Dr.

Shaoli Majumder

Stomach
The stomach is basically a dialation of the digestive tract for the temporary storage of food. The
stomach in mammals is transversely arranged and in most forms they take a saclike form.
It is divided into two regions, namely cardiac and pyloric. The cardiac part is adjacent to the
oesophagus and secretes mucus. The posterior part of the stomach leading into small intestine, is
called pyloric region. The point at which the esophagus enters the stomach may be thickened as a
result of the presence of a lot of smooth muscle in the outer muscular layer forming a cardiac
sphincter. Pyloric sphincter (Gr. pyloros, gatekeeper) is found at the posterior end of the stomach
where it joins the duodenum. The sphincter can contract to retain food in the stomach. The inner
concave side of the stomach is called lesser curvature and outer convex surface is called greater
curvature. The sac like bulge lies to the lateral of the cardiac region, called fundus. The inner
surface of the stomach is raised by a number of longitudinal folds, called rugae. The secretion of
the stomach is known as gastric juice. It contains mucus, hydrochloric acid, pepsin, renin and
gastric lipase.

 HISTOLOGY
(i) mucosa: it is the innermost principal layer and consists of a surface epithelium, a deeper
lamina propria, and a muscularis mucosa
(ii) Sub-mucosa layer: this is the second principal layer and is composed of loose connective
tissue containing nerves, capillaries, lymphatic ducts and nodules, and ganglia of the
parasympathetic nervous system.
(iii) Muscularis externa: Is formed of smooth muscles: outer longitudinal muscles, middle
circular and oblique muscles
(iv) Serosa: It is the outermost layer and is formed of squamous epithelium
Dr. Shaoli Majumder

Glands in the different regions:

The glands that are found in the stomach have been named according to their location. The
glands located on the cardiac part of the stomach are called cardiac glands. Those located on the
fundus and pyloric parts are called fundic and pyloric glands respectively. The fundic glands are
of greatest importance in digestion. There are three types of fundic glands.
They are mucous neck cells that produce a lot of mucus, chief or zymogen or peptic cells that
secrete pepsinogen in mammals and parietal cells that produce hydrochloric acid. In addition
there are G cells that secrete a hormone called gastrin, which stimulates the parietal cells and
overall gastric secretion.

 STOMACH IN DIFFERENT GROUPS:

1. Agnatha
Living agnathans have no definite stomach .The digestive tract one long tube from mouth to vest
exhibiting no gross differentiation of esophagus, stomach, or intestine. The epithelium of
agnathan tract is a single layer of cells including goblet cells that secrete mucus and flask shaped
cells that synthsise proteolytic (protein splitting)enzymes. The base of each cell is in contact with
the underlying vascularised layer of mucosa from which they receive nourishment.
2. Fishes
The stomach of fishes display a wide variety of shapes: it is usually either stiaght or bent into a J
or U shaped. The gar stomach is almost straight, sharks exhibit the more common j shaped. The
entire stomach of some teleosts is one large ceacum. Chimeras and lungfishes lack stomach since
they swallow only finely divided food.
3. Amphibian and Reptiles
The stomach remains simple and straight or gently curved in most most amphibians and reptiles,
but is rounded and very much muscular in crocodilians.
4. Avian
Birds face special challenges when it comes to obtaining nutrition from food. They do not have
teeth and so their digestive system must be able to process un-masticated food. The stomach of
birds has two chambers: the proventriculus or crop that is generally spindle-shaped, where
gastric juices are produced to digest the food before it enters the stomach, and the gizzard (also
known as ventriculus), where the food is stored, soaked, and mechanically ground. There is a
clearly defined constriction between these two structures. The gizzard is designed according to
Dr. Shaoli Majumder

the feeding habits of the bird and resembles a biconvex lens superficially. The wall of the gizzard
has a lot of smooth muscle.
5. Mammals
A true stomach is wanting in monotremes and is represented by a wide sac-like structure. The
inner lining is devoid of gastric glands. In platypus (Ornithorhynchus) the two parts, namely
cardiac and pyloric, are almost fused along the lesser curvature, and appear as a wide sac. In
kangaroos (Macropus), the stomach is a large, sacculated non-glandular chamber and the cardiac
chamber is divided into many but not as in ruminant artiodactyles. In many rodents, lagomorphs,
in some monkeys and in man, the cardiac and pyloric regions are marked by a constriction in
between them. Such stomach is known as hourglass stomach. In rodents and lagomorphs, the
food is passed twice through the alimentary canal (caecotrophy). In the blood sucking bat
Desmodus, the pyloric part has become elongated to form a caecum-like structure for storage of
sucked blood. In pangolins (Manis), the pyloric region of the stomach acts as the gizzard of birds
that contains a quantity of stones. The stomach of whales and hippopotamus is divided into
several chambers. The stomach of herbivorous mammals is comparatively larger and compli-
cated than the carnivores.

