after that age mainly occurs as fat deposition. Of the two fox species
the blue fox has a steeper growth curve than the silver fox and the latter
also deposits less fat. Data in Table XXXIV-4 indicate a considerable
decrease in energy supply per kg metabolic body size in mink and blue fox
in the late part of the growth period, whereas the decrease for silver fox is
moderate. Data for mink females are probably too high, which has also been
indicated in practice. The ME supplies recommended during the last period
of growth are fairly close to the estimates of MEm (Table XXXIV-2 and Table
XXXIV-3) as being the exception data for mink females.
34.1.2.3 Energy requirements for fur production
Mink and foxes are pelted when the winter fur is prime. Since the priming
process like the reproductive processes is regulated by photoperiod it occurs
in November or early December on the northern hemisphere. A complete
furring cycle comprises a kit fur, a summer fur, which in the mink starts to
grow at an approximate kit age of 6 weeks, and a winter fur which starts to
grow at about 16 weeks of age (DOLNICK, 1959).
1125
Table XXXIV-2. Estimates of MEm requirements in mink
1126
Page 1126
16:38
2008.09.03.
a Recalculated from BMR assuming k =0.75.

m
b Energy expenditure of animals kept under conditions denoted as maintenance.
c Discarded by the authors as too high.
Table XXXIV-3. Estimated MEm (MJ/W0.75) in blue foxes and silver foxes based on data
recalculated from HANSEN et al. (1991) and PERELDIK (1975)
Table XXXIV-4. Recommended daily ME supply per kg metabolic size for growing mink, blue
fox, and silver fox kits. Data recalculated from HANSEN et al. 1991
The winter fur growth starts during the period in which the kits are
still growing. Therefore, separation of the requirements for fur production
and growth is not easily accomplished. Hitherto, no estimates of energy
requirements and utilization for fur production have been published. When
considering the data on ME supply in Table XXXIV-4, it may be noted that
the period of most intensive winter fur growth in the mink extends ap-
proximately from 20-26 weeks of age, a period in which the recommended
energy supply is decreasing. Simultaneously, the main body weight gain
occurs as fat deposition, for which ME is more efficiently utilized than for
protein deposition. It can therefore be concluded that the energy require-
ment for fur production is moderate, and that it is counteracted by a
1127
decrease in energy requirement for growth.

34.1.2.4. Energy requirements for pregnancy
The mink has delayed implantation and therefore gestation length varies
from about 40 to 75 days with an average of 51 days. True gestation, from
implantation to parturition, lasts 30 days. Gestation length in the fox is 52-
53 days (see TAUSON and VALTONEN, 1992). Reproduction performance is
highly susceptible to changes in nutrient supply. Indeed, by using a flush
feeding regimen comprising a two week period of moderate restriction in
energy supply (about 80% of MEm) which then was followed by ad libitum
feeding from 3-5 days before the start of the breeding season litter size could
be improved by up to 2 kits (review by TAUSON, 1993). The improved repro-
ductive performance was probably mediated by changes in metabolic hor-
mones such as insulin, IGF-1, thyroid hormones and leptin (TAUSON et al.
2000, 2002).
The increase in weight in the pregnant uterus (y) in mink in relation
to day in pregnancy (t) has been described by the equation
y=0.6074+e0.0987t (TAUSON et al. 1994), which clearly demonstrates the
exponential growth of the conceptus. Despite the rapid increase in weight of
the uterus during the last days of pregnancy, the total amount of energy
deposited in maternal and foetal tissue has been estimated at less than
0.400 MJ close to parturition (TAUSON et al. 1994). This implies a moderate
energy requirement for pregnancy, even when assuming a low utilization of
ME for pregnancy, which is supported by the fact that HE does not increase
as an effect of pregnancy. ME intake increases from mating until after
implantation, after which it decreases to a low level towards the end of ges-
tation, during a period of hyper-leptinaemia. Indeed, despite the lack of
increase in HE and the moderate energy deposition in pregnancy products,
mink dams are in a clearly negative energy balance in the late part of ges-
tation, and energy is partly supplied by mobilization of body reserves
(TAUSON et al. 1994; TAUSON and ELNIF, 1994; TAUSON et al. 2004b). Attempts
to estimate efficacy of ME utilization for pregnancy have not yet given
acceptable results (own vain efforts). The quoted results contrast with WANG
et al. (1988) who claim that both energy consumption and HE is higher in
pregnant than in non-productive mink females.
34.1.2.5. Energy requirements for lactation
The neonatal mink kit is physiologically immature, being blind, almost hair-
less, without the capacity of thermoregulation, and with very limited loco-
1128
motor ability. They are almost devoid of mobolizable energy reserves, ha-
ving a fat content in their body of about 1% at birth, and liver glycogen
stores which can be estimated to sustain life for less than 1 h (TAUSON,
1994). Mink kits become functionally homeothermic at an age of close to six
weeks, and younger kits are unable to maintain their heat production, and
might enter a torpor-like stage if exposed to a temperature below prevailing
in the nest-box. This might be an adaptive mechanism in order to econo-
mize with their limited body energy reserves (TAUSON et al. 2005). Despite
being vulnerable at birth, mink kits have the capacity for a rapid growth
rate, with the maximum relative growth rate being 23% per day and an aver-
age over the first 42 days of life of 9% per day (TAUSON, 1994). Until about
25 days of age mink kits consume only mother’s milk and weaning usually
takes place at an age of 6 to 7 weeks.
Data on energy requirements for lactation is mainly derived from pro-
duction experiments, in which it has not been possible to separate the
requirements for maintenance and milk production. The energy require-
ment is strongly dependent on stage of lactation and litter size. TAUSON
(1988c) found that ME consumption increased 2.5 times from the first to
the 5th week f lactation for females with 5-7 kits. In spite of this increase in
ME consumption, a considerable weight loss in lactating females showed
that part of the energy requirement for lactation had to be supplied by mobi-
lization of body reserves. Furthermore, it was found that a dietary energy
concentration of 16.0 MJ/kg DM was required until the kits started to con-
sume solid food, and from then the energy density of 17.3 MJ/kg DM gave
satisfactory results regarding kit growth and female performance. Our more
recent data confirm the need for a rapid increase in energy intake during
lactation and that high yielding lactating dams are unable to sustain their
energy requirements solely by food intake (FINK et al. 2004; TAUSON et al.
2004b).
34.1.3. Protein and amino acids

Being carnivorous animals the mink and the fox species usually consume
diets with higher protein contents than their actual requirements, and the
excess protein is thus used as energy source (TAUSON, 1995). The animals
are well adapted to high protein diets but there are some indications that
very high protein levels may be detrimental to mink affected by plas-
macytosis, because those animals have an impaired ability to concentrate
1129
their urine (VALTONEN et al. 1982). Because diets for fur bearing animal are
usually based on animal by-products, protein quality may vary consider-
ably, which must be taken into account when formulating diets. The protein
supply is usually expressed as a percentage of ME, which is useful for mak-
ing sure to provide the animals with a specific amount of protein, and also,
when using diets with differing energy concentrations.
Determination of the minimum requirements of proteins and amino
acids are complicated by the fact that low protein diets are usually very
unpalatable and therefore less readily consumed. Furthermore, several N-
balance experiments have shown that the N-balance is usually over-esti-
mated owing to N-losses during urine collection. ELNIF (1992) evaluated
sources and the level of N-losses and found that evaporative losses were of
minor importance. By the use of osmotic mini-pumps a N recovery slightly
below 80% in fed adult female mink at maintenance conditions was demon-
strated (WAMBERG et al. 1996; TAUSON et al. 1997). Recommendations on
nutrient supply for fur bearing animals have been formulated by HANSEN et
al. (1991), in which life cycle stages are represented in four production peri-
ods, namely December to parturition, representing requirements for main-
tenance and pregnancy, parturition to 30 June, mainly covering the lacta-
tion period, 1 July to 31 August, representing the period of intensive kit
growth, and 1 September to pelting, combining the late growth period and
the furring period. In the followings the nutrient requirements will, when
possible, be stated for the separate life processes, but comparisons will be
made to the recommendations for the four production periods (HANSEN et al.,
1991; Table XXXIV-5).
34.1.3.1 Protein and amino acid requirements for maintenance
Strict minimum levels for protein requirements for maintenance are not
known for the fur bearing animals. For mink and blue fox, NRC (1982)
states 20% of ME, a level that probably is above the true minimum require-
ment. Data from adult silver foxes indicate that protein levels below 19-22%
of DM were detrimental to health and normal fur development, but that lev-
els of 13-16% of DM were sufficient to maintain body weight before priming
(RIMESLØTTEN, 1978). There is no available information on amino acid
requirements for maintenance.
34.1.3.2 Protein and amino acid requirements for growth
The major efforts to determine protein and amino acid requirements of fur
bearing animals have concentrated on the growth and furring periods.
1130
Because these two processes partly progress simultaneously the require-

