UNIT 1 NUTRITION, FEEDING,

DIGESTION
Structure
Introduction
Objectives
Nutrition
Proteins
Carbohydrates
Lipids
Vitamins.
Minerals and Trace Elements
Water
Feeding Mechanisms
Feeding on Small Particles
Fteding on Food Masses
Feeding on Liquids
Digestion
lntrllccllular Digestion
Digestive Tract
Dibestive Enzymes
Maintenance of Gut Lining
Coordination of Digestion
Absorption
Energy Metab-olism
Summary
Terminal Questions
Answers

1.1 INTRODUCTION
All organisms require a fairly steady supply of nutrient materials from the
environment to obtain energy in order to stay alive. You would recall from FST-1,
Unit 14 that animals are heterotrophs because they depend on already synfhesised
organic compounds from plants and other animals to obtain their food. Unlike
autotrophs (plants and chemosynthetic bacteria) animals have only limited
synthesising abilitiks.
In LSE-01, you have read that cellular metabolism provides energy for various
processes in organisms, like locomotion., excretion, osmoregulation, synthesis of new
materials for growth and maintenance and reproduction. To provide energy for these
processes raw material or nutrients are required which are supplied by food. In
addition animals require amino acids, vitamins and minerals which they cannot
synthesise. The study of nutrition involves both the need for food to provide energy
and the need for specific food components.
The process by which animajs acquire and ingest their food is referred to as feeding.
Diverse types of feeding mechanisms have been evolved by different groups of
animals. Virtually all foofwhether of plant or of animal origin has to be broken down
to simple compounds by the process of digestion. Digestion and absorption of food
constitute the essential link between nutrition and metabolism. In this unit we shall
first discuss the nature and components of food and the specialised feeding
mechanisms. There exists a relati~nshiphetween the nature of ingested food and type
of feeding mechanism used in acquiring the food. Then we will consider the digestion
and absorption of nutrients. Towards the end of the unit we shall be discussing the
energy metabolism in animals.
i
Objectives
After studying this unit you should be able to :
distinguish between essential and non-essential nutrients and explain why animals
exhibit differences in their essential food requirements
describe the various feeding strategies evolved by the animals in relation to the
available food
 distinguish between intracellular and extracellular digestion of proteins,
carbohydrates and fats and explain the role of gastrointestinal hormones
summarise the process of absorption of food from the alimentary canal
explain energy metabolism in animals relating it to oxygen consumption.

1.2 NUTRITION
As we have said earlier all animals are heterotrophs and require food from the
environment. What is this food made up of? If the food of a number of different
animals is broken down we find that it consists of proteins, carbohydrates, fats, water,
minerals and vitamins.
I
All animals require the above-mentioned broad categories of nutrients although in
different amounts. Some of these nutrients are ,used mainly as fuel (carbohydrates
and fats), while others'are required principally as structural and funational
components (proteins, minerals and vitamins). However, proteins, carbohydrates
and fats can all serve as fuel for the body's energy needs, but no animal can survive
on fuels alone. Therefore, a balanced diet is needed to meet all the requirements of
the body for energy, growth, maintenance, reproduction and physiological
regulation. Now.let us discuss the importance of these different classes of food in
relation to animal nutrition.

The terms essential and non-
m n t i a l ami* acids are not very
significant because the
non-essential amino acids are j b t
as important for the body. May hi
so important that the body cannot
leave them to be supplied
externally and so has mechanisms
to synthesise them.
1.2.1 Proteins
Proteins are continually synthesised in the cells as they are the principal component
required for growth. Proteins are composed of amino acids which are derived largely
from the diet and partly from the breakdown of protein available in. the body.
Generally all proteins are made from about 20 different amino acids in various
combinations. However, it is not necessary to supply.al1 the 20 amino acids. Some
can be formed in the body, using other amino acids but others have to be supplied
through diet because they are not formed in the body. The amino acids that are ,
synthesised in the body are called nonsssential amino acids while those that have to
be supplied through diet are known as essential amino acids.
The requirement of 'essential amino acids differs from organism to arganism. Some
bacteria require only one amino acid in sufficient quantities in the growth medium to
be able to synthesise the rest. In contrast mammals certainly cannot fulfil their protein
requirements by only one amino acid.
How can one determine wbich amino acid is essential and which is non-essential? The
nutritional requirements are determined by deletion experiments i.e. by removing a
single nutrient from the diet and then observing the growth and health of the animal.
By this method it was found out that 10 amino acids are essential for the growth and
well-being of rats (see Table 1.1).

Table 1.1 :Amino acids clssdCied according to dietary needs d hum- and rats

Essential Non-essentid
Rats Humans Rats Humans

Lysine Phenylalanine I Glycine Glycine

Tryptophan Lysine I Alanine Alanine
Histidine Isoleucine I Serine Serine
Phenylalanine Leucine I Cysteine Ty rosine
hucine Valine I Tyrosine Aspartate
Isbleucine Methionine -1 Aspartate Glutamate
Theronine Cystine Glutamate Proline
Methionine Tryptophan Proline Hydroxyproline ' ,,
Valine Theronine Hydroxyproline Citrulline .
Arginine Citrulline Histidine
Arginine
Absence of anjlone of these except arginine produces nutritional deficiency and even
death. Rats are able to synthesise arglnine but at such a slow rate that it does not
meet the demands of normal growth. To what extent the animal requires a particular
amino acid in diet depends on the synthetic ability of the body cells. Organisms with. '
marked synthetic ability, for example, bacteria (mentioned earlier) require a few
essential amino acids. Organisms like mammals, that require many essential amino
acids have a marked synthetic disability.

1.2.2 Carbohydrates
Fifty five to seventy per cent of the required energy in animals is derived from
carbohydrates. However, fats and proteins can also be broken down and used for
supplyin4 energy. In most animals this happens only when the dietary intake of
carbohydrates is low. In contrast, Drosophila uses only carbohydrates as a source of
energy for its flight muscles and when the supply is exhausted the insect cannot fly
even though it uses stored fat for other metabolic processes. Whereas, locusts are
known to use only lipids for their long migratory flights.
Most animals, however, use a variety of hexose sugars like glucose, fructose,
mannose, and galactose as interchangeable sources of energy. In this way no
particular carbohydrate is really considered essential in a way similar to amino acids.
But even if no carbohydrate is considered essential, growth of certain animals will be
better on one type of sugar than on another. This can be explained better by the
results of the following experiment. Young locusts showed that when dietary sugar
was maltose growth was maximum or optimum and growth was minimum when no
carbohydrate was given. Other sugars supported sub-optimal growth. What could be
the reason for this difference? One of the main causes is the difference in the rate of
movement of sugars across the gut wall into the blood. From the above experiment
we can conclude that certain insects have a preference for a certain carbohydrate
which can be called an essential or preferred nutrient. In the above experiment with
locusts, maltose was the preferred nutrient.

1.2.3 Lipids
All animal tissues contain lipids or fats as essential components of cell membrane. It
is also stored in certain tissues. Lipids are body's chief source of energy and are
essential for diverse functions such as insulation, padding, synthesis of steroid
hormones and carriers of fat soluble vitamins.
Many animals can live on little or no dietary fat because it can be formed from
proteins as well as carbohydrates. But the synthetic ability of many animals is limitec
in respect to certain unsaturated fatty acids and cholesterol. For instance, vertebrates
can synthesise cholesterol readily: In humans cholesterol is considered harmful in diet
because it is a major factor in the development of atherosclerosis or hardening of
arteries. On the other hand, insects cannot synthesise cholesterol from their
precursors. Therefore, it must be supplied in their diet. Studies on rats show that
three fatty acids - linoleic, Liolenic and archidonic acids are not synthesised.
Therefore, they are considered essential fatty acids.
Many insects, birds and some mammals also reveal such a dietary requirement of
fatty acids. It seems that animals in general have a better synthetic ability for lipids
than for amino acids.

