INTRODUCTION

Digestion includes all the activities of the alimentary tract and its associated glands in the preparation of the food for
absorption and in rejection of the residue.
• Most foods are consumed in a highly complex and insoluble form that they cannot be absorbed into the blood or
lymph without preliminary digestive changes. The majority of foods must undergo many changes before absorption.
Food
• Food is a mixture of substances included to meet the nutritive requirements of an animal.
• Food should contain the following substances: protein,carbohydrates, fats vitamins, inorganic salts and water.
• A single food may not contain all the ingredients in the right quality and quantity. Hence, it is necessary for an animal
to have a variety of foods to remain healthy.
• The higher animals are dependent on plants or other animals for their nutritive requirements.
Carnivorous animals (dog, cat, etc.) obtain bulk of their energy and nutrient requirements from animal sources. Most
carnivores are predators or scavengers. Their diet has high energy concentration and is easily digested. They have a
short and simple digestive tract. Stomach is spaciousand store large amount of food.
• Herbivorous animals (cattle, sheep, horse, goat, etc.) derive their food from vegetable sources. Their diet contains
less fat and low energy concentration. Energy in the plants is available as carbohydrates that cannot be broken
down by digestive enzymes of the host but can be broken down by microbial enzymes and the breakdown products
are utilized by the host animal. Hence, their digestive tract contains a voluminous extension to accommodate the
microbes. Microbial breakdown of diet in the absence of O2 is called fermentation; fermentative degradation is a
slow process and food must be retained in the fermentation place for longer duration. In ruminants the microbial
fermentation occurs in the forestomach and in monogastric herbivores like horses, pigs and rodents this occurs at
the large intestine.
Ruminants and mono-gastric herbivores can be divided into three functional groups based on their nutrient intake
Browsers – (animals that eat leaf and buds of plants) Animals of this group eat leaves, fruits, seeds, buds; these animals
are selective eaters and prefer easily digestible parts of the plants. E.g. rabbits, all the smallest ruminants, deer and
giraffes
Grazers – (animals that eat grass and fibrous plants) These animals eat large amounts of fibre-rich food of low
digestibility; they have voluminous fermentation chamber e.g. horses, cattle, sheep, elephants
Intermediate feeders – Animals of this group fall between the other two groups; they eat variety of plants according to
seasonal availability. E.g. goats, camels, reindeer
Omnivorous animals (man, pig, etc) take the foods of both animal and plant [Link] in this group eat fruits,
roots and other plant parts as well as food of animal origin.
Alimentary Tract (Digestive Tube)
• It extends from the lips to anus and includes mouth, pharynx, oesophagus, stomach, small and large intestine. The
length and complexity of the tract vary in different species.
• In carnivores, digestive tract is relatively short and simple.
• In some herbivores, (horse, rabbit) the stomach is relatively simple whereas the large intestine is much more
complex and voluminous which help in microbial fermentation of food material and these animals are known as
hind-gut (post-gastric) digesters.
• In certain herbivores, (ruminants) the stomach is extensively large and complex to help in microbial fermentation,
whereas the large intestine is relatively small. These animals are known as fore-gut (pregastric) digesters.

1
PREHENSION OF FOOD
• Prehension means seizing and conveying of food into the mouth.
• Animals fetch their feed and take it to their mouth using lips, teeth and tongue with the help of head and jaw
movements
Dogs and Cats: Food is held with the forelimbs but it is passed into the mouth by the head and jaw movements
Horse: The upper lip is the main prehensile organ and it is strong, sensitive and mobile. Food is collected by the
upper lip with the aid of the tongue. While grazing, the horse draws the lips back and uses the incisor teeth to
severe the grass.
Cattle: The tongue is large, strong, mobile and rough; it is capable of being protruded from the mouth. The tongue can
be readily curved around the forage, by an upward movement of the head, the grass is cut off.
Sheep: has clefted upper lip which permits close grazing. The incisor teeth and tongue are the principal prehensile
organs, but the tongue is not protruded out in grazing as in cattle.
Goat: Same as in sheep but the upper lip has no cleft.
Pigs: Under natural conditions, it digs up the ground with the snout and carries the food so obtained to the mouth
by the action of the pointed lower lip.
Drinking
• The way in which liquids are carried to the mouth varies greatly in carnivores and in herbivores.
• The dog and cat make a ladle of the free end of the tongue by which the liquid is carried to the mouth.
• Other domestic animals draw liquid into the mouth by suction. The lips are closed all around except for a small place
in front, which is under the water. The tongue moves backward to create a negative pressure in the mouth which
draws the water into the mouth.
• Birds fill the oral cavity with waterand then raises the beak and water flows into pharynx
• Sucklingis accomplished by the creation of negative pressure in the mouth largely by the action of the tongue. Milk
in the teat is in higher pressure than the atmospheric pressure; hence it is forced into the mouth because of the
action of the tongue where the pressure is lower than the atmospheric pressure.
MASTICATION (Chewing)
• It is the mechanical reduction of food that takes place in the mouth.
• The grinding occurs between the molar teeth.

2
• Importance of mastication
➢ Grinding the food reduces particle size; finely divided food present a greater surface than the coarse food for
the action of the digestive juices.
➢ Food well mixed with the saliva is more easily swallowed.
• In carnivores, mastication is imperfectly performed whereas in herbivores due to the coarse bulky nature of food,
mastication is of much greater significance.
• In carnivores, vertical movement of the lower jaw is important for mastication.
• In herbivores, lateral movement of the lower jaw and some to-and-fro movement are important, In ruminants, it is
only during remastication that the food is thoroughly ground.
• Birds do not have teeth
DEGLUTITION (Swallowing)
• Deglutition refers to the passage of food from the mouth through pharynx and oesophagus to the stomach.
• It commences as a voluntary act but becomes an involuntary reflex during its execution. It involves many muscles
and their motor nerves.

Act of Swallowing
It is divided into 3 phases
(1) From mouth to pharynx (voluntary act)
(2) Through pharynx into the oesophagus (reflex action)
(3) Down the oesophagus into the stomach (reflex).
• Once swallowing is initiated, it cannot be stopped. During swallowing, respiration is stopped momentarily.
• The muscular force necessary to drive the food bolus through the mouth and pharynx up to the oesophagus is
derived chiefly from the contraction of 20 to 30 muscles involved in the swallowing process. The muscles press the
tongue against the hard palate and the root of the tongue is drawn backwards. At the same moment, elevation of
the soft palate cuts off communication with nasal passage. The upper oesophageal sphincter relaxes (normally
closed). The tongue acts like a plunger driving the mass of food towards the oesophagus. The hyoid bone and the
larynx are now pulled forward and the entrance to the oesophagus is opened. The epiglottis closes the larynx shut.
The bolus of food enters into the oesophagus. The upper end of the oesophagus undergoes receptive relaxation.
• From the oesophagus, the food moves down to stomach by peristaltic activity of oesophagus.
Nervous Control of Deglutition
• Swallowing is a reflex act.
3
• The afferent impulses are carried by glossopharyngeal, vagus and trigeminal nerves to deglutition centre
(swallowing centre) located in the medulla of the brainstem
OESOPHAGUS
• The oesophagus extends from the pharynx to the stomach, crossing the thorax and perforating the diaphragm.
• It is made up of four layers of tissues with the muscle layer consisting of circular and longitudinal layers
• The oesophagus is normally closed at the pharyngeo-esophageal junction by an esophageal sphincter and the
opening of the oesophagus into the stomach is also normally kept closed.
• Cardia: The opening of esophagus into the stomach is called cardia.
• A sphincter of smooth muscle known as cardiac sphincter closes the cardia. The cardiac sphincter is closed except
during swallowing
STOMACH
Functions of stomach
• The stomach can be divided into two physiological regions:
➢ The proximal region at the oesophageal end serves storage function.
➢ The distal region serves grinding and sieving function.
• For the purpose of study of motility, the stomach may be considered to include three parts:
o fundus (proximal stomach)- the muscular activity is weak continuous contraction in this region and this
region is capable of adaptive relaxation as food enters the stomach; so the stomach can accept large
amounts of food without increase in intraluminal pressure. Thus, the fundus serves as a food storage area
and not much of mixing of food occur.

o The body or corpus (mid region) -of the stomach serves as a mixing vat for mixing the gastric juice with
food.
o Antrum- is the distal stomach: it shows intense slow wave potential change and muscular contractions. It
acts as a gastric pump and regulates the propulsion of food through the pyloric sphincter into the duodenum.
The antral contractions also serve to retropel the pyloric contents and thus delay the passage of solid
particles from the stomach.
• The stomach movements are peristaltic waves. The cells of Cajal that lie between the longitudinal and circular
muscles act as pacemaker cells and initiate muscular contractions of stomach. Reverse peristaltic waves are also
noticed in the stomach
Regulation of Stomach Movements
• The stomach muscles possess a high degree of automaticity.
• Although gastric muscles possess inherent rhythmicity (due to cells of Cajal), its movements are regulated by the
vagi and sympathetic nerves and hormones.
• Stimulation of vagus increases peristaltic activity of the stomach and suppresses muscular activity in proximal
stomach. Anticipation of eating increases vagal activity.
• Sympathetic nerves inhibit peristalsis.

4
• Gastrin increases gastric motility. Secretin and CCK suppresses motility.
Control of gastric emptying
• The rate at which food leaves the stomach must match the rate at which it can be digested and absorbed by the
small intestine.
• There are reflexes that regulate the gastric emptying.
The duodenal factors that inhibit gastric emptying are
1. low pH,
2. high osmolality,
3. increased intraluminal pressure in duodenum
4. high peptide and fat concentration
• Osmotic pressure and acidity of duodenal contents are detected by appropriate duodenal receptors and they alter
the vagal activity. Increase in osmotic pressure and decrease in acidity depresses gastric motility.
• The endocrine system controlling the gastric motility include
➢ secretin, produced by low duodenal pH;
➢ CCK released in response to fat in duodenal contents;
➢ GIP is released in response to carbohydrates;
• Low fat meal leaves the stomach in 3 to 4 h. The strongest stimulus inhibiting gastric motility through hormone
release is fat in duodenum. High fat content in meals delay the gastric emptying
VOMITING (Emesis)
• Vomiting is the forceful ejection of the contents of the stomach through esophagus and mouth.
• Vomiting is common in carnivores and omnivores.
• Herbivores or rodents seldom or never vomit. However, the basic mechanism of vomiting appears to be present
in all species.
• The uncommonness of vomiting in horse may be due to the highly closed cardia, and constricted terminal part of
the oesophagus. The esophagus enters the stomach at a sharp angle and when stomach expands, it closes the
opening. This anatomical arrangement of stomach in horse makes regurgitation extremely rare in these animals. In
other non-vomiters, their inability to vomit may be due to the absence or rudimentary development of vomiting
centre.
• Vomiting is a reflex act , Vomiting centre is present in the reticular formation of the medulla.
• Drugs which induce vomiting are known as emetics (morphine, CuSO 4)which may act through receptors in the
gastro-intestinal tract or may act on the chemoreceptor trigger zonepresent in the 4th ventricle of brain from which
impulses pass to vomiting centre to cause vomiting.
Act of Vomiting
• After a deep involuntary inspiration, the glottis is closed and nasopharynx is closed by elevation of soft palate. In
the succeeding expiratory movement with closed glottis, the intra-thoracic pressure increases and is accompanied
by contraction of abdominal muscles and diaphragm. Immediately following these events the oesophagus, gastro-
oesophageal junction and the body of stomach relax and the pylorus contracts firmly. Assisted by the pressure
exerted by contraction of abdominal muscles and the diaphragm, the stomach contents enter the oesophagus where
they are further propelled to and out of the mouth by raised intra-thoracic pressure.
• The stomach plays a passive role during the actual act of vomiting. There may be massive antiperistalsis of the
upper small intestine with reflux into the stomach.
• The significance of vomiting is to assist the animal in removal poisons taken in with food
• Vomiting leads to loss of fluids from the stomach and duodenum. The loss of H + ion from stomach can cause
metabolic acidosis. Loss of fluids can cause drop in blood pressure and other circulatoryproblems.
INTESTINALMOVEMENTS
The functions of the gastro intestinal movements are
• To mix the ingesta with digestive secretions
• To bring the digested products into contact with intestinal mucous membrane for absorption

5
 • To move the food masses from place to place in the intestine
• To expel the residue from the rectum through anus
• To assist in flow of blood and lymph through vessels of intestinal wall
• The gastrointestinal system is regulated at two levels. One level is control by the CNS and endocrine system.
• Second level of control is exerted by intrinsic nerves and endocrines located within the gut. This intrinsic control
system regulates the gut function automatically based on local stimuli like amount and type of food etc.
Gastrointestinal Smooth Muscles
• Intestinal wall is made up of four layers
• From lumen to outward they are mucosa(made up of epithelial cells consisting of absorptive cells, goblet cells,
endocrine cells and undifferentiated cells), submucosa(contains nerve fibres, blood and lymph vessels), muscular
and serosa layers.
• The muscular layer consists of two layers of smooth muscles - outer longitudinal and inner circular layers.
• These two layers are surrounded by a specialised type of smooth
muscle cells known as interstitial cells of Cajal. These ICC cells
exhibit rhythmic, because these cells are interconnected and also
connected with general smooth muscle cells by tight junctions the
electrical activity is propagated over large areas of smooth
[Link] arrangement permits the GI muscles to function as
syncytium (whole unit functioning as a single cell). The ICC thus
acts as the pace maker for GI slow waves of GI tract.
• The smooth muscles of the GI tract show conduction of impulse
from fibre to fibre and sensitivity to stretch.
• The smooth muscle cells of the gut have a resting membrane
potential of –55 to -60mV (inside negative to outside).
• This potential undergo changes in two ways: Slow waves and
Spikes
• Duodenum produces more frequent spontaneous, rhythmic
changes in membrane potential.
• In the small intestine of dogs, slow waves occur about 20 times /min.
• In stomach and colon they are less frequent – about 5/min.
• The nervous system cannot cause contraction of all the GI muscle cells in one segment to contract simultaneously,
because individual muscle cells do not receive nerve supply as in skeletal muscles. Hence, control of smooth muscle
contraction is achieved by the combined effect of intrinsic nerves and slow waves.
NEURAL REGUALTION OF THE INTESTINE
Enteric Nervous System (ENS)
• The enteric nervous system (ENS) is extensive and consists of
receptors and cell bodies, all of which lie on the gut [Link]
ENS includes sensory and motor neurons and it is connected
to the CNS through autonomic nerves but can also function
autonomously without ANS.
• The cell bodies of the ENS are arranged in two systems of
ganglia:
(1) The myenteric (Auerbach’s) plexus
(2) The submucosal (Meissner’s) plexus.
• The myenteric plexus consists of ganglia located between the
circular and longitudinal muscle layers.
• The submucosal plexus have their ganglia in the submucosal
layer.

6
• Neurons arising from myenteric plexus innervate the muscle layers of the intestine and are primarily motor in
function. The submucosal plexus supplies intestinal glands, endocrine cells, submucosal blood vessels and is
involved in the control of intestinal secretion.
• Sensory neurons arise from mechanoreceptors within the muscular layer and chemoreceptors within mucosa.
• Mechanoreceptors monitor distension of gut wall and chemorecptors monitor chemical conditions in the lumen of
gut.
• There are two types of efferent neurons in the ENS: stimulatory and inhibitory.
• Many of the stimulatory neurons are cholinergic with acetylcholine as neuroregulatory transmitter. Other
stimulatory transmitter known is substance-P.
• The inhibitory neurons produce peptides as neuroregulatory transmitters which include VIP, somatostatin and
nonpeptide regulators like NO, and ATP.

Extrinsic Nerves to Gut

• Parasympathetic and sympathetic nervous systems containing both sensory and motor components form a link
between the ENS and CNS.
• Parasympathetic nerves generally increase the activity of intestinal smooth muscles i.e. increases motility. Vagus
is the main parasympathetic nerve supplying most of the gut except colon which receive parasympathetic
innervation through pelvic nerve.
• Sympathetic nerves generally decrease the activity of intestinal smooth muscles (decreases motility) and produces
contraction of sphincters.
• The sympathetic fibres act through release of regulatory substance norepinephrine and some fibres act by releasing
somatostatin and neuropeptide Y (NPY).
ENDOCRINEREGULATION OF THE GUT
• The GI tract has variety of enteroendocrinecells, which are distributed diffusely throughout the gut. These cells
are located among other mucosal cells in both secretory and absorptive areas of the mucosa. The apex of these
cells is exposed to the lumen of the intestine where they can sense the lumen contents.
• The endocrine cells respond to changes in the lumen contents by releasing hormones into the submucosa; from
here they are absorbed into blood stream for producing effect in some other parts or diffuse through extracellular
fluid to have local effect (paracrineor autocrine activity).
• All the GI endocrine products are peptides and referred to as regulatory peptides. These regulatory peptides
influence various gut functions.
• A subgroup of GI endocrine cells called enterochromaffin cells function as paracrine cells. They have similar
structure to endocrine cells and secrete a regulatory substance serotonin (5-hydroxy tryptamine).
Major Gastrointestinal Hormones:
• On the basis of structure and function, GI hormones can be grouped into threefamilies –
(1) gastrin family that includes gastrin and CCK
(2) secretin family that consists of secretin, glucagon, VIP, GIP etc.
(3) The third group includeother gut hormones that do not belong to either of these two groups.

