Unit 3 Digestive System
Unit 3 Digestive System
UNIT 3
DIGESTIVE SYSTEM
Structure
3.1 Introduction 3.4 Digestive System in Non-
Objectives
Mammalian Vertebrates
Reptiles
Fishes
Birds
Amphibians
Reptiles
3.5 Digestive System in
Mammals
Birds
Trophic Groups in Mammals
Mammals
Digestive Tract
3.1 INTRODUCTION
In the previous unit you have studied about cartilage and bones of vertebrates.
In this unit you will study that all vertebrates possess an alimentary canal as
an organ for digestion and absorption of food materials. Digestion does not
occur in a particular region of the alimentary canal but takes place in different
regions so that digestion of food materials may be completed. All vertebrate
alimentary canals have a common basic organization and similar major
subdivisions. However, the differences seen in the anatomy in different animal
groups is correlated to their different feeding habits. We shall discuss here the
feeding mechanism in different groups of vertebrates such as fishes,
amphibians, reptiles, birds and mammals and correlate with the adaptations
seen in their alimentary canal.
We will start the unit with the study of structure and arrangement of teeth in
vertebrates and see that they can be classified in different ways. Then we will
discuss the feeding mechanisms and digestive systems of non-mammalian
and mammalian vertebrates.
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Block 1 Comparative Anatomy of Vertebrates-I
Objectives
After reading this unit you should be able to:
In most other vertebrates teeth develop partly from the epidermis and partly
from the underlying dermis. There is a covering of enamel with an underlying
layer of dentine. Fishes, amphibians and reptiles have simple pointed teeth.
Some fishes like the chimeras and lungfishes have plated teeth that have
rough or serrated ridges separated by grooves. In mammals each tooth is
lodged in a socket called alveolus in the jaw. The part of tooth developed from
epidermis is enamel. The remainder of tooth - dentine, cement and pulp - is
formed from the sub-adjacent mesodermal tissue (Fig.3.1). Amongst
vertebrates the teeth of lampreys are quite different from all others as they are
cornified epidermal structures that do not have either enamel or dentine.
(a) (b)
Fig. 3.1: Tooth structure (a) Tooth with single root, (b) Molar tooth with roots.
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Unit 3 Digestive System
Teeth can be classified according to their attachment with the jaws (see
Fig.3.2), appearance and nature of replacement as given in the Tables 3.1 and
3.2.
Fig. 3.2: Classification of teeth on the basis of attachment to the jaw bone. a) In
thecodont type the teeth are set in sockets in the jaw bone as seen in
mammals and alligators; b) In acrodont type the teeth are attached on
the surface of the jaw bone as seen in most non-mammalian
vertebrates; c) in pleurodont type the teeth are attached to the rim, on
the inner side of the jaw bone as seen in modern amphibians and
lizards.
The most specialized teeth among vertebrates are seen in mammals, the teeth
are not only specialized to capture or clip food, but they are also specialized to
chew it.
The heterodont dentition of mammals includes four types of teeth within the
mouth - i.e. incisor at the front, canine next to them, premolars along the
sides of the mouth, and molars at the back. Incisors are generally used at the
front of the mouth for cutting or clipping; canines for puncturing and holding;
premolars and molars for crushing or grinding food. It is often hard to
distinguish premolars from molars visually. The collective term used for both is
cheek or molariform teeth. Cheek teeth may be quite diverse, a reflection of
their many specialized functions.
The number of each type differs among groups of mammals. The dental
formula is shorthand expression of the number of each kind of tooth on one
side of the jaw for a taxonomic group. For example, the dental formula of dog
is:
3 1 4 2
I. ,C. ,Pm. , M.
3 1 4 3
This means that there are three upper and three lower incisors (I) one upper
and one lower canine (C), four upper and four lower premolars (Pm), and two
upper and three lower molar (M), 21 per side or 42 total (see Fig.3.3 also).
Sometimes, the dental formula is written as 3-1-4-2/3-1-4-3, the first four
numbers indicating the upper teeth and the second four numbers are the lower
teeth. The dental formula for mouse is 1-0-0-3/1-0-0-3. You notice that the
missing canines and premolars are indicated by zeros. Rodents do not have
canines their incisors are used for gnawing, scraping and nibbling. The
78 incisors are sharp and keep growing throughout life.