1.2.4 Vitamins
Animals cannot sustain a healthy life if they are fed on a diet having only
carbohydrates, fats and proteins. They also require vitamins in small quantities in the The physiological role of vitamin K
range of milligrams o r micrograms. These function as coenzymes in metabolic was discovered in birds fed a
reactions. You might find it useful t o reread Unit 21 in FST-01 for the list of vitamins, cholesterol free diet for the
putpose of studying cholesterol
their main functions and their dietary sources. Table 1.2 gives a more detailed list of synthesis. The birds developed
vitamins important in animal and human nutrition along with their diverse functions. severe bleedings which were
The synthetic ability for vitamins also varies among different animal species and those traced to a vitamin deficiency.
essential vitamins that the animal cannot synthesise must come from its dietary Dnly after this it was discovered
sources. For instance, most animals can synthesise ascorbic acid but humans cannot. that vitamin K is necessary for
mammals.
We also depend on intestinal bacteria to synthesise vitamin K and BI2.Fat sol 'rle
vitamins like A , D, E and K can be stored in the fat deposits of the body but vit -ts
 L Table 1.2 :The Vitamine and their chPRfterWcs
~ipM-so~ubIeviernlns:
A (CZoH,O) anti- Plants form (carotene,
xerophthalmic C&%) in green leaves,
carrots, etc; is changed in liver
to animal form (C&iWO),
present in fish-liver oil
(shark); both forms in
egg yolk, butter, milk
D (C,H,O), Fish-liver oils, especially
antirachitic tuna, less in cod; beef
fat; also exposure of skin
to ultraviolet radiation
E or tocopherol Green leaves, wheat-
(C2UH~f12), germ oil and other vege-
antlstenl~ty table fats, meat, milk
(C31%b02)r Green leaves, also certain
antihemorrhagic bacteria, such as those of
intestihal flora
Water-soluble vitamins:
B complex Yeast, germ of cereals
Thiamine (B,) (especially wheat,
(CIZHI~ON~S). peanuts, other legumi-
antineuritic nous seed), roots, egg
yolk, liver, lean meat
Riboflavin (B2) Green leaves, milk,
(CI~HZOOC.N~) eggs, liver, yeast
Nicotinic acid, or Green leaves, wheat germ, egg
niacin (C6HS02N), yolk, meat, liver, yeast
antipellagric
Eolic acid Green leaves, liver, soyabeans,
(C19H1906N7) yeast, egg yolk
Pyridoxine (B6) Yeast, cereal grains, meat,
(C,H,zOzN) eggs, milk, liver
Pentothenic acid Yeast, cane molasses, peanuts,
(C9H1703N) egg yolks, milk, liver
Biotin (vitamin H) Yeast, cereal grains, cane
(CIOHI~Q~N~S) molasses, egg yolk, livkr,
vegetables, fresh fruits
Cyanocobalamin (BIZ) Liver, fish, meat, milk, egg
- .(GHPONI~OI~P~)
.
yolk, uysters, bacteria and fer-
mentations of Srreptomyces;
synthesised only by bacteria
C, or ascorbic acid Citrus fruits, tomatoes, vege-
(cdi806) tables;.alsoproduced by animals
(except primates and
guinea pigs)
- -
.L. Usinger, R.C. Stebbins, and J.W.Nybakken, General Zoology, 6th ed., Mc:Graw-Hill, ~ e York,
Maintains integrity of epithelial
tissues, especially mucous
membrane; needed as part of
visual purple in retina
of eye
Regulates metabolism of
calcium and posphorus;
promotes absorption of
calcium*inintestine;
needed for normal growth
& mineralisation of bones
Antioxidative; maintains
integrity of membranes
Essential production of
prothrombin in liver;
necessary for blood
clotting
Needed for carbohydrate
metabolism; thiamine
pyrophosphate an essential
coenzyme in pyruvate
metabolism (stimulates
root growth in plants) ' growth; polyneuritis (nerve
Essential for giowth;
forms prosthetic group of
FAD enzymes concerned
with intermediate meta-
bolism of food and elec-
tron-transport system
Forms active group of nicotina-
mide adenine dinucleotide, which
functions in dehydrogenation
reactions
Essential for growth and forma-
tion of blood cells; coenzyme in-
volved in transfer of single-
carbon units in metabolism
Present in tissues as pyridoxal
phosphate which serve as
coenzyme in transamination and
decarboxylationof amino acids
Forms coenzyme A, which cata-
lyzes transfer of various carbo-
xylated groups and functions in
carbohydrate and lipid metabolism
Essential for growth; functions
in CO, fixation and fatty acid
oxidation and synthesis
Formation of blood cells, growth;
coenzyme involved in transfer
of methyl groups and in nucleic
acid metabolism
Maintains integrity of capillary
walls; involved in formation of
"intercellular cement"
-
7 .
R ~ o t ~ o q a
la hunmb, except M aotcd
Xerophthalmia (dry cornea, no tear
secretion), phrynoderma (toad skin)
night blindness, growth retardation,
nutritional croup (hoarseness)
in birds
Rickets in young (tqnes soft,
yielding, often deformed);
osteomalacia (soft bones), ,
especially in women of Asia
Sterility in male fowls and
rats, degeneration of testes
with failure of sperma-
togenesis, embryonic growth
disturbances, suckling para-
lysis and muscular dystrophy in
young animals
Blood fails to clot
On diet high in polished rice,
beriberi (nerve inflammation);
loss of appetite, with loss of
tone and reduced motility in
digestive tract; cessation of
inflammation)in birds ,
Cheilosis (inflammationand
cracking at corners of mouth),
digestive disturbances.
"yellow liver" of dogs, curled-
toe paralysis of chicks,
cataract
Pellagra in humans and monkeys,
swine pellagra in pigs, blacktongue
in dogs, perosis in birds
Anaeniia, haemorrhage from kidneys,
and s ~ r u (defective
e intestinal abson,
tion)'in humans; nutritional cytopen$
(reduction in cellular.elementsof
blood) in nionkeys; slow growth and
anaemia in chicks and rats
Anaemia in dogs and pigs; dermatitis
in rats; paralysis (and death) in pig,
rats and chicks; growth
retardation
Dermatitis in chicks rnd rats, graying
of fur in black rats, "goosestepping"
and nerve degeneration in pigs
Dermatitis with thickening of skin
in rats and chicks, perosis
in birds
Pernicious anaemia, slow growth
in young animals; wasting disease
in ruminants . .
Scurvy (bleeding in mucous
membranes, under skin, pnd into
joints) in humans and gulnea pigs
w 1979
 Nutrition. Feeding, Digestion
that are water soluble like B or C need to be supplied continually as they are lost
through urine.

1.2.5 Minerals
I
and Trace Elements
Oxygen,l,carbon, hydrogen and nitrogen are the most common elements that make
up 96%'of the total weight of a mammal. The next most abundant elements are
calcium, phosphorus, potassium, sulphur, sodium, chlorine and magnesium. These
make up nearly the remaining 4%.
Fifteen additional elements are required but their total combined amount in
mammalian body is less than 0.01% of the body weight. Of the known 90 naturally
occurring elements how many are essential for life? This is nat known for many
animals. In humans 26 elements are known to be necessary.
Table 1.3 : Approximate composition of human tissue

Element Per cent body weight

Oxygen 65.0
Carbon 18.0
Hydrogen 10,0
Nitrogen
Calcium
Phosphorus
Pot;~ssiuni
Sulphur
Sodium
Chlorinc ~.
Magnesium 0.05

Table 1.3 gives the approximate coinposition of human tissue. We all know that
carbon, oxygen, hydrogen are present in water and other organic building blocks of
the body which also contain nitrogen, sulphur and phosphorus. Calcium is an
important constituent of the skeletal structuves of animals and its role in physiological
processes such as muscle contraction will be studied later in Unit 6: If the level of
calcium concentration falls below half its normal value it leads to severe or fatal
tetanic cramps. Table j.4 gives the role of these important minerals.
able 1.4 : Physiologic~lr o l e 4 important minerals
Elements Deficiency Disease

Sodii~~n Main extracellular positive ion: Unknown on normal diet. Table salt, salt
(Na) Regulates plasma volume. acid- Secondary in illness or added to
hase halance: nerve and injury prepared food
muwlc Function
Potassium Major intracellular positive ion: Secondary t o illness.
(K) nervc and muscle function: injury or diuretic
acid hase balance therapy; paralysis,
mental confusion
muscular weakness
Calcium Component of bones, teeth; Children-rickets Dairy products,
(Ca)
, regulation of nerve, muscle Adults -ostmalacia beans, leafy
function; blood clotting vegetables
Phosphorus 9onG formation, part of DNA. Children-rick$ Phosphate f w d
(P) RNA, ATP, etc.; energy Adults -osiomalacia additives
metabolism
'Magnesium Bone and teeth: carbohydrate Secondary to mal- ' Leafy green
(Mg) metabolism ahsorption o r diarrhoea, vegetables
;~icoholism
Chlorine Major extracellular negative In infants fed on salt Table salt
(a) ion; osmotic and acidibase free formula; secondary
balance; stomach acid to vomiting. diuretic
therapy, renal disease.

