TOPIC # 8 DIGESTIVE SYSTEM

Animals obtain their nutrition by consuming other organisms. Depending on their diet, animals can be
classified into the following categories: plant eaters (herbivores), meat eaters (carnivores), and those
that eat both plants and animals (omnivores). The nutrients and macromolecules present in food are not
immediately accessible to the cells. There are a number of processes that modify food within the animal
body to make the nutrients and organic molecules accessible for cellular function. As animals evolved in
complexity of form and function, their digestive systems have also evolved to accommodate their various
dietary needs.

Herbivores are animals whose primary food source is plant-based. Examples of herbivores include
vertebrates like deer, koalas, and some bird species, as well as invertebrates such as crickets and
caterpillars. These animals have evolved digestive systems capable of handling large amounts of plant
material. Herbivores can be further classified into frugivores (fruit-eaters), granivores (seed eaters),
nectivores (nectar feeders), and folivores (leaf eaters).

Carnivores are animals that eat other animals. The word carnivore is derived from Latin and literally
means “meat eater.” Wild cats such as lions and tigers are examples of vertebrate carnivores, as are
snakes and sharks, while invertebrate carnivores include sea stars, spiders, and ladybugs. Obligate
carnivores are those that rely entirely on animal flesh to obtain their nutrients; examples of obligate
carnivores are members of the cat family, such as lions and cheetahs. Facultative carnivores are those
that also eat non
animal food in addition to animal food. Note that there is no clear line that differentiates facultative
carnivores from omnivores; dogs would be considered facultative carnivores.

Omnivores are animals that eat both plant and animal-derived food. In Latin, omnivore means to eat
everything. Humans, bears and chickens are example of vertebrate omnivores; invertebrate omnivores
include cockroaches and crayfish.

Invertebrate Digestive Systems

Animals have evolved different types of digestive systems to aid in the digestion of the various foods they
consume. The simplest example is that of a gastrovascular cavity and is found in organisms with only one
opening for digestion. Platyhelminthes (flatworms), Ctenophora (comb jellies), and Cnidaria (coral, jelly
fish, and sea anemones) use this type of digestion. Gastrovascular cavities are typically a blind tube or
cavity with only one opening, the “mouth”, which also serves as an “anus”. Ingested material enters the
mouth and passes through a hollow, tubular cavity. Cells within the cavity secrete digestive enzymes that
break down the food. The food particles are engulfed by the cells lining the gastrovascular cavity.

The alimentary canal is a more advanced system: it consists of one tube with a mouth at one end and an
anus at the other. Earthworms are an example of an animal with an alimentary canal. Once the food is
ingested through the mouth, it passes through the esophagus and is stored in an organ called the crop;
then it passes into the gizzard where it is churned and digested. From the gizzard, the food passes
through the intestine, the nutrients are absorbed, and the waste is eliminated as feces, called castings,
through the anus.

Vertebrate Digestive Systems

Through evolution, vertebrate digestive systems have adapted to different diets. Some animals have a
single stomach, while others have multi-chambered stomachs. Birds have developed a digestive system
adapted to eating unmasticated food.

Monogastric: Single-chambered Stomach

As the word monogastric suggests, this type of digestive system consists of one (“mono”) stomach
chamber (“gastric”). Humans and many animals have a monogastric digestive system. The process of
digestion begins with the mouth and the intake of food. The teeth play an important role in masticating
(chewing) or physically breaking down of food into smaller particles. The enzymes present in saliva also
begin to chemically break down food. The esophagus is a long tube that connects the mouth to the
stomach. Using
peristalsis, or wave-like smooth muscle contractions, the muscles of the esophagus push the food
towards the stomach. In order to speed up the actions of enzymes in the stomach, the stomach is an
extremely acidic environment, with a pH between 1.5 and 2.5. The gastric juices, which include
enzymes in the stomach, act on the food particles and continue the process of digestion. Further
breakdown of food takes place in the small intestine where enzymes produced by the liver, the small
intestine, and the pancreas continue the process of digestion. The nutrients are absorbed into the blood
stream across the epithelial cells lining the walls of the small intestines. The waste material travels on to
the large intestine where water is absorbed and the drier waste material is compacted into feces; it is
stored until it is excreted through the rectum.