Unit 3 Digestive System
The teeth of mammals are modified according to their feeding habits and can
be classified accordingly as given in Table 3.3. In herbivores the teeth are
hypsodont that can grind plant material to break tough cell walls. Their cusp or
occlusal surface is worn unevenly because the minerals which form the
surface-enamel, dentine and cementum-differ in hardness. Occlusal surfaces
are functionally important because they ensure that ridges and depressions
persist throughout life, thereby maintaining a rough grinding surface which
does not become smooth with continued use. The selenodont and lophodont
teeth have folds of enamel running deep in between areas of dentine, as the
teeth surfaces wear off the surface enamel is lost but the folds remain. The
exposed dentine wears off so that ridges of enamel are separated by grooves
of dentine thus forming good grinding surfaces.
Hypsodont
Brachyodont
Bunodont
Lophodont
Selenodont
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Unit 3 Digestive System
Fig. 3.4: Tusks arise from both upper incisors in the elephant, and from canines
in walrus. The teeth are absent in adult anteaters, and baleen whales.
SAQ 1
In the following statements, put a tick mark (√) on the correct ones and a cross
(×) mark on the incorrect ones in the given boxes.
i) The part of the tooth developed from the epidermis is the enamel. ( )
ii) Thecodont dention has teeth arising from sockets in the jaw bone. ( )
iv) Some mammals have indefinite number of teeth e.g. elephant and ( )
monkey.
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Block 1 Comparative Anatomy of Vertebrates-I
3.3 FEEDING MECHANISMS
Obtaining nutritional essentials is clearly a key to the success of any
animal/species. Much of the routine functioning of an animal is directed
towards this purpose. For example, the complex and sophistication of the
nervous system, evolved largely due to the selective pressure on obtaining of
sufficient food and on avoidance of becoming someone else's meal. Animals
use various strategies to feed. Some species search, stalk, pounce, capture,
and kill. Sessile animals, unable to move about, resort to more subtle means,
such as surface absorption, filter feeding or trapping. Vertebrate, have various
feeding devices that are described here in a systematic evolutionary
sequence. We will now discuss different devices employed by various groups
of vertebrates.
3.3.1 Fishes
One of the most important, successful and widely employed methods for
feeding that evolved is filter feeding. Majority of filter feeders use ciliated
surfaces to produce currents which can draw drifting food particles into their
mouths. Free-swimming filter feeders like herring and basking shark have the
advantage of being able to swim through their food and thus can be more
selective in their feeding. The water flows out through the gills leaving the food
behind (Fig. 3.5).
A second group of fishes are herbivores which eat flowering plants, algae and
grasses. Although plant eaters are relatively few in number, they are crucial
intermediates in the food chain especially in fresh water rivers, lakes, and
ponds which contain very little plankton.
Another group of fishes are omnivores feeding on both plant and animal food.
Finally there are scavengers which feed on organic debris and on the
82 parasites that suck the body fluids of other fishes.
Unit 3 Digestive System
Fig. 3.5: Herring and other filter-feeding fishes use gill rakers, which project
forward from the gill bars into the pharyngeal cavity to strain out
planktons. Herring swim almost constantly, forcing water and
suspended food into the mouth, food is strained out by gill rakers, and
water passes out of the gill openings.
3.3.2 Amphibians
Adult amphibians consume a wide variety of food. Anurans and salamanders
feed mostly on insects and other arthropods, while caecilians feed mostly on
earthworms and other small bugs. Aquatic salamanders lunge at their prey
with wide open mouths sucking it in with their expanded buccal cavity. Frogs
are carnivores like most other adult amphibians. They feed on insects, spiders,
worms, slugs, snails. millipedes or any thing else which moves and is small
enough to be swallowed whole. They snap at moving prey with their
protrusible tongue (Fig. 3.6) which is attached to the front of the mouth and is
free behind. The free end of the tongue is highly glandular and produces a
sticky secretion, that adheres to the prey. Teeth present on pre-maxillae, and
vomers are used to prevent escape of prey but not for biting or chewing them.
The larval stages of anurans (tadpoles) are herbivores, feeding on pond algae
and other vegetable matter.
Fig. 3.6: Feeding in frog showing the position of tongue in catching an insect.
The tongue in frog is highly mobile. It is attached to the front of the
mouth permitting the sticky organ to be flicked far out with
considerable speed and accuracy, the insect sticks at the end and the
tongue returns back in the mouth. The prey is then crushed against a
peculiar patch of teeth on the roof of the mouth and swallowed whole.