ments have usually been evaluated for both processes together. Based on
N-balance studies with mink kits from 10 to 30 weeks of age and fed diets
with digestible protein content varying from 17% to 54% of ME, GLEM-
HANSEN (1980a) concluded that 40-42% of ME was sufficient to cover the
requirements in the period of 10-17 weeks of age and 31-32% of ME met the
demands from 19 to 24 weeks of age. Contrasting to this, BERG et al. (1984)
found that 19% of ME gave maximum N-retention in 19 weeks old kits.
SKREDE (1977) found that 28% of ME from protein in a diet based on fish
meal gave a satisfactory growth rate but in another experiment with fish
and fish offal products as main protein sources (SKREDE, 1978b), 26-27% of
ME from protein gave sub-optimal results. It was furthermore found, that
N-balance decreased for protein levels below the lowest one giving satisfac-
tory growth in the period of July-August. A protein level of 27% of ME from
weaning to the end of August, and 23% from September to pelting gave a
normal growth performance and normal values for some haematological
parameters, but the content of plasma amino acids was higher in animals
fed the low protein diet compared with controls (TYÖPÖNNEN et al. 1986). In
a later study (TYÖPÖNNEN et al. 1987) 25% and 20% of ME from proteins in
the periods described for the previous experiment, was sufficient for growth
but elevated plasma levels of the branched amino acids (valine, leucine and
isoleucine) were observed. LUND (1983) has also reported a normal growth
performance at a protein supply of 25% of ME. Experiments with blue foxes
have indicated that a normal growth performance in the early growth peri-
od, from approximately 9 to 14 weeks of age, is supported by a protein level
of 28% of ME (RIMESLØTTEN, 1976; Tøng, 1982, 1983; BERG et al. 1982) or even
as low as 22% (TØNG, 1984). Based on the experimental data quoted above
the recommended minimum protein supply in the period from 1 July to 31
August is 30% of ME for minks and 28% of ME for fox (HANSEN et al. 1991;
Table XXXIV-5).
The knowledge regarding the essential amino acid requirements in
fur-bearing species is mainly connected to the sulphur-containing amino
acids cystine and methionine, because cystine makes up to a substantial
amount of the protein content in hair. GLEM-HANSEN (1992) states that mink
hair contains only 7-12% of the total deposited amount of protein, but
approximately 60% of the total deposited cystine, despite the fact that pro-
tein moulted with the summer coat was not included in the calculations.
1131
GLEM-HANSEN and HANSEN (1981) have shown that deposition of all amino
acids except cystine follows the deposition of nitrogen. Cystine which shows
a marked increase in deposition during the period from moulting of the
summer coat to priming. Experiments conducted by GLEM-HANSEN (1980b;
1982) indicate that optimum levels for sulphur-containing amino acids were
0.6-0.7 g/MJ (2.6-2.7 g/16 g N) from 10 to 19 weeks of age, provided the
amino acids originated from the feedstuffs or were supplied as L-methion-
ine or L-cystine, because D-methionine is very poorly utilized by minks.
Similar levels were also reported to fulfill the requirements when only nat-
ural feed sources were used (SKREDE, 1978b). There are some indications of
the tryptophan requirements: PERELDIK et al. (1970) suggesting a level of 0.1
g/MJ and SKREDE (1981) stating that digestible tryptophan requirements
are rather below that value. LEOSCHKE and ELVEHJEM (1959) found methion-
ine and arginine to be the first limiting amino acids for mink fed on a casein
diet, the arginine content of which was far below the level in diets based on
natural feedstuffs. Based on earlier investigations reviewed by GLEM-HANSEN
(1992) and more recent experiments by Børsting and CLAUSEN (1995), sug-
gested requirements for the essential amino acids for mink in the growing-
furring periods are given in Table XXXIV-6.
34.41.3.3 Protein and amino acid requirements for fur production
The process of furring involves a very high deposition chiefly of cystine in
the growing hair. GLEM-HANSEN (1980a) estimated that 31-32% of ME from
protein was sufficient in the early part of the furring period and that the
protein requirement in the period of 26 weeks of age until pelting was equal
or possibly higher than the above data, and a subsequent investigation
(GLEM-HANSEN, 1980c) indicated a deterioration in the fur quality when the
protein supply was below 32% of ME. SKREDE (1978b) found 31-32% of ME
during the total growth period to support the requirements for growth and
furring. TYÖPÖNNEN et al. (1986) found that 27% of ME from protein from
September to pelting supported a normal fur quality whereas 23% resulted
in impaired fur quality. If protein was made up to 20% of ME from
September to pelting, pelt quality was reduced and it was not improved by
methionine or lysine supplementation or by the combination of the two
amino acids (TYÖPÖNNEN et al., 1987). Results from LUND (1983) support the
above results by indicating a deteriorated fur quality when the protein level
was 25% of ME. For blue fox kits RIMESLØTTEN (1976) have found that 25% of
ME from protein is sufficient for supporting normal fur quality in kits aged
1132
from 16 weeks or more. The quoted experimental data have resulted in rec-
ommendations regarding a minimum protein supply of 30% of ME for minks
and 26% of ME for foxes in the period from September to pelting (HANSEN et
al. 1991; Table XXXIV-5). However, more recent findings by DAHLMAN (2003)
suggest that a level of 21-22% of ME from protein is sufficient for sustain-
ing growth and production for high quality fur, provided a dietary content
of 0.4 g digestible methionine/MJ ME) and that during the later stages of
the growing-furring period even a 15-16% supply of ME may be adequate,
provided the above methionine supply. As mentioned above the furring
process puts high demands on the supply of sulphur-containing amino
acids. GLEM-HANSEN (1980c) estimated the requirement for cystin and
methionine to be 0.9-1.0 g/MJ (3.7-4.1 g/16 g N) from 20 to 24 weeks of
age and 0.7-0.8 g/MJ (3.0-3.1 g/16 g N) from 26 to 30 weeks of age, where-
as SKREDE (1981) estimated 0.7 g/MJ to be sufficient. Estimates of amino
acid requirement based on GLEM-HANSEN (1992) and BØRSTING and CLAUSEN
(1995) are recorded in Table XXXIV-6.
Table XXXIV-5: Recommended supply of proteins, fats and carbohydrates as percentages of

ME for mink and foxes. Values for proteins are the minimum recommenda
tions, whereas fat is given as a recommended range and maximum recom
mended supply is stated for carbohydrates (HANSEN et al. 1991)
34.1.3.4 Protein requirements for pregnancy

The true protein requirement for pregnancy is not known, but it can be
assumed to be moderate owing to the quantitatively low protein deposition
in maternal and foetal tissue, the process of retention, however, being
reflected by a decreased rate of protein oxidation during the last part of
1133
pregnancy (TAUSON et al. 2004b). SKREDE (1978c) fed a protein level of 26-
27% of ME during winter and during pregnancy. An increased rate of early
postnatal kit losses mainly in yearling females indicated that the protein
supply might have been a border-line. Feeding a diet supplying 14% of ME
from protein during the true gestation period resulted in poor reproductive
performance with a high rate of barren females and high perinatal kit loss-
es, suggesting that the diet was deficient in protein (TAUSON, unpublished
results).
34.1.3.5 Protein requirements for lactation.
Provided a good protein quality, GLEM-HANSEN (1979) found that 42% of ME
from protein supported normal kit growth. At lower levels kit growth was
impaired, and could not be restored by an adequate protein supply after
weaning. SKREDE (1978c) found that 38-39% of ME from protein was suffi-
cient for female kits but that male kit growth performance in the lactation
period was improved when the protein supply was increased to 49-50% of
ME. Recent findings from experiments with different levels of protein sup-
ply to lactating mink suggest that, provided a good protein quality and using
carbohydrates with readily available starch, low protein diets (30% of ME
from protein) resulted in superior milk yield as measured by the water iso-
tope dilution technique, and superior kit growth, while dams maintained
body weight better than dams fed on high protein diets (60% of ME) (FINK et
al. 2004, 2005). By using a factorial approach authors estimated amino acid
requirements in high-yielding lactating minks at the values reported in
Table XXXIV-7 (FINK et al. 2005). For blue foxes RIMESLØTTEN (1976) found
that 35% of ME from protein secured an optimum milk yield. Hansen et al.
(1991) have concluded that the minimum protein supply to minks and foxes
is 40% and 37% of ME, respectively during the lactation period (Table
XXXIV-5).
1134
Table XXXIV-6: Estimated requirements for essential amino acids in growing mink kits from
1 July to pelting. Data from BØRSTING and CLAUSEN (1995) and GLEM-HANSEN (1992).
BØRSTING’S and CLAUSEN’S data state apparently digestible amino acids, whereas
GLEM-HANSEN has used true digestibility. GLEM-HANSEN states that the data pre
sented may not be the minimum requirement but levels that meet the requirement
aMethionine+cystine 0.48 g/MJ 1 July-15 August, and 0.72 g/MJ 16 August to pelting.
Table XXXIV-7: Estimated requirements of apparently digestible amino acids in lactating