Hormone Production Functions Stimulus Releasing

Gastrin Distal Stomach Stimulates Acid secretion from stomach Protein in stomach, high pH, vagal stimulation
glands
Secretin Duodenum Primarily stimulates HCO3 secretion from pancreas;
Acid in duodenum
also stimulates bile HCO3 secretion
CCK Duodenum to Primarily stimulates enzyme secretion from Proteins and fats in small intestine
ileum; highest pancreas; stimulates gall bladder
in duodenum contraction; inhibits gastric emptying
GIP Duodenum and Mainly inhibits gastric motility and gastric Carbohydrates and fats in small intestine
upper jejunum emptying and secretory activity; stimulates
insulin secretion

7
 Regulates motility pattern of gut during inter Acetyl choline; fasting; cyclic release every 1.5-
Motilin Duodenum and
upper jejunum meal period – regulates MMC 2h by neural stimuli

Other Regulatory Peptides

• In addition to the gut endocrine peptides, neuroregulatory peptides produced by the intrinsic nerves also have effects
similar to endocrine and paracrine substances; (e.g.) somatostatin, substance-P, VIP, neurotensin, opiods etc.
Movements of Small Intestine
• Motility of the small intestine occurs in two phases:
➢ One phase occur during digestive period following food intake and
➢ Second phase occur during interdigestive period, when less food is present in gut.

• In the digestive phase two patterns of motility occurs: propulsive and nonpropulsive.
o The propulsive pattern is referred to as peristalsis
o The non-propulsive pattern is segmentation.
Movement during Digestive Phase
Rhythmic Segmentation
• It is produced by contraction of circular muscles.
• During segmentation, a mass of food lying in a length of intestine (3-4cm long) is divided into smaller ovoid pieces
by constrictions caused by circular muscles. Within few seconds, the constricted portions relax and new areas
constrict.
• Segmentation may be taking place in many different areas of small intestine at the same time. The amplitude of
segmentation contractions varies and is strong after feeding.
• Segmentation occurs in dogs 12-18 times a minute in small intestine.
• The effect of segmentation is to mix the food material with digestive secretions and to expose the mixture to the
absorptive mucosa. Slight onward movement of ingesta also occurs during segmentation.
Peristalsis
• The main mechanism for the onward movement of semisolid intestinal content is peristalsis.
• It is achieved by creation of a moving ring of constriction produced by contraction of circular muscle, which pushes
the bowel contents to succeeding areas of relaxation produced by contraction of longitudinal muscles.
• A stimulus at any point in the intestine can cause contraction above and distension below. The wave of contraction
and relaxation moves along the intestine as a peristaltic wave, which carries the ingesta towards the lower end of
the tract. This movement is neurogenic and is carried out by intrinsic nerves.
• A combination of segmentation and slow wave movement by
brief peristalsis ensures ample opportunity for absorptive
process to be completed.
Movements during inter-digestive phase:
• Migrating myoelectric motility complexes (MMC) are waves
of peristalsis which sweep through the intestines in a regular
cycle during inter-digestive [Link] MMC serves to push
undigested materials out of the small intestine.
• The MMC begins in the duodenum as slow waves leading to
spike and muscle contraction.
Antiperistalsis
• Peristalsis occurring towards oral direction is designated as antiperistalsis or reverse peristalsis. It occurs throughout
the digestive tract, but it is not widespread or powerful as peristalsis.
• It helps to
➢ Delay the movement of food down the intestinal tract
8
 ➢ To ensure adequate mixing
➢ To regurgitate duodenal contents into stomach
Ileocaecal Sphincter

• The ileocaecal sphincter is at the junction of small and large intestine,it is a circular muscle that remains constricted
most of the time. It prevents back flow of colon contents into the ileum.
Movements of Large Intestine
• The movements of the large intestine are more sluggish than of small intestine. This helps the large intestine to
function as a reservoir of faecal matter and as a place for prolonged bacterial attack on cellulose and other
substances.
• In horse the digesta first enters the caecum from where it reaches the colon and retained in the large colon for a
prolonged period for microbial digestion. In horses microbial fermentation occurs in both caecum and colon.
• In ruminants the ingesta enters the colon first and then retropelled into the caecum for microbial digestion.
Functions of Large Intestine
• Large intestine helps in
1) absorption of water and electrolytes
2) storage of faeces
3) Fermentationof materials that escape digestion in the small intestine.
• Large intestine motility must be slow particularly in those species where microbial fermentationis extensive in large
intestine – horse, rabbit. The material entering large intestine in these animals is retained for several days
Colon
• Mixing is achieved by segmentation contractions and helps in absorption and fermentation functions.
• The large intestine of non-ruminant herbivores and omnivores is sacculated, while that of carnivores and ruminants
are not. These sacculations are the seat of active contraction.
• The longitudinal muscles in the large intestine of horse are not disbursed but arranged in discrete muscle bands
called taenia. The taeniaare shorter in length and hence the large intestine is divided into sac-like evaginations
called haustra. Segmentation in the colon of horse and pig is pronounced and results in the formation of sacculations
known as haustra.
• Reverse peristalsis is a normal feature in colon of ruminants and rodents. It helps in reflux of digesta from colon to
caecum. Caecum again contracts and forces the ingesta into the colon. This to-and-fro movement of the digesta
continue. In horse, antiperistalsis helps in delaying forward movement of ingesta in colon. It aids in rapid absorption
and selective retention of water and microorganisms essential for fermentation in large intestine. In addition, periodic
intensive propulsive motility occurs involving the entire colon, which are called “mass movements”.
• Colon of dog and cat consists of short caecum, ascending, transverse and descending colon.
Caecum
• This is small in size in carnivores and man.
• It functions as a reservoir in which cellulose and other dietary constituents undergo bacterial fermentation in
[Link] caecal movements are sluggish and becomepowerful producing mass movements to evacuate
caecal contents to colon.
• In herbivores, materials pass into caecum from ileum and distend it. Then partial emptying of caecum into colon
occurs by contraction of caecum followed by antiperistalsis of colon which pushes materials into caecum.
DEFECATION
• It is a complex reflex act in which the faeces are discharged from the terminal colon and rectum.
• Two sphincters guard the anal opening –
➢ an internal anal sphincter of smooth muscle (it is continuation of circular layer of rectum)
➢ anexternal anal sphincter of striated muscle.
• The internal sphincter remains tonically closed most of the time and receive parasympathetic innervation from sacral
spinal segments through pelvic nerves and sympathetic from lumbosacral segments through the hypogastric nerves.
• In most species, sympathetic nerve causes contraction of sphincter and parasympathetic nerve causes relaxation.
• The external sphincter in innervated by somatic motor nerves passing through pudendal nerve
9
Nervous Control of Defecation
• Distension of the terminal colon and rectum by faeces causes reflex relaxation of the internal sphincter and stimulate
the surrounding nerve endings in the anal region which set up a reflex called recto-sphincteric reflex.
• Afferent impulses enter the CNS through sacral nerves. This results in an inspiratory effort with closed glottis and
contraction of abdominal muscles increases intra abdominal pressure.
• Peristalsis of colon and rectum and relaxation of anal sphincter leads to defecation.
• In trained animals (dog, cat), voluntary constriction of external anal sphincter can delay act of defecation; due to this
voluntary act, rectum relaxes to accommodate more faecal matter.
• The nerve centre controlling the anal sphincter in found in lumbosacral spinal cord.
• Higher defecation centre is present in the floor of the IV ventricle, close to the vomiting centre; frightened animals
frequently defecate by facilitation of brain centres of defecation.
• Frequency of defecation: horse – 5-12 times/day; cattle – 10-24 times; carnivores – 2-3 times
SECRETORY FUNCTIONS OF DIGESTIVE TRACT
SALIVARY GLANDS
• The salivary glands provide digestive fluids for the mouth and stomach.
• There are three main salivary glands all of which are paired
• There are also numerous small glands found in the mucous membrane of the
mouth.
• The three main salivary glands are
➢ parotid,
➢ sublingualand
➢ sub-maxillary(mandibular) glands.
• Secretary glands in general are divided into serous, mucous and mixed types.
• Serous glands give rise to a thin watery secretion containing protein/enzymes. Eg. Parotid glands.
• Mucous glands produce a secretion containing glycoprotein–mucin but these glands don’t produce enzymes.
• Mixed glands function similar to both serous and mucous glands eg. Submaxillary and sublingual glands.
• The parotid glands of most animals are serous (but in some animals the secretions are devoid of enzymes).
• The submaxillary gland is mixed (man, dogs, ruminants, cats, etc.) in some species.
• The sublingual glands of horse, ox, pig, dogs, and cats are mixed.

Salivary Secretion
• The functional salivary unit is called salivon.
• The epithelial cells of the acini transport electrolytes from the ECF into the lumen of acini and water flows from the
extracellular fluid by osmosis. Mucus and proteins synthesized by glandular cells of acini are added into the lumen
by [Link] primary secretion is iso-osmotic to ECF.
• As this primary secretionis flowing through the collecting ducts, its composition is modified. The cells of the duct
epithelium reabsorb Na and Cl ion exchange for K or H and HCO3.
• The final product is saliva and it is hypotonic especially in nonruminants and has less Na and more K than the
extracellular fluid.
Regulation of Salivary Secretion
Nerve supply
• The salivary glands receive double efferent innervation (i.e.) parasympathetic and sympathetic nerves.
• The salivary secretary centres are located in medulla and the afferent and efferent fibres of salivary glands are
contained chiefly in the facial and glossopharyngeal nerves.
• Parasympathetic fibres are present in the facial and glossopharyngeal nerves. These nerve fibres end on salivary
acini and stimulate the cells by cholinergic receptors.
• Stimulation of the parasympathetic fibres causes copious secretion rich in water and HCO3 but is low in protein and
also produces vasodilatation in the gland.
• Stimulation of the sympathetic nerve (-adrenergic nerve fibres) produces a small amount of thick saliva rich in
protein and causes vasoconstriction in the gland.
• Anticipation of eating brings about parasympathetic response resulting in increased salivary secretion.
• Food need not enter the mouth to cause the flow of [Link] sight, odour or even the thought of food provide the
individual is hungry, elicits salivary secretion. Under these conditions the mouth is said to be watery.
• The afferent pathways involved in this reflex are optic and olfactory nerves. This reflex involves cerebral cortex and
is designated as Psychic reflex by Pavlov (1910).
10
• Psychic reflex secretion is slight or absent in horse and sheep and present in goats and pigs.
• When food enters the mouth, a copious secretion of saliva takes place by reflex stimulation of the salivary glands
through buccal receptors and secretary centres.
Amount of Saliva
• Quantity of saliva secreted:
a) Horse – 50mL/ min during mastication
b) Cow –100 to 200 L / day
c) Dog – 0.5 L/day in 20 kg dog
Composition of Saliva
• Saliva contains 98% water and the remainder consisting of ions and organic compounds
• Saliva contains mucin (glycoproteins that when mixed with water becomes viscous mucus that lubricate the food,
bind bacteria and protect oral mucosa), proteins and in some animals ptyaline (salivary amylase) enzyme. Inorganic
salts are same as in blood. Saliva has a lower concentration of Na and higher concentration of K than blood. Saliva
also contains immunoglobulin IgA and lactoferrin which are bacteriostatic
• Reaction of saliva in domestic animals is slightly alkaline;
• In ruminants saliva is distinctly alkaline and in man the pH of salivary secretion is 6.6.
Functions of Saliva
(1) It moistens the food and facilitates mastication and swallowing; dissolve some of the food providing stimulation to
the taste receptors. Because of the mucin content of saliva, the food particles are formed into a bolus for swallowing;
the swallowing passage is also lubricated.
(2) Saliva provides a proper media for the bacterial growth and activity in the rumen. It is done by neutralising the
organic acids produced by the bacterial action and by prevention of frothing of the rumen contents. Ruminant saliva
contains a high amount of bicarbonates, phosphates and urea than serum, which make them a good buffer for the
rumen. The salivary ureasupplies nitrogen and phosphates provide P for the bacterial growth in rumen.
(3) Saliva has an important protective function of oral mucosa, which is kept moist and lubricated and mouth is rinsed
by saliva. This is due to spontaneous secretion of saliva even when food is not present in the mouth and also during
sleep. During nausea before vomiting, there is increased salivary secretion and this protects the mucous membrane
during subsequent passage of the acid vomitus.
(4) Saliva of some animals and man contains an amylolytic enzyme termed ptyaline or salivary amylase. This enzyme
hydrolyses starch through dextrins to maltose and glucose. (Optimum pH for ptyaline activity is 6.2 though it will act
in a range of pH 4 to 9.
(5) Saliva of pigs possesses significant amylase activity. However, this activity is weaker than that of man. Horse cattle,
sheep and goats posses no amylolytic activity in saliva. Dogs and cats contain small amount of amylase. Young
animals like calves, kids and man have another enzyme called lingual lipase, a fat digesting enzyme which
disappears in adult animals.
(6) Saliva posses bacteriostatic properties; this is due to the presence of lysozymes, IgA and lactoferrins which have
antibacterial activity. However, this activity is mild.
(7) Salivary glands provide water, which readily evaporates from the buccal mucosa and assist in thermoregulation in
dogs and cats. In dry warm atmosphere, watery saliva is secreted which evaporates from the oral mucosa during
panting.
GASTRIC SECRETION
• After the swallowing of food, it is received by stomach where it is subjected to gastric digestion.
• As regards stomach structure and function, domestic animals fall into two general classes.
1) Ruminants – Cattle, sheep, goat, camel, buffalo
2) Non-ruminants – Horse, cat, dog, pig

• The stomach of nonruminants is relatively

simple consisting of only one
compartment and hence referred to as
simple stomach.

11
• The stomach of ruminants is more complex consisting of four compartments of which only one secretes gastric
juice.
• Simple stomached animals include carnivores, herbivores and omnivores.
Functions of the Stomach
(1) It is reservoir of food, where digestion of food materials occurs. It is involved in grinding the food to reduce their size
and also controls the rate of passage of food to the small intestine.
(2) It produces the intrinsic factor concerned in the absorption of vitamin B 12 from the intestine, and functions in
hematopoiesis.
Structure of stomach in simple-stomached animals
• The stomach is a hollow, sac like organ.
• Stomach is made up of four layers. From outside to inside the layers are serous, muscular, submucosa and mucosa.
• In domestic animals, the stomach mucosa is divisible into
➢ glandless oesophageal region
➢ glandular region.
Esophageal Region
• In cow, the stomach is enlarged and compartmentalised; in horse, the esophageal region is large making up 1/3 to
1/5 of the surface area of stomach. It is non-glandular covered with stratified squamous epithelium,In pig the
esophageal region is limited to a small area around the cardia. In dogs, the esophageal region is lacking.
Glandular Region
• The glandular mucosa of the stomach has many invaginatons or pitsknown as gastric pits. These pits extend into
the gastric mucosa as straight and branched tubules, forming gastric glands and the gastric glands open into the
lumen through narrow openings on the gastric mucosa.
• The surface area of the stomach and the lining of the pits are covered with surface mucous cells that secrete thick
mucus which helps to protect the surface epithelium from acidity. A marked decrease in the mucus secretion will
lead to stomach ulcers
o The cardiac zone contains cardiac glands (secrete mucus) only;
o The fundic zone containsparietal glands (secrete HCl and pepsinogen)
o The pyloric zone contains pyloric glands (secrete mucus and gastrin).
• The fundic or parietal glands are compound tubular glands and are the proper gastric glands. The fundic glands
contain three main types of cells.
a) Chief cells or peptic cells: These cells secrete enzymes.
b) Parietal or oxyntic cells: These cellssecrete HCl. The parietal cells also secrete the intrinsic factor.