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Block 1 Comparative Anatomy of Vertebrates-I
3.3.3 Reptiles
Reptiles are mostly carnivorous though some like the land turtles, tortoises are
vegetarian feeding on grass and other vegetable matter. Green iguanas and
Uromastix are also vegetarian while marine iguanas feed on sea weed.
Reptiles like chameleon fling out their tongues to catch insects from a distance
of several inches.
Fig. 3.7: Rattlesnake skull in a side view. (a) Partly open. (b) Open for striking.
The fangs are tubular for delivery of toxin and are hinged to facilitate
their storage between strikes.
Reptilian jaws or palate are provided with pointed teeth that help these
animals in holding, tearing or swallowing their prey. Non-mammalian teeth are
generally poorly differentiated from one another. One exception is found
among the poisonous snakes, such as vipers, cobras and rattle snakes, which
have modified teeth, called fangs which are used to inject venom (Fig. 3.7).
These fangs are either equipped with a groove or are hollow, very much like a
syringe needle to deliver venom at the site of bite. In rattle snakes, fangs fold
back against the roof of the mouth, but extend perpendicularly when the mouth
is opened to strike. Snakes cannot tear or chew their food. Captured prey is
swallowed whole, a surprising feat since the prey is often larger than the
snakes. The mouth is extremely flexible because of the arrangement of bones
in the head and jaw. The lower jaw is loosely attached to the quadrate bone
and can be disconnected while swallowing and even the bones of the palate
are movable, all helping to draw the prey into the gaping mouth. The
oesophagus and stomach can stretch considerably as can the body wall.
There is no sternum, so the ribs can move freely as the prey passes through
the gut. This enables a snake to swallow animals larger than the diameter of
its head.
3.3.4 Birds
Birds have no teeth, but instead have horny beaks which exemplify adaptive
radiation suited to a gastronomic (art of choosing, preparing and eating good
food) life style.
Birds consume a variety of other animal foods such as seeds, fruit, insects,
84 worms, molluscs, crustaceans, fish, frogs, reptiles, mammals as well as other
Unit 3 Digestive System
birds. A very large group of birds feed on nectar. Some birds are generalists
that is, omnivorous like crows and bluejays that eat whatever is seasonally
abundant. Others are specialists called stenophagous, or “narrow range
eating” species that focus on a specific type of food.
Beaks of birds are strongly adapted to their feeding strategies. Beaks may be
generalised as strong, pointed beaks of crows, to grotesque, highly
specialised ones in flamingoes, hornbills and toucans. Figure 3.8 shows some
of the specialized type of beaks that suit the birds feeding habits. The beaks of
small seed eaters are short, stout, pointed like that of a sparrow. The crossbill
is a unique type of seed eater, its upper and lower beaks do not align and are
crossed, it can use the beak to open pine cones. Seed eating birds eat their
food as whole, but may subject it to grinding in a muscular gizzard which
contains stones (gastroliths) pebbles which aid birds in the grinding process.
Insect eaters have thin, short pointed beaks. A well known insect eater is the
woodpecker which has a straight, hard, chisel like beak. Anchored to a tree
trunk with its tail serving as a brace, the woodpecker delivers powerful, rapid
blows (upto 20 times/sec) to build nests or expose the burrows of wood boring
insects. It then uses its long, flexible, barbed tongue (three times longer than
the bill) to seek out insects in their holes and tunnels. Woodpecker’s skull is
especially thick and fitted close to the brain to absorb shock.
Nectar eaters like the hummingbird has long, slender beak, that is a protective
covering for its tongue. The tongue has two grooves that draw water by
capillary action. They also supplement their diet by catching flying insects by
widening the gape of the mouth as their mandible can be actively flexed
downwards.
Birds of prey such as hawks, kites, owls, eagles etc., have interesting
adaptations. In some cases, the feeding habits are reflected in the feet as well.
Typically raptorial birds or birds of prey, have long, curved talons for grasping
prey which they tear with their strong hooked beaks. Ground foraging species
such as grouse and pheasants have heavy, strong feet for scratching the soil.
Flamingo uses a filter feeding apparatus (Fig. 3.9) to filter small organisms and
other morsels it finds in the muddy bottoms of its fresh water habitat.
