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Lawrie’s Meat Science
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Related Titles

New Aspects of Meat Quality

(978-0-08-100593-4)
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(978-0-08-100763-1)
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(978-0-08-101215-4)
Woodhead Publishing Series in Food Science,
Technology and Nutrition

Lawrie’s Meat Science

Eighth Edition

Edited by
Fidel Toldrá
Woodhead Publishing is an imprint of Elsevier
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Contents

List of Contributors xiii

Preface xv

1. Introduction
Jeffrey W. Savell
1.1 Meat and Muscle 1
1.2 Meat From Other Animals 2
1.3 Domestication of Livestock 4
1.4 Trends and Developments 7
1.5 Conclusions and Future Trends 15
References 16

2. Factors Influencing the Growth of Meat Animals

Aidan P. Moloney, Mark McGee
2.1 Introduction 19
2.2 Measurement of Growth and Body/Carcass Composition 20
2.3 Animal Influences on Growth of Farm Animals 22
2.4 Nonanimal Influences on Growth of Farm Animals 29
2.5 Interactions Between Animal and Nonanimal Influences
on Growth of Farm Animals 36
2.6 Future Developments 41
References 42

3. The Structure and Growth of Muscle

Peter P. Purslow
3.1 Introduction 49
3.2 Muscle Tissue 50
3.3 Generalized Skeletal Muscle Structure 51
3.4 The General Structure of the Striated (Skeletal) Muscle Cell 52
3.5 Muscle Fiber Types 59
3.6 Structure of the Individual Intramuscular Connective
Tissues 61
3.7 Variations in Intramuscular Connective Tissue Content
Between Muscles 64
3.8 Composition of Intramuscular Connective Tissues 64

v
vi Contents

3.9 Collagens in Muscle 64

3.10 Collagen Cross-Linking 67
3.11 Microfilaments and Elastin 68
3.12 Glycoproteins and Proteoglycans 71
3.13 CelleMatrix Connectors: Laminin and Fibronectin,
Dystrophin, and the Integrins 72
3.14 The Structure of Adipose Tissues 76
3.15 Muscle Development and Growth 77
3.16 Muscle Development in Embryogenesis and Prenatal
Growth 77
3.17 Adipogenesis and Fibrillogenesis 79
3.18 Fetal Development 81
3.19 Postnatal Muscle Growth 81
3.20 Muscle Cells Grow by Incorporating Satellite Cells 85
3.21 Distribution and Percentage of Fat and Lean Muscle
in the Growing Animal 85
3.22 Muscle Cuts 88
3.23 Conclusions and Future Trends 90
References 91

4. Chemical and Biochemical Constitution of Muscle

Clemente López-Bote
4.1 General Chemical Aspects 99
4.2 Biochemical Aspects 116
4.3 Factors Affecting Meat Quality 132
4.4 Conclusions and Future Trends 149
References 150

5. The Conversion of Muscle to Meat

Sulaiman K. Matarneh, Eric M. England, Tracy L. Scheffler,
David E. Gerrard
5.1 Introduction 159
5.2 Postmortem Metabolism 160
5.3 The Factors Controlling the Rate of Postmortem
Metabolism 167
5.4 The Factors Controlling the Extent of Postmortem
Metabolism 168
5.5 Abnormal Postmortem Metabolism 170
5.6 Preslaughter Stress 173
5.7 Development of Meat Quality Attributes 175
5.8 Postmortem Handling and Meat QualitydTemperature 178
5.9 Aging and Proteolysis 179
5.10 Conclusions 182
References 182
 Contents vii

6. Meat Microbiology and Spoilage

Monique Zagorec, Marie-Christine Champomier-Vergès
6.1 Introduction 187
6.2 Origin and Dynamics of Meat Microbial Contamination
Involved in Spoilage 189
6.3 Mechanisms Involved in Meat Bacterial Spoilage 191
6.4 The Main Microbial Contaminants of Meat Involved in
Spoilage 193
6.5 The Main Spoilage Manifestations and Their Microbial
Causes 197
6.6 Future Trends 199
References 200

7. The Storage and Preservation of Meat: IdThermal

Technologies
Youling L. Xiong
7.1 Introduction 205
7.2 Chilling 206
7.3 Freezing 216
7.4 Heating 220
7.5 Novel Thermal Procedures 224
7.6 Future Trends 226
References 226

8. The Storage and Preservation of Meat:

IIdNonthermal Technologies
Dong U. Ahn, Aubrey F. Mendonça, Xi Feng
8.1 Introduction 231
8.2 Ionizing Radiation 232
8.3 High Pressure 247
8.4 Freeze Dehydration 255
References 259