(Neck chief cells are the progenitor cells for gastric mucosa; when they mature, they migrate to all areas of the
gland differentiating into other types of gastric gland cells. The mucosa of parietal glands contains
enteroendocrinecells that secrete somatostatin and [Link] pyloric glands contain gastrin-producing
endocrine cells called ‘G’ cells. )

Composition and Functions of Gastric Juice

• It is a colourless fluid, often containing mucus.
• The pH is around 2 – 2.5.
• The organic substances include three enzymes
(1) Pepsin
(2) Rennin
(3) Gastric lipase
• The gastric juice contains a mucoprotein secreted by parietal cells known as intrinsic factorwhich is necessary for
the absorption of vitamin B12 required for haematopoiesis.
Pepsin
• It is a proteolytic enzyme and is synthesised in the peptic cells (chief cells) in an inactive form pepsinogen.
• HCl activates pepsinogen to pepsin, Optimum pH range for pepsin activity is between 1.5 and 3.
• Pepsin converts proteins to polypeptides.
Rennin

12
• It is the milk-coagulating enzyme which also possesses proteolytic activities
• It is present in the gastric juice of young animals (calves, lambs, piglet etc.). It is secreted as inactive prorennin, and
is activated by HCl.
• The rennin changes casein of milk to [Link] ions react with paracasein to form a gel, calcium
paracaseinate. This coagulated milk helps in delaying the passage through the stomach and the action of pepsin
on milk protein is prolonged.
Casein + Rennin ---> Paracasein (soluble) +Ca2+ Calcium paracaseinate (coagulum)
Gastric Lipase
• It is present in low concentration in the gastric juice of carnivores and absent in birds and ruminants.
• It hydrolyses fat into fatty acids and glycerol.
Hydrochloric Acid
• It is present in the gastric juice of all vertebrates, HCl is produced by the parietal cells of the fundic [Link]
production of acid secretion by the parietal cells involves expenditure of energy.
Secretion
• During active digestion, the concentration of HCO3 increases for a short time and increases the pH of blood which
is called post-prandial alkaline tide.
• Concentration of HCl in the stomach content varies with the nature of food, stage of digestion, amount of saliva etc.
Functions of HCl

1. It activates pepsinogen and prorennin to their active form

2. Helps the pepsin in the digestion of proteins by providing acidic environment
3. May bring about slight hydrolysis of sucrose
4. Acts as a stomach antiseptic – prevent fermentation of food in stomach by destroying microbes
Control of Gastric Secretion
• The rate of secretion increases when food is eaten.
• The gastric secretion may be divided into three phases
(1) Cephalic phase
(2) Gastric phase
(3) Intestinal phase

Cephalic Phase

13
• Stimulation of sensory nerve endings in mouth and pharynx or anticipation of eating can evoke this phase of
secretion. Presence of food in the stomach is not necessary for initiation of secretion.
Gastric Phase
• When food enters the stomach, there is more copious secretion of gastric juice; this constitutes the gastric phase of
secretary response.
• Gastric phase is caused by at least two kinds of stimuli.
(1) Mechanical stimuli
(2) Humoral or hormonal stimuli
Mechanical Stimulation
• When food or even inert substances come in contact with the stomach mucous membrane, gastric juice start to
flow.
• Distension of the stomach stimulates the intrinsic nerves which releases Ach; the Ach stimulate the G cells and
parietal cells. The G cells by producing gastrin further stimulate the parietal cells to release [Link], a protein
hormone produced by the ‘G’ cells of the pyloric glands stimulates the gastric acid secretion. Histamine is another
powerful stimulator of gastric acid secretion, Histamine acts through H 2 receptors and it is present in the gastric
mucosa. Vagus nerve can potentiate gastrin release and condition the parietal cells for the action of gastrin.
Intestinal Phase
• Addition of food to the intestine through a fistula will excite gastric secretion and this is due to a humoral mechanism
involved in the intestinal phase of secretion. This may be due to secretion of intestinal gastrin and pancreatic peptide
(PP) from duodenum during digestion of food and stimulating the gastric glands to activity.
• Relative importance of the three phases: Cephalic phase accounts for 45% of total daily secretion, gastric phase -
another 45% and intestinal phase for 10% or less.
Gastric Mucosal barrier
• The gastric juice is highly acidic and can cause tissue damage. But this damage does not happen in the stomach
because of mucosal barrier contributed by mucus and HCO3.
• The gastric mucus is continuously produced from cardiac and pyloric glands and neck chief cells of fundic glands
and surface epithelium of stomach.
• Gastric ulcers are produced by a special type of bacteria– Helicobacter pylori, alcohol, certain anti-inflammatory
substances like NSAIDS (aspirin)
• Ulcers can occur in many domestic animals and very common in dogs
• Ulcers occur both in stomach and duodenum.
• Anti histamine drugs (H2 blockers) greatly reduce H production and helps in healing of ulcers
HUNGER:
• A craving, desire, or urgent need for [Link] an uneasy sensation occasioned normally by the lack of food and
resulting directly from stimulation of the sensory nerves of the stomach by the contraction movement of the empty
[Link] can adapt to variable environmental conditions by adjusting their food intake.
• Two centres in the hypothalamus are involved in regulating feed intake
• Stimulation of appetite centre(hunger centre)located on the ventro-lateral part of hypothalamus causes an animal
to search for food and eat it voraciously. This centre tellsan animal when to eat.
• Stimulation of satiety centre in the ventro-medial hypothalamus makes an animal to stop eating. This centre makes
an animal when to stop eating; in other words how much to eat and it prevents overeating
• Fullness of stomach and duodenum after a meal stimulates stretch-sensitive neurons which transmit impulses
through vagi to the satiety centre which in turn stops eating. Some hormones released during eating and when the
stomach is full are also involved in stimulating the satiety centre
• There are many theories that try to explain the regulation of food intake. Three important ones are
o Glucostatic theory– availability of glucose at hypothalamus determines food intake – increase in blood glucose
reaching hypothalamus after a meal stimulatessatiety centre.
o CCK theory– CCK released in response to increased concentration of peptides and fatty acids in the small
intestine stimulatessatiety centre
o Lipostatic theory– increase in adipose tissue of the body releases a hormone leptinfrom adipose cells which
in turn stimulatesthesatiety centre; this is involved in long-term regulation of feed intake
• The feeling of hunger pangs in man is accompanied by marked rhythmic contractions of the stomachand is termed
as hunger contractions.

14
PANCREAS
• Pancreas consists of endocrine and exocrine portions and the former is made up of islets of Langerhanswhich
secrete insulin and glucagon into blood.
• The major part of pancreas is exocrine; it consist of acini responsible for production of pancreatic juice and ducts
which convey the juice to duodenum.
Secretion
• The enzymes of the pancreatic juice are synthesised and secreted by the acinar cells. The acini are similar to
salivary gland, made up of single layer of secretory cells surrounding a lumen
• The duct system (small intercalated ducts join to form major ducts that gradually increases in size, finally unite to
form a common duct that enters into the duodenum) is responsible for the secretion of water and electrolytes,
particularly the small intercalated ducts, which contain high level of carbonic anhydrase.
• The pancreatic alveoli, lobules and lobes are supported by connective tissues and the islets of Langerhans cells
are present in the connective tissue.
• In horse, dog and fowl there are usually two pancreatic ducts.
• In most species, the pancreatic duct empties into duodenum directly. In sheep, goats and humen, it joins the
common bile duct before entering duodenum.
• Carbonic anhydrase in the acinar cells produce H + and HCO3- from water and CO2. The H+ ion is transported into
blood by a Na-H exchanger and HCO3 passes through the apical cell membrane into the lumen through Cl-HCO3
exchanger. Na+ enters the lumen from blood and a net NaHCO3 secretion occurs.
• When this secretion passes through duct system, HCO3diffuses from the duct lumen into the blood in change for Cl-
.
• The concentration of HCO3 and Cl in the pancreatic juice depend on rate of secretion –at high rate of secretion,
HCO3level will be more in pancreatic juice (less time available for exchange with Cl-) and when rate of secretion is
low, Cl concentration will be greater in pancreatic juice
• The pancreatic enzymes are synthesized in acinar cells, packaged in zymogen granules and released by exocytosis.
Composition and Amount
• Pancreatic juice is a clear alkaline fluid made up of two separate secretions;
• One is aqueous phase containing a higher concentration of HCO3 and lesser amounts of Cl
• Second is organic phase, comprising pancreatic enzymes.
• In ruminant and horses, pancreatic secretion is continuous; in horses but not in ruminants, the rate of secretion
increases greatly after feeding
• The amount of juice secreted in different animals is as follows (in L/100Kg body weight/day):
➢ Horse – 10 to 12 Cattle – 3 to 5 Sheep – 0.5 to 1
Pancreatic Enzymes
• The HCO3 is important for partial neutralisation of the acid chyme from stomach and for maintenance of H + ion
concentration suitable for digestive activity of pancreatic enzymes.
• The enzymes of the pancreas are capable of digesting fats, carbohydrates and proteins.
• There are three major groups of enzymes – proteases, lipases and amylase. Other more specific enzymes are also
present.
• Proteolytic enzymes of pancreas are secreted as zymogen granules or proenzymes.
• They areTrypsinogen 2) Chymotrysinogen 3) Procarboxypeptidase A and B 4) Proelastase
• The proenzyme trypsinogen is converted to trypsin either by autocatalysis or by the action of enterokinase (an
enteropeptidase) which is an enzyme present in duodenal and jejunal juice.
• Chymotrysinogen, procarboxypeptidases and proelastases are activated by trypsin.

15
• Trypsin, elastase and chymotrypsin are endopeptidases;
• Carboxypeptidase is exopeptidase.
• Pancreatic amylase (amylopsin) is secreted in an active state. They require an optimum pH of 6 to 9 and Cl - ions
for their action. They act on starch and produce oligosaccharides and maltose.
• Pancreatic lipase (steapsin) is secreted in the active form, splits fats to free fatty acids and glycerol. Calcium ions,
polypeptides, peptides and bile salts enhance lipase activity.
• Cholesterol esterase and phospholipase act on cholesterol esters and phospholipids producing non-esterified fatty
acids, cholesterol and lysophospholipid.
• Ribonuclease and deoxy ribonuclease present in pancreatic juice reduces ribose nucleic acid and deoxyribonucleic
acids to mononucleotides.
Regulation of Pancreatic Secretion
Nervous Regulation
• When the cephalic phase of stomach secretion occurs, impulses are simultaneously transmitted along the vagi
nerves to pancreas; this results in secretion of large concentration of enzymes into the pancreatic acini but little
secretion flow through the duct to intestine because little water and electrolytes are secreted along with enzymes.
This phase of pancreatic secretion is called as cephalic phase.
• Entry of food into the stomach causes distension of stomach and this result in vagovagal reflex; this reflex stimulates
pancreatic secretion. This is called as gastric phase of pancreatic secretion.
• After the food enter the small intestine, distension of intestine results in production of enzyme secretion by the
pancreas. This stage is called as intestinal phase of pancreatic secretion, which is controlled by both nerves and
hormones.
• Neural control of intestinal phase is provided by the intrinsic nerves of the intestine and vagus. The acetylcholine
released through vagal action and local nerve reflexes sensitise the pancreas for the action of secretin and CCK.
• Hormonal Regulation
• Two hormones are important in regulating pancreatic secretion
• They are secretin and CCK.
• Secretin is present in the mucosa of duodenum and upper small intestine of many species. It causes copious
secretion of pancreatic juice rich in HCO3 but poor in enzymes and other proteins.
• Secretin is the first hormone discovered; In 1902, Bayliss and Starling discovered this hormone in dogs. It is a
polypeptide hormone, act directly on pancreatic duct cells and stimulates HCO 3 of pancreatic secretion; it also
increases bile secretion from liver.
• Stimulus for the release of secretin is presence of acid ingesta, peptides, soaps and amino acids in duodenum. It
also inhibits gastric secretion.
• The secretin is released when the duodenal pH falls below 4.0.
• The HCO3 neutralises the acidity of duodenal contents.
• Secretion of large quantities of thin watery pancreatic juice with high concentration of HCO3 and less or no enzyme
secretion which is produced by secretin are called as hydrolytic secretion.
• Cholecystokinin (CCK) is present in duodenal and small intestine mucosa. This hormone stimulates secretion of
enzymes from pancreas and this action is identical to vagal stimulation.
• The secretion resulting from CCK stimulation is rich in enzymes and this type of secretion is called ecbolic
secretion.
• CCK also causes contraction of gall bladder and delays gastric emptying.
• Vasoactive intestinal polypeptide (VIP), another intestinal hormone, stimulates pancreatic HCO3 secretion.
• Pancreatic polypeptide, an intestinal hormone, inhibits pancreatic HCO3 and enzyme secretion.
LIVER
Functions of Liver

16
 1. Secretion of bile
2. Metabolism of protein, carbohydrates and fat
3. Detoxification of harmful substances
4. Storage of vitamins
5. Destruction of erythrocytes
6. Formation of blood proteins – fibrinogen, prothrombin etc.
7. Formation and storage of glycogen and regulation of glucose level of systemic blood
8. Deamination of amino acids and formation of urea
9. Destruction of uric acid
10. Synthesis of fatty acids from carbohydrates and proteins, phosphorylation of fats, inter- conversion of fatty acids,
partial oxidation of fatty acids and formation of ketone bodies
11. Destruction of hormones, drugs etc.
BILE SECRETION
• Liver is composed of interconnecting plates of epithelial cells (hepatocytes) forming a continuous interconnecting
latticework around the vascular supply to the organ. Between the plates of liver are many interconnecting cavities
containing vascular sinusoids. The sinusoids are lined by kupfer cells which are part of mononuclear phagocytic
system (MPS) or RE system.

• The liver parenchyma cells known as hepatocytesproduce the bile.

• Between theindividual hepatocytes is the small canals – bile canaliculi that continue as biliary duct system that
finally form a large bile duct - hepatic bile duct
• The bile duct epithelium alters the composition of bile by adding HCO 3 and water.
• The hepatic bile ducts join the cystic duct of gall bladder and runs as common bile duct-ductus choledochus.
• The hepatic bile duct communicates directly with both duodenal lumen and gall bladder through cystic duct (in horse,
there is no cystic duct).
• Flow of bile is relatively constant in ruminants and pigs since the sphincter of Oddi is underdeveloped and it is
continuous in horse.
• The common bile duct opens through the duodenal wall and the opening of the duct is guarded by a sphincter
muscle called sphincter of Oddi. It is well-developed in carnivores than herbivores.
• Relaxation of the sphincter of Oddi is caused by CCK and passing peristaltic wave over duodenum.
• In man, dog, cat and horse the pancreatic and bile ducts open close together into the duodenum, whereas in man,
sheep and goat, the pancreatic duct opens into the common bile duct so that a mixture of bile and pancreatic juice
enters the duodenum.
17
• In pigs and ox, the opening of two ducts is some distance apart.
Gall Bladder
• Formation of bile is continuous but they are stored in gall bladder between periods of digestive activity and emptied
from gall bladder during digestion.
• The concentrated bile is discharged into the duodenum by gall
bladder contraction during digestion.
• Horse, rat, deer, mouse, giraffe, camel, elephant, pigeon and dove do
not posses a gall bladder and flow of bile in these species is continuous.
• Gall bladder contraction is under the control of nerves and
hormones.
• CCK secreted from the upper part of small intestine produces gall
bladder contraction. When food enters duodenum, it causes
release of CCK and bile enters duodenum.
• Once gall bladder is empty, bile flows directly into the duodenum
and its secretion is maintained by enterohepatic circulation of
bile salts.
Bile
• Bile is a digestive secretion and it helps in solubilisation and absorption of fat.
• Bile is also regarded as an excretory secretion because it contains lipids including cholesterol and break down
products of haemoglobin is removed from the body through bile.
• Bile is a viscid greenish or golden coloured liquid with a bitter taste and composition varying in different species.
• Bile contains 1) bile pigments 2) bile acids (salts) 3) cholesterol 4) lecithin 5) mucin 6) fats, soap and urea 7)
inorganic substances
Bile Pigments
• Bile pigments are biliverdin and its reduction product, the bilirubin.
• The colour of bile is due to bile pigments. Bile pigments are waste products of haemoglobin breakdown and are
formed in MPS cells of spleen, bone marrow and liver which are concerned with destruction of RBCs.
• Bile pigments are present in blood in low concentration.
• Because the bile is concentrated in gall bladder, cholesterol, bile pigments and Ca 2+ are prone to be precipitated
and can form gallstones.
• Jaundice is a condition in which there is excessive accumulation of bilirubin in tissues especially in fatty tissues
and in visible mucous membrane imparting a yellowish colouration.
Types of jaundice
I. Obstructive Jaundice it is caused by blockage to flow of bile. E.g. due to gallstones, bile ducts are obstructed
and conjugated bilirubin accumulates in blood.
II. Hepatic Jaundice is due to liver damage caused by disease or poison. In this condition, bilirubin is not
conjugated and accumulates in tissues and blood as free bilirubin. In new-born animals, mild jaundice occurs
during the first few days of life because liver is immature to excrete bilirubin (neonatal jaundice)
III. Haemolytic Jaundice occurs due to excessive production of bilirubin by RBC breakdown. Since excess
haemoglobin is broken down, it exceeds liver’s capacity to conjugate available bilirubin and so both free and
conjugated bilirubin level increases in plasma. Occurs in hereditary diseases (sickle-cell anaemia), parasitic
infections (babesiosis) incompatible antigen-antibody reaction (mismatched blood transfusion)
• Van den Bergh Test can be used to differentiate free and conjugated bilirubin and it is helpful to find out the type
of jaundice.
Bile Salts
• Bile salts form half the solids of bile. They are cyclopentano perhydrophenanthrene compounds
• Bile salts are derived from cholesterol.

18
• The bile acids produced in the liver are called primary bile acids; they are cholic, cheno deoxycholic acids. They are
conjugated with either taurine or glycine to form taurocholic or glycocholic acids(glycine is an amino acid and taurine
is sulphur containing nitrogenous substance derived from cysteine). These acids form salts with sodium.
• Secondary bile acids -deoxycholic and lithocholic acids -are formed from the primary bile acids by the action of
bacteria in colon; they are absorbed and excreted in bile.
• Conversion of insoluble cholesterol to bile acids results in formation of a molecule with a water-soluble (hydrophilic)
side and a lipid-soluble (hydrophobic) side. This combination of hydrophobic-hydrophilic (amphipathic) is
characteristic property of detergent. Because of this dual solubility, detergents make the lipids soluble in water.
Thus, the bile acids help to emulsify dietary lipids and solubilise the products of fat digestion.
Secretion and Regulation of Bile
• Bile secretion involves two components
1. Bile salt dependent flow
2. Bile salt independent flow
Bile Salt Dependent Flow
• The formation and secretion of bile by liver is an active process, carried out by hepatocytes. Bile salts are secreted
into canaliculi; presence of bile salts and Na+ in canaliculi draws water by osmosis from the cell into bile.
• Substances that stimulate bile secretion are known as cholerectics and the most important cholerectic is bile salts
themselves. The bile salts act directly on liver to increase secretion.
• Bile salts are synthesised by liver and secreted in bile. After entering the absorptive region of small intestine, they
are reabsorbed into the portal blood and passed back to liver; the absorption of bile salts is active and occurs in
ileum only. This recycling of bile salts is known as entero-hepatic circulation of bile salts. The reabsorbed bile salts
reaches liver and they are the most potent stimulant to bile secretion.
• About 90% of bile salts are reabsorbed in ileum.