3.3.5 Mammals
Mammals feed on a wide variety of food sources. Some mammals require
highly specialised diets, while others are opportunistic feeders which live on
diverse diet. In all mammals food habits and physical structure are intimately
linked. Specializations for finding, capturing, reducing, swallowing, and
digesting food determine the shape and habit of a mammal, Teeth, more than
any other single physical characteristic, reveal the life habit of a mammal (Fig.
3.10). All mammals have teeth, except certain whales, monotremes and
anteaters and their modifications are correlated with what they eat.
As mammals evolved, major changes occurred in the teeth and jaws during
the mesozoic era. Unlike the uniform homodont dentition of the reptiles.
mammalian teeth became differentiated to perform specialized functions such
as: cutting, seizing, gnawing, tearing, grinding or chewing (please refer to
section 3.2 again). Teeth differentiated in this way are called heterodont. The
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Unit 3 Digestive System
primitive tooth formula, which represents the number of each tooth type in one
half of the upper and lower jaw, was I,3/3, C 1/1, Pm 4/4, M 3/3. Members of
the order Insectivora e.g., shrews, some omnivores and carnivores come
closest to this primitive pattern (Fig.3.10). Unlike reptiles, mammals do not
replace their teeth continuously throughout their lives. Most mammals grow
just two sets of teeth, a temporary set called deciduous or milk teeth (set)
which is replaced by a permanent set when skull has grown large enough to
accommodate a full set. Only incisors, canines and premolars are deciduous;
molars are never replaced and the single permanent set must last a life time.
SAQ 2
Fill in the blank spaces with appropriate words from the text.
iv) The tongue of frog flicks out to catch insects because it is attached in the
____________of the mouth.
3.4.1 Fishes
As you can see in fig. 3.11 the alimentary canal of Scoliodon consists of
mouth, buccal cavity, pharynx, oesophagous, stomach, intestine and rectum.
Mouth is a ventral crescentric opening which leads into a spacious dorso-
ventrally compressed buccal cavity. The buccal cavity is lined with a thick
mucous membrane raised ventrally into a thick fold to form the so called
tongue which is non-muscular and non-glandular. The mucous membrane is
rough due to the presence of dermal denticles. Teeth are oblique and have
sharp more or less compressed cusps, the edges of which are smooth and
non-serrated. Teeth are all alike in shape, homodont, and are borne in
several parallel rows on the inner margin of the upper and lower jaws. Teeth
88 are used to catch prey and prevent its escape but not to crush or masticate it.
Unit 3 Digestive System
Though there are several rows of teeth (polyphyodont) yet only one row
functions at a time and the old row is replaced by a new one. There are no
glands in the buccal cavity comparable to the salivary glands of higher
vertebrates.
The buccal cavity opens into pharynx on either side of which lie the internal
openings of the spiracle and five gill-pouches. The cavity of pharynx is lined
with mucous membrane containing numerous dermal denticles. The secretion
of mucous glands in the pharynx has no digestive function but simply helps in
lubricating the passage of food.
The pharynx narrows posteriorly to form the short oesophagus which has thick
muscular walls with an internal lining of mucous membrane raised into
longitudinal folds.
The bursa entiana continues into the intestine. The intestine is a wide tube
running straight backward into the abdominal cavity and opens posteriorly into
the rectum. The internal surface of the intestine is increased by a
characteristic fold of the mucous membrane, the spiral valve, having one
edge attached to the inner wall of the intestine and the other rolled up
longitudinally on itself into a spiral, making an anti-clock wise spiral of about
two and a half turns. In a transverse section the spiral valve looks like a watch
spring. The spiral valve serves not only to increase the extent of the absorptive
surface of the intestine but also prevents rapid flow of food through the
intestine.
The rectum is the last part of the alimentary canal. The tubular rectal (caecal)
gland opens dorsally into the rectum. The rectum leads into cloaca into which
the alimentary canal as well as the urinogenital ducts open.
The glands of alimentary canal comprise a massive bilobed liver and a thin V
shaped gall bladder. The bile duct opens in the anterior part of the intestine
where the spiral valve starts. The pancreas is a pale compact irregular body
lying between the fold of the stomach and intestine.
In a very large majority of teleost species the teeth are small, conical, and
recurved, suitable for preventing the struggling prey from slipping out of the
mouth, but quite unfitted for either tearing or crushing. In some bony fishes
alimentary canal shows little differentiation into regions, but as a rule, gullet,
stomach, duodenum, ileum and rectum are more or less clearly
distinguishable histologically. The stomach is V-shaped but its cardiac region
may be prolonged into a blind pouch (Fig. 3.12). This is often very distensible,
allowing some of the deep-sea Teleostei to swallow fishes as large as
themselves. In many genera of several families stomach is entirely absent.