9. The Storage and Preservation of Meat: IIIdMeat

Processing
Fidel Toldrá
9.1 Introduction 265
9.2 Curing 266
9.3 Fermentation 273
9.4 Dehydration 277
9.5 Smoking 280
viii Contents

9.6 Processing Technologies for Cured Meat Products 281

9.7 Conclusions and Future Trends 292
References 292

10. Storage and Preservation of Raw Meat

and Muscle-Based Food Products: IV Storage
and Packaging
Joe P. Kerry, Andrey A. Tyuftin
10.1 Introduction 297
10.2 Impact of Microbiology on Fresh Meat Quality
Attributes 298
10.3 Common Technologies Used to Preserve Fresh Meat
Products and Assist in a Combined Manner
to Extend Product Shelf Life 299
10.4 Packaging Materials Used for Fresh Meat and
Muscle-Based Products 309
10.5 Conclusions 320
References 324
Relevant Websites 327

11. The Eating Quality of Meat: IdColor

Cameron Faustman, Surendranath P. Suman
11.1 Introduction 329
11.2 Myoglobin Concentration 329
11.3 Myoglobin Structure 332
11.4 Color Phenomena in Fresh Meat 336
11.5 Color in Cooked Nitrite-Cured and Salted Uncooked
Meats 343
11.6 Cooked Meat Color 345
11.7 Anomalies in Meat Color 347
11.8 Measuring Meat Color 350
11.9 Summary Statement 351
References 351

12. The Eating Quality of Meat: IIdTenderness

David L. Hopkins
12.1 Introduction 357
12.2 Measuring Tenderness 374
12.3 Conclusions and Future Trends 375
References 376
 Contents ix

13. The Eating Quality of Meat: IIIdFlavor

Mónica Flores
13.1 Aroma and Taste Compounds 383
13.2 Volatile Compounds Generation Reactions 384
13.3 Methodology for Meat Aroma Volatile Identification 392
13.4 Pre- and Postslaughter Factors Affecting Aroma 398
13.5 Off-Flavors 403
13.6 Meat Product Flavor 407
13.7 Conclusions and Future Trends 412
References 412

14. The Eating Quality of MeatdIV Water-Holding

Capacity and Juiciness
Robyn D. Warner
14.1 Introduction 419
14.2 Definition of Water-Holding Capacity and
Juiciness 419
14.3 Structural Influences on the Water-Holding Capacity
of Uncooked (Raw) Meat 422
14.4 Factors Influencing Water-Holding Capacity in Raw
Muscle 424
14.5 Changes in Water-Holding Capacity During Cooking
of Raw Meat 430
14.6 JuicinessdInfluencing Factors and Interactions With
Water-Holding Capacity 437
14.7 Factors Influencing Water-Holding Capacity of Meat
Products 438
14.8 Methods to Measure Water-Holding Capacity and
Juiciness 446
14.9 Conclusions and Future Trends 452
References 453

15. The Eating Quality of Meat: VdSensory Evaluation

of Meat
Rhonda K. Miller
15.1 Introduction 461
15.2 Why Sensory Evaluation of Meat Is Unique 462
15.3 Overview of How Sensory Is Perceived and Defining
Sensory Attributes 463
15.4 Sensory Controls for Meat 472
15.5 Sensory Techniques 478
15.6 Emerging or Underutilized Sensory Techniques 496
15.7 Conclusions 498
References 498
x Contents

16. Phenotyping of Animals and Their Meat:

Applications of Low-Power Ultrasounds,
Near-Infrared Spectroscopy, Raman Spectroscopy,
and Hyperspectral Imaging
Donato Andueza, Benoıˆt-Pierre Mourot,
Jean-François Hocquette, Jacques Mourot
16.1 Introduction 501
16.2 Principles of the Main Methods 502
16.3 Applications in Meat Quality Assessment 506
16.4 Conclusions 513
Acknowledgment 514
References 514

17. Meat SafetydI Foodborne Pathogens and Other

Biological Issues
Alexandra Lianou, Efstathios Z. Panagou, George-John E. Nychas
17.1 Introduction 521
17.2 Foodborne Illnesses Associated With the Consumption
of Meat and Meat Products 522
17.3 Bacteria and Bacterial Toxins 523
17.4 Viruses 538
17.5 Parasites 541
17.6 Prions 543
17.7 Current and Emerging Challenges to Meat Safety
Management 544
17.8 Concluding Remarks and Future Trends 546
References 546