mink (per W0.75) during weeks 1 to 4 post partum (FINK et al. 2005)
1135
34.1.4. Lipids
The minimum requirement for essential fatty acids is not known for the fur
bearing animals, but NRC (1982) suggests the minimum level of linoleic acid
to be about 0.5% of DM for adult animals and 1.5% for pregnant and lac-
tating females and young growing kits. Using conventional feedstuffs these
levels are often exceeded and cases of deficiency are not likely to occur. Both
fish and fish products being the major feed ingredients high dietary levels
of long-chained and unsaturated fatty acids are supplied, which may
involve a considerable oxidative stress. Care must therefore be taken to pre-
vent fat peroxidation and to give an adequate vitamin E supply. The fur-
bearing animal species are tolerant to high fat diets as demonstrated by
ALSTRØM and SKREDE (1995a, b), and fat being a cheap source of energy has
resulted in high dietary levels being used in practice. Both in the lactation
(TAUSON, 1988c) and the early growth period (SKREDE, 1983) a high dietary
energy concentration is a prerequisite for an optimal animal performance.
There are some indications that fat source may influence fur quality char-
acteristics. Hence, HILLEMANN (1982) and HILLEMANN and MEJBORN (1983)
found that fur quality was improved when at least 12-20% of the dietary fat
consisted of linoleic acid, which was achieved by substituting tallow and fat
from swine with poultry fat and vegetable oils. Similar findings were report-
ed by TAUSON and NEIL (1991), who found improved fur quality when slaugh-
ter house offal was substituted with fish oil or rapeseed oil. ROUVINEN (1987),
on the other hand, found no clear relation between the level of linoleic acid
and fur quality, but the incidence of fur defects gave some indications con-
cerning the positive effects of linoleic acid.
34.1.5. Carbohydrates
The fur bearing animals have no specific physiological requirement for
dietary carbohydrates, but many carbohydrate sources are used for energy
supply. Owing to low digestibility, carbohydrate feedstuffs often have an
energy density decreasing effect on the diet. Therefore, the amount of car-
bohydrates in fur bearing animal diets has to be limited, especially in peri-
ods when a high energy density is beneficial for animal performance.
Recommendations on nutrient supply (HANSEN et al. 1991; Table XXXIV-5)
have been formulated with the intention of covering protein requirements
and allow to a suitable dietary energy density and an appropriate ratio
between fats and carbohydrates.
1136
The above arguments have contributed to the limited amount of car-

bohydrate supply recommended during the lactation period, but as report-
ed above, lactating mink dams may benefit from a higher yield of highly
digestible carbohydrates in terms of milk yield and kit body gain (FINK et al.
2004, 2005). It has also been demonstrated that glucose metabolism was
almost unaffected over a wide range of protein/carbohydrate supplies.
When mink dams, that are fitted with jugular vein catheters, in their late
lactation were fed on diets with a protein content varying from 30 to 60% of
ME, containing either substantial amounts of carbohydrates or being car-
bohydrate free. FINK and Børsting (2002) and FINK et al. (2002a, b) showed
that quantitative glucose turnover was almost independent of dietary nutri-
ent supply. Based on this finding it was concluded that the mink, besides
its dependence on gluconeogenesis while fed on high protein diets, has a
high glycolytic capacity.
1137
34.2. Nutritional and Metabolic Disorders and Diseases in

Fur-Bearing Animals
© Andras Bersenyi
For certain nutrients, clinically observable conditions have been shown to

be related to chronic ingestion of either a deficient diet or excessive amounts
of compounds. Although the overall incidence of nutrient deficiencies and
toxicities in fur-bearing animals is largely unknown, several dietary syn-
dromes have been identified. Some of them are discussed below according
to the organs affected.
34.2.1. Dietary imbalances in carnivorous fur animals (ie. ferrets,

minks and foxes)
34.2.1.1. GENERALLY ABOUT NUTRIENT DEFICIENCIES

Depending on the degree of energy deficiency, retardation or cessation of
growth may be accompanied by varying stages of emaciation. The fur may
be dull and lack shine. Milk yield may be reduced in lactating females. As
protein requirements usually increase with higher energy density of the
diet, animals fed excessive dietary fat may risk protein deficiency. Protein
deficiency manifests as slow development rate in youngsters, low concep-
tion rates and failed lactation in breeding females and impaired immunity
in animals of all ages. Dietary arginine is required for optimum growth.
Arginine deficiency has been reported in ferrets, while taurin deficiency has
not. It manifests as encephalopathy. Carbohydrate deficiency includes low
growth, poor reproductive performance and loss of hair. The mink and fer-
ret requires dietary niacin because they can not metabolize sufficient pre-
cursor tryptophan to meet their niacin requirement. Signs of deficiency may
include anorexia, pellagra and ’blue tongue’.
34.2.1.2. REPRODUCTIVE ORGANS AND PERFORMANCE EFFECTED BY NUTRITION

Due to the damage of mucos membrane in testes and eggs, the reproduc-
tion performance in both sexes is reduced by vitamin A deficiency.
Moreover this condition may also manifest as retardation of growth, night
blindness, hyperkeratosis (ie. dry, scaly and rough skin) and death. Even
vitamin B6 (pyridoxine) deficiency affects the reproductive performance of
fur-bearing animals; in males the testes become atrophic, aspermatic and
1138
degenerative, while in females both embryo mortality and abortion increase,

and the litter size decreases.
34.2.1.3. SKIN PROBLEMS (IE. DRY SKIN, ITCHY SKIN) INFLUENCED BY THE
NUTRITION
If the animal’s skin has flakes or white powder on the surface of it, the skin
is too dry. The two most frequent causes of dry skin are poor nutrition and
bathing too often. (Ferrets, like cats, do not require routine bathing; no
more frequently than once a month.) Itchy skin may occur even if the skin
is not dry. Itchiness can come from parasites, nutritional disorders, allergic
reactions or diseases. Low levels of dietary fats and essential fatty acids
(ie. PUFAs) are the primary reasons for these dermatologic problems.
Furthermore, lack of proteins from meat sources can also be an occasion as
well as vitamin A deficiency or its excessive amounts after feeding liver.
Overdoses of vitamin A can lead to toxic conditions, making skin disorders
even worse.
34.2.1.4. NUTRITIONAL ANAEMIA

Iron may be of importance for normal pigmentation of the pelt in fur-bear-
ing animals. Certain other fishes, usually from the cod family, generate
formaldehyde or thrimethylamineoxide (TMAO), which interferes with IRON
metabolism and causes severe anaemia and fur depigmentation. Feeding
large amounts of raw marine fish of the cod (Gadidae) family, such as coal-
fish (Gadus virens), Atlantic whiting (G. merlangus) and Pacific hake
(Merluccius productus), may result in severe anaemia and whitish underfur
(„cottonfur”). Freezing the raw fish appears to increase the problem, while
heating (approx. 90°C) it destroys or inactivates the causative factor. Very
high levels of trimethylamine oxide (TMAO) are involved in such fish.
These N-containing compounds are broken down by an enzyme present in
the fish digestive tract, resulting in several products (eg. formaldehyde),
which are not utilized by animals and can inhibit the absorption of dietary
iron. Therefore the Hb concentration is reduced, causing anaemia. The
characteristic deficiency symptom is lack of underfur pigmentation (greying
or whitening), microcytic-hypochromic anaemia, severe emaciation and
growth retardation. Lactating females are affected with weakness, weight
loss and because of the decreased milk production the growth rate of pup-
pies tends to be slow and mortality increases until weaning.
1139
However, it has also been found that chelat-bind iron such as ferric gluta-
mate and ferrous fumarate are satisfactory supplements for preventing
dietary iron deficiency.
In addition, high dietary levels of phosphorus, zinc, copper, man-
ganese and cadmium can reduce the iron absorption in animals, but TMAO
content in certain fish species is of far greater importance.
The discolouration of the fur of ferrets, minks and foxes could be attributed
to deficiency of copper (Cu), riboflavin (vitamin B2), vitamin B12 and
pantothenic acid, reducing the concentration of Hb. Moreover, the conse-
quences of vitamin B12 deficiency may be the limited utilization of vitamin
B1 and B2. Pantothenic acid deficiency also manifests itself as alopecia.
34.2.1.5. BIOTIN DEFICIENCY

Grey underfur and loss of guard hair (alopecia) also occurs when high lev-
els of uncooked eggs, particularly turkey eggs, are fed. A typical clinical sign
in fox is the achromatrichia and spectacle eye (CHEEKE, 1999) Raw eggs,
which are otherwise an excellent protein source, contain a substance called
avidin that binds with and inactivates dietary biotin. The consequent biotin
deficiency causes achromotrichia and alopecia in severe caseses. When
enzymes cause these nutritional problems, they can be inactivated by heat-
ing, however, this adds expense to the diet preparation. Cooking at 91oC for
at least 5 minutes inactivates avidin. Alternatively, diets known to contain
these interfering substances can be supplemented with the deficient nutri-
ent(s). Avidin, an antinutritional factor present in eggs, inactivates biotin
required for pigmentation and hair growth. Biotin deficiency can be pre-
vented by heat treating the eggs. Peroxides of the rancid feed may also
destruct biotin.
34.2.1.6. URINARY CALCULI (BLADDER STONES OR UROLITHIASIS)