Avian
Birds face special challenges when it comes to obtaining nutrition from food. They do not have teeth and
so their digestive system must be able to process un-masticated food. Birds have evolved a variety of
beak types that reflect the vast variety in their diet, ranging from seeds and insects to fruits and nuts.
Because most birds fly, their metabolic rates are high in order to efficiently process food and keep their
body weight low. The stomach of birds has two chambers: the proventriculus, where gastric juices are
produced to digest the food before it enters the stomach, and the gizzard, where the food is stored,
soaked, and mechanically ground. The undigested material forms food pellets that are sometimes
regurgitated. Most of the chemical digestion and absorption happens in the intestine and the waste is
excreted through the cloaca.

Ruminants are mainly herbivores like cows, sheep, and goats, whose entire diet consists of eating large
amounts of roughage or fiber. They have evolved digestive systems that help them digest vast amounts of
cellulose. An interesting feature of the ruminants’ mouth is that they do not have upper incisor teeth. They
use their lower teeth, tongue and lips to tear and chew their food. From the mouth, the food travels to the
esophagus and on to the stomach.

To help digest the large amount of plant material, the stomach of the ruminants is a multi-chambered
organ. The four compartments of the stomach are called the rumen, reticulum, omasum, and
abomasum. These chambers contain many microbes that break down cellulose and ferment ingested
food. The abomasum is the “true” stomach and is the equivalent of the monogastric stomach chamber
where gastric juices are secreted. The four-compartment gastric chamber provides larger space and the
microbial support necessary to digest plant material in ruminants. The fermentation process produces
large amounts of gas in the stomach chamber, which must be eliminated. As in other animals, the small
intestine plays an important role in nutrient absorption, and the large intestine helps in the elimination of
waste.

Pseudo-ruminants

Some animals, such as camels and alpacas, are pseudo-ruminants. They eat a lot of plant material and
roughage. Digesting plant material is not easy because plant cell walls contain the polymeric sugar
molecule cellulose. The digestive enzymes of these animals cannot break down cellulose, but
microorganisms present in the digestive system can. Therefore, the digestive system must be able to
handle large amounts of roughage and break down the cellulose. Pseudo-ruminants have a
three-chamber stomach in the digestive system. However, their cecum—a pouched organ at the
beginning of the large intestine containing many microorganisms that are necessary for the digestion of
plant materials—is large and is the site where the roughage is fermented and digested. These animals do
not have a rumen but have an omasum, abomasum, and reticulum.

Gut structure and function can also vary with the habitat and other physiological characteristics of a
species. The digestive tract of fish has adaptations for a marine or freshwater environment. The hindgut
of terrestrial vertebrates plays an important role in the conservation of the fluids, electrolytes, and nitrogen
in the digestive secretions, and in the urinary excretions of reptiles and birds. Fish, amphibians, and
reptiles are ectotherms, whose body temperature, metabolic rate, and nutritional requirements vary with
their ambient temperature. The endothermic birds and mammals require a higher and more continuous
input of nutrients for the maintenance of a constant body temperature.

The digestive tract may be described as a tube extending from the lips to the anus, constructed with such
dilations and constrictions throughout its length as to form a number of compartments, each of which has
a specific function in the digestive process. The contents of the tube, that is, the foods taken in, or
ingested, by the animal, are propelled by wavelike movements known as peristalsis. In addition, to
moving the food along the tract until it is eventually expelled, the movements serve to mix the food with
the digestive juices, to promote absorption of digested material, and to enhance the flow of blood and
lymph through the intestinal vessels.

The digestive process begins with the prehension, or taking up of food; continues with its mastication
(chewing), insalivation (mixing with the salivary secretions), deglutition (swallowing), and the digestion,
absorption, and assimilation of those food elements that are of nutritive value; and terminates with
defecation, or the excretion of the remaining ingested matter not capable of being absorbed or utilized as
nourishment. To perform these functions, the digestive tract of the animal is equipped with mouth, teeth,
tongue, pharynx, esophagus, stomach, intestine, and anus. Even among the various common
domestic species, differences exist in certain portions of the digestive tract to accommodate the special
requirements of the species.