18. Meat Safety: II Residues and Contaminants

Marilena E. Dasenaki, Nikolaos S. Thomaidis
18.1 Introduction 553
18.2 Chemical Contaminants and Residues 554
18.3 Risk Assessment 567
18.4 Analytical Methods 568
18.5 Future Trends and Perspectives 577
References 578

19. Meat Authenticity and Traceability

Luca Fontanesi
19.1 General Overview 585
19.2 Intrinsic Characteristics of the Meat 587
19.3 Extrinsic Characteristics of the Meat 618
19.4 Conclusions and Future Trends 622
References 623
 Contents xi

20. Meat Composition and Nutritional Value

Jeffrey D. Wood
20.1 Introduction 635
20.2 Meat Consumption Patterns 635
20.3 Composition of Meat 636
20.4 Effects of Cooking on Nutrients in Meat 653
20.5 Conclusions and Future Possibilities 654
References 655

21. Meat and Health

Kerri B. Gehring
21.1 Introduction: Nutrients Supplied From Meat 661
21.2 Meat in Healthy Nutrition and Diet 663
21.3 Recommended Meat Intakes 668
21.4 Functional Muscle Foods 670
21.5 Problematics With Toxins and Residues 671
21.6 Conclusions 674
References 674

22. Edible By-products

Herbert W. Ockerman, Lopa Basu, Fidel Toldrá
22.1 Introduction 679
22.2 Main Edible By-products 680
22.3 Nutritional Value of Edible By-products 684
22.4 Products Resulting From Edible By-products 691
22.5 Added Value Products Obtained From Edible
By-products 693
22.6 Conclusions and Future Trends 694
References 694

Index 697
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List of Contributors

Dong U. Ahn, Iowa State University, Ames, IA, United States

Donato Andueza, INRA, UMR1213 Herbivores, Saint-Genès-Champanelle, France;
Clermont Université, VetAgro Sup, UMR1213 Herbivores, Clermont-Ferrand,
France
Lopa Basu, University of Kentucky, Lexington, KY, United States
Marie-Christine Champomier-Vergès, UMR1319, MICALIS, INRA, Université
Paris-Saclay, Jouy-en-Josas, France
Marilena E. Dasenaki, University of Athens, Athens, Greece
Eric M. England, The Ohio State University, Columbus, OH, United States
Cameron Faustman, University of Connecticut, Storrs, CT, United States
Xi Feng, Iowa State University, Ames, IA, United States
Mónica Flores, Instituto de Agroquı́mica y Tecnologı́a de Alimentos (CSIC), Valencia,
Spain
Luca Fontanesi, University of Bologna, Bologna, Italy
Kerri B. Gehring, Texas A&M University, College Station, TX, United States;
International HACCP Alliance, College Station, TX, United States
David E. Gerrard, Virginia Polytechnic Institute and State University, Blacksburg,
VA, United States
Jean-François Hocquette, INRA, UMR1213 Herbivores, Saint-Genès-Champanelle,
France; Clermont Université, VetAgro Sup, UMR1213 Herbivores, Clermont-
Ferrand, France
David L. Hopkins, Centre for Red Meat and Sheep Development, Cowra, NSW,
Australia
Joe P. Kerry, University College Cork, Cork City, Ireland
Alexandra Lianou, Agricultural University of Athens, Athens, Greece
Clemente López-Bote, Universidad Complutense de Madrid, Madrid, Spain
Sulaiman K. Matarneh, Virginia Polytechnic Institute and State University,
Blacksburg, VA, United States
Mark McGee, Teagasc, Grange, Dunsany, Co. Meath, Ireland
Aubrey F. Mendonça, Iowa State University, Ames, IA, United States
Rhonda K. Miller, Texas A&M University, College Station, TX, United States