Mineral requirements are met by a varied intake of adequate amounts of w h o l ~grain cereals, legumes,
leafy green vegetables, meat and dairy products.
 The additional 15 elements that make up less than 0.01% of the body weight occur
in such small amounts that they are called trace elements. Table 1.5 lists some of these
trace elements thatare considered essential. Although present inminute amounts the
essential trace elements are just as necessary as the essential amino acids. For
instance, cobalt is needed in the specific form of B12and its deficiency leads to severe
anaemia. In ruminants'vitamin B12is formed in the rumen'by bacteria provided a
sufficient amount of cobalt is present in the diet.
Table 1.5 : Physiological role of essential trace elements

Element Physiological Role Deficiency Disease source'

Iron Component of haem group in Anaemia Iron cookware '

(Fe) haemoglobins, cytochromes

Copper Needed to make haemoglobin, Anaemia; secpndary to
(cu) bone, part of cytochrome malnutrition, Menke's
syndrome
Iodine Component of thyroid Children: cretinism lodised salt,
(1) hormone Adults: goitre, hyper- seafood
thyroidism, myxedema
Manganese Needed in urea formation, Unknown in humans
(Mn) protein metabolism, glycolysis,
citric acid cycle
Cobalt Constituent of vitamin B I Z , B I Zdeficiency Foods of
(Co) RBC formation animal origin
Zinc Essential constituent of many Hypogonadism, growth
(zn) enzymes, needed for normal failure, impaired wound
senses of smell and taste healing, decreased taste
and smell
Molybdenum Constituent of some enzymes Secondary to parenteral
(Mo) nutrition
, Flourine Hardness of Dental caries Drinking
(F) teeth water
Selenium Needed in fat Marginal deficiency where
The first conclusive evidence that salt content is low,
(se) metabolism
cobalt is an essential trace secondary to parenteral
element came from Australia ' nutrition
where a serious disease of cattle
and sheep developed. Addition of Chromium Needed in glucose Impaired glucose
small amounts of cobalt, (Cr) metabolism tolerance
prevented this disease. Each Trace element requirements are met by a varied intake of whole grain cereal, legumes, leafy green
sheep is made to swallow a vegetables, meat and dairy products
ceramic coated cobal ball that
remains in the mmer and slowly Some of the trace elements are essential for complex organisms while others may not
releases the cobalt o\ifr several be so essential. Flourine, for example, is an essential element for normal growth of
years. rats but is not strictly essential for humans, though we know that it has a well-defined
role in prevention and treatment of dental caries. It is difficult to find out the role of
each and every trace element and further research will be needed to increase our
knowledge.
1.2.6 Water
Water is the most important constitdent of all living tissue. It forms up to 95% of the
fresh weight of some animals. We all know that water is lost through sweat, excretion
and evaporation from the respiratory surfaces. It must therefore, be replenished by
drinking, through food and in small quantities by metabolic processes of the body
like synthesis and oxidation of fats, proteins and carbohydrates.
SAQ 1
a) Match the words in column A with descri~tionsin column B.
Column A
I
i) Synthetic ability a) Amino acids that are required for growth
but have to be supplied through diet.
ii) Synthetic disability b) Most bacteria can produce all the required
amino acids from only one nutrient present
in their growth medium.
iii) Essential nutrient c) Humans can synthesise cholesterol but
insects cannot, from non-sterol precursors.
b) Why don't domestic cats and dogs need fruit in their diet while humans do? ,

1.3 FEEDING MECHANISMS

All animals have evolved successful methods for extracting their required nutrition
from the environment. Thus we find a diversity of feeding mechanisms or strategies
according to the nature of food that an animal can obtain. Table 1.6 lists the major
feeding methods in-animal groups based on the type of food available. It would not
be possible to discuss each food gathering device in detail but in a brief discussion
we shall consider the basic principles on which the different feeding mechanism
operate. From Table 1.6 you will note that taxonomically different animal groups
living in the same habitat obtain food in a similar manner. For example, many marine
animals (annelids, molluscs, crustaceans) may be filter feeders though the organs
concerned with the process of filtration may not be anatomically similar.

Table 1.6 : Feeding methods classified according to type of food

Type of food Method of feeding Anlmab using the method

Small particles Digestive vacuoles Amoeba, Radiolarians

Use of cilia Ciliates, Sponges,
Bivalves, Tadpoles
Mucous traps Gastropods, Tunicates
Tentacles Sea cucumbers
Filter feeding Small Crustaceans, Herrings, Baleen
Whales, Flamingoes, Petrels

Large food masses Ingestion of inactive Detritus feeders, Earthworm

masses
Scraping, chewing, boring Sea urchins, Snails, Insects,
Vertebrates
Capture and swallowing Coelentrates, Fishes, Snakes, Bats,
of prey Birds
Fluid or soft tissue Sucking plant sap, nector Aphids, Bees, Humming-birds
Ingestion of blood Leaches, Ticks, Insects
Vampire bats
Sucking of milk or Young Mammals, Young Birds
Similar secretions
External digestion Spiders
Uptake from body surface Parasites, Tapeworm
Dissolved organic Uptake from dilute Aquatic invertebrates
solution solution
Symbiotic supply of Intracellular symbiotic Paramecium, Sponges, Flatworms,
nutrients algae Corals, Hydras, Clams.

1.3.1 Feeding on Small Particles

r
Microscopic algae and bacteria can be taken in directly into the cell by the digestive
vacuoles. But one of the most successful methods of feeding on sma 1 particulate
matter is filter feeding or suspension feeding. Particulate matter incl des detritus,
living and dead plankton. Most filter feeders use ciliated surfaces to p oduce currents
- that draw drifting food particles into the mouth. The animal extracts the suspended
food particles by means of structures that act as filters often aided by secretion of
, mucous which traps the food particles. In sponges, the flagella of the choanocytes,
I
Anlaul phrddogy -I the cells that line the body cavity, create internal water currents. The body wall has
numerous pores called ostia and the water is drawn in across the flagellated chamber
...
(Fig. 1.1) into the body cavity. Food particles are trapped by the flagella and directed
to the surface of choanocyte, which ingest the particles by phagocytosis.

v Water currents

ngocoel (body cavity)

Choanocyte mucus

move water)
. Amebocyte Amebocyte receiving
food vacuole

Fig. 1.1 : Sponges obtain their fdod by filtering seawater. Food particles pass down the collar cells or
choanocytes.andenter them through pbagocytosis.

Moreelaborate methods of Qlter feeding are seen in tube dwelling'polychaetes which

use teiltacles to entangle the food particles. Figure 1.2 shows some of the filter feeders
and their feeding mechanisms. Filter feeders include both sessile and free 'living
forms. The sessile forms generally accept what they get, though, some can selectively
choose their food according to size. For instance in Sabella a tube living polychaete,
while large sand particles are-rejected small food particles enter the food groove. Free
rakers swimming forms are selective feeders. Examples are, many of the microcrustaceans,
fishes such as herring, menhaden and bbking sharks, certain birds such as flamingo,
pelican and the largest of all animals the baleen whale.