Parts of the Digestive System

ORAL CAVITY

The oral cavity, or mouth, is the point of entry of food into the digestive system. The food consumed is
broken into smaller particles by mastication, the chewing action of the teeth. All mammals have teeth and
can chew their food.

Prehension or the taking up of food, is accomplished in most animals by means of the lips, teeth, and
tongue. In the horse, prehension is performed by the strong and flexible upper lip and incisor teeth. When
grazing, this animal employs lips and incisor teeth, cutting or tearing the vegetation seized by jerking
movements of the head or neck. The ox is not endowed with such mobile lips or with upper incisor teeth.
Instead it uses its long, muscular tongue to pull the grass or hay into its mouth, cutting it off between the
lower incisors and the upper gum by an upward movement of the head and neck. The sheep seizes its
food in much the same manner, but it makes use of its mobile lips rather than its tongue in gathering food
into the mouth. The native habit of the pig of burrowing or rooting to find its food has survived the
domestication of this species. Prehension, however, is accomplished by the movements of the pointed
lower lip and is assisted by the teeth and tongue. In the dog and the cat, the food is seized by the incisor
and canine teeth and brought into the mouth by jerking movements of the head and jaws. The organ of
prehension in fowls is the beak. The fowl picks up its food in its toothless beak and passes it to the base
of the tongue preparatory to swallowing.

Drinking is accomplished by the horse, ox, sheep, and pig by sucking the fluid into the mouth by the aid
of the tongue and pharynx. The lips form a small opening which is dipped beneath the surface of the
water, and the tongue moving back and forth like a piston in a cylinder creates a negative pressure in the
mouth which causes water to pass into the mouth with the backward thrust of the tongue and to be
swallowed with the forward thrust. The pharynx serves as a valve in this pumping process. The dog and
the cat take up liquids by lapping with the tongue. In the fowl, drinking consists merely scooping up the
fluid into the lower beak and then elevating the head to permit it to flow by gravitation into the crop or
stomach. The pigeon's method of drinking, however, more nearly resembles that of the horse or ox than
that of other birds. This bird thrusts its beak deep into the water and by a very rapid movement of the
tongue sucks in the liquid.
The extensive chemical process of digestion begins in the mouth. As food is being chewed, saliva,
produced by the salivary glands, mixes with the food. Saliva is a watery substance produced in the
mouths of many animals. There are three major glands that secrete saliva—the parotid, the
submandibular, and the sublingual. Saliva contains mucus that moistens food and buffers the pH of
the food. Saliva also contains immunoglobulins and lysozymes, which have antibacterial action to
reduce tooth decay by inhibiting growth of some bacteria. Saliva also contains an enzyme called salivary
amylase that begins the process of converting starches in the food into a disaccharide called maltose.
Another enzyme called lipase is produced by the cells in the tongue. Lipases are a class of enzymes that
can break down triglycerides. The lingual lipase begins the breakdown of fat components in the food. The
chewing and wetting action provided
by the teeth and saliva prepare the food into a mass called the bolus for swallowing. The tongue helps in
swallowing—moving the bolus from the mouth into the pharynx. The pharynx opens to two passageways:
the trachea, which leads to the lungs, and the esophagus, which leads to the stomach. The trachea has
an opening called the glottis, which is covered by a cartilaginous flap called the epiglottis. When
swallowing, the epiglottis closes the glottis and food passes into the esophagus and not the trachea. This
arrangement allows food to be kept out of the trachea.

Mastication

After the prehension of food, the next step, except of course in the toothless fowl, is mastication.
Mastication, or chewing, of food is accomplished by means of grinding teeth so situated in the upper and
lower jaws that they can be employed to crush and divide the food substance preparatory to swallowing.
The mechanism of mastication involves the use of the two jaws, the tongue, and the cheeks. The upper
jaw is a rigid portion of the head, while the lower jaw is hinged to the skull so that it can be moved in a
vertical and to some extent in a lateral or diagonal plane against the upper jaw. These movements are
brought about by the muscles of the jaws. There is a pronounced sidewise or lateral movement of the
lower jaw in the ruminants (ox and sheep), slightly less in the horse, and still less in the omnivorous pig.
In the carnivores (dog and cat), there is very little lateral movement.