Urinary calculi in fur-bearing animals occur mainly in pregnant and lactat-
ing females as well as in male kits because of increased mobilization of min-
erals, and mortality from the disease may be very high.
The main constituent of the stones is magnesium ammonium phosphate
hexahydrate (MgNH4PO4·6H2O), and their formation appears to be favoured
by alkaline urine. These calculi are quite soluble at pH below 6. The pH of
urine in carnivora ranges from 5.5 to 7.5 according to the feed given. Ferrets
fed diets (e.g. cat foods) containing mainly plant protein are particularly
1140
susceptible to bladder stones. The products formed from the digestion of

plant proteins make the urine alkaline (urinary pH rises above 6.5), for-
mating struvit stones. The incidence of urinary calculi is low among animals
where fish is the princepal feed and more common when horsemeat is the
main ingredient in the diet. Other predisposing factors may be the contam-
ination of food, decreased water intake or increased ash intake.
34.2.1.7. WET-BELLY DISEASE (URINARY INCONTINENCE)

Wet-belly disease is a non-fatal condition that occurs sporadically, mostly
in obese males, with an incidence in the late summer and autumn (from
September to November). It is characterized by dribbling of urine and con-
sequently both the skin and fur become soaked and discoloured around the
urinary orifice. Since affected areas of the pelt must be discarded, the con-
dition is of economic importance. The cause is unknown, but at least 3 fac-
tors, including genetic strain, high dietary fat level and obesity, appear to
have the greatest influence on incidence. The disorders is usually associat-
ed with bacterial infection; Proteus organisms may be isolated in the uri-
nary tract.
34.2.1.8. CHASTEK PARALYSIS

This disease is eventually a deficiency of vitamin B1 caused by inactiva-
tion of vitamin B1 in the diet. This condition is associated with the feeding
certian raw fish that contain the enzyme thiaminase. Some fresh material
fed to mink may contain interfering enzymes that would be inactive in the
dried feeds given other classes of livestock. A well-known example is thi-
aminase, an enzyme found in the flesh of a number of species of fish,
including bullheads, carp, herring and ocean smelt), which destroys dietary
thiamine and causes anorexia and Chastek paralysis in mink. The list of
fish containing thiaminase see below in Table XXXIV-8. Affected animals
gradually lose their appetite and weight, and die after convulsion and paral-
ysis (star-gazer opistotonus). The most significant pathological findings are
brain lesions. The occurrence of Chastek paralysis can be prevented by
cooking the fish or giving the raw fish on alternate days.
1141
Table XXXIV-: Occurence of Thiaminase in Fish
34.2.1.9. YELLOW FAT DISEASE (NUTRITIONAL STEATITIS)

This conditions occurs in young, rapidly growing animals as a result of
excessive rancid unsaturated fatty acids (PUFAs) and/or a deficiency of
vitamin E in the diet. The high level of dietary PUFAs is associated with
excess of oily marine fish (eg. cod liver oil and herring oil). Polyunsaturated
fats are highly susceptible to oxidation within the feed as well as within the
host’s tissue. Vitamin E is a critical nutritional component in protecting tis-
sue against lipoperoxidases. The disease manifests as anorexia, hemolytic
1142
anaemia and progressively impaired gait leading to paralysis followed by

death. Necropsy findings include a yellow discolouration of body fat and
fatty degeneration of liver. In order to avoid the condition a tocopherol to
PUFA ratio (mg:g) of at least 0.5 has been recommended.
Other possible effects of feeding excessive dietary fat see above in protein
deficiency. (see also: VESD in Chapter XII).
34.2.1.10. TAURIN DEFICIENCY

Based on some case reports and on the similarity of metabolism of the tau-
rin, it is now suggested to be a requirement in the ferret, mink and foxes,
too. In the commercialized feed mixtures for ferrets the recommended tau-
rin concentration is of 0.23 to 0.25% in dry matter (LEWINGSTONE, 2000). In
case of deficient supply, heat treatment (Maillard reaction) and gut flora
abnormalities the expected deficiency symptomes are similar to those of
cats: retinal and neural damages, impairad immune system, reproductive
troubles and retarded kitten growth.
34.2.1.11. NURSING SICKNESS

This disease appears in the 5-6th weeks of lactation. There is reduced
appetite (anorexia), weakness, rapid dehydration, weight loss (wasting) and
staggering movement following death. It is said to be a salt deficiency dis-
order and recommended treatment is 0.3 to 0.5 per cent NaCl in the diet.
34.2.1.12. SORE PAWS

The occurrence of sore paws in young foxes a few days after birth could be
observed with a high rate of mortality in untreated animals. The incidence
of this condition is likely to be a consequence of magnesium deficiency
because sore paws has been reported in Mg-deficient dogs, too. Puppies of
young females deprived of Mg during pregnacy and early lactation are par-
ticularly affected.
34.2.1.13. NUTRITIONAL HYPERPARATHYROIDISM (OSTEODYSTROPHIA FIBROSA)

Exclusively feeding meat or fish results in an unbalanced diet and causes
nutritional diseases. Nutritional (alimentary) secondary hyperparathy-
roidism is usually associated with feeding all-meat diets that are low in
calcium, thus creating an imbalance in the ratio of calcium to phosphorus.
All ages are susceptible, but rapid growth predisposes young animals to the
1143
hyperphosphorosis. The bones are soft and pliable (rubber-like), and fruc-
tures may be present, thus, individuals are unable to stand. Microscopically
bones are osteoporotic with typical lesions of osteodystrophia fibrosa. The
disease can be prevented by ensuring adequate calcium intake and a prop-
er calcium:phosphorus ratio (1.2-1.7 to 1).
The condition must be differed from true rickets in young animals.
Rachitic changes and abnormal bone development may occur when the diet
is deficient in vitamin D, calcium or phosphorus.
34.2.1.14. NUTRITIONAL MUSCULAR DYSTROPHY

Animals in good condition die suddenly without any obvious clinical signs
as a consequence of myocardial necrosis. Distrophy of myocardium (heart
muscle) and skeletal muscle is a consistent finding in animals with seleni-
um (Se) and vitamin E deficiency. Etiologic factors in the development of
these lesions include low dietary levels of Se and vitamin E as well as sul-
fur-containing amino acids; high dietary concentrations of polyunsaturated
fats; exposure to prooxidant conditions (toxicity by O2, O3, Fe, etc.); and
intake of Se antagonists (eg. Cu, Zn, Co, Cd, Sb, Ag and Te).
34.2.1.15. OVERFEEDING OF NUTRIENTS (BRIEFLY)

Excessive energy intake may lead to obesity as well as wet-belly disease
detailed above, and dyspepsia and diarrhoea caused by high dietary carbo-
hydrate level (>35-40%).
34.2.2. Dietary imbalances in herbivorous fur animals (ie.

chinchilla)
The state of health with chinchillas is indicated well by faeces. The normal
faeces are 3-5 mm in size and oil green. The quality of faeces can be
changed by feeds. The diarrhoea may be caused by a rapid change of feeds,
mycotoxins (eg. T-2, F-2), highly fermentable feeds (ie. cabbage, lettuce) and
large quantities of moisture, green plants. Otherwise, mouldy hays and
high-protein feeds may result in obstipation. Leguminous plants (ie. tri-
foils) bloat easily the chinchilla.
1144
SOME SPECIAL CONDITIONS IN CHINCHILLAS