Among the fishes, herring have gill rakers (Fig. 1.2b) that function as a sieve to catch
plankton. The basking sharks feed exclusively on plankton and can filter up to 200
tons of water in 1 hour.

The flamingo also a plankton eater uses its beak to strain small organisms from the
water (Fig. 1 . 2 ~ )However,
. the baleen whale is specialised for filter feeding. Its
filtering apparatus cons,jsts of a series of horny plates attached to the upper jaw
(Fig. 1.2d). As the whale swims, water flows between the plates retaining the
planktan.

+'- 1.3.2 Feeding on Food Masses

(4
1 digestive tract.
Fig. 1.2 : Some filter feeders and
their feeding mechanisms
Unlike filter feeders which feed in water, animals that obtain and eat solid food are
not restricted to the aquatic environment, and show a great variety of adaptations
related to their feeding habits. Animals like earthworms ingest the medium in which
they live and digest the organic material ih it as the material passes through the
I
Many animals use methods for chewing and sctaping to obt'ainlfood. Their food is
often of ~ l a norigin.
t Numerous insects and other invertebrates as well as herbivorous
vertebraies use Gese methods of feeding. A rahp-like structurs known i s radula is
used by gastropods to scrape algae from roqks of;t9 rasp through vegeta.tiio (Fig. 1.3).
 R a d a retractor Radula protractor
-
(b') (4
(a)
Flg.l.3:a) S s s i t E . l & ~ b a d + ~ p W s b o w h g t ~ d u L P w W ~ ~ t o ~ p ~
vegetatba.
b) RohctkmdradulP
c) Retrrtlon d red&
(a)
However, most carnivorous animals simply seize and swallow their prey whole. Jaw
Relatively few invertebrates feed in this way but an interesting example is the
carnivorous polychaete Nereis which has a muscular pharynx armed with chitinous tentacles
jaws that can be everted to capture prey (Fig. 1.4). Once a prey is caught the pharynx
is retracted and the prey is swallowed. The teeth of lower vertebrates (fishes,
amphibians, reptiles) are mainly used to grip the prey and prevent its escape till it is Rmtomiu
swallowed. Snakes are familiar examples that are well adapted to this kind of feeding
\
\

Fig. 1.5 : Snakes cannot tear or chew food. They swallow prey whole. The mouth is extremely flexible
because of thearrangement of bones in the head and jaw. Lower jaw is loosely attached to
-
quadrate bone and bones of the palate are movable. The help to draw is prey into the gaping
i mouth.

Fig. 1.4 :Narlr v h , an e m ~ t

pdy-,
a) antdw end with
everted Jaw to
apturn Prcr
b ) e x t m u l ~ ,

15

Spiders provide an interesting
example of fluid feeding. Their
preybare usually larger in size and
covebed by a hard chitinous
covering. Spiders, therefore,
pierce the covering by their
hollow jaws and pump digestive
juices into the prey's body. These
liquify the tissues and then the
'kpider'sucks the prey empty.
True mastication i.e. e&of food is found only in mammals. Their teeth are
adapted for this specific function. Mammals have basically four types of teeth
(Fig. 1.7) each adapted for different type of feeding. Incisors are adapted for biting
and cutting and stripping; canines for seizing and piercing; premolars for crushing;
and the molars for crushing and grinding. The number and size of these teeth varies
according to the type of foad eaten.,
as M premolars A ~c~~
Fig. 1.7;,Mammaliandentitlon - teeth of (a) generalised mammal, (b) squirrel, (c) AMcan Uon, (d) OX
1.3.3 Feeding on Liquids
Animals feeding on liquids are generally highly specialised for their feeding habits.
Certain protozoa, endoparasites and aquatic invertebrates take up nutrient molecules
through their integument from the- medium in which they live. For example,
endoparasites, which include parasitic protozoa, tapeworms, flukes, certain molluscs
and crustaceans are surrounded by host tissue or alimentary canal fluids which are
highly nutritive. These parasites lack a digestive system of their own.
All of us are familiar with insects that have well-developed piercing and sucking
organs. Mosquitoes, bedbugs and lice and leaches among annelids are some
examples. They use anticoagulant to prevent blood from clotting as it leaves the blood
vessels ruptured by their piercing or rasping jaws.
I
SAQ 2
a) You.must have observed a squirrel, a cow and a dog feeding. What kind of
differences would you expect to find in their dentition?
b) Match the type of feeding apparatus in column A with the kind of food in column
B.
Column A Column B
1)- Radula a) Blaod, plant sap
2) Cilia b) Detritus in mud
3) Mucous Sheets c) Large chunks of food
4) Sucking mouth parts d) Algae on rocks
5) Teeth e) Suspended particles
 ,1.4 . DIGESTION
i In the earlier sections we considered the nutritional requirements and the various
ways used by heterotrophic organisms to obtain nutrition. Whether food is used to
give energy or to build the body, the large molecules of food have to be broken down
into simpler constituents before they can be used by the body. The process by which
the food is broken down into simpler molecules is known as digestion. This breakdown
is achieved with the aid of enzymes and can take place inside the cell -intracellular
digestion or outside the cell -extracellular digestion often in a specialised digestive
tract.
Let us first consider intracellular digestion and see how it is different from
extracellular digestion.

We all know that unicellular organisms do not have a separate alimentary canal
system. All the functions of life are carried out inside a single cell. Food is taken in
directly into a cell by phagocytosis/endocytosisand then with the help of enzymes
digested in a food vacuGle. Fig. 1.8 shows the process of endocytosis in Amoeba.

Nucleus
molecules
<v'"
.I
Lysosome 3:
.'.-..",..*"";
..:. .
.
...._
4

Fig. 1.8 :Digestion in amoeba .

Similar intracellular digestion occurs in sponges, some coelentrates, ctenophores and
turbellarians. Although the process is called intracellular digestion, the food material
is actually separated from the rest of the cellular material by a membrane which it
can cross after digestion. In organisms such as cnidarians and platyhelmintbs, a gut
or enteron is present and here along with extracellular digestion where enzymes are
secreted into the cavity, intracellular digestion also takes place within the cells that
line the ca2ity. However, in annelids and molluscs more extracellular than
intracellular digestion takes place. Digestion is entirely extracellular in nematodes,
,"
insects, echinoderms and vertebrates.

1 1.4.2 Digestive l'rict

Extracellular digestion takes place in a tubular cavity that extends tiroughout the
length of the organism. All animals after flatworms have-a tubular alimentary
organisation open at both ends. The. development of extracellular digestion freed
many animals from feeding continuously on small particles. They could now quickly
1 ingest a few large chunks of food. The overall tubular organisation of the digestive

I tract also allows the food to travel in one direction passing through regions of
digestive specialisation (Fig. 1.9).

1 In general the digestive system of metazoans is divided into 4 major functional regions
of:
reception
conduction and storage
: I* digestion and absorption
ce?duction and. formation of faeces.
I *

I
I : Storage
\ ,' (Some spccies)
The region for reception is associated with devices for mastication or chewing of food
(like teeth); for paralysing the struggling prey (toxic enzymes from saliva); initiating
digestion and lubricating the food with mucous.
The oesophagus of chordates and some invertebrates serves to conduct the bolus
(mass of chewed food) by peristaltic movement from buccal cavity. In some animals
this fegion has a crop for storage. The crop ip birds is also used to ferment mildly or
digest food. This is later regurgitated by pare'nt birds for their nestlings. The storage
region allows the animals to store food and use it when it is not easily available. For
example, leaches take in infrequent large meals of blood and digest it slowly over a
month. The herbivore animal spends hours masticating the food it takes in hurridly
and stores it in its stomachsfor further use.
In the third region or digestive region the enzymes reduce the food to a forin that

II (Add)

Digestion
can be absorbecfby the body of the organi:m. As the food is digested, the absorbable
food is passed to the blood stream and the unabsorbed material is stored briefly in
the final section of the alimentary canal where further removal of excess water and
consolidation of undigested material into faeces takes place, before it is expelled out
of the body. In vertebrates this function is carried out in the large.intestine.
In higher vertebrates, each area of the gut is specialised for a certain activity,
digestive enzymes are produced in glands as well as in the wall of the gut. Absorption
,

1
occurs in the intestine predominantly.