The masticatory teeth, or molars, which are common to all species, are located laterally in both upper and
lower jaws. Food material brought between the opposing rows of teeth by the movements of the cheeks
and tongue are comminuted, or reduced to fine particles, by the crushing or grinding movements of upper
and lower molars as they are brought together during the chewing process. In the horse, ox, and sheep,
the grinding surface of the molars slopes sidewise, those of the upper jaw presenting an acute angle on
the outer side and those of the lower jaw being pointed on the inner side. In the pig, dog, and cat, the
grinding surface is more nearly at right angles to the long axis of the tooth.

Mastication in the horse, pig, dog, and cat is a single process preparatory to swallowing. The dog and cat
chew very little, usually bolting their food in relatively large pieces.

In the ruminants, represented by the ox and sheep, this process is divided into two phases: (1)
Preliminary or incomplete mastication when the food is first taken, and (2) complete mastication, which is
postponed for more leisurely performance.

Salivary Secretion

Mastication and the presence of food in the mouth cavity stimulate the secretion of saliva in the mouth in
all mammals. In man the principal constituent of this secretion is a substance known as ptyalin, or salivary
amylase, which functions in the digestion of starchy substances, but of the domestic animals only the pig
possesses enough ptyalin in its saliva to be of importance in digestion. The salivary secretion in most
domestic animals serves mainly to lubricate the food for swallowing. In ruminants the presence of saliva in
the paunch assists in the regurgitation of partially masticated food for re-chewing. In digestion by ptyalin or
salivary amylase, the starch is first changed to soluble starch and finally to maltose.

Deglutition / Swallowing
Deglutition is the act of swallowing the food that has been taken into the mouth. It is accomplished by a
series of muscular movements involving principally forcing the food to the rear of the mouth cavity by
elevation of the fore part of the tongue and depression of the root of the tongue, followed by the dilatation
of the pharynx. The bolus then passes through the pharynx and esophagus into the stomach. The initial
phase of the act of deglutition, that relating to its passage into the esophagus, is entirely a voluntary
process, but the completion of the act is involuntary.

The quantity of food material that can be swallowed at one time is comparatively small in the horse on
account of the narrow lumen, or space between the walls of the pharynx and the esophagus. A much
larger bolus can be swallowed by the ox, whose deglutitive organs are of ample proportions.
Nevertheless, ruminants frequently choke on apples, beets, turnips, and other pieces of food that have
not been chopped fine enough for ready swallowing with little chewing. There is no special or unusual
factor involved in the swallowing process on the part of the pig, dog, cat, or fowl.
Fluids are swallowed in much the same way as solids, except that they are passed into the esophagus
and stomach with extreme facility and rapidity as compared with solid boluses.

ESOPHAGUS

The esophagus is a tubular organ that connects the mouth to the stomach. The chewed and softened
food passes through the esophagus after being swallowed. The smooth muscles of the esophagus
undergo a series of wave like movements called peristalsis that push the food toward the stomach. The
peristalsis wave is unidirectional—it moves food from the mouth to the stomach, and reverse movement is
not possible. The peristaltic movement of the esophagus is an involuntary reflex; it takes place in
response to the act of swallowing.

A ring-like muscle called a sphincter forms valves in the digestive system. The gastro-esophageal
sphincter is located at the stomach end of the esophagus. In response to swallowing and the pressure
exerted by the bolus of food, this sphincter opens, and the bolus enters the stomach. When there is no
swallowing action, this sphincter is shut and prevents the contents of the stomach from traveling up the
esophagus. Many animals have a true sphincter; however, in humans, there is no true sphincter, but the
esophagus remains closed when there is no swallowing action. Acid reflux or “heartburn” occurs when the
acidic digestive juices escape into the esophagus.

STOMACH

A large part of digestion occurs in the stomach. The stomach is a saclike organ that secretes gastric
digestive juices. The pH in the stomach is between 1.5 and 2.5. This highly acidic environment is required
for the chemical breakdown of food and the extraction of nutrients. When empty, the stomach is a rather
small organ; however, it can expand to up to 20 times its resting size when filled with food. This
characteristic is particularly useful for animals that need to eat when food is available.