Globe fishes can inflate the gullet with air or water, as a result of which they
can float upside down. A spiral valve is very well developed in Polypterus and
Sturgeons, vestigial in Lepisoteus and Amia, and absent or vestigial in all
Teleostei, except possibly in Chirocentrus (Isospondyli). A trace occurs in the
herring. Liver is usually large. Pancreas may be present as a compact gland,
or may be widely diffused between layers of the mesentery, or in part
surrounded by the liver. Pyloric Caeca are commonly present, and vary in
number from a single one to two hundred. Anus is always distinct from, and in
90 front of the urinogenital aperture.
Unit 3 Digestive System
3.4.2 Amphibians
The mouth opening is a wide slit. Teeth, which are ankylosed to the bones, are
present upon the premaxillae, maxillae and vomers, They are absent in Pipa
and some toads. Tongue is immovable in Urodela, movable and free behind in
Anura, in which it is used as a prehensile organ. Salivary glands are not
present. In many oesophagus, stomach, small intestine and rectum are
present.
The pharynx leads into a broad tubular part of the alimentary canal called
oesophagus. This part of alimentary canal is very short due to the absence of
neck. But the oesophagus is highly distensible as its inner lining is thrown into
a large number of longitudinal folds that allow its expansion during the
passage of the ingested food to the stomach. The stomach comprises two
parts, the anterior expanded cardiac part and the posterior short narrow
pyloric part. The stomach opens into long tubular part the intestine consisting
of two parts the small and large intestine. The small intestine leads to a short,
broad colon or large intestine called rectum. The hind end of the rectum is
called the cloaca and possesses a median ventral appendage, the urinary
bladder. The urinary and generative ducts open into cloaca. The cloaca opens
to the exterior by the anus. The digestive glands such as liver and pancreas
are present and the liver has a gall bladder. 91
Block 1 Comparative Anatomy of Vertebrates-I
3.4.3 Reptiles
Teeth are usually present on the premaxillae, maxillae and dentary, and
frequently on the palatine and pterygoid. They are continually replaced, and
are pleurodont, acrodont, or thecodont. Teeth are conical or hooked and are
adapted for prehension not for mastication except in some extinct forms. In
Chelonia teeth are absent, being replaced by the horny epidermal beak-like
covering of jaws.
In this section to make you understand the digestive system of reptiles we will
describe the digestive system of a Uromastix lizard (Fig. 3.14). The alimentary
canal is a long and convoluted tube. It can be divided into mouth, buccal
cavity, pharynx, oesophagus, stomach, duodenum, ileum or small intestine,
colon or large intestine, rectum and cloaca. The mouth opening is a wide gap
bounded by the upper and lower immovable and muscular lips. The mouth
opens into the buccal cavity which has a well developed muscular tongue on
its floor. Tongue is attached to the floor of the buccal cavity along the median
ventral line. The tongue is long, bifid and protrusible having voluntary muscles,
taste buds and mucous glands. In the upper jaw, teeth are present on
premaxillae and maxillae, whereas in the lower jaw the dentary bears teeth.
The teeth of Uromastix are pleurodont which means teeth are attached to the
outer border of the bones of jaw.
The pharynx lies posterior to the tongue. The lining of the pharynx is thrown
into longitudinal folds. The pharynx leads posteriorly into the oesophagus. It is
capable of great distension and it opens into long cylindrical sac like structure
the stomach which is wider than the oesophagus. It has thick muscular walls
and lies in the left half of the body and is curved having the appearance of U-
shape. The stomach is differentiated into two parts; the anterior part is known
as cardiac stomach which lies dorsal to the left lobe of the liver, and the
92 posterior part is known as pyloric stomach lying slightly to the right side. The
Unit 3 Digestive System
wall of the stomach is much thicker than that of the oesophagus and of the
intestine. Numerous well developed longitudinal folds of mucous membrane
are seen in the lumen of both cardiac and pyloric stomach. Stomach is the
place where digestion occurs. The pyloric wall is in the form of a muscular ring
lining of the inner wall of the posterier extremity of the pyloric stomach.