Jl
(Alkaline)

1.4.3 Digestive Enzymes

Absorption and
1 Assimilation
J
Storage of waste
+
Defecation
FLa.L.9: Ccncrdbed dlgwtlvt
trod. Om way pslrs~ge
of food allows sequtntlal
stag- In dlgcstlon.
. Duhcd Ilna represent
crop Lo m m . . n l d s .
Now let us consider the general principles of digestion that are applicable t6all
animals. We will start with the digestive enzymes that breakdown the large food
molecules into smaller soluble component units. This breakdown involves the uptake
of water and is called hydrolysis. Before reading the following sub-sections, you
would find it useful to read Units 9 and 10 of LSE-01 to recapitulate the nature and
properties of enzymes in general. However, digestive enzymes differ in the following
ways:
a) Digestive enzymes are not as narrowly specific as other enzymes rather they show
group specificity. For example, enzymes that digest carbohydrates can digest
polysaccharides of both animal and plant origin,
b) Even though enzymes performing similar functions in different animals are given
same names, they are not identical chemically. For example, trypsin (an enzyme,
that hydrolyses proteins) in humans is not identical to that fouqd in fish.
Temperature and pH for optimum activity is also different. For example, trypsin
from vertebrate pancreas acts best in the pH range of 7-9 but in silkworm the
pH range is 6.2-9.

c) Digestive enzymes from pancreas parti'iularly those that digest proteins are
secreted in an inactive form.

Thethree major classes of digestive enzymes are:

i) Proteases.that hydrolyse peptide bonds in proteins,
ii) Carbohydrases that hydrolyse glycosidic bonds in carbohydrate,

iii) Lipases that hydrolyse ester bonds in fats

. . Protein Digestion
Enzymes that digest proteins are divided into two groups endopeptidases and
exopeptidases according to site of their action in the protein molecule. Endopeptidases
confine their attack to the interior of the protein molecule so that the large peptide
chain is broken into smaller fragments/. This provides many sites for action of
exopeptidases that attack only peptide bonds at the end of a peptide chain releasing
amino acids, dipeptides and tripeptides. There are several types of endopeptidased
and exopeptidaseg. They are listed-in Table 1.7.
 Active form Preferred Peptide
Inactive form Link Attacked
a -

Endopepticlaw
HC1 Link to amino group of aromatic
Pepsinogen , pepsin' amino acid (tyrosine and
pepsin
phenylalanine)
entemkinase ' Link to carbxyl end of arginine
Trypsinogen trypsin
trypsin ' or lpine
Link to czubxyl group of aromatic
GYrn-in~gen w *chymotrypsin amino acid (tryptophan, tyrosine,
' phenylalanine) and also bonds adjacent
to methionine and leucine when'they
are present

Link to terminal anjno acid with

h e amino group
Carboxpeptidase (H+)
(trypsin) Link to terming amino acid with
free carboxyl group
Bonds between pairs of
amino acids

From the table youcan see that these exopeptidases and endopeptidases attack specific
peptide bonds depending on the chemical group near them. Theeinactive forms need
activators and autocatalysts to convert them into active forms. For example pepsinogen is
secreted by the vertebrate stomach. The stomach also secretes HCL which makes the
medium acidic (pH 2). This activates pepsinogen into pepsin. Pepsin specifically hydrolyses
peptide bond between a dicarboxylic and an aromatic amino acid (Fig. 1.10a). In this way
short fragments of polypeptide chains are formed. Invertebrates seem to lack pepsin arid
their main endopeptidase is more like trypsin. Look at Fig. 1.10a again, chymotrypsin also
attacks a peptide bond involving' aromatic amino acid but on the carboxyl terminal end of
the molecule while pepsin attacks on themnino terminal end.

FHa .pepsin Chymotrypsin

-
CH,
I
HN-CH-CO

Clutamic acid
b'
HN-CH-CO
I
- v
NH-CHz-CO
(CHz)3
NH-CH-CC
I
-
6
,
NH-

Clycine Lysine

Tyrosine

H3CVCH 3

CHa
CH
I
Am~nopeptldase Carboxypeptidase
4$
.. /

N Termlnal
NH,-CH-CO
I
;
H N -CHt-CO
*
1"'
HN-CH-COOH
1
Leucine Clycine
Tyroslne
I (b) .

I
I
mg. 1.10 :Rotdn d i g d o g enzymm:
a) a ~ s p c d l l e p e p t i d e b o m b i o a p r o t e ~ ~ t
 T w i n is secreted by the pancreas in an inactive form trypsinogen. It is activated by
enterokinase secreted by the glands in the intestinal wall. As trypsin is formed, it activates
more trypsinogen to be converted into trypsin. This is autocatalytid activation. Trypsin acts
in an alkaline medium betwcen pH 7-9. It breaks a peptide bond next to basic amino acid
like arginine or lysine.
The polypeptide fragments are further digested by the exopeptidases. Carboxypeptidase
require the presence,of zinc ion and trypsin.-Otherexopeptidases are secreted in active form
but need metal ions as cofactors to increase their activity.
Fig 1.10(b) illustrates the action of aminopeptidase which removes terminal amino
acids having free amino groups and carboxypeptidase which removes terminal amino
acids possessing a free carboxyl group. In this w'ay these two enzymes remove
peptides from each end until a dipeptide fragment consisting of only two amino acids
remains. Bonds between these pairs of amino acids are split by dipeptidases releasing
free amino acid.
The amino acids now, may be absorbed through the cells of the indstinal wall.

Carbohydrate Digestion
Simple sugars like glucose and fructose can be absorbed and metabolised directly but
disaccharides such as sucrose or lactose and polysaccharides such as starch and
glycogen have to be broken down to monosaccharides before they can be used in
metabolic pathways. Carbohydrases, that digest carbohydrates can .be grouped into
two categories:
i) Polysaccharases that split polysaccharides into disaccharides or trisaccharides.
ii) Glycosidases that break up the disaccharides or trisaccharides to monosaccharides.
The digestion of carbohydrates, like proteins also occurs in steps. These along with
the enzymes responsible for digestion are given in Table 1.8.

,
Table 1.8 : Digestion of carbohydrates . . .. . ; ~

Poly-harides P"'ys;lcchar- ~ i ~ ~ Glyccwidaws

~ ~ h r~Monosaccharide
i d ~ ~ .
(c6111005)x (C~~H220~~) (CnHZnOn)

Glycogen Amylases Maltose

M ~ I ~ ~ S C , , .
(animals) . .
Amylases 'Maltase ' .
Starch Maliose * Glucose
(Plants)