Fat overfeeding may cause fatty degeneration of liver and low quality of fur
because of irregular development of hair. In case of choline and/or Met
deficiency the transformation of beta-carotene to vitamin A is limited.
Consequences may be the so called yellow ear disease caused by excess of
carotene granules as well as reduced reproductive performance by vitamin
A deficiency. If the feed intake is stopped for a short period of time (1-2 days)
around parturition because of injury to forelegs, dusty diet and reduced
water intake, considerable amount of fat is mobilized, resulting in hepatic
hyperlipidosis (hunger ketosis) in obese females. This condition also
occurs in obese ferrets, cats and ponies.
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Tauson, A-H., Fink, R., Forsberg, M., Lagerkvist, G. and Wamberg, S. (2000): LH release in
mink (Mustela vison). Pattern of the LH surge and effect of metabolic status. Reprod.
Nutr. Dev. 40, 229-247.
Tauson, A-H., Fink, R., Hansen, K.B., Hansen, N.E. and Chwalibog, A. (2004a): Utilization
of milk energy by suckling mink kits. Arch. Anim. Nutr. 58, 181-194.
Tauson, A-H., Forsberg, M. and Chwalibog, A. (2002): Plasma concentrations of leptin mir-
ror changes in body weight but do not influence the pattern of the preovulatory
luteinizing hormone surge in mink (Mustela vison). J. Nutr. 132, 1790S-1792S.
Tauson, A-H., Forsberg, M. and Chwalibog, A. (2004): High leptin in pregnant mink
(Mustela vison) may exert anorexigenic effects: a permissive factor for rapid increase
in food intake during lactation. Brit. J. Nutr. 91, 411-421.
Tøng, L. (1982): Resultat av Helves Stiftelses utfodringsförsök 1981. Finsk Pälstidskrift 16,
287.
Tøng, L. (1983): Resultat av 1982 rs försöksverksamhet. Finsk Pälstidskrift 17, 274.
Tøng, L. (1984): Resultat av 1983 rs försöksverksamhet. Finsk Pälstidskrift 18, 269.
Työppönen, J., Berg, H. and Valtonen, M. (1987): Effects of dietary supplement of methio-
nine and lysine on blood parameters and fur quality in mink fed with low-protein
diets. Acta Agric. Scand. 37, 487-494.
Työppönen, J., Valtonen, M. and Berg, H. (1986): Low-protein feeding in mink: effects on
plasma free amino acids, clinical blood parameters, and fur quality. Acta Agric.
Scand. 36, 421-428.
Valtonen, M., Mäkelä, J. and Eriksson, L. (1982): Njurens koncentrationsförm ga hos fris-
ka och plasmacytotiska minkar. XIV Nordiske Veterinärkongress, Copenhagen,
Denmark, 372-373.
Wamberg, S. (1994): Rates of heat and water loss in female mink (Mustela vison) measured
by direct calorimetry. Comp. Biochem. Physiol. 107A, 451-458.
Wamberg, S., Elnif, J. and Tauson, A-H. (1996): Assessment of the accuracy of quantita-
tive urine collection in mink (Mustela vison) using osmotic pumps for continuous
release of p-aminohippuric acid and inulin. Lab. Anim. 30, 267-272.
Wang, P., Lu, H. and Zhao, W. (1988): Energy metabolism of Mustela vison during preg-
nancy and lactation. Acta Ther. Sin. 8, 139-145.
Wenzel, U.D. (1982): Pelztiergesundheitsdienst. VEB Gustav Fischer Verlag, Jena
Wenzel, U.D. (1990): Das Pelztierbuch. Ulmer, Stuttgart
1149
JAV_List_Abbr_VND_FSGy:MINTAÚJ.qxd 2008.09.03. 16:41 Page 1151
LIST OF ABBREVIATIONS
AAFCO =American Association of Feed Chemistry Officials

ADF = acid detergent fibre
ADL = acid detergent lignin, 72% sulphuric acid soluble lignin
ALP = alkali phosphatase
ALT = alanine-aminotransferase
AMEn = apparent, zero N-balance corrected metabolisable energy
A.O.A.C. = American Association of Analytical Chemists
approx.= approximately
ARC = Agricultural Research Council (UK)
AST = aspartate-aminotransferase
AV = anisidine value
AVMA = American Veterinary Medical Association
BCS = body condition score
BE = basis excess
BE = brute energy (synonym: GE = gross energy)
BFS = bacterially fermentable substance (bakteriell fermentierbare
Substanz)
BSE = bovine spongiform encephalopathy
CAB = cation-anion balance, or DCAB= dietary cation anion balance
CFU = colony forming unit
CL = corpus luteum
CLps = corpus luteum pseudogravidity
CT = computerised tomography
CV = variance (standard deviation in %)
CVN= Dutch Central Bureau for Animal Nutrition
DAS = diacetoscirpenol
DC = digestion coefficient
DE = digestible energy
D(E)XA = dual-energy X-ray absorptimetry
df = discount factor
dg = degradability (in rumen)
DLG = Deutsche Landwirtschaftsgesellschaft
DM = dry matter
DOS = deoxynivalenol; deuterium oxide space
1151
dUA = dietary undetermined anion

EAA = earth alkali alkalinity
EB = electrolyte balance
EBW = empty body weight
ESR=erythrocyte sedimentation rate
ESVCN = European Society of Veterinary and Comparative Nutrition
ESVIM = European Society of Veterinary Internal Medicine
FCE/FCR = feed conversion efficiency/ratio (kg/kg)
FDA = Feed and Drug Administration
FELASA = Federation of European Laboratory Animal Science Association
FFDM = fat-free dry matter
FID= flame ionisation detector
PMN= neutrophyls polymononuclearis (cells)
FOS = fructose-oligosaccharides
GGT, g-GT = gamma-glutamyl transferees
GIT= gastro-intestinal tract
GMO = genetically modified organism
GnRH = gonadotropic releasing factor
GRAS = generally accepted as safe
GSH-Px = gluthation-peroxydase
GTF = glucose tolerance factor
hCG = human choriogonadotropin
HD = high degradable crude protein
HDL = high density lipoprotein
HU-value = Hounsfield unit
IAEA = International Agency of Atomic Energy
IAPP= islet-associated polypeptide
IFN = International Feed Number
INQ = Index of Nutritional Quality
INRA = Institut National de la Recherche Agronomique
k = partial utilisation of ME to NE (NE/ME × 100)
LD = low degradable crude protein
LH = luteinising hormone
LW = live weight
MDA= malondialdehyde
ME = metabolisable energy
MOS = mannose oligosaccharide
1152
MP = metabolisable protein
MPS = mononuclear phagocyte system
MUN = milk urea nitrogen
NDF = neutral detergent fibre
NE = net energy
NEFA = non-estherified fatty acid
NEg = net energy gain
NEl = net energy lactation (NEL)
NEm = net energy maintenance
N-f.e. = nitrogen free extracts
NFC =nonfibre carbohydrates (NFC=100% of DM-%NDF-%CP-%-%ash-
%ether extract) and non-structural carbohydrates
NMR = nuclear magnetic resonance
NPN = Non Protein Nitrogen
NPR = Net Protein Ratio
NPU = net protein utilisation
NRC = National Research Council (USA)
OMMI = Országos Mezőgazdasági Minősítő Intézet (Hungary)
P/E ratio = protein to energy ratio
P4 = progesterone
PCR = polymerase chain reaction
PDI = protein digestible (in) intestine
PEM = protein energy malnutrition (syn. Protein Calorie Malnutrition)
PER = Protein efficiency Ratio
POTZ= preferred optimum temperature zone (reptiles)
PPAR-peroxysome proliferators-activated receptor
ppb = μg/kg
ppm = mg/kg
PUFA = poly-unsaturated fatty acid
PUN = (blood) plasma urea nitrogen
PV= peroxide value
q = metabolizability [ME/BE × 100)
RDP = rumen degradable protein
RUP = rumen undegradable protein syn. UDP, bypass protein)
(S)EUROP = (Super) European (carcass qualification system)
SET= standard environmental temperature (fish)
T3 = triiodotironine
1153
T4 = thyroxine
TBA= thiobarbituric acid
TDN = total digestible nutrients
TME = true metabolisable energy (poultry)
TOBEC = total body electrical conductivity
TOH = tritiated water
Totox = 2PV+AV
TRIG = triglyceride
UDP = (rumen) undegradable protein (syn. RUP, bypass protein)
UEL = unité encombrement lait (fulfilment unit for dairy cow)
UEM = unité encombrement mouton, syn., fulfilment unit, FU for sheep))
UFC = unité fourragère cheval (horse net energy unit, France)
UFP = urea fermentation potential. g
VFA = volatile fatty acid
VLDL = very low density lipoprotein
vs. = versus
W0.75 = metabolic body mass/size
1154
JAV_SEQ_VND_FSGy:MINTAÚJ.qxd 2008.09.03. 16:42 Page 1155
SELF-EVALUATIONG QUESTIONS
A) ESSAY OR DEFINITIONS TYPE QUESTIONS
After the hypotheses what may have been the first

type of digestion at the ancestor mammalians?
Compare CT and DEXA methods (analogies and dif-
ferences).
Compare the body composition of plants and ani-
mals.
Consequences of fibre deficiency in the rabbit dietet-
ics
Consequences of subacute T–2 toxin intake by rab-
bits
Describe in a few sentences the principles involved
in determining each fractions of the proximate
analysis.
Describe the intensive, semi intensive and extensive
breeding system of rabbits.
Describe the mechanism of caecotroph production,
fate of globules in the stomach, and the regula-
tion of their intake.
Describe the pathomechanism of the antibiotics-
treatment enteropathy
Give the characteristics of the pregnancy needs of
the pregnant rabbit
Give the meaning of the following terms: bomb
calorimetry, chromatography, pH, NDF, ADF,
ADL.
Give the nutrients requirements of the contemporar-
ily pregnant and suckling doe.
How can be applied the Archimedes Law in predict-
ing body composition?
What are dioxins and how the total amount of differ-
ent dioxins can be expressed? (TEQ:…..)
1155
SELF EVALUATION QUESTIONS
How can we get the N.f. e. content of a feedstuff?