You can see from Fig. 3.14 a that ileum opens into a large intestine which
comprises a proximal colon and distal rectum. The caecum pouch arises from
the junction of ileum and colon. Rectum opens into cloaca that in turn opens to
the exterior by the cloacal opening. The cloaca consists of three chambers,
the coprodaeum, the urodaeum and the proctodaeum. Different chambers of
cloaca serve for reabsorption of water both from faeces and urine. The
digestive glands that are associated with the alimentary canal are gastric
glands, liver, pancreas and intestinal glands. 93
Block 1 Comparative Anatomy of Vertebrates-I
The salivary glands are usually absent in all reptiles. There is a sub-lingual
gland in Chelonia. Both upper and lower labial glands and palatal and lingual
glands are always present. The poison glands of snakes are upper labial
salivary glands.
3.4.4 Birds
Inspite of great differences in the mode of nourishment the avian digestive
organs present a fairly uniform structure; their unusual structures in the
digestive system are adaptations for flight. The jaws are covered by a hard
horny sheath (rhamphotheca) and transformed into the beak. The
rhamphotheca is often composed of several pieces (compound). True teeth
are entirely absent, at least in living birds. While the upper beak is formed by
the fused premaxillae, the maxillae and the nasal bones, the lower beak
corresponds to the two rami of the lower jaw, the fused extremities of which
are known as the myxa (Fig.3.15). The form and development of the beaks
vary extremely according to the special mode of subsistence (see Fig. 3.8
again).
Tongue which is always movable, lies on the floor of buccal cavity. It consists
of horny or fleshy covering of two cartilages attached to the anterior end of the
hyoidbone and serves for pushing the food back, and frequently for seizing
food. Oesophagus is muscular, longitudinally folded and the length generally
depends on the size of the neck. It frequently possesses a dilation known as
crop especially in birds of prey, and also in granivorous birds, in which the
food is stored and softened (Fig. 3.16a). In pigeons crop bears two small
rounded accessory sacs.
SAQ 3
Match the statement given in column B with the chordate group given in
column A.
Column A Column B
Fig. 3.17: Digestive System of an insectivore showing short intestine. Note there
is no caecum.
Herbivorous mammals which feed on grasses and other vegetation form two
main groups; the browsers and grazers, such as the ungulates, that are,
hooved mammals including horses, deer, antelope, cattle, sheep and goats,
and the gnawers such as the rodents which include rabbits and hares. In
herbivores, the canines are either reduced in size or may be absent, while the
molars that are adapted for grinding, are broad and usually high-crowned.
Rodents have sharp chisel-shaped incisors which grow throughout life and
must be worn away to keep pace with their continuous growth. If the incisors
do not grow continuously, the cutting surface of the teeth will get eroded soon
by its excessive use due to its continuous gnawing habit.
In some of the herbivores like horses, zebras, rabbits, hares, deer, elephants
and many rodents, gut has a spacious side pocket or diverticulum, called a
caecum that serves as a fermentation chamber and absorptive area (Fig. 3.18
& 3.19). Hares, rabbits and some rodents often eat their faecal pellets
(coprophagy) giving the food a second pass-through the fermentating action
of the intestinal bacteria. Coprophagy also provides an opportunity for the
animal to obtain vitamins produced by the bacteria lodged in the caecum. 97
Block 1 Comparative Anatomy of Vertebrates-I
Ruminants like cattle, bison, buffalo, goats, antelopes, sheep, deer, giraffes
etc. have a huge four chambered stomach (Fig. 3.19). You will learn more
about the specialized ruminant stomach a little later in this section.
Fig. 3.20: Digestive system of a carnivore showing short intestine and colon,
small ceacum.
Generally carnivores lead more active life than do the herbivores. Since a
carnivore must find and catch its prey, there is a premium on speed and
intelligence; many carnivores, such as cats are known for their stealth and
cunningness in hunting prey. This has lead to a selection of herbivores
capable either of defending themselves or of detecting and escaping
carnivores. Thus for the herbivores, there has been a premium on keen
senses and agility. Some herbivores, however, survive by virtue of their sheer
size e.g. elephants or by defensive group behaviour for example North
American musk oxen.
Omnivorous mammals live on both plants and animals for food. Examples
are pigs, racoons, rats, bears and most primates including human beings.
Many carnivore forms also eat fruits, berries and grasses when hard pressed.