The ruminant is unique among animals in being equipped with a partitioned stomach consisting of four
compartments. These are known as the rumen, paunch, or first stomach; the reticulum, honeycomb, or
second stomach; the omasum, many plies, or third stomach; and the abomasum or rennet, the true or
fourth stomach. The food material, which is usually of a coarse nature, after being partly chewed enters
the paunch or rumen, from which it is later regurgitated or returned to the mouth and re-chewed in
individual boluses during repose. This process of re-chewing is known as rumination. Each bolus of food
brought up for rumination is chewed for somewhat less than a minute. Grains form a part of the bolus only
to the limited extent that they are accidentally caught in the roughage. Consequently, all un-chewed grain
passes through the animal and appears in the feces in an intact condition. In cattle a certain quantity of
whole grain is not crushed during eating and rumination, and on this account grain is commonly ground
for cattle. This is rarely done for sheep, which chew most grains very thoroughly. In ruminants the mucous
membrane lining the cavity of the mouth is thicker than that of the horse and is furnished with many horny
protuberances, known as papillae, which facilitate the manipulation of food during mastication.

The ruminant digestive system is found in cattle, sheep, goats, and deer.
Digestion

The stomach is the major site for protein digestion in animals other than ruminants. Protein digestion is
mediated by an enzyme called pepsin in the stomach chamber. Pepsin is secreted by the chief cells in the
stomach in an inactive form called pepsinogen. Pepsin breaks peptide bonds and cleaves proteins into
smaller polypeptides; it also helps activate more pepsinogen, starting a positive feedback mechanism that
generates more pepsin. Another cell type—parietal cells—secretes hydrogen and chloride ions, which
combine in the lumen to form hydrochloric acid, the primary acidic component of the stomach juices.
Hydrochloric acid helps to convert the inactive pepsinogen to pepsin. The highly acidic environment also
kills many microorganisms in the food and, combined with the action of the enzyme pepsin, results in the
hydrolysis of protein in the food. Chemical digestion is facilitated by the churning action of the stomach.
Contraction and relaxation of smooth muscles mixes the stomach contents about every 20 minutes. The
partially digested food and gastric juice mixture is called chyme. Chyme passes from the stomach to the
small intestine. Further protein digestion takes place in the small intestine. Gastric emptying occurs within
two to six hours after a meal. Only a small amount of chyme is released into the small intestine at a time.
The movement of chyme from the stomach into the small intestine is regulated by the pyloric sphincter.

The action of bacteria upon food taken into the stomach of the horse, pig, or dog is short-lived owing to the
rapid production of hydrochloric acid there. In the ox and other ruminants, however, considerable bacterial
action is exerted upon the enormous amount of material taken into the capacious rumen prior to its
regurgitation, re-mastication, and digestion. This bacterial action is favored by the slightly alkaline reaction
of that portion of the stomach and its freedom from gastric secretions, as well as by the prolonged sojourn
of food matter therein. The changes brought about by bacteria inhabiting the alimentary tract include the
breaking up of cellulose and fats, the production of organic acids such as acetic, butyric, and lactic, and
the splitting of the carbohydrates known as polysaccharides. The gases, methane, carbon dioxide,
hydrogen, etc., are formed during bacterial action in the rumen, but they are of no value to the animal and
are excreted as waste.

When digesting protein and some fats, the stomach lining must be protected from getting digested by
pepsin. There are two points to consider when describing how the stomach lining is protected. First, as
previously mentioned, the enzyme pepsin is synthesized in the inactive form. This protects the chief cells,
because pepsinogen does not have the same enzyme functionality of pepsin. Second, the stomach has a
thick mucus lining that protects the underlying tissue from the action of the digestive juices. When this
mucus lining is ruptured, ulcers can form in the stomach. Ulcers are open wounds in or on an organ
caused by bacteria (Helicobacter pylori) when the mucus lining is ruptured and fails to reform.

A pseudo-ruminant is an animal that eats large amounts of roughage but does not have a stomach with
several compartments. The digestive system does some of the same functions as those of ruminants. For
example, in the horse, the cecum ferments forages. An animal with a pseudo-ruminant digestive system
can utilize large amounts of roughages because of the greatly enlarged cecum and large intestine, which
provide areas for microbial digestion of fiber. Pseudo-ruminants often eat forages as well as grains and
other concentrated feeds. Besides horses, examples of pseudo-ruminants are rabbits, guinea pigs, and
hamsters. The stomach of the horse is comparatively small.