How can you describe the voluntary feed intake of
the breeding doe during the different stages of
reproductive life?
How to express the rabbit’s energy requirements?
How to feed the female replacement growers?
If the INQ of milk for calcium is 4, how many milk
will cover the daily requirement of a man?
Is the allotriophagia a special type of trophallaxis?
It is not necessary for registration of new feed addi-
tives
Overfeeding of those plants may cause calcinosis
(Solaneum malacoxygeum, ergot)
Rules for the use of antibiotics, hormones and GMO
in US and EU are the…..(same/not same.
Significance of the alkaline value of feed mixture
Special features of the feeding and nutrition of the
breeding bucks.
Specific features of breedings bucks in energy – sex-
ual cycle – libido
State the main functions of water in the animal
body.
What are the consequences of fibre deficiency
(and/or Carbohydrate Overload of the Hind Gut)
What are the differences (zoology, taxonomy, biology)
between hare and rabbit?
What are the features of a good indicator material
during a dilution method for measuring body
composition? Give some examples.
What is the disadvantage of the rice bran?
What are the maintenance requirements of the
breeding doe’s of different categories?
What is the principle of the Body Condition Scoring?
What is the relationship between the extinction of
the dinosaurs and the development of mammals?
1156
Describe the basic mechanisms of the nutrige-

nomics.
Why are rabbits extremely against mycotoxin (e. g.
T–2 toxin peroral intake)?
Why has addition of lactic acid to the drinking water
a preventive effect against rabbit diarrhoea?
B) MULTIPLE CHOICE QUESTIONS
1. The Keshan disease in China is

a) a cardyomyopathy caused by Se deficiency
b) a cervical paralysis
c) a curled toe paralysis
d) an allergy with skin syndromes
2. Which of the following statements concerning aging is

true?
a) Dogs of various breeds age at the same rate.
b) Aging is associated with increased susceptibility to injury
and decreased capacity to resist or repair injury.
c) A group of elderly dogs is likely to be more homogeneous
than a group of young dogs.
d) Aged dogs always require more dietary protein than pup-
pies do.
e) Chronologic age is a reliable indicator of functional age.
3. In the short term, weighing patients frequently is most

useful in evaluating the status of
a) water b) protein
c) carbohydrate d) calcium
4. A body condition score of 5 of 5 BCS system indi-

cates
a) emaciated body condition. b) thin body condi-
tion.
c) obesity d) moderately fleshy body condition.
1157
5. The protein requirements of old dogs

a) are always decreased from the maintenance level for
young adult dogs.
b) are always greater than the maintenance level for young
adult dogs.
c) must be individualized
d) must be met only with foods of animal origin.
6. Continuous grazing is less effective than

a) rotational grazing b) strip grazing
c) both of them (a+b)
d) neither of them
7. It can help to prevent the human osteoporosis

a) vitamin Ab) vitamin D c) vitamin E d) vitamin K
8. Liberation of ammonia during heat processing of feeds

indicates that there are
a) reactions between protein molecules
b) reactions between proteins and carbohydrates
c) reactions between proteins and fats
d) reactions between carbohydrates and fats
8. If the quality of corn silage is good, it contains this acid

in the highest amount
a) acetic acid b) propionic acid
c) lactic acid d) butyric acid
9. By a mega dose of vitamin C the harmful effect of aflatox-

in B1 can be reduced in
a) dairy cow b) beef cattle c) sheep d) poultry
10. Its crude fibre content is usually higher than 20% on

DM basis
a) meadow hay b) alfalfa hay
c) both of them d) neither of them
1158
11. The secondary alimentary hyperparathyroidism (all meat

syndrome) in cats can be caused by the overfeeding of
a) Ca b) P c) Mn d) Zn
12. Tables created according to the new protein evaluation

system for ruminants express the protein content of the
feeds in the form of
a) MPN b) MPE
c) both of them (a+b) d) neither of them
13. The optimum nitrogen: sulphur ratio in dairy diets is

a) 20:1 b) 15:1 c) 10:1 d) 7:1
14. Its presence is necessary for the carotene – vitamin A

conversion in dairy cows
a) P b) Zn
15. The substitution number (SN) of corn is 0.3. It means,

that by adding 1 kg DM of corn to the forage, the DM
intake from the forage
a) decreases by 30% b) decreases by 0.3 kg
c) increases by 30% d) increases by 0.3 kg
16. Its crude protein content is between 20 and 30% on DM

basis
a) corn silage b) green alfalfa
c) meadow hay d) meadow grass
17. Atypical form of milk fever can occur when

a) the absolute Ca level is low in the diet
b) the relative blood Ca level is high related to the level
of Mg and P
c) the absolute Mg level increases in the blood
d) the relative blood Ca level is low related to the level of
Mg and P
1159
18. They are storage fungi

a) Alternaria genus c) Cladosporium genus
b) Fusarium genus d) Aspergillus genus
19. The INQ value for vitamin A of liver is 100 for human. It
means that
a) related to the requirements liver provides 100 times
more energy than vitamin A for human.
b) the absolute energy content of the liver is 100 times
higher than its vitamin A content
c) the absolute vitamin A content of the liver is 100
times higher than its energy content
d) related to the requirements liver provides 100 times
more vitamin A than energy for human.
20. One of its deficiency syndromes is distortion of breast-

bone in poultry
a) vitamin E b) vitamin D
21. If the by-pass category of a feedstuff is low, it means

that the most of its protein content
a) will be degraded in the rumen
b) can go through the rumen without degradation
c) will not be degraded neither in the rumen nor in the
large intestine
d) can go back into the rumen in amino acid form by the
rumino-hepatic cycle
22. Tables can contain not only the total phosphor content
of the feeds but also the amount of
a) digestible P b) available P
c) precipitated P d) inorganic P
23. Its protein requirement in the practice may be expressed

in crude protein
a) Angora rabbit b) growing pig
c) laying duck d) riding horse
1160
24. The supply of dairy cows can be well evaluated accord-

ing to its concentration in the milk
a) sodium b) iodine
25. By giving 4-8 times more from the group of vitamin B

than the normal requirement, the lean growth can be
improved in
a) swine b) broiler chick
c) growing lamb d) neither of them
26. One of its deficiency syndromes is opisthotonus

a) vitamin B1 b) Mn
27. By increasing of the lignin content of the feeds

a) the feed intake increases
b) the feed intake doesn’t change essentially
c) the feed intake decreases
d) the nutrients’ digestibility increases
28. For determination of the nutrients’ digestibility by a

total collection method
a) registration of the feed intake and collection of the
faeces are required
b) registration of the feed intake and collection of the
faeces and urine are required
c) only the feed intake must be reiterated
d) only the faeces and urine must be collected but the
registration of feed intake is not necessary
29. Body condition score (BCS) is

a) the same as body weight
b) a physical measure that allows a definitive statement
on nutritional health
c) an estimatation of the fat cover of the body and an
indicator of body energy reserves or the lack of them
d) the degree of physical activity in an aged pet
1161
30. One of its deficiency syndromes is skipping of the

Achilles tendon in beef bulls
a) Ca b) P c) Cu d) F
31. The approximate dry matter content of fresh pasture

grass is
a) 20-25% b) 45% c) 60% d) 85%
32. The energy density expresses how many

a) crude protein is in the feedstuff related to 1 MJ NE
(CP/NE)
b) crude protein is in the feedstuff related to 1 fulfilment
unit (CP/UE)
c) fulfilment unit is in the feedstuff related to 1 g crude pro-
tein (UE/CP)
d) net energy is in the feedstuff related to 1 fulfilment unit
(NE/UE)
33. One of its deficiency syndromes in sheep may be

anaemia
a) vitamin K b) vitamin K, Fe and Cu
c) vitamin K and Fe d) vitamin K, Fe, Cu and Co
34. It published feeding recommendations

a) INQ b) ARC c) NSC d) MPE
35. Its deficiency syndrome is grass tetany

a) K b) Cl
36. Its most important antinutritive factors are saponins

a) soybean b) barley c) alfalfa d) rapeseed
37. Biological value (BV) index is

a) the proportion of nitrogen intake that is retained
b) weight gain per crude protein intake
c) proportion of ingested nitrogen that is not excreted in the
faeces
d) the proportion of absorbed nitrogen that is retained
1162
38. The discount factor of corn silage is 0.053. It means,

that by increasing the feeding level by 1 unit
a) the digestibility of the corn silage will decrease by 5.3%
b) the digestibility of the corn silage will increase by 5.3%
c) the NEl content of the corn silage must be multiplied by
0.053
d) the NEl content of the corn silage must be raised to the
0.053 power
39. They are chelates

a) S-Cu-Mo b) Met-Zn
40. Its determination is made by direct chemical analysis

a) organic matter b) N-free extract
41. NDF minus ADF equal to….