Bile Salt Independent Flow

• This phase involves ductular epithelium. Na+ ions are actively transported from ductular cells into the lumen
accompanied by HCO 3-, Cl- and water. HCO3 concentration in bile is higher than that blood.
• This phase is under the control of secretin and it results in HCO3- rich secretion. Vagal stimulation also provokes
bile secretion.
HormonesRegulating Bile secretion
• Secretin stimulates bile secretion. It increases HCO3- secretion from the biliary duct cells. Acidic duodenal contents
releases secretin and HCO 3 helps toneutralize the acidic content
• The CCK causes relaxation of sphincter of Oddi and contraction of gall bladder and also increases flow of bile.
Functions of Bile Salts (Functions of Bile)
• Due to the presence of bile salts, bile is useful in the digestion and absorption of nutrients in the following ways.
1) It activates pancreatic lipase
2) Assists in fat emulsification
3) Increases solubility of higher fatty acids which are insoluble in water and aids in their absorption.
4) Bile assists in absorption of fat-soluble vitamins
Other functions of bile in the intestine
5) It is a reservoir of alkali and thus assists in maintaining optimal reactions in intestine
6) Mucin and mucin-like substances of bile act as stabilisers of fat emulsion in intestine.
7) Bile has antiseptic properties and regulates bacterial growth in bowel. When bile does not enter the intestine,
fat absorption is diminished and other constituents of food become coated with fat. Hence, their digestion is
limited and proteins putrefy. Therefore, the faeces develop an offensive odour.
8) Bile has a mild laxative effect.
Reactions in Intestine

19
• Gastric juice is acidic in reaction. Duodenal juice, pancreatic juice, bile and intestinal juice are alkaline.
• pH at different regions of digestive tract
Stomach Duodenum Jejunum Ileum Caecum Large colon Rectum
2– 2.5 7– 7.4 7.5 7.55 7.24 7.09 6.24
DUODENAL SECRETION
• In the duodenum, a number of mucous glands known as Brunner’s glands lie in the submucosa. Their ducts open
into crypts. The Brunner’s glands occupy only a short distance in carnivores and they extend over most of the
duodenum in herbivores.
• The secretion of Brunner’s glands is thick, clear mucous fluid with a pH between 7 and 8 has a high HCO3 content
and possesses enzymic activities.
• Secretion of duodenal glands is known as duodenal juice.
• It contains enzymes amylase, lipase, sucrase and lactase.
• Secretin and vagal stimulation increases secretion.
• The duodenal juice helps to neutralise the acidic gastric contents as they enter duodenum and thus protects the
duodenal mucosa from acid contents.
INTESTINAL SECRETION
• The small intestine is composed of four layers – outer serosal, muscular, submucosa and the innermost mucosal
layer.
• Mucosa is the functional layer where digestion and absorption occurs.
• The mucosa of the small intestine is thrown intomany circular folds called plica circulares
• The surface epithelium of the mucosa projects from the surface into the lumen of intestine and these are called as
villi;
• The luminal surface of the intestinal epithalialcells exhibits numerous microvilli called brush border
• Located over the entire surface of the small intestine between the villi are small pits called crypts of Lieberkuhn and
they are the proper intestinal glands.
• The intestinal glands contain goblet cells (secrete mucus and HCO 3), enteroendocrine cells(produce many types of
chemical messengers) and large paneth cells(produce enzymes including enterokinase).
• The cells of the crypts secrete fluid and enzymes which form the intestinal juice or succus entericus.
• The surface of the villi is lined by tall columnar epithelial cells called enterocytes which are the absorptive cells.
Regulation of Intestinal Secretion
• Mechanical stimulation of intestinal mucous membrane by the digesta causes secretion of intestinal juice
and it is the most important stimulus for intestinal secretion.
• Nervous regulation is limited and not clear. Vagal stimulation increases duodenal secretion and
sympathetic stimulation inhibits secretion.
Digestive enzymes

Enzyme Source Substrate Products

Saliva

Salivary amylase Salivary glands Starch and glycogen Maltose (disaccharide),

maltotriose
(trisaccharide), and a-
dextrins

20
 Lingual lipase Gland in the tongue Triglycerides and other Fatty acids and
lipids diglycerides

Gastric secretions

Pepsin Chief cells Proteins Peptides

Gastric lipase Chief cells Short-chain Fatty acids and

triglycerides monoglycerides

Renin Chief cells Milk casein Coagulation of milk

casein

Pancreatic Secretions

Trypsin Pancreatic acinar cells Proteins, Peptides

chymotrypsinogen,
procarboxypeptidase

Chymotrypsin Pancreatic acinar cells Proteins Peptides

Elastase Pancreatic acinar cells Proteins Peptides

Carboxypeptidase Pancreatic acinar cells Terminal amino acid at Peptides and amino acids
carboxyl end of peptides

Pancreatic lipase Pancreatic acinar cells Triglycerides Fatty acids and

monoglycerides

Ribonuclease Pancreatic acinar cells Ribonucleic acid Nucleotides

Deoxyribonuclease Pancreatic acinar cells Deoxyribonucleic acid Nucleotides

Brush Border Enzymes

a-dextrinase Plasma membrane of microvilli a-dextrins Glucose

Maltase Plasma membrane of microvilli Maltose Glucose

Sucrase Plasma membrane of microvilli Sucrose Glucose and fructose

Lactase Plasma membrane of microvilli Lactose Glucose and galactose

Enterokinase Plasma membrane of microvilli Trypsinogen Trypsin

Aminopeptidase Plasma membrane of microvilli Terminal amino acid at Peptides and amino acids
amino end of proteins

Dipeptidase Plasma membrane of microvilli Dipeptides Amino acids

Nucleosidase Plasma membrane of microvilli Nucleotides Nitogenous bases,

pentoses, and
phosphates

Phosphatase Plasma membrane of microvilli Nucleotides Phosphate ions

DIGESTION IN THE RUMINANT STOMACH

• Ruminants are animals that regurgitate and remasticate their food.
• The principle feature of digestive physiology in the ruminants is that fermentative digestion or microbial digestion
occurs on a massive scale in the first two parts of the stomach.
21
• Ruminants include four families of animals – Cervidae, Giraffidae, Antilocapridae, Bovidae –consisting of about 155
species of animals
• Ruminant-like fermentation of food also occurs in the three chambered stomachs of camel, Llama and in the
stomach of marsupials and in hippopotamus.
• Some fermentation occurs in the large intestine of many animals and is most prominent in equidae.
Foregut and hindgut digesters
• The important anatomical and physiological features that allow fermentation of food to occur within the alimentary
tract are –
o Large capacity, slow passage of food, buffered fluid environment and removal of the end products of
fermentation.
• These conditions are found in rumen and reticulum and also in the large intestine of horses.
• In pregastric (foregut) digesters (ruminants) both soluble and insoluble dietary constituents are fermented by
microorganisms before enzymatic digestion. The end products of carbohydrate fermentation are VFAs (SCFA) and
glucose does not reach the intestine for absorption. Hence ruminants synthesize glucose from VFAs to maintain
blood glucose.
• In postgastric (hindgut) digesters (horse, rabbit, pig), soluble carbohydrates are digested before microbial
fermentation and glucose is absorbed from gut.
• The fermentation of cellulose is a slower process and a marked delay in the forestomach of ruminants allows efficient
fibre digestion.
• Fermentation in large intestine of horse is confined to the constituents of plant fibres, but the digestion of cellulose
and hemi-cellulose is not as complete in the large intestine of horse as in the rumen since passage rate is
comparatively faster in the large intestine of horse and fibre digestion is less efficient.
• In ruminants, dietary proteins are fermented and end products are utilized to produce microbial protein which then
reaches the small intestine for digestion and absorption. The advantage of this is that NPN substances are utilized
to synthesize microbial protein.
• The bacteria and protozoa of the large intestine of horse are not absorbed into the host and do not appear in faeces
but they die and disintegrate as the gut contents pass caudally.
•
Relative Capacity of the Digestive Tract (%)

Animal Stomach Small Intestine Caecum Colon &Rectum

Cattle 71 18 3 8
Sheep & Goat 67 21 2 10
Horse 09 30 16 45
Pig 29 33 6 32
Dog 63 23 1 13
Anatomy of the Ruminant Stomach
• The ruminant stomach consists of 4 compartments-rumen, reticulum, omasum and abomasum.
• The first three compartments are known as fore stomach and abomasum as true stomach.
• Dietary constituents are fermented in the forestomach by microbes to generate ATP for their growth;
• Fermentation end products are absorbed by the ruminants to be used as metabolic substrates.
• Capacity of the ruminant stomach varies greatly with age and size of the animal.
• In medium sized animals, the capacity is about 250L (cattle) and in sheep it is 25 to 30L.
• The relative size of the 4 compartments varies with age; in new-born calf, the first 3 compartments are small and as
the animal grows, these compartments grow in size. The abomasum is the largest compartment of the new-born
stomach.

22
• The forestomach development is related with roughage intake. Young ruminants begin ingesting limited roughage
when they are 1-2 weeks old. Accelerated development of the fore stomach takes place from 3 to 8 week’s period
and during this time the microbes’ establish, ruminal papillae develop and omasal leaves are stimulated by the
SCFAs.
• As the animal grows, the rumen and reticulum grows much faster than abomasum; in the adult, the ruminoreticulum
constitutes 69% of the total stomach capacity, abomasum 23% and omasum 8%.
• When the
rumen is full, it
extends from the
diaphragm up to
pelvis
• The rumen
has dorsal and
ventral sacs, both
communicating
freely with each
other and the
sacs are
surrounded by
muscular pillars.
• Rumen communicates with reticulum over the rumino-reticular fold.
• From the cardia to the reticulo-omasal orifice extends a groove called the reticular (esophageal) groove.
• The mucous membrane of the rumen is stratified squamous epithelium and non-glandular and covered with
numerous papillae.
• The reticulum lies against the diaphragm and liver. It is small and flask shaped. The esophagus enters the reticulum
at cardia.
• The reticulum communicates with the omasum through reticulo-omasal orifice and with rumen through the reticulo-
ruminal fold.
• The reticular mucous membrane is non-glandular and is thrown into folds which resemble honeycomb.
• Omasum lieson the right of the [Link] somewhat kidney shaped.
• It communicates with reticulum through reticulo-omasal orifice and with abomasum through omaso-abomasal
orifice.
• Omasal groove or sulcus is a groove that extends downwards from the inlet to the outlet of the omasum.
• The interior of the omasum shows numerous laminae of different sizes which are studded, with numerous papillae;
• Certain ruminants have no omasum (e.g.) camel.
• In sheep and goats omasum is not well developed.
• Semi-liquid materials enter the omasum through the reticulo-omasal orifice.
• The functions of omasum are:
➢ absorption of SCFAs, water and electrolytes through the papillae in the omasal leaves;
➢ regulate onward passage of digesta from reticulorumen to abomasums;
➢ it is a minor site of fermentation.
Abomasum
• The abomasum is the glandular compartment of the ruminant stomach. It communicates with omasum and with
duodenum through pylorus.
• Abomasum is divided into fundic, body and pyloric regions and has fundic and pyloric glands. The fundic and body
mucosa is thrown into about 12 large spiral folds. The pyloric region is same as in other animals.
• Abomasum is the only part of the ruminant stomach that secretes gastric juice.
• Abomasum receives a continuous flow of materials from the fore stomach.
23
• Abomasal secretions from the fundic region contain pepsin and HCl and pyloric secretion is scanty.
• The fundus is usually quiescent body, pylorus show numerous contractions and relaxations.
• Gastric juice in the fundus region has a pH close to 2.0 and has constant peptic activity. Pyloric secretions are
slightly alkaline, and have little peptic activity.
• Abomasal secretions are dependent on volume of material flowing into it; pyloric distension, rise in abomasal pH
and SCFA level stimulates gastrin and HCl secretion. Parasympathetic nerves control abomasal secretion.
DEVELOPMENT OF RUMINANT STOMACH
The development of ruminant stomach can be considered in four phases
1) The newborn phase (0 to 24 h of birth): The forestomach is small in size (39% of total stomach), contains no
microbes and papillae and is non functional. Abomasum also does not secrete acid or pepsinogen and the
immunoglobulins of the colostrum pass to the intestine without being digested.
2) The prerumiant phase (1 day to 3 weeks age): Principal food is milk, sucking promotes salivation which
contains pregastric esterase that hydrolyses milk fat. Passing of milk through pharynx stimulates closure of
reticular groove and milk bypasses reticulorumen to reach the abomasums.
3) Transitional phase (3 to 8 weeks age): Along with milk the animal starts to ingest roughage which initiates
growth of salivary glands and reticulorumen. Microbes from feed, water and from other ruminants establish in
the young animal. Microbial fermentation begins, VFAs are produced which stimulates rumen papilla growth.
Gases are produced and expelled out by eructation. Intermediary metabolism shifts from glucose based to VFA
based.
4) Preweaningand postweaning phase (8 weeks to adult age): Reticular groove reflex becomes less functional.
Forestomach grows and reaches adult proportions.
MOTILITY OF RUMEN AND RETICULUM
• The walls of the reticulo-rumen are muscular, posses intrinsic nervous system and are capable of complex co-
ordinated motility patterns and there are two patterns of motility which are–
• Primary or mixing contractions also called primary cycle or A sequence
• Secondary or eructation contractions also known as secondary cycle or B sequence
• The primary cycle: It consists of
1) a biphasic contraction of reticulum
2) contractions of dorsal sac
3) contractions of ventral sac of rumen.
• The primary cycle of rumino-reticular contractions begin from the reticulum.
• The reticular contractions are biphasic contractions in which the first contraction known as mixing contraction makes
the reticulum to half its size which then relaxes and it again contracts with strong force; the second contraction is
called as evacuation contraction.
• The contractions of the rumen start during the second contraction of the reticulum from the cranio-dorsal sac and
moves caudally. The dorsal sac contraction is followed by caudally moving ventral sac contractions; this is
succeeded by cranial moving contraction of dorsal and then ventral sacs. This sequence of contraction is called the
primary cycle.
➢ The function of the primary contraction is to mix ingesta
➢ Helps in the separation of large and small particles.
• Secondary cycle: Secondary contractions when occur follow the primary contraction and it is cranially moving
contraction beginning from the caudodorsal blind sac and moving over the dorsal sac.
➢ The function of the secondary contraction is to expel gas out of the rumen.
• The reticulo-rumen contractions are increased during eating and by coarseness of the diet.
• Number of contractions per 5 min – during feeding 5-8, during rumination and rest 4-5; during sleep-no contractions
• During rumination, triphasic contractions occur in reticulum in which the biphasic contraction is preceded by an extra
contraction. It is stronger than the first two and referred as regurgitation contraction because regurgitation of bolus
24
 occurs during this phase. During rumination, ventral sac movement of the primary wave is absent and ventral sac
movements of secondary wave are strong and prolonged.
Functions of rumino reticular contractions
(1) Direct liquid flow caudally to the cranial sac of the rumen
(2) Direct low density ingesta to the dorsal sac
(3) Assist in regurgitation
(4) Regulate ingesta flow from reticulo-rumen to omasum
Nervous Control of Ruminant Stomach
• Major contractions of rumen and reticulum are dependent on vagi.
• Total vagotomy abolishes rumen and reticular contractions, rumination, eructation and reticular groove reflex. Vagus
is also the motor nerve to abomasum.
• Tension receptors and epithelial/mucosal receptors are sensory receptors of fore-stomach and they send impulses
through vagus. Tension receptors monitor tension in the muscular wall; epithelial/mucosal receptors are stimulated
both by mechanical stimuli (coarseness of rumen material) and chemical stimuli - decrease in pH, increase in VFA
reduce motility.
• Sympathetic nerves are inhibitory to ruminant stomach but their actions are not very important.
• Even though the intrinsic nerves of reticulorumen can cause contractions, they are weak and are not cyclical and
cannot compensate for extrinsic vagal contractions.
RETICULAR GROOVE (Esophageal Groove)
• The reticular groove begins at the cardia and extends up to the reticulo-omasal [Link] reticular groove continues
as omasal canal which extends up to abomasum
• The reticular groove is a gutter-like invagination. When stimulated, the muscles in the lips of the groove contract
causing it to shorten and twist. The twisting causes the lips to close together forming a tube. In young ruminants the
reticular groove functions by reflex closure thereby enabling to conduct milk and water from the oesophagus directly
to the reticulo-omasal orifice.
• The closure of the groove is a reflex act initiated by stimulation of receptors in the mouth and pharynx. Sucking from
teats, stimulation of chemoreceptors in the mouth and pharynx by milk constituents causesreflex closure of the
groove.
• Although groove closure is an unconditioned reflex, the manner in which the animal takes milk has an influence on
the groove's response. When an animal is eager for milk, rapid and effective closure of the groove occurs. Water
consumption does not typically initiate groove closure.
• The reflex becomes almost non-functional when the young animal stops sucking
Effect ofChemicals on the Reflex Closure of the Reticular Groove
• Some chemicals when administered can cause closure of the groove and they are recommended before orally
giving drugs to ruminants which are not intended for rumen or reticulum but to reach abomasum.
• In young healthy cattle up to two years of age, the reflex can be evoked by administration of sodium chloride, sodium
bicarbonate and sugar. NaHCO3 (60ml of 10% solution) stimulates closure in calves.
• Copper sulphate causes reflex closure of the groove in sheep.
Stratification in rumen
• The rumen contents are stratified and segregated by the effects of gravity and motility.
• In forage fed animals there are four major zones or phases –
➢ a gas zone at the dorsal rumen created by the fermentation gases,
➢ a solid zone also known as rumen mat consisting of intermixed particles of fermenting forages. This phase is
floating due to the air trapped in the feed particles and gas bubbles of fermentation gases.
➢ At the bottom of the rumen in the ventral sac is the liquid zone and
➢ In between the solid and liquid zones is the slurry zone. Microbes are found attached with particles in the rumen
mat and free floating in the liquid zone.