xiii
xiv List of Contributors

Aidan P. Moloney, Teagasc, Grange, Dunsany, Co. Meath, Ireland

Benoı̂t-Pierre Mourot, INRA, UMR1213 Herbivores, Saint-Genès-Champanelle,
France; Clermont Université, VetAgro Sup, UMR1213 Herbivores, Clermont-
Ferrand, France; Valorex, Combourtillé, France
Jacques Mourot, INRA, UMR 1348 PEGASE, St-Gilles, France; Agrocampus Ouest,
UMR1348 PEGASE, Rennes, France
George-John E. Nychas, Agricultural University of Athens, Athens, Greece
Herbert W. Ockerman, Ohio State University, Columbus, OH, United States
Efstathios Z. Panagou, Agricultural University of Athens, Athens, Greece
Peter P. Purslow, National University of Central Buenos Aires Province, Tandil,
Argentina
Jeffrey W. Savell, Texas A&M University, College Station, TX, United States
Tracy L. Scheffler, University of Florida, Gainesville, FL, United States
Surendranath P. Suman, University of Kentucky, Lexington, KY, United States
Nikolaos S. Thomaidis, University of Athens, Athens, Greece
Fidel Toldrá, Instituto de Agroquı́mica y Tecnologı́a de Alimentos (CSIC), Valencia,
Spain
Andrey A. Tyuftin, University College Cork, Cork City, Ireland
Robyn D. Warner, Melbourne University, Parkville, VIC, Australia
Jeffrey D. Wood, University of Bristol, Bristol, United Kingdom
Youling L. Xiong, University of Kentucky, Lexington, KY, United States
Monique Zagorec, UMR1014 SECALIM, INRA, Oniris, Nantes, France
Preface

When I was contacted by the publisher to prepare the eighth edition, my first
thoughts were focused on the enormous responsibility that I was assuming. In fact,
and after talking with many colleagues, I think that most of the meat scientists
worldwide have learned about meat with any of the earlier editions of Lawrie’s
Meat Science book. This is the type of book that meat scientists must have on hand
in their personal library. I personally learned a lot about meat science from
the previous editions of this book. This is why I am so grateful to Professor
Ralston Lawrie for his initiative in writing this nice book and publishing its first
edition in 1966 and the following editions and also to Professor Dave Ledward for
the work done in updating and expanding the recent editions of the book.
The main goal of the book is to provide the reader with a comprehensive
resource, covering the wide field of meat science. This means from the production
of animals, the structure of the muscle, its conversion into meat, the different
technologies used for preservation and storage and the eating and nutritional
quality and safety of meat through the processing industry and distribution until
reaching the consumer.
The book includes leading-edge technologies (i.e., nanotechnology, novel
preservation technologies) and techniques (i.e., proteomics, genomics, metab-
olomics) in chapters not only related with meat quality, nutritional value, and
meat safety but also in other relevant issues such as traceability and authenticity,
which have a strong demand from all sectors involved in “farm to fork” and have
been the focus of some recent scandals not only in the European Union but also
in other areas of the world.
After 50 years since the book was first published, this eighth edition is facing
new times and has got considerable changes in relation to the previous editions.
The most relevant change that you will immediately notice is the format of the
book that is now an edited book with multiauthored chapters. This means that
each chapter is authored by well-known scientists having an excellent expertise
on the respective topic. These authors have contributed to get the chapters
completely rewritten and updated. Another relevant change is the breakdown
into more specific chapters. Some chapters remain with similar titles although
substantially revised and updated in content, such as the introduction, the factors
affecting the growth and development of meat animals, the structure and growth
of muscle, the chemical and biochemical constitution of muscle, the conversion
of muscle to meat, meat microbiology and spoilage, and meat composition and
nutritional value. Other chapters have been completely changed like the set of

xv
xvi Preface

chapters on storage and preservation of meat, which are now split into four
chapters: thermal and nonthermal technologies, processing and packaging, and
storage. The eating quality of meat is split into six chapters: color, tenderness,
flavor, water-holding capacity and juiciness, sensory evaluation, and latest
technologies for assessment of quality. There are two new chapters under meat
safety: foodborne pathogens and other biological issues, and residues and
contaminants. Finally, the book also includes new chapters on topics of current
relevant interests such as authenticity and traceability, edible by-products, and
meat and health.
I sincerely hope that readers will find this book of interest and providing
useful information. I wish to thank all the contributors for their hard work and
good job done with the delivered chapters and making this book possible.
I also wish to thank the production team at Woodhead Publishing, especially
Mrs. Karen Miller, the Editorial Production Manager, Lisa Jones, the Senior
Project Manager, and Mr. Robert Sykes, the Acquisitions Editor, for their
dedication during the preparation and elaboration of the chapters and during the
publication of this book.

Fidel Toldrá
Editor
Chapter 1

Introduction
Jeffrey W. Savell
Texas A&M University, College Station, TX, United States

Meat science is a discipline that requires a complete understanding of the

complexities of antemortem and postmortem factors that impact the final
product for the consumer. Subsequent chapters in this book will delve into
these factors more deeply, but an overview of some of the background and
current issues related to meat production is an important starting point for this
journey.