a) cellulose b) hemicellulose c) lignin d) ash
42. One of its deficiency syndromes is oedema of eyelids

a) vitamin C b) vitamin E c) vitamin B1 d) pyridoxin
43. Bulky feeds have

a) low dry matter content related to the whole volume
b) high digestible organic matter content related to the
whole volume
c) low crude fibre content related to the whole volume
d) low dry matter content related to digestible organic mat-
ter content
44. To the overheating of the feeds this amino acid is the

most sensitive one
a) Lys b) Met c) Cys d) Arg
1163
45. It is chemically determined according to its nitrogen

content
a) N-free extract b) crude protein
c) true protein d)a+b+c together
46. Characteristic of roughages

a) They are bulky feeds
b) Their fat content is higher than 8%
c) Their CF content is lower than 10% on DM basis
d) They are concentrated energy feeds
47. They belong to the group of nutriceuticals

a) antibiotics b) hormones
c) steroids d) organic acids
48. „Sneaker” syndrome caused by biotin deficiency is fre-

quent at
a) turkey b) boars c) sows d) mink
49. The protein overeating

a) reduces the ovulation rate b) causes worse nidation
c) improves nidation d) increases the body weight at birth
50. For expressing the energy requirement of beef cattle and

small ruminants for maintenance, the following term is
used in Germany
a) NEm b) ME c) NEg d) neither of them
51. They can be used as silage additives

a) cereal grains b) molasses
52. Which of the following measurements provides a defini-

tive statement on nutritional health?
a) history and diet assessment
b) physical examination
c) clinical chemistry
d) none of the above alone
1164
53. Ration of beef cattle can be supplemented by urea when

a) the UFP value is positive
b) the rumen protein balance is negative
54. PSE muscle by VESD in living animal is present

a) broiler chicken b) beef cattle
c) newborn lamb d) growing-fattening pig
55. One of its deficiency syndromes is night blindness

a) vitamin A b) Zn
55. Which biogenic amine can cause food/feed allergy with

skin syndromes?
a) tiramine b) histamine c) cadaverine d) triptamine
56. The most significant risk of drug-induced nutrient defi-

ciencies occurs in old dogs with
a) gastric torsion.
b) marginal nutrient intake and chronic disease
c) acute disease
d) subnormal temperature.
57. Green chops

a) are more digestible than hays
b) are fed dried to livestock
c) are less digestible than hays d) b + c
58. Homeostasis of Ca is regulated only on the level of

excretion in
a) dairy cowb) cat c) dog d) rabbit
59. In case of fall of hair caused by biotin deficiency in cats

one has not to differentiate it from
a) IDDM, hypothyreosis b) taurine deficiency
c) Cushing syndrome d) Zn deficiency
1165
60. The inheritable imperfect zinc absorption is more fre-

quent in
a) husky and malamute b) Puli
c) German shepherd d) Golden retriever
61. The inheritable copper retention trouble is more fre-

quent at
a) Dwarf Schnauzer b) Badlington terrier
c) Dachshund d) Hungarian Vizsla
62. Put in increasing order according the energy level of

cat’s, dog’s and cow’s milk
a) cow, cat, dog b) dog, cat, cow
c) cat, dog, cow d) cow, dog, cat
63. In case of secondary renal hyperparathyroidism the

following clinical signs will not occur:
a) opisthotonus b) anaemia
c) uraemia d) osteodystrophia fibrosa
64. The slimming diet of dog and cat may contain higher
level of
a) fibre b) low degradable starch
c) silica d) sawdust
level of
a) high degradable starch b) L-carnitine
c) sawdust d) ascorbic acid
level of
a) high degradable starch b) silica c) vitamin A
level of
a) high degradable starch b) silica
c) trivalent (organic) chromium d) NSC
1166
68. The maintenance energy requirement of dog could be

calculated by using the
a) metabolic body weight (W0.75) b) live weight
c) body length d) hearth girth
69. The acidification of the cat’s urine helps to prevent the

formation of
a) cystine stone b) oxalate stone
c) struvite stone d) urate stone
70. Forced and quick weight reduction program in obese

cats may cause
a) chronic renal failure b) dilated cardiomyopathy
c) hepatic hyperlipidosis d) cerebrocortical necrosis
71. In case of congestive heart failure one has to choose

feeds of
a) low in potassium b) high in potassium
c) low in sodium d) high in sodium
72. The cat’s capacity to transform carotene into vitamin A

is
a) 20% b) 10% c) zero % d) 50%
72. Lack of this compound in the cat’s diet may cause

dilated cardiomyopathy and central retina degeneration
a) dl-methionine b) L-carnintine
c) taurine d) thiamine
73. When using the method of direct calorimetry this

parameter will be also measured
a) O2 consumption c) both of them (a+b)
b) CO2 production d) neither of them
74. A dog is considered to be geriatric when it reaches how

many years of age?
a) 10 b) 5 c) 6
d) none of the above (Because canine aging is a heteroge-
neous process, no single age applies to all dogs.)
1167
75. One of its deficiency syndromes in birds is the cervical

paralysis (stretched neck)
a) biotin b) pantothenic acid
c) folic acid d) niacin
76. For determination of digestible energy by experimental

way we calculate with the
a) gross energy content of the feed and that of the faeces
b) net energy content of the feed and the gross energy con-
tent of the faeces
c) gross energy content of the feed, faeces and urine
d) gross energy content of the feed, faeces, urine and gases
77. The most important factor influencing the onset of the

first fertile oestrus in dairy heifers
a) energy b) protein c) P d) Mn
78. Micro mineral having similar skin and mucous mem-

brane effect as the vitamin A
a) Zn b) Fe c) Cu d) Mn
79. Mode of action of the intestinal bacterial defence mecha-

nism:
a) competitive exclusion c) bile acid binding
b) antibiotic degradation d) toxic amine production
80. UFP can be calculated as follows

a) MPE-MPN b) fat content c) CP-MPN d) dg, TDN
81. The most important regulator of the long-term control of

feed intake is
a) the adipose tissue b) hunger-satiety centre
c) hypothalamus d) b + c together
1168
82. Forages generally have

a) higher P content than Ca content
b) about the same level of Ca and P
c) higher Ca than P content
d) higher P content than that of the seeds
83. What is the meaning of “indicator bacteria”?

a) These bacteria will produce red colour in acidic cir-
cumstances.
b) Their presence allows us to suppose the (faeces) con-
tamination with pathogens
c) Bacteria used to produce methyl-orange indicator
d) Neither of them
84. AgY is
a) an antigen in the egg yolk having IgA effect
b) an immune deficiency syndrome in rats
c) a feeding standard
d) a gene responsible for high body fat level
85. As age increases, canine lean body mass generally

a) decreases. b) increases.
c) stabilizes d) does’nt change its ratio to body fat.
86. Chemical score (CS) means

a) weight gain per crude protein (CP) intake
b) retained nitrogen per absorbed nitrogen
c) retained nitrogen per nitrogen intake
d) limiting amino acid in the feed protein per the same amino
acid in the standard protein
87. It can cause secondary Cu deficiency

a) S b) Mo c) both of them (a+b) d) neither of them
88. The “k” value means

a) ME/GE b) ME/MEm c) GE/NE d) NE/ME
1169
89. It has high buffering capacity

a) corn silage b) alfalfa silage
90. It has effects typically on the females’ ovaria

a) Cu b) Zn c) Mn d) Mo
91. The unit of measurement of TDN is

a) MJ/kg DM b) kJ/g DM c) g/MJ d) g/kg DM or %
92. The first limiting amino acid for milk production

a) Lys b) Met c) Cys d) Arg
93. The protein: energy (P: E) ratio of a feedstuff means

a) the amount of protein in gram related to 1 MJ energy
b) the amount of energy in MJ related to 1 g protein
c) the energy content of 1 g protein in the feed in MJ
d) The amount of protein in gram related to 1 g carbohy-
drates
94. The premix is a mixture of

a) microelements only
b) vitamins and micro elements
c) vitamins, microelements, Ca and P
d) vitamins, microelements, Ca, P and amino acid supple-
ments
95. Metabolizable protein means

a) the digestible microbial true protein + digestible UDP in the
small intestine
b) the digestible crude protein + digestible RDP in the small
intestine
c) the digestible microbial true protein + digestible UDP in
the rumen
d) the digestible crude protein + digestible RDP in the
rumen
96. In case of its deficiency the ruminal protein synthesis

can decrease
a) Se b) S c) Mg d) I
1170
97. The equation for predicting dry matter intake of dairy

cow we calculate with the
a) body weight and environmental temperature
b) metabolic body weight and milk production
c) milk production and NEm content of the feedstuff
d) milk production and environmental temperature
98. If the ration of dairy cows is deficient both in Ca and P

a) first you eliminates the Ca deficiency and after that the P
deficiency by supplements
b) first you stop the P deficiency and after that the Ca defi-
ciency by supplements
c) first you increase the amount of grains to give more P
than you increase the amount of forages to give more Ca
d) first you increase the amount of forages to give more Ca
than you increase the amount of grains to give more P
99. It is an omega-6 type fatty acid

a) oleic acid b) arachidonic acid
c) linolenic acid d) neither of them
100. The meaning of “Earth alkali alkalinity” is the follow-