The fox which usually feeds on mice, small rodents and birds, eats frozen
apples, beech nuts, and corn when its natural food is scarce. For most
mammals, searching for food and eating occupy most of their active life. Some 99
Block 1 Comparative Anatomy of Vertebrates-I
migrate to regions where food is in plenty, while others hibernate and sleep
during the winter months. But there are many provident mammals which build
up food stores during period of plenty. This habit is most pronounced in
rodents such as squirrels, chipmunks, gophers and certain mice.
(i)
101
Block 1 Comparative Anatomy of Vertebrates-I
(ii)
The oesophagus opens into the rumen close to its junction with the reticulum.
The rumen is much larger than the rest. Its mucous membrane is beset with
numerous short villi. The reticulum, is much smaller than the rumen, has its
mucous membrane raised up into a number of anastomosing ridges. These
are capable of closing together in such a way as to convert the groove into a
canal. The mucous membrane of omasum is raised up into numerous
longitudinal leaf-like folds. The abomasum, smaller than the rumen but larger
102 than the reticulum, has a smooth vascular and glandular mucous membrane.
Unit 3 Digestive System
In some ruminants omasum is absent. In the rumen and reticulum of deer,
cattle, sheep etc., there exist a dense population of protozoa and bacteria that
attack and break down cellulose which forms the major part of the diet.
Fermentation produces acetic, butyric and propionic acids which are
neutralized by sodium bicarbonate secreted in the saliva. As a result methane
and CO2 are produced and belched out.
Food residues, fluid and micro-organisms move down the alimentary canal. In
the omasum fluid is absorbed. In the abomasum, the protozoa and probably
the bacteria, are destroyed by secreted HCI. The abomasum also produces
digestive enzymes. In camels stomach is not as complex as in the more
typical ruminants. There is no distinct omasum, and the rumen is devoid of
villi. Both rumen and reticulum have connected with them a number of pouch
like diverticula, the openings of which are capable of being closed by sphincter
muscles. In Cetacea stomach is also divided into compartments. In porpoises
(sea animals like a dolphin or a small whale) oesophagus opens into a
spacious rumen, the cardiac compartment of the stomach, which has a
smooth, thick, mucous membrane. This is followed by a second median
chamber of considerably smaller dimensions. This has a glandular mucous
membrane, which is thrown into a number of complex folds. A long and narrow
third or pyloric, compartment follows terminating in a constricted pyloric
aperture. Beyond this beginning of the small intestine is dilated into a bulb.
Absorption of nutrients takes place in the small intestine which is the longest
part of the alimentary canal in most vertebrates (see the figures of digestive
tracts in the previous sub-section). The inner surface of the intestine is highly
folded with finger like projections that increase the absorptive surface. In
humans (see Fig. 3.23) the intestines are divided broadly in three regions: the
duodenum which connects to the stomach, the jejunum where most of the
carbohydrates and amino acids are absorbed and the third part is known as
ileum, where vitamins and bile salts are absorbed into the blood stream.
The rectum is the terminal part of the large intestine as seen in Fig.3.23. Its
primary function is to store faeces until defecation through the anus. The
Prototherians resemble reptiles, birds and Amphibia and differ from most
mammals in the retention of a cloaca. Into this not only rectum but the urinary
and the genital ducts open. In marsupials a common sphincter muscle
surrounds both anal and urinogenital apertures. In female there is a definite
cloaca. In nearly all eutherians the apertures are distinct, and separated from
one another by a considerable space the perinaeum.
The most common oral glands in mammals are the salivary glands.
There are usually three primary pairs of salivary glands, named for their
approximate positions: mandibular (submandibular or submaxillary),
sublingual, and parotid. They form the saliva, which is added to food in
the mouth. These three pairs of glands lie at the angle of the jaws,
usually at about the juncture between the head and the neck, but they
104 are positioned superficial to the neck musculature. Ducts from the
Unit 3 Digestive System
mandibular and sublingual glands run anteriorly and release secretions
into the floor of the buccal cavity. The duct from the parotid gland opens
into the roof of the buccal cavity. In some species, additional salivary
glands may be present. In dogs, cats, and some other carnivores, a
zygomatic (orbital) gland is present, usually beneath the zygomaticarch
(Fig. 3.24). Like most digestive secretions, saliva contains mucous, salts,
proteins, and a few enzymes, most notably amylase, which initiates
starch digestion. Saliva also aids swallowing by lubricating food.