25
Accumulation of Foreign Matter in Rumen
• Cattle may consume some foreign matter with their food and these may be retained in stomach, sometimes causing
serious injury to the wall and to the adjacent structures producing traumatic gastritis, traumatic pericarditis and other
inflammatory conditions.
• The reticulum and abomasum are places of accumulation of foreign matter. The foreign materials seen in reticulum
are nails, wire, glass and sand; in abomasum small pebbles, cinders etc; some foreign matter can also be seen in
rumen but never in omasum.
RUMINATION
• The process of bringing food materials back from the ruminant stomach to the mouth for further mastication is known
as rumination (chewing the cud).
• It is composed of four phases:
➢ 1) Regurgitation
➢ 2)Remastication
➢ 3) Reinsalivation
➢ 4) Reswallowing.
• These four phases with a slight pause after swallowing make up a cycle of rumination.
• Rumination is confined to ruminants only.
• Ruminants consume their food hurriedly with little mastication during feeding. In the capacious rumen, the food is
stored and undergoes thorough mixing, maceration and fermentation. At intervals, portions are returned to mouth
for rumination.
Regurgitation
• The food that returns to the mouth comes primarily from the liquid part of reticulum. The regurgitated mass consists
of small particulate matter highly mixed with liquid and in advanced stage of fermentation. Recently eaten forages
whose particle size is too great to be suspended in the rumen fluid are not included in the material that is
regurgitated.
• Entrance of ingesta into the oesophagus is accomplished by an inspiratory effort with tongue and soft palate raised
to close buccal and nasopharyngeal orifice. This effort produces a drop in intra-thoracic and intra-oesophageal
pressure. Simultaneously, extra-reticular contractions occur before the biphasic contraction. The cardia (caudal
oesophageal sphincter) open; a cud of soupy reticular content is drawn into the oesophagus due to negative intra-
thoracic pressure. An antiperistaltic wave of oesophagus carries the ingesta to mouth, the glottis closing briefly
when the material crosses pharynx.
• The lack of involvement of the abdominal muscles in increasing intra-abdominal pressure in the ruminant is a major
difference between regurgitation in ruminants and vomiting in other species
Remasticationand Reinsalivation
• Immediately after the bolus arrives in the mouth, the liquid is squeezed from it, which is swallowed. The remaining
solid mass is chewed with slow, regular chewing movements for about 40-seconds. During this process saliva is
added and several swallows occur. When chewing is completed, the remaining solid mass along with saliva is
swallowed.
• During reinsalivation, parotid glands are more active.
Redeglutition
• The reswallowed food returns to rumen, enters reticulum and passes quickly to other compartments.
• Time spent in rumination varies in different animals and with different rations.
• Average daily rumination time in cattle is 10 hon hay diet.
• The proportion of grain and roughage in ration influences rumination time. With low roughage diets or when
roughage is finely ground, total rumination time may be 3 h/day or less. Highest incidence of rumination occurs
during afternoon and in the middle of the night.
NervousControl of Rumination

26
• The rumination is a reflex act even though it can be influenced by voluntary control. A rumination centre exists in
medulla.
Eructation
• It is the expulsion of fermentation gas that has accumulated in the rumen.
• For eructation, the primary stimuli are the presence of gas in the dorsal sac.
• Volume of gas produced in rumen of dairy cow is ½ to 1 L/min (2000 – 4000 L/day). Most of the gases produced
are eliminated by eructation. Eructation occurs at about 1 to2/ min.
• Before eructation, the gas layer is moved cranially by the secondary cycle contraction of the dorsal sac and the
cardia is cleared of fluid. The cardia and lower oesophageal sphincter open, and gas enter into the relaxed
oesophagus. An anti peristaltic wave occurs in the oesophagus and soft palate is elevated.
• Part of the gas is expelled through the mouth and part of the gas from pharynx enters trachea then to lungs and
eliminated or absorbed into blood.
• Receptors for eructation reflex are tension receptors located in the reticulum, cardia, and cranial rumen sac which
are stimulated by accumulation of gas. Afferent and efferent fibres are in vagus. Centre for eructation is present in
medulla.
• When motility of reticulo-rumen is depressed or inhibited or if gas production is abnormal, gas accumulates in the
rumen; this condition is known as bloat or tympany. Feeding lush growing alfalfa results in dietary bloat due to the
formation of stable foam, which traps gas in the rumen. Oils and non-absorbed surfactants are used to treat this
condition.
SALIVARY GLANDS OF RUMINANTS
• Parotid, submaxillary and sublingual are the major salivary glands of ruminants. In addition, sheep and cattle have
two inferior molar glands and numerous small glands called buccal and labial glands found in cheek and lips.
Additionally, palatine glands are present in hard and soft palate and pharyngeal glands in the pharynx and roof of
the tongue.
• Parotid and inferior molars are serous glands. Buccal, pharyngeal and palatine glands are mucous glands. Sub-
maxillary sublingual and labial glands are mixed glands.
• Flow of saliva in cow is 60 to 160 L/day; in sheep 6.0 to 16 L/day
• Ruminant saliva is rich in HCO3, HPO4, and urea. Ruminant saliva is isotonic to blood plasma but has higher
concentration of Na, K, HCO3, HPO4 and lower concentration of Cl. At high secretory rate, K and HPO4
concentration falls slightly with corresponding rise in Na, HCO3 and Cl.
• The high HCO3 and HPO4 makes the ruminant saliva distinctly alkaline with pH upto 8.1 and this high level of HCO3
and HPO4 help to neutralise the VFA in the fore-stomach and helps to maintain normal rumen pH.
• Salivary urea is a source of nitrogen for microbial protein synthesis.
MICROBIOLOGY AND FERMENTATION IN RUMEN
• About 70 to 85% of digestible dry matter of the diet is digested by microorganisms present in the rumen; this results
in the production of volatile fatty acids (VFAs or short-chain fatty acids-SCFAs), CO2, CH4, ammonia and microbial
cells.
• Rumen microbes digest all major carbohydrates (CH 2O) of ruminant diet such as cellulose (not digested by
mammalian enzymes), starch and other CH2O.
• The microorganisms can synthesise proteins and B complex vitamins required for their growth and metabolic
activities from carbohydrates, organic acids, NH3 and minerals and these microbes pass out of the rumen and get
digested in the host animal’s digestive tract. Thus the ruminant animal can be maintained on diets free of essential
amino acids.
• The gases produced in the rumen are CO 2 and CH4 with small amounts of N2, H2, and O2. The fluid portion of the
rumen (rumen liquor) contains NH3, SCFA, CO2 or HCO3.
• The pH of the rumen fluid ranges from 5.8 to 7.0, which decrease after feeding.
• The following conditions in rumen help in the development of rumen microbes.
1. Frequent intake of food by the animal, providing a regular supply of substrate for micro-organisms

27
 2. Soluble products of microbial activity are readily absorbed through the rumen wall and therefore do not
accumulate and inhibit the microbial growth.
3. Temperature of the rumen is maintained at 38-42C.
4. Volume of rumen contents is regulated by passage at intervals of liquid materials to the omasum through retculo-
omasal orifice.
5. Ruminants secrete a large volume of saliva which is rich in HCO 3 and HPO4. Saliva maintains rumen fluid
volume and steady pH through buffering action.
• Rumen contains many species of micro-organisms.
• Anaerobic ciliate protozoa and non-spore forming anaerobic bacteria and anaerobic fungi are the major microbes
in rumen.
• The microbes have a volume of 3.6% of strained rumen fluid and this volume contains 50% ciliate protozoa and
50% bacteria.
• Metabolic activity of bacteria is far greater than protozoa though total volume of small bacteria might be same as
ciliate protozoa, which is due to greater surface area provided by the bacteria.
Life span No. / ml rumen fluid % of total microbial mass
Bacteria 20-30 min (amylolytic)
1 – 1010 50-90
18 h (cellulolytic)
Protozoa 6-36 h 4 – 105 10-50
Fungi 24 h 1 -104 5-10
• Most of the rumen bacteria and protozoa are strict anaerobes, although some facultative bacteria are also present
which are found attached to the rumen wall.
• Microorganisms exist in the fluid phase and attached to feed particles and some attached with rumen epithelium
• Number of bacteria is higher in animals maintained in green pasture than those fed dry rations. When ciliate protozoa
are absent, viable bacterial count increases. Rate and method of feeding also affects bacterial count. Bacterial count
is also affected by frequency of feeding, time after feeding, composition of the diet, species of the animal, individual
differences etc.
Classification of Rumen Bacteria
1) Major Cellulolytic Species
1. Fibrobacter succinogenes
2. Ruminococcus albus & R. flavifaciens
3. Butyrivibrio fibrisolvens
2) Major Hemicellulolytic Species
1. Butyrivibrio fibrisolvens
2. Ruminococcus sp. &
3. Prevotella (Bacteroides) ruminicola
3) Major Pectinolytic Species
1. Butyrivibrio fibrisolvens
2. Bacteroides ruminicola
3. Succinivibrio dextrinosolvens
4) Major Amylolytic Species
1. Ruminobacter(Bacteroides) amylophilus
2. Streptococcus bovis &
3. Succinomonas amylolytica
5) Major Ureolytic Species
1. Succinivibrio dextrinosolvens
2. Selenomonas sp. & Butyrivibrio sp
3. Bacteroides ruminicola
6) Major Methane producing Species
1. Methanobrevibacter ruminantium

28
 2. Methanobacterium formicium
3. Methanomicrobium mobile
7) Major Sugar-Utilising Species
[Link] bryanti &
2. Lactobacillus sp.
8) Major Acid-Utilising Species
1. Megashaera elsdeni
2. Selenomonas ruminantium
9) Major proteolytic Species
1. Bacteroides amylophilus & B. ruminicola
2. Butyrivibrio fibrisolvens
3. Streptococcus bovis
10) Major Ammonia Producing Species
1. Prevotella ruminicola
2. Megashaera elsdenii
3. Peptostreptococcus anaerobius
11) Major Lipid-Utilising Species
1. Anaerovibrio lipolytica
2. Butyrivibriofibrisolvens &
3. Micrococcus sp.
• Amylolytic bacteria act on starch and soluble carbohydrates in the feed but unable to breakdown structural cell
wall carbohydrates like cellulose; produce acids like lactic acid which depresses rumen pH but these bacteria are
more tolerant to acidic pH.

• Cellulolytic bacteria break down cell wall carbohydrates–cellulose, hemicelluloses, fructosans and pectin; mainly
produce SCFAs. These bacteria are sensitive to low pH and their number is reduced at pH of below 6

• Proteolytic bacteria degrade feed proteins to peptides and amino acids which are engulfedby these bacteria to
produce VFAs and ammonia

• Methanogenic bacteria reduce CO 2 to CH4

Rumen Protozoa
• Many different species of protozoa are active in rumen.
• All of the protozoa are strict anaerobes and found only in ruminants.
• Ciliates form the majority of protozoa and flagellates are present to a very limited extent.

Family 1) Isotrichidae (holotrichs) 2) Ophyroscolecidae (oligotrichs)

Genus a) Isotricha b) Dasytricha a) Entodinium b) Diplodinium c) Epidinium d) Ophyroscolex

Isotrichidae - the entire body surface is covered by cilia; Orphyroscolecidae - most of the body
surface is naked apart from regions where cilia are concentrated in tufts or syncillia
• The ciliates belong to two families.
• Most of the rumen protozoa are found attached with feed particles which prevent their wash out to intestine.
• The protozoa obtain energy for their growth by fermentation of CH2O (ferment major plant constituents including
cellulose, hemicellulose, pectin, soluble sugars, starch) with production of acetic, butyric and lactic acids and CO 2,
H2 gases. Small amounts of propionate are also produced.
• They also hydrolyse lipids, hydrogenate unsaturated fatty acids and degrade proteins;
• They feed on bacteria by engulfing the bacteria for obtaining N for growth.
• Protozoal number is affected by pH (<5.5), type and composition of diet, season and frequency of feeding. Highly
digestible diets increase their number.
29
• Protozoa store large quantities of reserve starch-like polysaccharide which is used when exogenous energy supply
is exhausted. Most of the protozoa are associated with fibrous raft in the rumen.
Role of Protozoa
• Defaunation (removal of protozoa) does not affect the performance of ruminants adversely. The protozoa have the
ability to slow down the digestion of rapidly fermentable substrates like starch and soluble proteins. Protozoa ingest
starch, proteins and PUFA and store them protected from bacterial action. These substrates are digested by the
intestinal enzymes of the host when protozoa are washed out of the rumen to the lower digestive tract. Thus the
protozoa delays or prolongs the digestion of starch and soluble proteins.
• The protozoa feed on rumen bacteria and help to check bacterial over-proliferation especially during feeding of diets
rich in starch.
• By ingesting PUFA, the protozoa protect the PUFA from hydrogenation by bacteria and the PUFA are made
available to the host when protozoa are washed out of rumen.
• Mixed protozoal-bacterial protein is far better quality than bacterial protein alone in contributing essential nutrients
to ruminant animal.
Rumen Fungi
• Organisms originally thought to be flagellate protozoa are now found to be a stage in the life cycle of Phycomycete
fungi.
• The fungi attach to and invade plant particles and may be more important than is suggested by their low
concentrations in the fluid phase of ruminal contents.
• Many anaerobic fungal zoospores are found in rumen.
• Several are flagellate organisms.
• Neocallimastix frontalis, Sphaeromonas communis and Piromonas communis are identified in rumen.
• The fungi are cellulolytic, found attached with plant particles andmay contribute in the digestion of plant cell wall;
• Their number ranges from 105 to 107 per gram of rumen contents. Feeding of high roughage diet increases their
number.
Functions of Rumen Micro-organisms
1. Ferment the dietary carbohydrates including the cell wall constituents, which are not digestible in mammals and
produces SCFA and gases. The SCFAs absorbed from the rumen is the major energy source for the ruminant
animal.
2. Dietary proteins are broken down by the microbes to produce ammonia and branched-chain VFAs. The SCFAs
and NH3 are utilised by the microbes for their growth
3. Triglycerides are hydrolysed to glycerol and fatty acids. The glycerol is converted to propionic acid and
unsaturated fatty acids are hydrogenated
4. Microbes synthesise K and B complex vitamins
5. Microbes passing out of the rumen are utilised by the host animal as a source of protein called microbial protein
and it is an important source of protein to ruminants
Establishment of Bacteria in Young Ruminants
• Development of bacterial flora in young ruminants occurs at a very early age. The nature and rate of development
is affected by type of diet fed and degree of isolation of young from adult animals.
• Under normal conditions, bacteria of adult type establish at about 6 to 8th week of age in young animals. The ciliates
may not become established in young animals unless they are maintained in close contact with animals harbouring
them or are inoculated.
FERMENTATION OF CARBOHYDRATES
• Nearly all dietary proteins and carbohydrates are subjected to fermentative digestion.
• Microbes obtain energy by fermentative breakdown of structural carbohydrates, oligosaccharides and sugars.
• Speed of CH2O fermentation varies with their availability and solubility; soluble sugars are rapidly fermented and
starches are less rapidly fermented. Cellulose and hemi-cellulose are slowly fermented.