ing.
a) in growing animals a surplus of alkaline metabolites is
produced which should be buffered by feeds having
acidic characters
b) in growing animals a surplus of acidic metabolites is pro-
duced which should be buffered by feeds having alkaline
characters
c) the chelate forming compounds of the feed must be in
biologically active form
d) the homeostasis of minerals can be regulated by feeds
neutral characters
101. The energy intake goal for geriatric pets is

a) 20% restriction b) 20% enhancement
c) energy balance d) increasing crude fibre intake
to decrease energy density
1171
102. Sum of adipocytes are

a) organs for fat deposit b) endocrine organs
103. This is the horse energetic feed unit in France:

a) ME b) MP c) UFC d) UFP
104. It a gastric hormon controlling feed intake

a) leptin b) ghrelin c) NPY d) galanin
105. They have high copper content in their milk

a) ruminants b) strict carnivores
c) omnivores d) mules
106. Metallothioneins are

a) proteins chararterized affinity to metals
b) steroids c) enzymes d) feed additives
107. Cannibalism may be caused by deficiency of

a) sodium b) potassium c) manganese d) silica
108. Iron level in milk is low, because

a) mammary gland cannot excrete iron
b) iron is an anti vitamin E
c) to prevent mastitis
d) biglets can get it from the dam’s faeces
109. The idiopathic hypercalcaemia is not uncommon in

a) pigs b) turkeys c) cats d) dairy cow
110. How can we express the total amount of different diox-

ins as
a) TDN b) TEQ c) INQ d) ADR
110. Rules for the use of antibiotics, hormones and GMO in

US and EU are
a) different b) the same c) depend upon the animal
species d) exept apes are the same
1172
111. The modern food pyramide comprises……, too

a) exercise (physical activity) b) feed additives
c) aflatoxin intake d) avoiding of smoking
112. The central organ of the food safety in Europe is the

a) ICLAS b) FELESA c) EFSA d) UNESCO
113. It cannot provoke a ketoacidotic state
a) diabetes b) lactacidaemia
c) ketosis d) antiepileptic diet
114. Oxidation of this compound acidifies

a) Met b) Lys c) Arg d) Trp
115. In order to acidify the cat’s urine, one can use

a) Met tablets b) aluminium hydroxide
c) sodium chloride d) silica
116. Acidifying diet may not be good (benevolent)

a) dry cow b) periparturient breeding sow
c) peritonitis d) urolith (struvite) prevention
117. Total absence of Zn absorption may cause death,

because of
a) impaired growth b) perosis c) anaemia
d) lack of superoxide dimutase (SOD) activity
118. The hair analysis is a very good indicator of ……status

in organism
a) P, S, Mg b) Si, mAl, B
c) Zn, Cu, Mn, Se d) CAB (cation anion balance)
119. They are very important in the immune response and

cancer prevention
a) Si, As b) Cd, Pb, Al c) Zn, Se d) F, K
1173
120. Clinical signs of its toxicosis may be similar to the

swine mouth-and-foot disease (MFD)
a) nitrit, nitrate b) vitamin E
c) Se d) Cu
121. The calcitriol is a(n)

a) hormon b) active vitamin
c) both d) neither
122. „Kincky back” of turkeys is

a) Ca deficiency b) vitamin D deficiency
c) vitamin C deficiency d) genetic failure
123. One of these abbreviations is not a scientific organ

a) INRA b) DLG c) ARC d) NEL
124. Its overdosage may cause toxicosis

a) thiamin b) niacin c) folic acid d) axerophtol
125. Its overdosage may cause toxicosis

a) panthotenic acid b) biotin
c) L-carnitine d) cholecalciferol
126. Its manifestation is both genetically and environmen-

tally determined
a) scurvy b) rickets
c) cancer d) haemophilia
127. Immunostimulant effect of a vitamin can be evaluated

by
a) lymphocyte proliferation (mitotic index)
b) hematocrit c) RDR d) liquor analysis
128. It may help in differentiating avian nephritis and vita-

min A deficiency in chicken
a) visceral gout b) inclusion body (histopathology)
c) lameness d) blindness
1174
129. Pirrolo-quinoloquinon (PQQ), the recent dicovered vita-

min can be found in
a) orange b) carotte c) papaya d) alfalfa
130. It has no gulonolacton-oxidase enzyme

a) turkey b) turtle c) pig d) bat
131. It has no gulonolacton-oxidase enzyme

a) cattle b) dog c) rat d) shrim
132. It improves (allows, facilitates) the effect of vitamin D

a) biotin b) Si c) vitamin C
d) GHSH-Px (glutation peroxydase)
133. Its deficiency in egg may cause typical embryonal mor-

tality at the chicken
a) B2 b) B12 c) neither of them d) both of them
134. Taurine is a vitamin-like substance only for ........

a) insects b) turkeys
c) pigs d) cats
157. It may increase the sow’s litter size

a) panthotenic acid b) folic acid
c) neither of them d) both of them
1175

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JAV_Kolofon_VND_FSGy7:MINTA J.qxd 2008.09.03.

Veterinary Nutrition and Dietetics

NUTRITION AND DIETETICS

Sándor Gy. Fekete, DVM, PhD, DSc

“Pro Scientia Veterinaria Hungarica“

“Pro Scientia Veterinaria Hungarica“ Fooundation

First Paperback Editon 2008

Reviewers of the Hungarian Version

Reviewers of the English Version (selected chapters):

The Publisher: “Pro Scientia Veterinaria Hungarica“ Foundation

Printing office: OOK-Press Kft, Veszprém, Hungary.

his English-language textbook is intended for veterinary students, vet-

T erinary practitioners and other professionals working in the field of

Prof. Dr. habil. László BABINSZKY, PhD (Wageningen)

Assoc. Prof. Dr. András BERSÉNYI, DVM, PhD (Budapest

Assoc. Prof. Dr. Géraldine BLANCHARD, DVM, PhD (Alfort)

Assoc. Prof. Dan L. BROWN, PhD (Cornell, NY)

Prof. Dr. Tony C. A. BUFFINGTON, DVM, PhD (Davis, CA)

Prof. Dr. habil. Burghard JILGE, DVM, PhD

Prof. D. Carlos CASTRILLO Gonzales, PhD (Leon)

Prof. Dr. habil. Éva CENKVÁRI, MSc, CSc

Prof. André CHWALIBOG, DSc (University of Copenhagen)

Prof. Dr. Sándor György FEKETE, DVM, MSc, PhD (Moscow-Budapest),

Prof. Dr. Jose Antonio GUADA Vallepuga, DVM, PhD

Prof. Dr. Gyula HUSZENICZA, DVM, PhD (Budapest),

Prof. Dr. Christine IBEN, DVM, PhD (Vienna)

Prof. Dr. hab.dr h.c., dr h.c. Dorota JAMROZ

Prof. Geert P. J. JANSSENS, PhD (Ghent)

Prof. Dr. Ǻshild KROGDAHL, PhD (Oslo)

M.Sc. eng. Janusz KUBIZNA

Prof. Dr. Dr. h.c. mult. Josef LEIBETSEDER, DVM

Dr. Richard C. NAP, DVM, PhD (Utrecht)

Dr. Fatma Karakas OGÜZ, DVM, PhD (Burdur)

Prof. Dr. Merel RITSKES-HOITINGA, PhD, Dipl ECLAM

Bart SAVENIJE, PhD (Groningen)

Prof. Ewa SAWOSZ, Dr. habil.

Prof. Anne-Helene TAUSON, PhD

Dr. Angela TROCINO, PhD (Padova)

Prof. Gerolamo XICCATO, PhD (Padova)

Prof. Dr. Annette ZEYNER, Habil. (Göttingen)

XI. Minerals in the animal nutrition (Sandor Gy. FEKETE)

B) APPLIED ANIMAL NUTRITION

XIX. Evolution of digestion (Sandor Gy. FEKETE)

27.1. ”Standard” European goat

The evolution of veterinary nutrition and clinical dietetics into

sponding to the recommendations, proper animal care, the ration-

has developed into an independent discipline separately for all

1.1. Definition of feed

CHAPTER I: BASIC NOTIONS - IMPORTAQNCE-ROLE OF WATER

CHAPTER I: BASIC NOTIONS - IMPORTAQNCE-ROLE OF WATER

1.2. Main features of feedstuffs (pyramid-conception)

CHAPTER I: BASIC NOTIONS - IMPORTAQNCE-ROLE OF WATER

1.2.8. Taste, flavour and smell.

1.3. Importance of protein

*pig, cattle, poultry and fish **except butter

replaceable by the vegetable ones. On average, in the nutrition of the world

Conversion of feed protein: if the efficiency of production is expressed

CHAPTER I: BASIC NOTIONS - IMPORTAQNCE-ROLE OF WATER

Based on these data one can conclude that – on population level –

1.4. Water, electrolyte and acid-base balance

uge area of the Earth’s surface (363000 km2) is covered by water.

H Without water, there would be no life on Earth. Approx. 70% of the

CHAPTER I: BASIC NOTIONS - IMPORTAQNCE-ROLE OF WATER