Fig. 3.24: Salivary glands of a mammal, dog. Note the locations of the main
salivary glands (sublingual, mandibular, and parotid) along with
their ducts leading to the buccal cavity. All mammals possess
these three salivary glands. In dogs and cats, a zygomatic gland
is also present.
ii) Liver : Like all vertebrates, mammals possess a liver. The liver is the
second largest organ in the body only exceeded in size by the skin. It
functions in a wide variety of roles. Early in foetal life, the liver is directly
involved in the production of red blood cells, and later it is involved in the
destruction of old blood cells. It consists of two parts or main divisions
(i.e. right and left) completely separated from one another by a fissure
termed the umbilical, owing to its marking the position of the foetal
umbilical vein. Throughout life, liver detoxifies and removes toxic
substance from the blood. Majority of mammals have a gall-bladder.
When it is present, it is attached to or embedded in, the right central lobe
of the liver. The gallbladder is absent in cyclostomes, most birds, and a
few mammals, but otherwise is present throughout vertebrates. Gall
bladder is absent in the Perissodactyla (horse, tapirs rhinoceros), the
hyracoids and some rodents. Bile is manufactured by hepatocytes in the
liver and collected in the bile ducts, stored in the gallbladder, and
emptied into duodenum the via the common bile duct to emulsify fats, or
break them up into smaller droplets.Carbohydrates, proteins, and fats
are stored and metabolized in the liver. The liver is one of the most
heavily vascularized organs of the body, being supplied with arterial
blood via the hepatic artery. However, it is also supplied with venous
blood via the hepatic portal vein which runs directly from the intestines
and spleen to the liver, and deliversblood rich in nutrients absorbed
products of digestion.
Whether one or two, the ducts empty into the duodenal portion of the
intestine and release an alkaline exocrine product, pancreatic juice,
composed of the proteolytic enzyme trypsinogen, which is converted
within the intestine to the active protease, trypsin. Amylases for
carbohydrate digestion and lipases for fat digestion are also secreted.
Embedded in the pancreas are small pancreatic islets (islets of
Langerhans) that produce the hormones insulin and glucagon, both of
which regulate the level of glucose in the blood. The pancreas is thus
both an exocrine gland, producing pancreatic juice, and an endocrine,
gland, producing insulin and glucagon.
SAQ 4
Tick out (√) the correct answer in the box.
(i) Which organ in herbivores serves a fermentation chamber and provides
absorptive area?
(a) Liver
(c) Pancreas
(d) Caecum
(a) carnivores
(b) ruminants
(c) rodents
(d) Insectivores
(iii) In which mammal caecum has fluid storing and digestive functions?
(a) Kangaroo
(b) Lion
(c) Horse
(d) Man
(a) Insectivores
(b) Protherians
(c) Carnivores
(d) Cetacians
106
Unit 3 Digestive System
3.6 SUMMARY
Let us sum up what we have learnt in this unit:
Teeth are present in all vertebrates except birds at some stage in their
lives. Teeth develop partly from the epidermis and partly from the
underlying dermis. Teeth can be classified according to their way of
placement in the jaw, nature of replacement and their appearance.
Mammals have two distinct sets of teeth, the deciduous (milk) and the
permanent dentition. The number of various categories of teeth in the
jaws in mammals is conveniently expressed by a dental formula in which
the kind of teeth such as incisor, canine, premolar and molar are
indicated by the initial letters as i, c, pm, m.
The oral cavity receives the food and oesophagus conducts the food by
peristaltic motion and connects to the stomach. In birds the oesophagus
may have a diverticulum called crop to store grain. The stomach is
amuscular chamber which is the site of storage and digestion. It shows
increasing specialization from fishes to amphibians to reptiles. Birds
have a proventriculus (glandular stomach)and ventriculus (muscular
stomach or gizzard).
The small intestine is the longest part of the alimentary canal. It is the
site of absorption of digested food. The colon is the large intestine and
the site of absorption of water.
107
Block 1 Comparative Anatomy of Vertebrates-I
The caecum is simple in monotremes, absent in sloths, some cetaceans
and in a few carnivores. In humans and a few other animals like civets,
some rodents, monkeys the distal end of caecum has degenerated into
an appendix vermiformis. In some herbivores caecum is enormous.
Associated with alimentary canal there are various digestive glands like
oral glands in mammals, liver and pancreas found all vertebrates
including mammals.
3.8 ANSWERS
Self-Assessment Question
1. i) correct ii) correct
(iv) front
Amphibians: x)
Terminal Question
1. Refer to Section 3.2 of the unit.