Cellulose
(Plants & animals)
-Cellulases
- Cellobiose
Cellobioses
r Glucose

Trehalase
Trehalose r Glucose
(insects and
some plants)
Lactase
Lactose r Galactose
Glucose
Between 2-18 per cent of lnvcrtase
Sucrose r Fructose
caucasians loose the capacity to Glucose
produce lactase and between
95-1W per cent oriental and
native ~ f r i c a nraces loose the . Carbohydrate digestion in vertebrates and invertebrates is very similar. All the
ability td produce lactase as they enzymes shown in Table 1.8 are not required by all animals. The enzymes present
grow older. They can no longer are related to the food habits of the animal. However, amylase and maltase are of
digest milk which ferments in
their gut and produces diarrhoea universal occurrence. Amylase is secreted in the saliva of man and in larger amounts
and related problems. Interestingly by the pancreas. Enzyme production in some animals is also influenced by genetic
yoghurt and cheese do not create characteristics and enzyme induction. For example, production of maltase and
any problems as these contain less sucrase by the intestinal villi depends on the amount of ingested sugar. If a high
than 2 per cent lactose due to
.
maltose or sucrose diet is taken it induces the villi to produce more maltase and
action of bacteria.
sucrase within 2-5 days. Lactase production declines in humans as gut develops
after infamy. It ceases in some individuals so that they can no longer hydrolase this
 sugar. Now let us consider the digestion of cellulose, the most important structural
material of plants and a major component of the diet of herbivores. Very few animals
possess the enzyme.cellulases. Then how do animals that feed on plants breakdown
this carbohydrate? Cellulases enzymes are synthesised by many bacteria and Symbiotic flagaates from
protistans which live symbiotically in many herbivores and insects. Cellulose digestion termites are obligate anaerobic
is carried on by the help of these symbiotic microorganisms. The microorganisms live organisms. Because of this
sensitivity these flagellates can be
. . in the s t o h c h of the ruminants (i.e. cow, sheep, etc.) and breakdown the cellulose. removed from termite gut by
The breakdown pioducts are then utilised by the host. In some invertebrates like exposing termites to oxygen at 3.5
silver fish (Ctenolepisma lineata) true cellulases have been reported but the insect atm pressure. The protozoa are
cannot survive on an only cellulose diet. Some other invertebrates also have some selectively killed within half an
cellulases that partly digest cellulose but none show conclusive evidence of a complete hour and the termites survive.
breakdown of cellulose into glucose without the help of symbionts. Such treated termites do not
survive when fed on wood though
they stiU have bacteria in the gut.
This shows that anaerobic
Lipid Digestion . . .. . . p-rotozopsrather than bacteria, are
Digestion of fats is also similar inboth invertebrates and vertebrates. Lipases are the responsible for cellulqse digestion
in temiites.
- enzymes that hydrolyse fats. A single l\pise can catalyse many steps in the break down
'
of fat. The vertebratepancreas secrete an enzyme lipase but before it breaks down
fat, some detergent-like action is needed to emulsify the fat droplets. Bile salts from
the liver, lecithin and cholesterol form miscelles and do this job. They reduce the
surface tension at the fat-water interphase in a slightly alkaline medium and tiny
emulsific-ation droplets of fat are formed. Then the lipase begins to digest the
emulsified droplets. The resultant'fatt'y acids and monoglycerates are kept in solution
by help of bile salts again and are finally absorbed.
Glycerol is water soluble and easilyiabsorbed and metabolised. Fat like butter
is absorbed directly through the intestinal epithelium without hydrolysis.

1.4.4 ~aintenanceof Gut Lining

After studying the digestive enzymes you would'wonder why the gut linings are not
digested themselves. This is because animals have several mechanisms that protect
their gut lining from autodigestion. The mucous membranes of vertebrates secrete a
slightly'alkaline mucous that lubricates the food and protects the lining'cells from
' corrosive secretions. 1n addition, the lateral surfaces of exposedepithelial cells are
joined by tight junctions that prevent the secretions from penetrating between them.
Careful studies have also revealed that the entire lining of the gut is renewed every
third day in rats and every 2-6 days in humans. Similar mechanisms are present in
invertebrates also. In insects, the fore-gut and hind gut are lined by cuticle. This lining
is known as intimg. Only in the midgut, the epithelial cells are exposed, where most
of the digestion occurs. The midgut is lined by a delicate lining the peritrophic
membrane in some insects. This correspends to the mucous lining of vcrtebrates.

1.4.5 Coordination of Digestion

You have learnt that digestion is a process in which large food mplecules arc bipken
down step by step into their constituents. In primitive metazoans that are continuous
feeders, the enzyme producing cells secrete continuously. In higher animals more '
precise controls are needed to regulate the~eleaseof food from stomach to intestine.
and also the release of digestive enzymes at the proper time. The interplay of nervous
and hormonal control is beautifully illustrated when we study the &ordination of
digestive activity. t- .

In the mammalian mouth, control of salivary gland secretion is entirely nervous;

gastric secretions are under hormoaal.and neural control; and intestinal secretions
are slower and are primarily under hormonal control.
I Gastrointestinal secretion is largely under the control of gastrointestinal hormones
I secreted by endocririe glands of gastric and intestinal mucosa.

I
Gastrointestinal ~ o i m o n e s
I
The thiee Atlain mammalian gastrointestinal hormones are secretin, gastrin and
i chdecy&kinh (CCK). There are several other hormones, all peptides. The
physiology of only. thrce'major hormones is listed i i Table 1.9.
 .
I"- a.r . ,..uulururu -uu",-uuu .M--9 - ..- m
,".". I OY".YU..VY,

+++ .hormdne nuwe important than other two

Gastrin Seeretin

Secreted by Stomach Duodenum Duodenum

Stimulus for Peptides Acid (HCI) Amino acids
release fatty acids
Parasympathetic
Nerves
Effect on :
Gastric Motility
Gastric HCI
Secretion
Pancreatic secretion
bicarbonates
enzymes

Gastrin secretion is responsible for control of HCI volume; the presence of HCI in
turn inhibits further gastrin secretion.
Secretin is released under acidic conditions (low pH); digested fat or bile initiates
production of pancreatic juice low in enzymes but rich in salts important in
neutralising the acid chyme. CCK is secreted when partially digested proteins or HC1
(to a lesser extent) are present. It induces the flow of pancreatic juice rich in enzyme.
Fig. 1.11 summaries the action of GI hormones.

Gallbladder
I Stomach

Fig.l.11 : Action of several gastrointestinai hormones. Gastrin is secreted in response to intragastrk

proteins, stomach distention and stimulation by vagus nerve. Gastrin from the lower stomach
stimulates HCI seeretion and pepsin from secretory cells. Cholecystdrlnln (CCK) stimulates
pancreas to secrete digestive enzymes and bases to nwtrallse and digest chyme. It also Induces
contractlon of gall bladder to secrete bile salts. CCK is secreted in response to arrival of emlno
acids and fatty acids In deodenum from stomach.

These two hormones inhibit stomach motility. Arrival of fat from the stomach
initiates the release of CCK by intestinal mucosa, this ,causes gall bladder to release
bile which aids in fat digestion.

SAQ 3
a) What are the main advantages of having a digestive tract with a mouth and anus?

b) Choose the correct answer.

Digestion is brought -about by
i) acids, ii) enzymes, iii) alkaline solutions, iv) vitamins and minerals
 c) Fill in the blanks with appropiiate words.
In. the process of digestion proteins are converted into ........................
carbohydrates to .......................fats to .......................and ......................
d) Three hormones stirnulafe the release of digestive materials, .......................
stimulates release of gastric juices ....................... stimulates release of
bicarbonate ion. ......................stimulates release of bile and pancreatic
enzymes.

The monosaccharides, amino acids and other products of digestion must be passed
on to other tissues to be useful for the organism. The process by which the digested
material from the alimentary canal enters the blood stream is known as absorption.
In intracellular digestion the same cells are concerned with digestion and absorption
but in higher multicellular animals there are usually separate tissues and areas of gut
for enzyme production, digestion and absorption.
In this section we wi.1 mainly be concerned with absorption of amino acids, sugars
and fats released during extracellular digestion in vertebrates. In all vertebrates most
of the absorption is localised in the intestine. As you already know the wall of the
vertebrate intestine is folded and ridged to increase the absorptive surface. These
ridges or folds are covered by a velvet like pile of minute absorptive villi (Fig.1.12).
These are highly specialised absorptive organs with a core containing a network of
capillaries derived from blood vessels in the gut wall. Each villus also contains a
central lymph capillary known as 4acteal which begins blindly at the tip of the villus
and drains into the main lymph channels of the gut wall. Lipids pass mainly into the
lacteals while sugars and aminoacids are absorbed directly by the blood capillaries.
The villi and intestinal folds contain smooth muscles that contract to bring the villi
in contact with the food in the intestine; and also maintaining the circulation in
lacteals, lymphatics and small blood vessels.