1.1 MEAT AND MUSCLE

The basic definition of meat is the flesh of animals used for food. For the most
part and for most societies, meat comes from domesticated livestock with the
primary species being cattle, hogs, and sheep. Although skeletal muscle makes
up the greatest proportion of the products produced and consumed, various
organs and other offal items are important food components for many nations
and often contribute greatly to the export markets for those countries that
produce more than what can be consumed domestically.
One example of a technical definition of meat can be found at U.S.
Department of Agriculture (2016a):
Meat. (1) The part of the muscle of any cattle, sheep, swine, or goats which is
skeletal or which is found in the tongue, diaphragm, heart, or esophagus, with or
without the accompanying and overlying fat, and the portions of bone (in bone-in
product such as T-bone or porterhouse steak), skin, sinew, nerve, and blood
vessels which normally accompany the muscle tissue and that are not separated
from it in the process of dressing. As applied to products of equines, this term has
a comparable meaning.
1. Meat does not include the muscle found in the lips, snout, or ears.
2. Meat may not include significant portions of bone, including hard bone and
related components, such as bone marrow, or any amount of brain, trigeminal
ganglia, spinal cord, or dorsal root ganglia.

Lawrie’s Meat Science. http://dx.doi.org/10.1016/B978-0-08-100694-8.00001-7

Copyright © 2017 Elsevier Ltd. All rights reserved. 1
2 Lawrie’s Meat Science

Regulatory authorities within governments must define what constitutes

“meat” for its citizens as a way to ensure proper labeling and prevention of
adulteration, and it is expected that this definition will vary from country to
country. This definition from the United States has been updated since the
occurrence of bovine spongiform encephalopathy in the mid-1980s as reflected
by the reference to the absence of items now considered as “specified risk
materials” (U.S. Department of Agriculture, 2016b).