30
• In grains, most of the CH2O (starches, fructosans and simple sugars) are nonstructural and intra-cellular. Starches
are glucose polysaccharides with -1, 4 glucose linkage. The fructosans are polysaccharide of fructose units with 
linkage.
• In roughages, most of the CH2O is structural and found in plant cell wall. Plant cell wall is made up of cellulose,
hemicellulose and pectin.
• Three steps are involved in the degradation of CH2O.
Extra-cellular degradation
• The rumen liquor is the best source of bacteria and protozoa and contains several enzymes, which are secreted by
the micro-organisms. These are intra-cellular enzymes that come out of the microbial cell bodies and get mixed with
rumen liquor. The food substances that are soaked in the rumen fluid are acted by these enzymes and are degraded
to short chain oligosaccharides and sugars.
Intra-cellular Degradation
• The short chain oligosaccharides, disaccharides and simple sugars enter the bacterial cell and are further
metabolised by the microbes.
• The oligosaccharides and disaccharides are hydrolysed to simple sugars. The sugars are metabolised by microbes
by phosphorylative cleavage by intracellular enzymes and this leads to the formation of pyruvate, phospho enol
pyruvate, SCFA and CO 2 and CH4. During this process, 2 NAD are reduced to NADH.
• Starch is degraded by bacteria to maltose and some glucose; this maltose is further fermented to glucose.
Fermentation of glucose and other monosaccharides occurs mainly by EM pathway.
• Conversion of hexose to two molecules of pyruvate yields two ATP which is the main energy source for growth and
maintenance of bacteria.
• Cellulose is converted by cellulose enzymes to glucose and then to pyruvate by a complex process. Cellulose exists
in amorphous and crystalline forms; crystalline form is the most difficult to degrade in rumen.
• Hemicellulose is degraded by cellulases similar to cellulose.
• Pectins are degraded to galacturonic acid, methyl esters of galacturonic acid and other sugars. Pentoses produced
from hemi-cellulose and pectin is finally converted to SCFA.
• Pyruvate is the intermediate through which all CH2O must pass before being converted to SCFA, CO 2 and CH4.
• During the process of formation of SCFA from pyruvate, ATP are formed which is utilised by the microbes for their
activities like growth, multiplication etc.
• The SCFA, CO 2 and CH 4 are the by-products that are formed and microbes do not use them; but the SCFA thus
formed are the major energy source for the host animal- ruminants.
Acetic Acid Formation: It is formed by two pathways
1. Oxydative Decarboxylation of Pyruvic Acid
• Pyruvic acid is converted into acetyl-CoA by removal of CO2 and H2, in the presence of thiamine pyrophosphate
(TPP) and lipomide. The acetyl-CoA yields acetic acid.
2. Phosphoclastic Split
• Two molecules of pyruvic acid yield one molecule of acetic acid and formic acid. The formic acid is converted to
CO2 and H2.
Propionic Acid Formation: This is produced by two pathways
1. By CO2 Fixation

31
• CO2 combines with pyruvic acid to form oxaloacetic acid (CH2COOHCOCOOH) which is reduced by hydrogenation
to malic acid; malic acid on removal of water molecule is converted to fumaric acid (CHCOOH CHCOOH). This

leads to the formation of succinic acid by addition of H2; this is decarboxylated to yield propionic acid
(CH3CH2COOH).
2. By acrylate pathway
• Pyruvic acid on dehydrogenation (-2H) forms lactic acid (CHCHOHCOOH) which on removal of water is converted
to acrylic acid (CH2CHCOOH). Acrylic acid on hydrogenation (+2H) yields propionate.
Butyric Acid
• Two molecules of acetyl-CoA condense to form acetoacetyl-CoA and 4 atoms of H. Aceto acetyl-CoA is converted
to  hydroxy butyrl CoA, which produces crotonyl CoA; this gives rise to butyrl COA and ATP. The butyrl COA yields
butyrate (CH3CH2CH 2COOH).
• Formation of propionate leads to regeneration of NAD (required for glycolytic cycle to produce pyruvate from
glucose) and also oxidises the excess NADH [formed during the production of acetate and butyrate] to NAD.
• The production of acetate leads to generation of ATP and formation of excess NADH. NAD is regenerated with
release of free hydrogen and this H is used to reduce CO 2 to CH4 and H2O. Thus there is a direct relationship
between acetate and CH4 production; when more acetate is produced from pyruvate, more CH4 is produced.
• There is a reciprocal relation between propionate production and CH 4 formation; as more pyruvate is directed to
propionate production, CH4 formation is reduced.
End products of Carbohydrate Fermentation
• Due to carbohydrate fermentation in rumen, the final end products formed are
➢ Short Chain Fatty Acids (SCFA / VFA) – acetic, propionic and butyric acids.
➢ Some minor but metabolically important VFAs produced in the rumen are valeric, isovaleric, isobutyric and 2-
methylbutyric acids.
32
 ➢ Gases produced are CO2, CH4 and H2
• The CH4 accounts for nearly 30 to 40% of the total rumen gas production and CO2 accounts for 50 to 60%.
• The SCFA are available as salts such as acetate, propionate and butyrate after they are neutralised by Na salts.
• H+ is one of the end products of carbohydrate fermentation; accumulation of H+ will depress rumen pH leading to
destruction of many microbes thereby affecting fermentation
• the excess H+produced in the rumen is removed by
➢ reduction with CO2 to form CH4 by methanogenic bacteria;
➢ by hydrogenation of unsaturated fatty acids
➢ by propionate production.
• In cattle fed a mixed diet, the proportions of VFAs are
➢ acetate 60-65%,
➢ propionate 15-20%
➢ butyrate 10-15%.
• The ratio of A: P: B ranges from 70:20:10 for high forage diets to
60:30:10 for high grain diets.
• The normal total VFA content of the rumen ranges from 60 to
120mmol/L.
• The total value and percentage of individual VFAs varies due
many factors. The most important are the type of diet, quantity and
quality of feed, health of the animal etc. Propionic acid decrease
in animals fed on hay. Total VFA production is higher in animals fed high starch diet than high fibre diet.
• The rumen ecosystem adopts itself well to the diet and the type of bacteria that is predominant or active is based
on the type of diet.
• If the animals are fed on large quantities of easily digestible carbohydrates like starch or sugar or when animals are
changed rapidly from high roughage diet to high starch diet, lactic acid formation is increased. The increased lactic
acid formation reduces rumen pH to a very low level and suppresses the growth of other bacteria leading to rumen
dysfunction known aslactic acidosis withdehydration, lameness, coma and death.
Uses of SCFA (VFA) in Ruminants
➢ The propionate is glucogenic and contributes glucose through gluconeogenesis by entering the Kreb’s cycle at
the level of succinate.
➢ Acetate and butyrate contributes to the energy needs of the ruminant animal by entering Kreb’s cycle as acetyl-
CoA. They are also ketogenic and leads to the formation of ketone bodies –acetone, acetoacetic acid and -
hydroxy butyric acid. The ketone bodies can serve as energy source in certain tissues like CNS and heart.
➢ The acetate is the precursor for milk fat synthesis.
PROTEIN DIGESTION IN RUMINANTS
• The rumen contents have proteolytic activity which is contributed by the rumen microbes
• The diet of ruminants contains proteins and non-protein nitrogenous (NPN) substances such as amino acids,
ammonia, nitrates, urea etc.
• Of the total proteins entering the rumen, about half of the dietary protein will be degraded to ammonia
• That part of protein degraded in the rumen is called as rumen degradable protein (RDP).
• The fraction not degraded in the rumen “escape” or “by-pass” the rumen digestion, which is called as rumen
undegradable protein (UDP); this fraction reaches the small intestine for digestion.
• Bacteria are important in proteolytic activity and protozoa metabolize mainly the bacterial protein.
• When proteins enter the rumen, the following important events occur by microbial metabolism.

33
• As proteins enter the rumen, the RDP fraction is attacked by extracellular microbial proteases;majority of these
enzymes are endopeptidases that
produce short-chain peptides as end
products.
• The peptides produced extracellularly are
absorbed into the microbial cell bodies.
• Within the microbial cells, the peptides
can either be used for the formation of
proteins to be used microbial growth and
maintenance or further degraded for the
production energy through VFA
pathways.
• To enter VFA pathway, the amino acids
are deaminated to yield ammonia and a
carbon skeleton; the carbon structures of
many amino acids fit directly into various
steps of the VFA pathways, leading to the
formation of three major VFAs.
• Three branched-chain amino acids lead
to the formation of branched-chain VFAs
as follows
Valine + 2 H 2O→ isobutyrate + NH3 +
CO2
Leucine + 2 H2O→isovalerate+NH3 +
CO2
Isoleucine + 2 H2O → 2-methyl butyrate + NH3 + CO2
• These branched-chain VFAs are also called as isoacids, are important growth factors for many species of bacteria.
• Some proteins are converted to amino acids extracellularly. These amino acids are absorbed into the microbial cells
and are utilised by the bacteria; they are directly incorporated into the microbial protein or microbial cell walls or in
their nucleic acids.
• Some of the amino acids are destroyed by fermentative deamination with the production of NH 3, CO2 and SCFAs.
Ex. R-CH-NH2-COOH +2H 2O → RCOOH + CO 2 + NH3 + 4H
• Rumen fluid has urease activity; urea entering the rumen from saliva/blood and dietary source is hydrolysed to NH3
and CO2.
• Compounds such as nitrates are reduced to NH3
• Ammonia is utilised by the bacteria in the synthesis of microbial cell proteins. ATP obtained from CH 2O fermentation
is used by the microbes for this synthetic process.
• The NH3 from the rumen can also be absorbed through the rumen wall into the blood. Just like simple-stomached
animals, NH 3 can also arise from deamination of amino acids in the liver. The NH 3 is converted to urea (from general
detoxification process in all animals) and this may be excreted through kidneys or pass on to rumen via blood directly
into rumen or via saliva
• Ammonia is the principal soluble nitrogenous constituent of the rumen fluid.
• Normal range of NH3 in animals fed mixed ration varies from 5 to 25mg/100ml rumen liquor.
• If the concentration of NH 3 is in excess of the utilisable level, it is wasted and if the concentration is very high, it may
be toxic to the animal (urea toxicity).
• Concentration of NH3 in the rumen fluid is influenced by various factors.
➢ By the quantity of dietary protein
➢ By the solubility or quality of protein
➢ The quantity of urea that enters rumen through saliva

34
 ➢ The quantity of urea that diffuses into the rumen through the rumen wall
➢ Rate of ammonia absorption
➢ Quantity and quality of carbohydrates in the diet – synchronising energy and nitrogen release in rumen
• The microbial cell bodies, after death, passes out of rumen along with the fluid portion and also as attached with
particulate matter leaving rumen. In the intestine of the ruminant animal, these microbial cell bodies are digested
just like any other protein. This microbial proteinis a very important source of protein supply to ruminant animals.
• The microbial protein formed from amino acids in the rumen is highly digestible and has a high biological value.
• The microbes build their body protein, grow and multiply utilising energy obtained from carbohydrate fermentation.
Hence, the amount and quality of dietary carbohydrates play a vital role in the synthesis of microbial protein.
• Many rumen bacteria prefer to use NH3 even when peptides and amino acids are present, showing that quality of
dietary protein to ruminant animals is not very important for growth and multiplication of microbes. The conversion
of NH3 to microbial protein is the most important activity of the microbes. Ammonia as main nitrogen source is
utilised to synthesise amino acids by reverse catabolic process.
RCOOH + CO2 + NH3 + 4H →R-CH-NH2-COOH +2H 2O
• Urea is hydrolysed to NH3 and this NH3 is also utilised by the rumen microbes for its microbial protein synthesis
when sufficient energy is available. Thus, urea can be fed to ruminant animal up to 30% of total N. Urea feeding is
an effective way to increase the protein value of the ration when the feed protein content is very low; to provide
readily accessible energy for utilizing the NH3 from urea, addition of molasses has been found to be efficient
DIGESTION OF LIPIDS IN RUMEN
• Dietary lipids occur as structural lipids in forages and storage lipids in oilseeds.
• In forages, less than 50% of total lipids are free fatty acids and the majority is phospholipids. In oilseeds, 65-80% of
lipids are free fatty acids.
• Both rumen bacteria and protozoa possess lipase activity and hydrolyse the dietary lipids to glycerol and free fatty
acids. The glycerol is converted by the microbes to propionic acid.
• The dietary free fatty acids and the liberated fatty acids are not further broken down;
• The unsaturated fatty acids linolenic (18:3), linoleic (18:2) and oleic (18:1) are hydrogenated by bacteria into trans
unsaturated fatty acids and saturated acids (stearic acid 18:0).
• The bacteria are capable of synthesising long-chain fatty acids which are in trans form (most of the dietary fatty
acids are in cis form) and phospholipids from SCFAs.
SYNTHESIS OF VITAMINS
• Vitamin K and many B complex vitamins are synthesised in rumen (e.g.) thiamine, riboflavin, nicotinic acid,
pantothenic acid, cyanocobalamine, pyridoxine and biotin. They are absorbed from the rumen after their production.
• When a diet is deficient in cobalt, synthesis of vitamin B12 will be inadequate.
ABSORPTION FROM THE FORESTOMACH
• The epithelium lining the rumen, reticulum and omasum is stratified and sqamous. Absorption in the first three
compartments is more rapid than in the [Link] ruminal papillae increase the absorptive surface area.
• The SCFAs are absorbed from rumen, reticulum, and omasum and in case of monogastric animals from caecum
but not from the abomasum and small intestine. 70-85% of the VFAs are absorbed from the reticulorumen and most
of the remaining acids are absorbedform the omasum; only a very little of VFAs reach abomasum
• The rate of absorption of acetic, propionic and butyric acids increases as the rumen pH is decreasing (i.e. as rumen
acidity increases, so does the rate of VFA absorption). The rate of absorption of butyric acid is > propionic, which is
> acetic acid.
• Lactic acid absorption is only about 10% of the absorption rate of VFAs. Hence, large amounts of lactic acid that is
produced during excessive feeding of cereals or foods rich in sugars accumulates in the rumen, lowers the rumen
pH leading to acidosis
• Most of the acetate is absorbed unchanged with a small amount metabolized to CO2.
• All of butyrate is converted to -OH butyrate in the ruminal epithelial cells before absorption.

35
• About 30% propionate is converted to lactate in rumen epithelia cells
• In the liver, boththe lactate and propionate are converted to glucose.
• Ammonia is also absorbed from rumen;
Probiotics
• Probiotics are dietary supplements of live microorganisms thought to be healthy for the host organism. Lactic acid
bacteria (LAB) and bifidobacteria are the most common types of microbes used as probiotics in human and animals
(dogs). In ruminants,cultures of yeast - Saccharomyces cerevisiae are used as probiotic. The yeasts absorb starch
molecules thereby reducing the availability of starch for lactate-producing bacteria. Less lactate is produced, ruminal
pH does not fall very low, cellulolytic activity is not affected; the added yeast also increased protein mass in rumen
– all these activities favour increased VFA formation and microbial protein synthesis and efficiency of feed utilization
is improved
Buffers
• Addition of buffers to ruminants diet especially in grain based diet modify rumen fermentation, resists changes in
ruminal pH and increases outflow rate from rumen; these activities increases fibre digestion
• Sodium bicarbonate, calcium carbonate, bentonite (aluminium silicate clay) and magnesium oxide are generally
added as buffers in ruminants feed.
RUMEN DYSFUNCTION
Disorders of motility
• High levels of ruminal acidity (pH 5.8 or lower) as that occur in ruminal acidosis during excess grain feeding reduces
reticulo-ruminal motility even leading to stasis and this is due ot depressionof epithelial receptors in rumen by the
acidity.
• During severe bloat or ruminal impaction motility is depressed
• Loss of vagal innervations greatly depresses motility called vagal indigestion; this may occur due to accidental
ingestion of sharp objects like nails, needle etc that gets lodged in the reticulum and due to normal reticular
contractions,
• During the initial stages of milk fever, rumen motility is depressed
Ruminal acidosis
• Feeding high level of grain or concentrate to ruminants reduces rumination activity which results in lesserexcitation
of buccal receptors for reflex salivary secretion; accumulating VFAs in the rumeninhibits acid-sensitive rumen
epithelial receptors – both of these activities reduces ruminal motility. Accumulation of acid products and absence
of salivary buffers finally lead to ruminal stasis.
• When rumen pH falls below 6.2, propionate-producing bacteria are inactivated, lactic acid (stronger acid)
accumulates in the rumen, and rumen pH falls progressively lower. When pH drops to less than 5.5, amylolytic
bacteria are also inhibited and at this pH, Lactobacilli which produces lactic acid, becomes the dominant bacteria in
the rumen which further depresses rumen pH to less than 4.6
• The accumulated end products increase osmolality of ruminal fluid which drawswater into the rumen leading to
tissue dehydration and diarrhoea. Higher rumen acidity give rise to metabolic acidosis and also breaks down rumen
epithelium allowing anaerobic bacteria to enter into the systemic circulation; this may cause liver abscesses or
systemic toxaemia.
• This condition is clinically called as ruminal acidosis.
Ketosis
• Ketosis is a condition in which there is there is an increase in the concentration of ketone bodies especially acetone
and aceto-acetic acid in the body fluids including blood, milk and in the expired air

• It typically occurs in dairy cows especially high producing animals in early lactation near their peak production i.e.
4-6 weeks postpartum and is characterized by partial anorexia and depression and progressive decline in milk yield
• Ketosis is accompanied by hypoglycaemia; this occurs in starvation(starvation ketosis), loss of appetite (secondary
ketosis due to anorexia), late pregnancyalong with underfeeding (twin-lamb disease or pregnancytoxaemia of ewes)
and during peak lactation in high producing cows (acetonemia)

36
• During early lactation, the high producing animals are in negative energy balance which leads to adipose
mobilization and lactose of milk synthesis creates a high glucose demand. Adipose mobilization is accompanied by
high blood serum concentrations of nonesterified fatty acids (NEFA).