Fig. 1.12 :Lining of mnmmpli.n smPU intestine

a) Villus covered wlth digestive epithelium which consists of absorptlvecell and occasional goblet
cells.
b) An absorptive ceU. The apical surface bears a brush border of microvilli.

Now let us examine the processes involved in actual absorption of the digested food.
You could recall from LSE-Ol Unit 7 the transport processes across membranes. We
can summarise them as follows:
Diffusion A material passes down its electrochemical gradient; at the end
(passive) of the process the concentration inside the cell is equal to
that outside.
23

Glu T, is one of the 5 gluwse
transport molecules known to
carry gljlwse across cell.
membranes. These transporters
are broadly similar in structure
and function and numbered in
order of their discovery. These
transporters change their
configuration, one shape binds
glucose on the extracellular side
and the other shape binds glucose
in the intracellular side. This
binding and transportation in a
flip-flop manner is very rapid.
24
Mediated Transpon
Facilitated A specific membrane component is invplved in the transfer of
the substrate along the concentration gradient but the
concentration inside the cell never exceeds that of outside.
Active A specific membrane component is involved in the transfer of a
substrate, which can be accumulated inside the cell against the
electrochemical .gradient.
All animal groups make some use of these three processes. The transport of sugars
and of amino acids takes place through transport molecules that depend on the action
of sodium pumps and can be blocked by metabolic inhibitors such as cynide.
Some sugars for example, fructose are carried down their concentration gradient by
facilitated transport. This process requires no energy other than the one provided by
the concentration gradient of the diffusing substance. Other sugars like glucose and
galactose as well as amino acids are absorbed through special transporter molecules
in the membrane of the cell and depend on the Na' gradient between the lumen of
the gut and cytoplasm of the epithelial cells.
Let us first consider the absorption of glucose from the'gut lumen. (Refer to LSE-01,
Units 7 and 8). The molecule involved in absorption of,glucose is known as
cotransporter because it couples the transport of a glucose molecule with that of a
sodium ion. The energy needed is provided by the movement of sodium ion along its
gradient. The cotransporter enables cells lining the lumen of intestine to absorb even
quite small traces of glucose from food even though the epithelial cells may already
have high concentrations of glucose inside them.
Once inside the cell, the sodium ion is pumped out by ATP energised active transport
and the glucose molecule is transferred to the blood stream through another
transporter molecule, Glu Tz, along its concentration gradient (Fig. 1.13). Glu Tz
transports glucose in propdrtion to the sugar concentration present in the blood. If .
more glucose is present in the blood, transport is slowed and if glucose content of
blood is low then transport is accelerated.
Fig. 1.13 :Suggested mechanism for absorption of glucose. Na' and glucose are transpofled together
through carrier molecule or cotransporter located in the membrane. Inside the cell, sodium moves
i
1
out by ATP pump and glucose is taken by a transporter molecule to the blood.
Experimental evidence shows that atleast 4 transport processes for amino acids
occur in the mammalian gut. Two for neutral amino acids, one for basic and one for ,
acidic amino acid. Another separate transport system exists for dipeptides and
tripeptides. Once inside the cell, these breakdown into constituent amino acids bp
intracellular peptidases. The sugar and amino acids reach the circulatory system from
where similar mechanisms use the sodium gradient to transport amino acids and
glucose to the various tissues of the body.
-
Absorption of lipids is quite different from the absorption of monosaccharides and
amino acids. Fig. 1.14 shows the process. m e free fatty acids, monoglycerides and
lysolecithi?~leave the miscelles and pass through the membrane of the microvilli to
enter the epithelial cell. The miscelles may also be transported intact into the
epithelipl cells. In ~ i t h e event,
r these products are used to resynthesise triglycerides
anc$Jphc$pholipidswithin the epithelial cells. Triglycerides, phospholipids and
cholesterol combine with protein inside the epithelial cells to form small particles
called chylomicrons that are collected by the lacteals in the intestinal villi. Absorbed
lipids thus reach the venous blood through the lymphatic system.

duct
aod blood

Fig. 1.14 :Fatty acids and monoglycerides from the mlscelles within the small intestlne are absorbed by the
epithelial cells and resyntheslsed into triglycerldes. These a n then coated by protein to form
chylomicrons which enter the lacteals of the small intestine.

SAQ 4
In the figure of the human digestive tract and the associated glands given below:
a) indicate the sites of digestive enzyme activity.
b) indicate the enzymes responsible for digestion of carbohydrates, proteins and fats.

Salivary gland
Pharynx
Trachea
klrlRw8a-I
1.6 ENERGY METABOLISM
..
In the p ~ c e d i n gsections of the unit, you studied how the products of digestion of
food, viz: amino acids, sugars and fatty acids are absorbed and transported to the
body tissues. p e bxidation of these compounds yields virtually all the chemical
energy requiraaioy animals and the use of this chemical energy is referred to as their
energy metabolism. You also learnt in Section 1.2 that generally carbohydrates and
fats are the fuel which'provide energy but other organic compounds are within wide
limits interchangeable in energy metabolism.
How do we measure the rate of metabolism or the actual amount of energy liberated
during oxidative metabolism? One way could be to measure the total heat produced
by the organism per unit df time. This is the metabolic rate of the organism. The
metabolic rate can be determined by using the formulation:
rate of energy intake
. .
- rate of energy loss per unit time = metabolic rate
. Energy intake is the chemical energy content of ingested food over a given period.
Energy loss is the chemical energy that remains in faeces and urine produced by the
animal over the same period. The energy content of food and wastes is found out by
burning them in .a bomb calorimeter (Fig. 1.15). The material to be tested is placed
and burned with the aid of oxygen in a chamber surrounded by ajacket of water. ':..
,The heat produced is determined by the rise i" temperature of the surroundingv+ti$' . .
'Table 1.10 provides the caloric value of the common food stuffs estimaied in b@b
.
;.-
calorimeter and in the body. .
. . . . .

. .
,

. .

. .
heat from the hod

& .

These values are essentially average values. During oxidative degradation in animal
body, carbohydrates and lipids are fully oxidised to carbon dioxide and water just as
in a bomb calorimeter but proteins are not degraded because the major end-product
of protein metabolism, is urea which still possess some energy. Accordingly, the value
is lower in the body (4.1 kcaUg) as you can see from Table 1.10. Energy derived
from one gram of fat is much more than that derived from 1 gram of protein or of
carbohydrate.

Tabk 1.10 : Fuel Cootent of Food Materinls

+
Food Kiloeolorksper gram

In Bomb ,In Body I

Calorimeter
Carbohydrates 4.1 4.0
Lipids 9.4 9.0
Proteins 5.6 4.1
k * .

The heat broduced during the metabolic activities of the body helps in maintaining
the body temperature. Generally, warm blooded animals like birds and mammals
' have specific regulatory. mchanisms bv which heat production is either increased or
 *
dissipated in relation to the environmental conditions. You will learn more about nutrl*, Fqdlol, ,-
temperature regulation in Unit 7 of Block 2.
The most commonly used measure to determine metabolic rate is to find the oxygen
consumption of the animal. Oxygen consumption is a good indicator of metabolic
rate because heat produced for each litre of oxygen used in metabolism is the same
irrespective of the food stuff oxidised (see Table 1.11). The highest figure of 5 . U c a l
per litre of oxygen is only 10% more than the lowest f i g u r e s M i t r e of oxygen
for proteins. O n an average we consider metabolic rate to be equal to 4.8 kcalllitre
of oxygen used.