The true stomach of the ox or sheep is also comparatively small and receives food gradually in a well
macerated or ground-up condition from the first three stomachs. The stomach of the pig, dog, or cat is
sufficient in size to contain all the food that may be consumed at a meal, and food may be retained in the
stomach for a considerable time. In all mammals the food is mixed and softened in the stomach by
muscular movements, assisted by the action of the gastric juice. The softer portions of the food material
thus prepared are then passed into the intestine, but food not ready for intestinal digestion is not allowed
to pass from the stomach, with the exception of whole grain in cattle.

Fowl, having no teeth, must swallow their food whole or at best only coarsely broken by the beak at
prehension. Such material passes through the pharynx and esophagus into the crop, which in most avian
species is nothing more than a dilated portion of the esophagus. The purpose of the crop of the fowl, like
that of the rumen of the ox and sheep, is largely bulk storage of food. This material then passes by slower
stages into the proventriculus, or glandular stomach, a specialized gastric organ of small capacity whose
wall is furnished with secretory glands supplying pepsin and hydrochloric acid. The food material after
coming into contact with these digestive fluids continues on its way into the gizzard, or muscular stomach.
That organ is lined with a tough membrane and is so constituted that it retains a certain portion of the grit
normally swallowed by the fowl. The food substance is somewhat softened by the time it reaches the
gizzard, where it is subjected to crushing contractile movements stimulated by the presence of food. The
food substance is reduced to a very fine consistency by the rubbing together of the grit and the tough,
corrugated fining of the organ. This process corresponds to mastication. Strictly speaking, there is no true
gastric digestion in the fowl, such as is common to mammals. The food material after being finely divided
in the gizzard is passed on into the first portion of the small intestine, where it is more thoroughly mixed
with and digested by the juices derived from the proventriculus as well as the other digestive enzymes
emptying into the intestine. The alimentary tract of the chicken is for the most part representative of those
of the majority of species of domesticated fowl, but there are some minor differences.
The pigeon is peculiar in not possessing a gall bladder, and in having ceca that are small and poorly
developed as compared with those of the duck and chicken. Neither the duck nor the pigeon possesses a
true crop comparable to that of the chicken, but in these species there is a specialized enlargement of the
esophagus which serves the purposes of the crop. The small intestine of the pigeon is shorter in
proportion to the size of the bird than that of the duck or the chicken.

In the horse, ox, sheep, and pig, well-ground and softened food in the stomach undergoes gastric
digestion through the action of the pepsin and hydrochloric acid of the gastric juice. In the dog and cat, in
which mastication has been eliminated to a large extent, the food in the stomach is subjected to the same
process as in the other animals but for a longer period. Digestion by the gastric juice is essentially the
same in all animals. The pepsin of the gastric juice in the presence of hydrochloric acid acts upon proteins
of either animal or vegetable origin and converts native protein into the simpler products, proteoses and
peptone. The latter substances are not yet ready for absorption but require further conversion into amino
acids. This step in protein digestion remains to be completed in the intestine.

Another digestive enzyme sometimes known as rennin is present in the gastric juice of young mammals.
This enzyme coagulates milk so that the casein is prevented from too rapid passage through the stomach
and the pepsin is enabled to convert it into proteoses and peptone. In all species, following gastric
digestion the semiliquid substance is passed into the small intestine; at this point it is known as chyme.
Digestion by
pepsin occurs largely in the stomach, but it is continued to some extent in the first part of the small
intestine. SMALL INTESTINE

Chyme moves from the stomach to the small intestine. The small intestine is the organ where the
digestion of protein, fats, and carbohydrates is completed. The small intestine is a long tube-like organ
with a highly folded surface containing finger-like projections called the villi. The apical surface of each
villus has many microscopic projections called microvilli. These structures are lined with epithelial cells on
the luminal side and allow for the nutrients to be absorbed from the digested food and absorbed into the
blood stream on the other side. The villi and microvilli, with their many folds, increase the surface area
of the intestine and increase absorption efficiency of the nutrients. Absorbed nutrients in the blood are
carried into the hepatic portal vein, which leads to the liver. There, the liver regulates the distribution of
nutrients to the rest of the body and removes toxic substances, including drugs, alcohol, and some
pathogens.