• In ketosis high serum concentrations of NEFA and ketone bodies and low concentrations of glucose are observed.
In contrast to many other species, cattle with hyperketonemia do not have concurrent acidemia. The serum ketone
bodies are acetone, acetoacetate, and β-hydroxybutyrate (BHB).
• Pregnancy toxaemia occur just before parturition in ewes that carry more than one fetus and its occurrence is
precipitated by stress like underfeeding or severe cold
• The hypoglycaemia leads to decrease in insulin and increase in glucagon secretion which enhances lipolysis in
adipose tissue with β-oxidation and gluconeogenesis in liver
• Glucagon activates hormone-sensitive lipase of adipose tissue which increases NEFA in blood
Bloat
• Bloat is a condition wherein there is accumulation of gas in the reticulo-rumen and it mainly occurs due to failure of
eructation
• There are two forms of bloat – simple (free gas) bloatin which gas is present as a separate layerand frothy bloat
where gas is trapped in the rumen liquor as gas bubbles
• Simple bloat can occur when there is an obstruction to esophagus by foreignobjects or compression of esophagus
by some tumors. Absence of rumino-reticular motility as occurring during ruminal acidosis, impaction, abomasal
displacement etc also causes simple bloat
• In frothy bloat, gas is trapped in the fluid forming stable foam. This can occur in animals consuming fresh, young
legumes (clover, alfalfa) – leafy, young, fresh legumes are more prone to produce stable foam than stem, old or
wilted legumes.
CHEMICAL DIGESTION OF NUTRIENTS IN DOMESTIC ANIMALS
• Chemical digestion of each major nutrient is achieved by hydrolysis.
• Hydrolysis is splitting of a chemical bond by insertion of a water molecule.
• Glycosidic bonds in carbohydrates, peptide bonds in proteins, ester bonds in fats and phophodiester bonds in
nucleic acids are all cleaved by hydrolysis during digestion.
• Hydrolysis in the digestive tract is catalysed by the action of enzymes.
• There are two classes of digestive enzymes:
➢ Those that act within the lumen of gut
➢ Those that act at the surface of epithelium.
• Enzymes acting within the lumen originate from GI glands (salivary, gastric and pancreas). The secretions of these
glands are mixed with ingesta and produce their effect in the lumen of gut associated with their location. Thus the
actions they catalyse are referred to as luminal phase of digestion. This phase of digestion results in incomplete
hydrolysis of nutrients and produces short-chain polymers of original macromolecule.
• The enzymes bound with surface epithelium of small intestine break the short-chain polymers produced from luminal
phase of digestion into monomers which can be absorbed across the epithelium; this final phase occurring at the
surface membrane of epithelium is referred to as membranous phaseof digestion. This phase is followed by
absorption.
• Enzymes that hydrolyse substrates in the membranous phase of digestion are chemically bound to surface
membrane of the intestine; hence, the substrate must be brought in contact with the membrane-bound enzymes
present in the epithelium. These enzymes are synthesised within the enterocytes and transported to the apical
membrane. They project from the apical membrane into the glycocalyx layer; the membranous phase of digestion
occurs in this quite unstirred layer.
• The products from membranous phase of digestion never diffuse back into the lumen of gut; instead they are
absorbed soon after formation into the epithelial cells.
Carbohydrate Digestion

37
• Dietary carbohydrates are mainly starches, disaccharides, monosaccharides and fibrous carbohydrates (cellulose,
hemicellulose).

• Glucose, galactose and fructose are the most important monosaccharides in animal diets.
• Dietary disaccharides include lactose and sucrose; other sugars like maltose, isomaltose and maltotriose are
seldom present as preformed in diet but formed in the gut during carbohydrate digestion.
• Starch is present as amylose and amylopectin;
• Amylose contains glucose monomers linked by -1, 4 glycosidic linkage.
• Amylopectin also contain glucose linked by -1, 4 glycosidic linkage but chains are branched, having -1, 6
glycosidic linkage at branch points.
Luminal phase of digestion
• In luminal phase of digestion, -amylase hydrolyses starches to yield oligosaccharides (alpha dextrins, maltotriose,
maltose but no free glucose); fibrous carbohydrates are not digested by mammalian enzymes.
• -Amylase arises from pancreas of all species and salivary glands of some species. Hydrolysis of amylopectin
produces branched-chain oligosaccharides, known as limit dextrins and -1, 6-linked disaccharide, known as
isomaltose.
Membranous Phase of Digestion:
• The oligosaccharides and di- and tri-saccharides are hydrolysed in the outer part (luminal side) of the membrane of
the microvilli by membranous phase enzymes – maltase, isomaltase, sucrase and lactase. These specific
saccharidases in the glycocalyx of enterocytes hydrolyse the oligosaccharides to monosaccharides (glucose,
galactose, and fructose).
• Monosaccharides are absorbed into the portal blood and carried to liver although lymph stream removes some
sugars from the alimentary canal.
Protein Digestion
• Proteins (dietary, endogenous, microbial and shed mucosal cells) are hydrolysed by proteolytic enzymes present in
gastric, pancreatic and intestinal juices to amino acids.

38
• Hydrolysis occurs in the lumen of GI tract by endopeptidases and at the surface of mucous membrane by

Enzyme Action Source Precursor Activator

Pepsin Endopeptidase Gastric Pepsinogen HCl, pepsin
glands
Chymosin (Rennin) Endopeptidase Gastric Prorennin HCl
glands
Trysin Endopeptidase Pancreas Trypsinogen Enterokinase, exopeptidases.
trypsin • In luminal
Chymotrypsin Endopeptidase Pancreas Trypsinogen Trypsin phase of
digestion, gastric
Elastase Endopeptidase Pancreas Pro-elastase Trypsin and pancreatic
Carboxypeptidase A Exopeptidase Pancreas Procarboxypeptidase A Trypsin proteases yield
short-chain
CarboxypeptidaseB Exopeptidase Pancreas Procarboxypeptidase B Trypsin peptides and
some amino
acids. These oligopeptides are hydrolysed by oligopeptidases in the glycocalyx to produce amino acids, di- and tri-
peptides.
• Most proteolytic enzymes are endopeptidases (break proteins at internal peptide points, resulting in short chain
peptides but do not produce free amino acids).
• Exopeptidases act at the ends of peptide chain to release free amino acids
Luminal Phase of Digestion
• The proteolytic enzymes are secreted from gastric glands and pancreas as inactive zymogens which are activated
in the stomach or intestinal lumen (if secreted in their active form, they would digest the cells in which they are
synthesised).
• Activation of zymogens occurs in gut lumen.
• Pepsinogen and prorennin (chymosinogen) from stomach are activated by HCl. Pepsin also autocatalytically
activates pepsinogen.
• Trypsinogen from pancreas is activated by enterokinase,an enzyme produced from duodenal mucosa. The active
enzyme, trypsin activates other proteolytic pancreatic enzymes and also autocatalytically activates trypsinogen. The
HCl of stomach also has hydrolytic properties on protein.
Membranous Phase of Digestion
• Membranous-phase digestion of peptides is carried outby peptidases present on the enterocyte apical membrane.
These enzymes hydrolyse the peptides produced from luminal phase of digestion, yielding free amino acids.
• Some long-chain peptides are incompletely digested leading to di- and tri-peptides.
• The di- and tri-peptides are easily absorbed by the epithelial cells; these di- and tri-peptides are subsequently
hydrolysed by the intracellular peptidases, forming free amino acids. Thus, free amino acids are produced at two sites:
on the surface of the enterocytes and within the cell
Lipid Digestion
• Lipids make up a large portion of diet of carnivores, whereas they form only a minor portion of diets of herbivores.
• Primary dietary lipid is triglyceride (mainly contain long-chain fatty acids–16-18C); other lipids include cholesterol
and cholesterol esters from animal sources, waxes from plant sources and phospholipids from both plant and animal
sources.
• Lipid digestion occurs in four phases;

39
 ➢ emulsification,
➢ hydrolysis,
➢ micelle formation
➢ absorption
• Emulsification is reducing lipid droplets to a smaller size so that they form suspension in water. In the gut, this phase
begins in the stomach as lipids are warmed to body temperature and subjected to mixing and agitating actions of
distal stomach. Due to this action, lipid globules are broken down to droplets.
• Emulsification is completed in the small intestine due to the detergent action of bile acids and phospholipids.
• These bile acids reduce the surface tension of the lipid droplets which become further reduced in size.
• The pancreatic lipase can breakdown fat only when it comes in contact with lipid molecules. Lipase is water-soluble
but triglycerides are lipid-soluble and degradation can occur only at the interface between water and fat.
• Bile salts contain both fat-soluble portion and anegatively charged water-soluble portion. The lipid soluble part of
bile salt is dissolved in the fat droplet and the water soluble part protrudes from the surface of the droplet into the
water phase. The fat droplets are broken-down to small droplets by the segmentation contraction of the intestineand
are then covered by bile salts. These small droplets remain small because they cannot coalesceto form large
droplets due to negative charge of the bile salts repel each other
• Hydrolysis: The lipase moves through the water phase of the small droplet to the interface between fat and water
and breakdown the triglycerides
• The pancreatic co-lipase enzyme secreted along with lipase “make a pathway” through the bile-coated emulsified
lipid droplet, giving access to the lipase to reach the underlying triglyceride.
• Lipase cleaves the fatty acids from the end of triglyceride molecule (can not attack central fatty acid), resulting in
formation of two free, or nonesterified fatty acids and a monoglyceride from each triglyceride molecule.
• Other lipid digesting pancreatic enzymes are cholesterol esterase and phospholipase. The products of these
enzymes are nonesterified fatty acids, cholesterol and lysophospholipid.
• Micelles are aggregations of bile acids arranged with their lipid-soluble part oriented towards the centre and water-
soluble negatively-charged part directed outwards. The fatty acids, monoglycerides etc of the lipid digestion combine
with micelles and taken towards enterocytes for absorption. Micelles are considerably smaller than emulsified lipid
droplets. The soluble micelles allow the lipids to diffuse through glycocalyx and into close contact with absorptive
surface of the enterocytes.
ABSORPTION
• Absorption is the process whereby the digested foodstuffs are transferred from the lumen of the gastrointestinal
tract to the blood and/or lymph.
Sites of Absorption
• There is no absorption of food or end products of digestion in the mouth and oesophagus. Certain drugs may be
absorbed from these places (e.g. strychnine)
• In the stomach of monogastric animals, absorption is very limited under normal conditions since the food substances
are not ready for absorption. Certain drugs (e.g. ethanol) are absorbed from stomach. In the rumen, absorption of
SCFA and NH3 occurs.
• Small intestine is the chief site of absorption in all species of animals.
• The large intestine as an organ of absorption is of limited importance in carnivores and man except in the initial part
of colon where electrolytes and water absorption occurs.
• Large intestine of herbivores helps in the absorption of short chain fatty acids and NH 3.
Absorptive Surface
• Surface area of the small intestine is enormously increased to facilitate contact between mucosa and lumen contents
at three levels:

40
 ➢ First, the mucosa of intestine contains numerous folds known
as plica circularesand these folds increase the intestinal
surface area but they are present in some animals but not in all
species.
➢ Second, mucosal surface area is covered with finger-like
epithelial projections called as villi; they increase the surface
area by about 10 to 14 fold. They are seen in all animals
➢ Third, the villi are covered with brush like surface membrane
known as brush border, which are composed of microvilli that
further increase the surface area.
➢ The overall absorptive surface area of intestine in man is about
200 m2.
• At the base of the villi is gland-like structure known as crypts of Lieberkuhn. The villi and crypts are covered with a
layer of epithelium.
• The epithelial cells covering the villi and the crypts are called as enterocytes.
• The cell surface of enterocytes facing the lumen is called apex and covered with apical membrane. This membrane
contains the microvilli (brush border).
• Covering the microvilli is a jelly-like layer made of
mucopolysaccharides (mucus) and glycoproteins
known as glycocalyx. Digestive enzymes and other
proteins are attached to microvilli and project into the
glycocalyx.
• That part of enterocyte not facing the lumen is called as
the basolateral membrane (the base and sides of cell).
• Nutrients absorbed into enterocytes exit the cell
through basolateral membrane before entering into
blood.
Routes of Absorption
• Small intestine has extremely well developed blood and
lymphatic system which function in absorption of
products of digestion.
Lymph
• In the core of the villus, a large lymph capillary known
as lacteal is present. This capillary begins near the tip
of the villus and enters a plexus of lymph vessels on
the submucosa.
• The lymph capillaries arising from the villus of the
intestine drain into large lymph vessels of submucosa
which empty into the lymph vessels of mesentery;
these mesenteric vessels are connected with mesentric
lymph nodes. The lacteal vessels of mesentery proceed to and empty into the cisterna chyli. This vessel is continued
forward as thoracic duct, which empties into the venous system anterior to heart.
• Monoglycerides, long chain fatty acids, cholesterol and certain proteins - particularly the immunoglobulins during
the first 24 hours of life- is absorbed by the lymphatic system. The rate of lymph flow increases after a meal.
Blood
• Each villus contains several small arteries which enter the base of the villus and form a dense capillary network
immediately under its epithelium. Near the tip of the villus, one or two veins arise from a capillary network and run
downward.
• The blood capillaries of mucous membrane of the intestine including those of the villi unite to form venules and veins
which drain into the portal vein.
• The portal vein enters the liver where its blood is mixed with that of hepatic artery.

41
• The hepatic vein conveys the blood of liver to the posterior vena cava.
• Materials absorbed by blood includes amino acids of protein digestion, monosaccharides of carbohydrate digestion,
free glycerol of fat digestion, water, inorganic salts and short chain fatty acids. The rapid flow of blood allows efficient
absorption and rate of blood flow increases after a meal, but the increase is less than that of lymph.
Mechanism of Absorption
• The possible mechanisms of absorption are broadly classified into three groups:
1) Non-carrier Mediated transport – it is the passive diffusion that depends on the osmotic pressure and electro-
chemical gradient. It occurs through specific ion channels or transport pathways. The transport pathway involves
tight junctions [where epithelial cells join together] which are freely permeable to water and small inorganic ions.
The movement of materials through the tight junctions is called paracellular (around the cells) absorption.
Substances are also absorbed through the apical cell membrane which is called transcellular(through the
cells)absorption. Non-carrier mediated process aids in the absorption of water, short chain fatty acids, inorganic
salts and lipid soluble compounds.
2) Carrier mediated transport – includes
➢ Facilitated transport – which occurs in the direction of higher to lower concentration
➢ Exchange diffusion – Na+/H+ exchanger: transports H+ out and Na+ into the cell
➢ Active transport – occurs against concentration difference and energy dependent
• The carrier-mediated process may help water-soluble materials to pass the lipid layer of the cell membrane.
Glucose and amino acids are absorbed by active transport.
3) Pinocytosis – transport of intact luminal materials in vacuoles into the mucosal cells. Pinocytosis is important
for absorption of intact proteins and intact triglycerides.
Absorption of Lipids

• As the micelles [aggregates of bile salts] come in contact with emulsified fat droplets, they take-up monoglycerides,
medium- and long-chain fatty acidsand dissolve them at the middle of the micelle. These micelle with fatty acids
move at the space between the microvilli,
come in close contact with surface of
enterocytes,, thelipid components diffuse
through the glycocalyx to the apical
membrane and the long-chain fatty acids are
transported by binding with special fatty acid
binding proteins; these binding proteins
transport the long-chain fatty acids across the
cell membrane. Other components in the
micelle such as monoglycerides, cholesterol
and vitamin A diffuse into the apical
membrane (cell membrane composed of
phospholipids and products of lipid digestion
pass through them easily).
• The bile salts of the micelle remain in the gut,
and available for transport of additional
molecules of fatty acidsfor absorption. In the
ileum, bile acids remain in a free state devoid
of other lipids.
• In the ileum, specific bile acid transport system is available, which is a Na-cotransport system. This results in nearly
complete absorption of bile salts.
• After absorption, the bile salts are transported to liver by the portal blood. The liver extracts bile acids and maintains
the normal concentration of bile acids in systemic blood. The extracted bile acids are recirculated into bile; this
process [enterohepatic bile salt circulation] occurs repeatedly.
• Glycerol is absorbed by passive diffusion to enter the mesentric venous blood.
• Short chain fatty acids up to C10 are water-soluble and are absorbed into mesentric portal blood.
• Monoglycerides and long chain fatty acids enter the microvilli and pass on to the lacteal by simple diffusion.

42
• Cholesterol is readily absorbed from the small intestine. Presence of hydro cholesterol and or plant sterols inhibits
cholesterol absorption. In epithelial cells, cholesterol is reesterified before their transfer to lacteals.
• Phospholipids are hydrolysed by phosphoilpase enzymes of pancreas and intestinal epithelium to free fatty acids
and lysophospholipids and absorbed as such.
• Within the epithelial cells, long chain fatty acids are converted into fatty acyl-CoA involving co-enzyme A and ATP.
The fatty acyl co-enzyme reacts with monoglycerides to form di and triglycerides. The newly formed triglycerides
may differ from dietary fat. Glycerol PO 4 derived from glucose metabolism provides glycerol residue for the
triglyceride synthesis. In addition, phospholipids and cholesterol esters are produced in the epithelial cells. Small
amounts of proteins are added to the lipid droplet before their transfer from epithelial cells to lymph. The finished
products are called chylomicrons (containing high amount of triglycerides, low level of phospholipids, cholesterol
esters and proteins), which leave the cell by reverse pinocytosis and enter the lacteals.
• Lipid absorption begins in the distal duodenum and completed in the proximal jejunum. The absorbed fat is in the
form of an emulsion and imparts a milky appearance to the lymph, which during absorption is called “chyle”. The fat
in the chyle are the “chylomicrons”.
Absorption of Carbohydrates
• Monosaccharides especially glucose and galactose are absorbed by active transport by expending energy which
involves a carrier and sodium pump. Glucose absorption is taken as an example of absorption of monosaccharides.
Absorption of Glucose
• As glucose is produced by enzymatic digestion, it is attached to specific transport proteins that lie on the luminal
side of the enterocytes.