Table 1.11 : H q t pduetlon and respiratory quotient for foodstuff types

Heat produced Kcal

Per litre Per litre C02formed
of O2 0.f COz RQ =
consumed formed . 0, used

Carbohydrates
Fats
Proteins

Respiratory Quotient
Table 1.11 also shows the ratio of thevolume of carbon dioxide evolved to that of
the amount of oxygen consumed during oxidation. This is the respiratory' quotient or
RQ. It is an important concept in energy metabolism. From the table you can see
that RQ is usually between 0.7 and 1.0. However, RQ near 0.7 shows that fat is being
metabolised and RQ near 1.0 suggests carbohydrate metabolism. RQ in between 0.7
and 1.0 could indicate either protein or a mixed diet metabolism.
Quite often animals cannot utilise the entire food value because not all the food they
consume is fully digested. Also some portion is excreted as urea or ammonia. In
general, it has been observed that animals have a higher intake of food than what is
indicated by their oxygen consumption data so that their body weight is kept steady'.
The oxygen consumption per unit weight/per unit time mm30,1g/hr tends to decrease with
higher body weight of aaimals. In other words, small sized animals $ke mouse, shrew,
etc., have a higher metabolic rate than a large sized animal (an elephant) as evidenced
by their oxygen consumption. Accordingly smaller animals have a need to feed
constarTtly. This would also mean that an elephant can survive without food for a
much longer period of time than a mouse.

Energy Storage
As we said above, food intake and energy expenditure for animals is approximately
equal. If energy expenditure exceeds food intake, th,en the excess energy is taken up
by utilisation of body fat. However, if food intake is excess, then the surplus is stored
as fat irrespective of the kind of food eaten.
Excess carbohydrates are changed to fats and accordingly RQ exceeds 1. This is
because fats contain relatively less oxygen and the excess oxygen of carbohydrates is
used in the metabolism. This reduces the oxygen uptake and the respiratory carbon
dioxide, oxygen ratio is increased.
For this reason fat is ideal storage material for energy. It is much lighter and yields
twice as much energy as carbohydrates. Migratory birds that may have to fly more
than 1000 km non-stop, carry fat as 40% to 50% of their body weight.
Nonetheless, some carbohydrates are important in energy storage. Glycogen a
starch-like carbohydrate polymer is stored as granules in the skeletal muscles and
liver of vertebrates. During heavy muscular exercise when blood does not deliver
sufficient oxygen to meet demands, glycogen provides the energy. It is broken down
directly into glucose-6-phosphate, pToviding fuel fbr carbohydrate metabolism more
directly than does fat. . ,
On the other hand, many animals that do not move about, also store glycogen as
excess energy source. For example, clams, oysters and many intestinal parasites like
Ascaris use glycogen as the storage material. These animds have to face anaerobic
conditio,ns and in such situations glycogen breaks down to acetic acid to yield energy.

SAQ 5
A person on a diet does not consume any fat but only rice and sweets. After some
time he finds that he is still gaining weight. Why?

1.7 SUMMARY
All hetesotrophic organisms require a balanced diet of nutrients to survive, grow
and meet their energy demands. Nutrients that must be supplied to the animals in
diet are called essential nutrients and the requirement for these essential nutrients
indicates the animal's synthetic ability. Animals also have a quantitative
requirement for nutrients and if they don't get the required amount, deficiency
syndromes and suboptimal growth results.
Food is obtained by animals in different ways, including absorption through body
surface, endocytoiis, filter feeding, mucous trapping, sucking, biting and chewing.
The feeding strategies hate evolved according to the nature of food required.
Digestion is the breakdown of complex food molecules into simpler constituents
and two broad categories of chemical digestion are seen - intracellular and
extracellular. Intracellular digestion is characteristic of primitive animals and
extracellular digestion of the higher forms. Digestion of proteins, carbohydrates
and lipids is a step by step process in which larger molecules are broken down by
the action of specific enzymes. Lipases are not as specific as carbohydrases and
proteases.
Secretion of digestive juices as well as the motility of smooth muscles are under
neural and hormonal control. All gastrointestinal harmones are peptides. Both
direct activation by food in the gut and neural activation stimulates the endocrine
cells of the alimentary canal to secrete hormones.
The products of digestion are taken up by the absorptive cells of the intestine and
transferred to the blood capillarJes and lymphatic system. Transport of some sugars
occurs by facilctated diffusion which does not require'metabolic energy. Most
sugars and amino acids are transported with Na+ and a carrier molecule requiring-
energy. T k products of fat digestion are absorbed by diffusion across cell
membranes.
The absorbed nutrients provide fuel for body metabolism. The chemical energy
used in metabolism can be measured as heat energy. A given class of food
molecufes will liberate the same amount of heat and require the same amount of
oxygen when oxidised to water and carbon dioyide. The heat production and '
oxygen consumption of animals gives their metabolic rate and the respiratory
quotient is useful in determining the proportion of carbohydrates, fats or proteins
metabolised

1) What are essential nutrients? Why is that some nutrients are essential for some
animals and non-essential for others?
 ..
2) Make a table that shows the characteristics of intracellular and extracellular
digestion. List out some advantages of extracellular digestion from that
information.

.........................................................................................................
'

, 3) What prevents the walls of the digestive tract from being digested in animals?
.......................................................................................................
d

4) What ar,e the end-products'of food that can be absorbed by the body? Explain
I
how absorption of fats diffkrs from absorption of proteins and-sugars.

- --- -
1.9 ANSWERS
Self-assessment Questions
1) a) i) matches b)
ii) .matches c)
iii) matches a)
b) Because these animals can synthesise vitamin C while humans and some fruit
eating mammals have lost the ability to synthesise vitamin C.
2) i) Squirrel has chisel-like incisors used especially for gnawing. Cow has molars
that are used in a side to side grinding motion for mastication of vegetation.
Dog has pointed dagger-like canines for piercing and tearing food.
ii) 1) matches d)
2) matches e)
3) matches b)
4) matches a)
5) matches c)
3) a) It permits different parts of gut to specialise for performing different
digestive functions.
b) ii)
C) amino acids, monosaccharides, fatty acids and glycerol.
d) gastrin, secretin, cholesystokinin.
4) a) 1) Salivary glands
2) Stomach .
3) Pancreas
4) Gall bladder
5) Liver
6) Small intestine
b) Carbohydrates -amylase, sucrase, maltase, lactase
Proteins -
pepsin, trypsin, chymotrypsin, carboxypeptidase
. -
Lipids Lipase.
5) Excess carbohydrates that are not used for metabolism are converted to fats and
stored in the body. This increases the weight.
Terminal Questions
1) Essential nutrients are materials the animals need to sustain life but cannot
synthesise in their cells. The nutrients that are essential'differ for each type of
animal because the ability to synthesise is genetic and species specific. For
example, most vertebrates can synthesise vitamin C but human and some other
fruit eating species have lost this synthetic ability. Therefore, vitamin C is
essential for us but non-essential for some others.

2) Intracellular Extracellular

Ingestion of small particles Ingestion of large food masses

Digestive enzymes enclosed
in a small area. One kind of
enzyme acts at one time in
digestive vacuole
Suited to continuous feeding
Usually well-developed digestive tract that '
allows secreted enzymes to act on food at
different parts at the same time, digestion
occurs in phases
Suited t o discontinuous feeding
Two openings, a mouth and anus in complex
animals so that food passes in one direction

For more complex freely moving organisms extracellular digestion is more

advantageous because they don't have to feed continuously and a divi'sion of
labour in the digestive tract occurs so that only a few cells are devoted to the
digestive processes. Digestion can occur in different phases at the same time.
3) A lining of mucous secreted by the goblet cells and the fact the most digestive
'enzyme especially protease are present in inactive form.
4) Carbohydrates are hydrolysed to monosaccharides. Proteins are hydrolysed to
amino-acids and fats are hydrolysed to the free fatty acids and glycerol.
i) Amino acids and glucose enter the intestinal epithelial cells by a carrier or
cotransporter protein molecules that depend on the action of a sodium ion
pump. Free fatty acids or monoglycerides form miscelles with bile salts before
entering the intestinal epithelium through diffusion.
ii) Amino acids and glucose are passed on to the blood stream through anothe.
transporter molecule while fatty acids and monoglycerides form
chylomicrons which enter the lacteals from where they enter the blood
stream.