In the intestines the chyme undergoes further digestion by the action of pancreatic juice, intestinal juice,
and bile. The pancreatic juice contains three important enzymes—trypsin, steapsin, and amylopsin. To
a limited extent trypsin, like pepsin, converts native protein into proteoses and peptone. Principally,
however, it continues the digestion of protein where the pepsin leaves off, that is, breaks the proteoses
and peptones into the amino acids, which can be absorbed. The steapsin breaks down or hydrolyzes
fats into fatty acids and glycerol, in which forms they are ready for absorption. Some of the fatty acids
resulting from steapsin digestion combine with the available alkali in the intestinal juice to form soaps
which assist in emulsifying any remaining undigested fat. The amylopsin converts or hydrolyzes starch
and the dextrines into maltose. Maltose requires further breaking up by other digestive juice before it is
ready for absorption.

The intestinal juice contains a number of digestive enzymes, for example, enterokinase, erepsin,
maltase, sucrase, lactase, lipase, amylase, and nucleinase. Enterokinase is largely concerned in the
activation of the enzyme trypsin of the pancreatic juice. Erepsin, which is a proteolytic enzyme—that is,
one that has the power to break down proteins into simpler diffusible substances—converts peptones and
peptids into amino acids. Maltase converts the maltose resulting from the action of amylopsin on starch
into glucose, in which form it can be absorbed. Sucrase breaks down or hydrolyzes sucrose into glucose
and fructose, in which forms they are assimilated. Lactase, which is present only in young mammals
ingesting milk, converts lactose (milk sugar) into glucose and galactose. This enzyme is lacking in
mature animals that do not consume milk as food. However, it is reported that lactase can be made to
recur in adult animals fed milk or lactose. Lipase is the same as steapsin and converts fats into fatty
acids and glycerol. Amylase hydrolyzes starch. Nucleinase converts nucleoproteins into protein and
nucleic acid.

The human small intestine is over 6m long and is divided into three parts: the duodenum, the jejunum,
and the ileum. The “C-shaped,” fixed part of the small intestine is called the duodenum. The duodenum is
separated from the stomach by the pyloric sphincter which opens to allow chyme to move from the
stomach to the duodenum. In the duodenum, chyme is mixed with pancreatic juices in an alkaline solution
rich in bicarbonate that neutralizes the acidity of chyme and acts as a buffer. Pancreatic juices also contain
several digestive enzymes. Digestive juices from the pancreas, liver, and gallbladder, as well as from
gland cells of the intestinal wall itself, enter the duodenum. Bile is produced in the liver and stored and
concentrated in the gallbladder. Bile contains bile salts which emulsify lipids while the pancreas produces
enzymes that catabolize starches, disaccharides, proteins, and fats. These digestive juices break down
the food particles in the chyme into glucose, triglycerides, and amino acids. Some chemical digestion
of food takes place in the duodenum. Absorption of fatty acids also takes place in the duodenum.

The second part of the small intestine is called the jejunum. Here, hydrolysis of nutrients is continued
while most of the carbohydrates and amino acids are absorbed through the intestinal lining. The bulk of
chemical digestion and nutrient absorption occurs in the jejunum.

The ileum is the last part of the small intestine and here the bile salts and vitamins are absorbed into
blood stream. The undigested food is sent to the colon from the ileum via peristaltic movements of the
muscle. The ileum ends and the large intestine begins at the ileocecal valve. The vermiform, “worm-like,”
appendix is located at the ileocecal valve. The appendix of humans secretes no enzymes and has an
insignificant role in immunity.

Absorption

The digestive processes by which carbohydrates and fats are broken down into soluble or emulsified
forms ready for absorption have been described. The inorganic salts, water, and vitamins contained in
food require no breaking down but are absorbed as such. The end products of protein digestion are
amino acids. Carbohydrates are broken into simple sugars (monosaccharides), principally glucose.
The end products of cellulose digestion are probably the simple fatty acids (acetic and butyric) and
glucose. The fatsare broken down into fatty acids and glycerol. The absorption of all of these food
elements occurs chiefly in the small intestine, though some absorption occurs in the large intestine also,
particularly in the horse.