• These transport proteins have binding sites for both glucose and sodium. Once glucose and Na+ occupy the binding
sites, the transport protein moves across the cell membrane by active transport (expending energy)and dislodges
glucose and Na+ into the cell. In this process glucose is transported against concentration gradient
• The transport of glucose will not occur unless Na+ is present. Hence, this process is referred to as sodium co-
transport (symport). The carrier that helps the glucosetransport is the sodium dependent glucose transporter (SGLT)
• When glucose enters the enterocytes, its concentration increases within the cell and it moves down concentration
gradient from within the cell through the basolateral membrane by facilitated diffusion, to transcellular space and
into blood.
• Monosaccharides after reaching the liver are converted and stored as glycogen.
• Rate of absorption of different sugars from intestine is variable.
• Galactose is absorbed more rapidly than glucose; fructose absorption is slower than glucose absorption. Fructose
absorption is by facilitated diffusion and not energy dependent. Hence, fructose can not be absorbed against
concentration gradient. Mannose, xylose and arabinose are poorly absorbed by diffusion.
• Short chain fatty acids are absorbed by blood. The SCFAs are found in the blood draining the caecum of sheep and
horse and colon and caecum of pigs.
• Maltose, sucrose and lactose as such are absorbed very slightly. Disaccharides do not generally enter the blood
stream because of the presence of disaccharidases in the brush border of mucosa, which converts them to

43
 monosaccharides. When disaccharides appear in blood as in during mucosal injury, they are eliminated unchanged
in urine.
Absorption of Proteins
• The free amino acids are readily absorbed chiefly by active energy-requiring Na-cotransport system. There are
differences in the rate of absorption among individual amino acids.
• Three types of carriers are involved in transporting amino acids – one for acidic, one for basic and one for neutral
amino acids.
• Some di- and tri- peptides are also absorbed. Intracellular peptidase hydrolyses these peptides to amino acids.

• Intracellular amino acids diffuse across the basolateral membrane to reach the circulation via portal blood.
• Under certain circumstances native proteins may be absorbed due to some defect in intestinal epithelium associated
with age.
• Immunoglobulins from colostrum are absorbed by pinocytosis immediately after birth particularly in lambs, piglets,
kids, calves and pups. The immunoglobulin absorption decreases with time after birth and ceases after 24-36 h.
• Absorption of intact proteins involves the lymph pathway.
Absorption of Salt and Water
• A very large volume of fluid containing enzymes and electrolytes are added to the lumen of gastrointestinal tract by
the accessory glands, stomach and intestine to aid in digestion and absorption. The water and electrolytes must be
recovered from the GI tract to maintain fluid balance of the body.
• The amount fluid secreted into the lumen of GI tract is greater in herbivores than omnivores and carnivores
• In carnivores and omnivores, about 60% of total added fluids are reabsorbed before ileum, and the rest in the distal
ileum and large intestine, whereas in the post gastric fermenting herbivores most of the fluid is reabsorbed in the
distal ileum and large intestine.
• The salt and water are absorbed by two pathways
o One is transcellular pathway that involves moving the salt and water across the epithelial cell membrane into
the cells and then through the basolateral membrane into the extracellular fluid; this pathway involves active
transport, facilitated transport and osmotic drag of water
o The second pathway involves diffusion of salt and water through the paracellular pathway through tight junctions
on the apical cell membrane. Osmotic, hydrostatic and electrochemical gradients favour movement of salt and
water
• Water is absorbed through paracellular or transcellular pathway by osmosis. As electrolytes (Na+, K+, HCO3, Cl-)and
nutrients (monosaccharides, amino acids) are actively absorbed, water is drawn along passively from the lumen of
the intestine to capillaries.
• Na+ ions are involved in water transport, transport of monosaccharides, amino acids, pyramidine and bile salts.
• Na+ is transported in three ways- 1) Na+-cotransport 2) Cl- coupled transport in which Cl- attaches to a carrier just
like glucose 3) Diffusion by electro chemical gradient.
• Chloride ions are transferred in three ways; 1) Cl - coupled Na+ transport 2) in association with Na+ co-transport of
glucose through the paracellular pathway due to electrical charge 3) exchange with HCO3
• K+ is absorbed by passive diffusion through paracellular route due to concentration gradient.
44
• HCO3 absorption is by active transport and exchange with Cl-
• Sodium, K+ and Cl- ions are almost completely absorbed from the intestine
• Intestinal absorption of Ca2+occurs by two processes
• Active transcellular absorption occurs in duodenum in which Ca2+ enters the enterocytes by facilitated diffusion,
binds with a calcium binding protein called calbindin and transported out of the enterocytes into the blood. Vitamin
D induces Ca2+ binding protein and increases Ca 2+ absorption. Passive paracellular absorption occurs in the jejunum
and ileum in which the Ca2+ enters through the tight junctions of the enterocytes into the paracellular space and
then into the blood.
• Mg2+ is absorbed by active transport but its absorption is poor. Absorption of phosphorus is as phosphate and it is
active process related to cotransport with Na+.
• Iron is absorbed in the proximal duodenum and requires acidic pH. Dietary ferric iron (cannot be absorbed) is
reduced to ferrous state by brush border enzyme ferrireductase and enter the enterocytes. Vitamin C increases iron
absorption (reduces ferric to ferrous form)
• Absorption of Fe2+ is related to its level in mucosal cell. When the body requires iron, the absorbed iron enters the
blood and transportedbound with transferring(plasma iron transport protein). When the body’s reserve iron is
adequate, the iron binds with a protein called ferritin (synthesized by enterocytes) and is excreted through faeces.
• Cu2+ is absorbed in small amounts. Co 2+ and Mn2+ are readily absorbed.
• Fat-soluble vitamins A, D, E, K pass through mucosa passively as also water-soluble vitamins. Vitamin B12 requires
an intrinsic factor secreted by stomach for its absorption by active transport.
FUNCTIONS OF LARGE INTESTINE
• Materials that escape absorption in small intestine are gradually propelled through the ileocaecal valve into the large
intestine. The nature of the material varies greatly in carnivores, herbivores and so the significance of large intestine
varies in different species.
• The digestive processes are practically complete in the small intestine.
• Large intestine functions in returning to blood of water and electrolytes that has been poured out by digestive glands.
• It also acts as reservoir for waste material that constitutes the faeces, which are expelled at intervals from the bowel
by the act of defecation
• The large intestine consists of caecum, colon and rectum. The shape and size of large intestine varies considerably
among species.
Large Intestine of Carnivores
• Microbial fermentation in large intestine do not contribute greatly to the energy supply in these animals and hence
their large intestine is small
• Colon is short and nonsacculated. Caecum is poorly developed. Intestinal glands are present throughout large
intestine but there are no villi.
Bacterial Action
• No bacterial action occurs in small intestine of carnivores, since the conditions are not favourable for it.
• Bacteria present in the large intestine act on the ingesta. The bacteria are putrifactive (proteolytic) and cellulolytic
in type.

45
LargeIntestine of Herbivores
• The large intestine of omnivores and ruminants is not very capacious as in rodents and equines. Large intestine is
of great importance in non-ruminant herbivores.
• In all
herbivores, a
digestive tract with
a roomy
compartment
somewhere in its
course is
necessary for
maceration,
fermentation and
solubilisation of
fibrous food. In
ruminants, such a
compartment is
the rumen. In
simple stomached animals, the enormous caecum and colon meet this requirement and hence they are called as
hind gut digesters.
• Digestion of cellulose and other indigestible products, synthesis by bacteria and absorption of end products occur
in large intestine of simple-stomached herbivores similar to rumen fermentation.
Digestive process
• A considerable part of food of simple-stomached herbivores reaches the large intestine in a form not ready for
absorption.
• Digestive changes are brought about by the enzymes carried down from the small intestine and by bacteria and
protozoa. Enzyme action in large intestine is same as action elsewhere.
• The hind-gut maintains conditions favourable for microbial fermentation, which includes substrate availability, control
of pH and osmolality, anaerobiosis, retention of substrate and removal of end products of fermentation. There is
extensive urea recycling in caecum and colon and this supplies N2 for microbial growth.
• Bacterial action in large intestine is mainly on structural and non-structural carbohydrates and proteins. The
digestion of cellulose in large intestine yields products similar to those produced by fermentation in rumen viz. VFA,
CO2 and CH 4. Ammonia is produced from proteins and urea and it is absorbed from large intestine of horse.
• In ruminants, enzyme digestion occurs after microbes have attacked the food material and the host animal also
utilises the microbial cell bodies.
• In simple-stomached herbivores, fermentation digestion follows enzymatic digestion and only fermentation products
and not the microbial cell bodies are available for absorption by the host. The VFAs absorbed from the large intestine
contributes to the energy needs of the host animal. In horses, up to 75% of energy requirements can be met from
VFAs produced from large intestine.
• Volatile fatty acids – acetic, propionic and butyric acids are found in the large intestine of ruminants and horse.
Appreciable quantities are also produced in pigs, rabbit and fowl. Very small amounts are found in large intestine of
cat and dog. Bacterial synthesis of B complex vitamins occurs in caecum and colon.
FAECES
• Faeces refer to the waste matter voided from the bowel through the anus by the act of defecation.
• It is composed of water and undigested residues, bile acids, bile and mucin, shed cells from intestinal mucosa,
numerous bacteria, inorganic salts and products of bacterial fermentation – indole, skatole, etc
➢ Water content of the faeces varies with species; sheep – 68%, horse – 75%, cow – 83%, pig – 80%. Terminal
part of the colon is better developed in sheep than in cow and so it has less water content in faeces.
• Reaction of faeces in horse is acidic–pH 6.5-6.8
• The amount of faeces varies with amount and kind of food
• Amount of faeces is greater in herbivores than in carnivores.

46
• In carnivores and man, the amount of food residue excreted is small and volume of faeces is low since digestion
and absorption are nearly complete.
• In herbivores and pigs, the faeces contain large amounts of food residue and are therefore bulky. Most of the
undigested material is made up of crude fibre.
➢ Amount of faeces voided [in kgs]: Horse – 15-28; Sheep–1-3; Cattle–13-35Swine–0.5-2.5
AVIAN DIGESTION
• The alimentary canal of birds differs from that of mammals in the presence of crop in the oesophagus and in the
presence of gizzard.
• Even among birds, structural variations occur in the digestive tract. Birds that eat seeds and birds that eat leaves
and other plant parts have large caeca to allow fermentation of fibre-rich food. Birds of prey and owls eat feeds high
in energy and have a relatively simple digestive tract and small caeca which may also be absent in some birds.
Structure of digestive tract
• The mouth and pharynx are not sharply delimited in birds, there is no soft palate, teeth are absent in birdsand their
function is accomplished by the horny beak and gizzard.
• The beak in seed-eaters help to crush the feed while in birds of prey, they help to tear the feed similar to molars of
carnivores
• Feed is swallowed immediately after being eaten
• Salivary glands are present in the mouth
• Taste buds are presenton the tongue which are sensitive to salt, bitter and sweet.
• Crop is the sac-like dilatation of the oesophagus and present in most species although absent in some (insectivorous
birds and owl). A sphincter is present at the opening of the crop which opens only after the gizzard is filled.
• The glandular stomach or proventriculus functions primarily in secretion of gastric juice.
• The muscular stomach (gizzard / ventriculus) is highly specialised for grinding and mixing of digestive secretions
with food. The mucosa of the gizzard is covered by a tough coating known as koilin-composed of polysaccharide-
protein complex and it protects the mucosa during grinding.
• The small intestine of birds has a duodenum but the jejunum and ileum are not delimited. The vestige of the yolk
sac (Meckel’s diverticulum) may be found midway in the small intestine. The small intestine is longer in herbivorous
than in carnivorous birds. Villi are like in mammals except that they are taller, more slender and more numerous in
birds.
• The villi contain blood capillaries and there are no lacteals.
• Colon and rectum are very simple. Rectum ends in cloaca which is common passageway for digestive, urinary and
reproductive tracts.
• Located at the junction of the small and large intestine are the caeca which are usually paired. Large intestine is
relatively short.
• Caeca are absent in parrots and pigeons.
• Liver is bilobed and relatively large in most birds, the left hepatic duct communicates directly with the duodenum,
and the right duct sends a branch to the gall bladder.
• The gall bladder gives rise to bile ducts which empty into the duodenum.
• Gall bladder is present in chicken, duck and goose but absent in pigeon.
• The pancreas lies within the duodenal loop and it consists of three lobes and its secretion reaches the duodenum
via three ducts.
Motility
• During swallowing the tongue and hyoid apparatus actively move the food or fluid from mouth into the oesophagus.
Food is moved through oesophagus by peristalsis.
• Crop serves storage function. When chicken eats, the first food ingested goes directly to ventriculus and when
ventriculus is filled with food, the subsequently swallowed food goes to crop.

47
Gastro Duodenal Motility
• The muscular stomach has rhythmic contractions which are neurogenic and the glandular stomach and duodenal
part are dependent upon the intrinsic neural connections of muscular stomach.
• Gizzard helps in allowing the passage of small particles to pass to duodenum and retains large particles for further
grinding. Peristaltic and antiperistaltic contractions occur in the G.I. tract. Birds can vomit.
• Proventriculus and ventriculus undergo 2-3 contractions per min. Ventriculus contraction pushes the ingesta to
duodenum or into proventriculus.
• The gastro intestinal tract of birds is innervated by vagus which is the principal motor nerve to these organs;
stimulation increases motility.
• The gut also receives sympathetic fibres.
• Gritand small stones arepresent in the gizzard which is used for grinding hard food. Grit is not essential for normal
digestion but digestibility decreases without it.
Ileal, Colonic and Caecal Motility:
• Peristalsis and segmentation contractions are present in ileum. Frequency of contractions in ileum is 4/min. Inherent
myogenic slow waves are present in ileum and
regular MMC are also observed.
• Anti peristalsis is most powerful in colon; this
helps (1) the movement of urine from cloaca into
colon and then into the caeca for water
reabsorption (2) for filling of the caeca. Both
peristalsis and anti peristalsis that are myogenic
occur in caeca.
Secretion and Digestion
• Salivary glands of most birds have only mucus
secreting cells. However, serous cells are
present in few species and amylase is present in
poultry. The number and arrangement of
salivary glands vary. Function of saliva is
lubrication of digestive passage up to stomach.
• Crop contains mainly mucus secreting cells
which secret mucous; starch may undergo
digestion in crop either by bacteria or by amylase
of salivary gland.
• Pigeon crop secretes amylase and invertase
(sucrase) and produce fat cells which slough
off during feeding of their chicks. This is called
as pigeon milk.
• Stomach: Mucus secreting glands and compound
glands that secret mucus, HCl and pepsinogen
are present in proventriculus.
• Acid proteolysis occurs mostly in glandular
stomach where mechanical digestion also
occurs. pH of gastric juice is 0.5 to 2.5.
• Gizzard has the function of allowing small particles to pass to duodenum and retaining large particles for further
processing
• Intestinal, Pancreatic and Biliary digestion:Luminal phase of intestinal digestion is carried out by pancreatic
enzymes.
• Membranous phase of digestion of saccharides and peptides is done by enzymes attached with intestinal epithelial
brush border.
• Pancreas secretes trypsin, chymotrypsin and elastase which hydrolyze protein molecules to oligopeptides and
dipeptides. Carboxypeptidase A and B from pancrease aminopeptidases and dipeptidases attached with
enterocyte brush border releases free amino acids.
48
• Luminal digestion of starch (primary carbohydrate in poultry feed) by pancreatic α-amylase releases maltose,
maltotriose and limit dextrins which are then hydrolysed to glucose by membrane bound enzymes (maltase,
isomaltase and sucrase). Lactase is absent in birds.
• Pancreatic juice and bile are rich in HCO 3 which help to neutralize the acid chime in duodenum.
• Bile salts from liver emulsify fats which are hydrolysed by pancreatic lipase. Bile contains amylase. Bile salts are
reabsorbed in the ileum for recirculation through bile.
• Caecal Function: Birds can survive without caeca. Still, caeca helps in microbial digestion of cellulose.
Monosaccharide absorption occurs in proximal caeca. However, nutritive benefit to host from this digestion is
doubtful.
• Absorption of water from caecal contents is important.
• Urine is moved from cloaca into colon from which it passes into caeca for water absorption. Urea and uric acid of
urine helps in microbial growth in caeca. Vitamins are synthesised in caeca but they are not absorbed by the host.
Absorption
• Ileum is the chief site of absorption of fat, carbohydrates and protein.
• Bile is absorbed in the lower ileum.
• Glucose, galactose and fructose absorption is active, which is a Na+ dependent carrier system.
• Amino acids are actively absorbed.
• Fats are absorbed into blood.

49