Stomach of Ruminating Mammals:

Ruminants are mainly herbivores like cows, sheep, and goats, whose entire diet consists of eating
large amounts of roughage or fiber. They have evolved digestive systems that help them digest
vast amounts of cellulose.
 COW (Bos sp.)
The ruminat stomach of cow is very complex. It is divided into the nonglandular fore stomach
(rumen, reticulum, omasum) and the terminal glandular stomach, the abomasum.
 Origin:
The first three chambers — rumen, reticulum and omasum arise from the oesophagus and only
the fourth chamber, the abomasum is the actual derivative of the stomach.
Dr. Shaoli Majumder

 Anatomy
Rumen:
The rumen is the first and the largest (by volume) compartment of the ruminant forestomach. It
has low muscular folds and mucus membrane is beset with numerous short villi. The internal
lining is non-glandular and contains stratified squamous epithelium. The rumen does not contain
a muscularis mucosa.
The primary function of the rumen is as a storage compartment to facilitate microbial (bacterial
and protozoal) fermentation of ingesta. Volatile fatty acids are biproducts of this fermentation
process and are readily absorbed across the ruminal mucosa into the circulation, where they
serve as a directly accessible energy source. In addition, microbial digestion of fibrous ingesta
begins the digestion necessary to access nutrients from ingesta that will undergo further
breakdown in more distal parts of the gastrointestinal tract (e.g. abomasum).
Reticulum:
The chamber is much smaller than the first. The mucosal surface of the reticulum is composed of
long primary and shorter secondary folds that form a grossly apparent honeycomb-like pattern.
As in the rumen, the reticular mucosa is composed of a keratinizing stratified squamous
epithelium. The primary folds (large papillae) of the retriculum contain bundles of smooth
muscle within the lamina propria, also referred to as the muscularis mucosa. It largely acts to
further mechanical dissociation of digesta following microbial breakdown.
Omasum or Psalterium:
The mucosal surface of the omasum is composed of large, thin, leaf-like structures called
laminae. The microscopic structure of the omasum is similar to the reticulum in that it too
contains a muscularis mucosa that extends into the laminae. In addition, the muscularis externa
of the omasum extends into the laminae, which is unique to this compartment. Omasum is also
lined by a stratified epithelium whose thin layer is keratinized.
Abomasum or reed or true stomach or rennet:
The terminal chamber of the stomach of ruminants, the abomasum, is functionally and
microscopically analogous (identical) to that of the simple stomach of carnivores. The chamber
is smaller than the rumen but larger than the reticulum. It possesses a smooth vascular and
glandular mucous membrane with peptic glands. This chamber is the true stomach for its nature.

Process of rumination:
Rumination is the re-mastication of plant material together with microbial fermentation in
ruminants. The rumen serves mainly as storage organ. The cattle feed fairly rapidly and fills the
rumen with grain, grass and other herbage. The food within the rumen is churned by muscular
contraction and undergoes certain bacterial fermentation during its stay in the rumen.
Food from the rumen passes by degrees into the reticulum or directly to the oesophagus and
hence to the mouth by a process of retro-peristalsis. Such food regurgitated into the mouth is
called cud or bolus. When the cud is well-masticated and thoroughly mixed with the secretion of
the salivary glands, it again passes into the rumen. A new bolus is again formed and chewed and
sent back to the rumen. The process is repeated several times and when all the food is well-
masticated it passes to the reticulum and then to the omasum and abomasum. The abomasum is
provided with gastric glands. This chamber secretes a highly acidic, gastric juice whose pH is
1.05 to 1.32 and kills the microorganisms and initiates the process of digestion.
Dr. Shaoli Majumder

 Camel’s Stomach:
The Camel’s stomach is different in anatomy and histology with the stomach of ruminants, but
their rumination and fermentation are the same like the true ruminants. In camels the stomach is
three-chambered, the omasum being absent. Their openings are guarded by sphincter muscles.
C1: This first compartment acts as a 'hopper-like' container and is most comparible to the rumen,
although with some notable differences. Dorsal regions have stratified squamous epithelium and
are non-keratinized whilst ventral regions (glandular saccules) are lined by mucinous epithelial
cells. There are no papillae present in C1, rather provided with pouch-like water cells. C1 is
thought to be involved in absorption of water and solutes, but is not thought to play a role in
bicarbonate formation. The contents of C1 are homogenous and fibrous and there is no dorsal gas
cap as can be seen in ruminants. A single walled oesophageal groove is present.
C2: It is small and only partially separated from the first compartment and is separated from the
third compartment by a tubular sphincter. This compartment is comprised of mostly glandular
epithelium except for the area around the oesophageal groove. C2 also contains numerous
glandular divisions forming a retiform pattern in a similar manner to the reticulum.
C3: This compartment is entirely glandular and the terminal 1/5th portion contains gastric
glands. The mucosa of the glandular area is reddish brown whilst the non-glandular area is pink.