The principal avenues of absorption of food are the myriads of microscopic protuberances from the
mucosa, or lining membrane of the intestinal wall, known as villi. These minute, fingerlike structures are
composed of an outer layer of absorbent cells, a wall containing many blood vessels, and in the center a
lacteal or lymphatic vessel. Being in constant contact with the food in a digested state, the villi take up the
digested fats and pass them through the lymph channels (commonly known as lacteals) into the blood
stream. The lymph within the lacteals is a milky substance heavily laden with the digested fat and is
known as chyle. The end products of protein and carbohydrate digestion, inorganic salts, water, and
vitamins enter the blood vessels of the villi directly and are transported through the liver into the blood
system of the body. These absorbed food substances are carried to the body tissues, where after having
undergone complex metabolic processes they are utilized for growth, repair, and energy.

LARGE INTESTINE

The large intestine reabsorbs the water from the undigested food material and processes the waste
material. The human large intestine is much smaller in length compared to the small intestine but larger in
diameter. It has three parts: the cecum, the colon, and the rectum. The cecum joins the ileum to the
colon and is the receiving pouch for the waste matter. The colon is home to many bacteria or “intestinal
flora” that aid in the digestive processes. The colon can be divided into four regions, the ascending
colon, the transverse colon, the descending colon and the sigmoid colon. The main functions of the
colon are to extract the water and mineral salts from undigested food, and to store waste material.
Carnivorous mammals have a shorter large intestine compared to herbivorous mammals due to their diet.

Rectum and Anus

The rectum is the terminal end of the large intestine. The primary role of the rectum is to store the feces
until defecation. The feces are propelled using peristaltic movements during elimination. The anus is an
opening at the far-end of the digestive tract and is the exit point for the waste material. Two sphincters
between the rectum and anus control elimination: the inner sphincter is involuntary and the outer
sphincter is voluntary.

Accessory Organs

The organs discussed above are the organs of the digestive tract through which food passes. Accessory
organs are organs that add secretions (enzymes) that catabolize food into nutrients. Accessory organs
include salivary glands, the liver, the pancreas, and the gallbladder. The liver, pancreas, and
gallbladder are regulated by hormones in response to the food consumed.

The liver is the largest internal organ in humans and it plays a very important role in digestion of fats and
detoxifying blood. The liver produces bile, a digestive juice that is required for the breakdown of fatty
components of the food in the duodenum. The liver also processes the vitamins and fats and synthesizes
many plasma proteins.

The principal constituents of bile are bile pigments, bile salts, and cholesterol. The bile salts only are
concerned in the digestive process. They aid digestion by activating the pancreatic lipase, enhancing the
action of pancreatic amylase, emulsifying fats, and increasing the solubility of fatty acids and their soaps
so that they may be ready for absorption. Bile also is necessary for the absorption of the fat-soluble
vitamins. Bile being a reservoir of alkali helps to maintain the alkaline reaction so necessary for pancreatic
intestinal digestion.

The pancreas is another important gland that secretes digestive juices. The chyme produced from the
stomach is highly acidic in nature; the pancreatic juices contain high levels of bicarbonate, an alkali that
neutralizes the acidic chyme. Additionally, the pancreatic juices contain a large variety of enzymes that are
required for the digestion of protein and carbohydrates.

The gallbladder is a small organ that aids the liver by storing bile and concentrating bile salts. When
chyme containing fatty acids enters the duodenum, the bile is secreted from the gallbladder into the
duodenum.

In addition to the enzymes secreted by the various organs, there are, particularly in the digestive tracts of
herbivores and omnivores, bacteria which break down cellulose into substances that can be absorbed
and used by the animal organism. This bacterial activity is of considerable importance, since the
vegetable substance consumed in large quantities by the horse, ox, sheep, and pig contains a large
proportion of cellulose, which is not digested by the enzymes in the various digestive juices. This bacterial
digestion makes available for utilization by the body large quantities of otherwise indigestible food
material.