1.2 MEAT FROM OTHER ANIMALS

Throughout the world, there are many other animals used for primary or
secondary sources of meat for consumption. The buffalo (Bubalus bubalis) is
an important source of draft power, milk, meat, and hides in many Asian
countries, with the greatest numbers present in India, China, Pakistan, and
Nepal (Nanda and Nakao, 2003). Desert camels (Camelus dromedarius), in
addition to their historic use as a transporter, their drought tolerance, and their
ability to adapt to harsh arid and semiarid zones, provide food for parts of
Africa (Kurtu, 2004; Yousif and Babiker, 1989) and the Middle East (Elgasim
and Alkanhal, 1992; Kadim et al., 2006).
The goat (Capra aegagrus hircus) is a great contributor to the development
of rural zones and people (Dubeuf et al., 2004) and historically has been a
great source of meat, milk, fiber, and skin. Dubeuf et al. (2004) stated that
goats are found on all continents, with the greatest numbers being located in
Asia (especially China and India), Africa (especially Nigeria and Ethiopia),
Europe (especially Greece and Spain), and the Americas (especially Mexico
and Brazil). For species such as goats, sometimes meat production is sec-
ondary to that of milk or fiber, which often diminishes the value of meat in the
marketplace.
The horse (Equus ferus caballus) is used as a source of human food in
some cultures, with the majority of horse meat production/importation
occurring in Asia and Western Europe (Gill, 2005). Gill (2005) also stated that
the Western European countries with the greatest amounts of horse meat
produced, exported, and/or imported were Italy, Belgium, France, and the
Netherlands. Gade (1976) stated that the acceptance of horse meat in France as
a food item for humans would be one of the few documented cases of a change
in attitude from aversion to that of acceptance and was probably driven by
food-shortage crises of the past.
For many years, the United States slaughtered horses with the majority of
the more valued cuts destined for Western Europe and the less valued cuts
remaining for use in pet food manufacturing or use in zoos. In 2005, the first
successful attempt by the US Congress to find a way to stop horse slaughter
was through an act that prevented federal monies from being used to pay the
salaries or expenses of inspectors. Even though this bill expired several years
later, the US budget passed in early 2014 reinstated the ban on the use of
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Nikolaefsk
Nikolai
Nikolaieff
Nikolas
Nil
Nile
Niles
Nima
Nimrod
Nimrûd
Nin
Nine
ninepence
nineteen
nineteenth
Ninety
Nineveh
Ninfa
ning
Ningpo
Ninguta
Ninib
ninth
Ninthly
nipped
NIPPUR
nitrates
nitro
nitrogen
nitrous
Niu
Niuchuang
NIUCHWANG
Nizam
no
Nobel
nobility
Noble
nobleman
noblemen
nobles
noblest
nobly
nobodies
nobody
Nochistongo
noes
Nogales
noise
noisemaker
noises
noisily
Noisy
nomad
nomadic
nomads
NOME
nomenclature
nominal
nominally
nominate
nominated
nominating
nomination
Nominations
nominee
nominees
Non
nonacceptance
noncommissioned
Nonconformists
noncumulative
None
nonentities
nonfulfillment
nonpayment
nonreceipt
nonrecognition
nonrecurring
nonreservation
nonresident
nonsense
noon
noose
nor
Nordenfeldt
Nordenfelt
Nordenskïold
NORFOLK
normal
normally
Norman
Norris
Norrland
Norroy
Norte
north
Northeast
northeasterly
northeastern
northeastwardly
northerly
northern
Northumberlands
Northward
northwardly
northwards
NORTHWEST
Northwestern
northwestwardly
Norton
NORWAY
Norwegian
Norwegians
nose
noses
Not
notabilitles
NOTABLE
notables
notably
Notarbartolo
notarial
notary
Note
noted
Notekeeper
Notes
noteworthy
nothing
nothingness
Notice
noticeable
noticed
notices
noticing
Notification
notified
notify
notifying
noting
notion
notions
notoriety
notorious
notoriously
notulenhouder
Notwithstanding
nourish
nourishing
nourishment
Nous
Nouvelle
nouvelles
Nova
Novarum
novel
Noveleta
novelist
novels
novelty
NOVEMBER
now
Nowadays
nowhere
nowise
noxious
Noyes
Nozaleda
nozzles
Ntonda
Nubar
Nubas
Nubian
nuclear
nucleus
Nueva
NUFFAR
nugatory
nuggets
nuisance
nuisances
null
nullement
nulli
nullification
nullified
nullifies
nullify
nullifying
nullity
numb
number
numbered
numbering
Numbers
numerals
numerical
numerically
NUMEROUS
numerously
Nun
Nunciature
Nuncio
nuns
NUPÉ
Nupé
NUREMBERG
nurse
nursed
nursemaids
nurses
nursing
nurtured
nutrition
nutritive
Nyanza
Nyasa
Nyasaland
Nyassa
Nyassaland
Nâsr
o
Oahu
oak
Oakey
Oakland
oars
Oath
oaths
oats
Oaxaca
Ob
obedience
Obedient
obeisance
obelisk
obey
obeyed
obeying
obeys
Obi
object
objected
objecting
Objection
objectionable
objections
objective
objects
obligated
obligation
obligations
obligatory
oblige
obliged
obliges
obligingly
obliquely
obliterate
obliterated
obliterating
Oblivion
obnoxious
obscure
obscured
obscurities
obscurity
obsequies
observable
observance
observation
observations
OBSERVATORY
observe
observed
observer
observers
observes
observing
obsidian
obsolete
obstacle
obstacles
obstinacy
obstinate
obstinately
Obstruct
obstructed
obstructing
obstruction
obstructionist
obstructionists
obstructions
obstructive
obstructs
obtain
Obtainable
obtained
obtaining
obtains
obviate
obviated
obviating
obvious
obviously
occasion
Occasional
occasionally
occasioned
occasions
occult
occupancy
occupant
occupants
occupation
occupations
occupied
occupier
occupies
occupy
occupying
occur
occurred
occurrence
occurrences
occurring
occurs
ocean
oceanic
oceans
Oct
octave
OCTOBER
octroi
oculist
odd
odds
Odell
Oder
odes
Odessa
odious
odours
Oesterreichische
of
Off
offence
offences
offended
offenders
offending
offense
offenses
offensive
Offer
offered
offering
offers
Office
officeholders
officer
officered
OFFICERS
Offices
official
officially
officials
officio
offset
offshoot
offspring
offsprings
Oficial
Ofoo
oft
often
oftener
Ogaden
Ogadens
Ogden
Oh
OHIO
Oil
oils
Okhotsk
Oklahoma
Okuma
OLD
older
oldest
olds
oligarchy
Oliphant
olive
Olivecrona
Oliver
Olmsted
OLNEY
Olosenga
Olutanga
Olympia
Olympian
Olympic
Omaha
OMDURMAN
ominous
omission
omissions
omit
omits
omitted
omitting
omnipotent
omniscient
Omoso
Omri
Omsk
On
once
One
onerous
ones
onesided
Onesti
onlooker
Only
Onor