Fig: Stomach of Bos sp. Fig: Stomach of camel

 Role of microorganisms in rumination

The rumen can be viewed as an anaerobic and methanogenic fermentation chamber that contains
microorganisms that have the ability to utilise, and increase the productivity of, cellulolytic feeds
(i.e. straw, hay, silage and grass). The rumen microbiome, is characterised by its high population
density, extensive diversity (encompassing bacteria, archaea, protozoa and fungi) and complexity
of interactions. The continuous fermentation carried out by these microorganisms leads to
ingested compounds being broken down into their subcomponents. A rumen microorganism is
anaerobic or facultatively anaerobic, and produces end products that are either utilised directly
by the host or by other microorganisms as energy. The pH of the rumen is kept relatively
constant, typically 6–7, but may vary depending on diet.
Bacteria:
Rumen bacteria account for 1010 organism/mL of rumen fluid accounting for about 50% of the
microbial community. The bacteria work together: some breakdown certain carbohydrates and
Dr. Shaoli Majumder

proteins which are then used by others, some require certain growth factors, such as B-vitamins,
which are made by others, some bacteria help to clean up the rumen of others’ end products,
such as hydrogen ions, which could otherwise accumulate and become toxic to other organisms.
This is called ―cross-feeding‖.
 Cellulotytic bacteria - Ruminococcus albus, Ruminococcus flavefaciens, Fibrobacter
succinogenes
 Amylolytic Bacteria - Bacteriodes ruminocola, Streptococcus bovis
 Lactate-Using Bacteria - Megasphaera elsdenii
 Hydrogen-Using (or Methane) Bacteria: Methanobacterium ruminantium
Methanogenic archaea:
Rumen archaea are strictly anaerobic and are the only known microorganisms present in the
rumen capable of producing methane. Such archaea are referred to as methanogens. Archaea are
found in the rumen in the range of 106-108 organism/mL, accounting for less than 4% of the
microbial community. The 90% of rumen methanogens belong to the following genera;
Methanobrevibacter, Methanomicrobium, Methanosphaera, ―Rumen Cluster C‖, now referred to
as Thermoplasma and Methanobacterium. Most methanogens remove hydrogen gas by reducing
CO2 with hydrogen gas to form methane.
Protozoa:
Ciliate protozoa are organisms larger than bacteria and account for 106 organisms / mL of rumen
fluid, however they still make up to 50% of the total microbial biomass. They have various
activities:
 Cellulolytic and hemicellulolytic can digest plant particles
 Different protozoa have a positive role in digesting starch (more slowly than bacteria)
 Others protozoa can consume lactic acid, thereby limiting the risk of acidosis
 Some types of protozoa are able to remove oxygen so they have a stabilizing effect
upon anaerobiosis.
However, most of them degrade proteins very efficiently and release ammonia, so they can waste
dietary protein. These proteins represent around 25% of the microbial protein available for the
animal. Ciliate protozoa produce large amounts of hydrogen, which is a substrate for
methanogens. The ciliate species are predators of other rumen microbes. In fact, a single
protozoal cell can swallow up to several thousand bacteria in an hour so they play a very
important role in rumen microbial population stability.
Fungi:
Rumen fungi comprise up to 8-10% of microbial biomass and are strictly anaerobic. They play
an essential role in fiber digestion due to the production of filamentous rhizoids which invade
plant tissues, and their efficient enzymatic activities. This physical action to plant cell walls, can
facilitate access to more digestible tissues and help release polysaccharides, which are linked to
lignin increasing the pool of digestible energy for the rumen microflora. Rumen fungi (103–106
zoospores/ml) are anaerobic, falling into the class Neocallimastigomycetes, consisting of 6
previously recognised genera (Anaeromyces, Caecomyces, Cyllamyces, Neocallimastix,
Orpinomyces and Piromyces) with 21 known species. Although bacteria are the most prominent
microorganisms in the rumen, fungi are the best degraders. They produce high levels of
cellulases and hemicellulases, as well as possessing the ability to break down xylan due to the
production of xylanases. Fungi appear to initiate the feed breakdown process, indicating that
anaerobic fungi may be pivotal for feed utilization efficiency and animal growth and production
in pasture-fed ruminants.