10

DIGESTIVE SYSTEM

The digestive system consists of a long hollow tube or tract (alimentary canal)
that extends through the body from the oral cavity to the anus. Associated with this
tube are accessory organs such as the salivary glands, liver, and pancreas. These
organs produce numerous secretions that are delivered to the digestive tract through
the excretory ducts. These secretions facilitate the digestion and absorption of nutrients
from the ingested material.
The main function is to break down the ingested food into smaller units that can
be absorbed into the tissues and utilized for the maintenance of the organism. The
digestive tract includes the oral cavity, oral pharynx, esophagus, stomach, small
intestine (duodenum, jejunum, ileum) and large intestine (caecum, colon, rectum and
anus).

General Structural Features

The walls of each organ consist of four concentric layers: (mucosa, submocosa,
muscularis externa, and serosa or adventitia.

1. Tunica mucosa (mucous membranes). The wet epithelial lining of the digestive tube
and of other internal passageways that open to the surface constitutes a barrier
between the body proper and the outside environment. The characteristics of
mucous membranes are their absorptive capacity and its ability to secrete mucous.
This layer borders the lumen and has three parts:
 The epithelium (mucous membrane) derived from the endoderm, is a stratified
squamous in the oral cavity, oral pharynx, esophagus and anal canal; it is
simple columnar in the stomach, intestines and rectum.
 The lamina propria is a layer of loose connective tissue beneath the
endothelium; it contains small blood and lymphatic cells. In some organs, it
contains glands that are referred to as mucosal glands because they are
confined to the mucosa.
 The lamina muscularis (muscularis mucosae) is a thin layer of smooth muscle
bordering the submucosa.

2. Tela submucosa is a dense, irregular connective tissue layer that contains blood
and lymphatic vessels and the submucosal (Meissner’s) plexus of nerves. Some
organs are characterized by glands (submucosal glands) and lymphoid nodules in
this layer.

3. Tunica muscularis is a layer of muscle responsible for the movement of ingesta

through the tract. This consists of two layers of smooth muscle - an inner circular and
an outer longitudinal - through most of the tract. Between them lies the myenteric
(Auerbach’s) plexus. The muscle around the oral cavity is skeletal; where it is absent
(e.g. hard, palate, gingiva) the submucosa binds tightly to the bone. In the upper
esophagus, this layer contains mainly skeletal muscle, which is replaced by smooth
muscle in the lower portion. The stomach’s muscularis externa has three layers;
C.T. Reyes, DVM, MS   Lecture Guide in Veterinary Histology 105
106
Digestive System  Lecture Guide in Veterinary Histology

outer longitudinal, middle circular, and inner oblique. The colons outer longitudinal
layer is gathered into three bands, the taeniae coli. The smooth and skeletal
muscles encircling the anal canal form involuntary and voluntary sphincters,
respectively.

4. Tunica serosa (serous membrane) and adventitia. The tract’s outer covering differs
by location. The esophagus and rectum are surrounded and held in place by a
connective tissue and adventitia like that around blood vessels. Intraperitoneal
organs (jejunum, ileum, and colon) are suspended by mesenteries and covered by
a serosa composed of a thin layer of loose connective tissue covered by a simple
squamous epithelium (mesothelium). Retroperitoneal organs (duodenum,
ascending and descending colon) are bound to the posterior abdominal wall by
adventitia and covered on their free (anterior) surfaces by serosa.

THE ORAL CAVITY

The oral cavity is the initial portion of the digestive system. Here, food is
ingested, masticated (chewed), and lubricated for swallowing. The major structures of
the oral cavity are the lips, tongue, oral lining, and the salivary glands. Since food is
broken down physically in the oral cavity, this cavity is lined by a protective, non-
keratinized stratified squamous type of epithelium. The same type of epithelium extends
over and covers the inner surface of the lips.

Lips

The junction between the integument and the digestive system occurs on the lips.
They are covered outside by skin containing hair follicles, sebaceous glands and sweat
glands; and inside by a mucous membrane.
 The epithelium is covered with a layer of dead cells, like that of the skin but
believed to contain high percentage of relatively transparent eleidin (white flakes
or wind burn).
 The labial mucosa is covered by a stratified squamos epithelium that is keratinized in
ruminants and horse but nonkeratinized in carnivores and pig.
 Aggregates of labial glands, usually serous or seromucous are distributed in the
propria submucosa.
 The tunica muscularis consists of striated muscle fibers of the orbicular muscles
(orbicularis oris).

Cheeks

The cheeks, like the lips, are composed of an external covering - the skin, a middle
muscular layer, and a mucous membrane lining.
 Buccal mucosa is lined with stratified squamos epithelium that may or may not be
keratinized, depending on the particular area or species. This is the type of
epithelium that is characteristically found in wet epithelial surfaces where there is
considerable “wear and tear” and from which no absorption occurs. In ruminants, it
is studded with conical papilla.
 107
C.T. Reyes, DVM, MS  Digestive System

 Buccal glands are minor salivary glands, compound tubuloacinar and varied
secretions depending on the location and the species.
 The lamina propria consists of a fairly dense fibroelastic tissue and extends into the
epithelium in the form of high papilla.

Hard Palate

 The mucosa has keratinized stratified squamos epithelium.

 Thick in ruminants and forms the dental pads (pulvinus dentalis).
 Branched tubuloacinar mucous and seromucous glands are located in the caudal
part of the hard palate in all domestic animals except the pig.

Soft Palate

The soft palate is formed by a fold of mucous membranes of the oral and nasal
cavities, with a core of striated muscle fibers.
 The oral surface is covered by a stratified squamos epithelium, whereas the nasal
mucosa has a ciliated, pseudostratified columnar epithelium.
 The propria and submucosa contain branched tubuloacinar mucous and
seromucous glands.

Tongue

The tongue is a muscular organ covered by a mucous membrane. It is important

in prehension, mastication and swallowing of food.
 The epithelial lining is stratified squamos with varying thickness of the stratum
corneum. The tongue consists of a core of interlacing bundles of skeletal muscle
fibers covered by a mucous membrane. The epithelium of the lower surface of the
tongue is smooth, whereas its dorsal surface is irregular and with numerous epithelial
elevations called macroscopic papillae.
 The functions of the tongue during digestion are to perceive taste and to assist in
chewing and swallowing food (bolus).
 Sensation of taste is detected by specialized cells located in numerous taste buds
found in the stratified squamous epithelium of the fungiform, circumvallate and
foliate papillae on the tongue.
 In humans, the foliate papillae are rudimentary. In addition to the tongue, where
the taste buds are most numerous, the taste buds are also found in the mucous
membrane of the soft palate, pharynx, and epiglottis.

Macroscopic papilla of the tongue

The filiform, conical and lenticular papillae facilitate the movement of ingesta
within the oral cavity. The fungiform, vallate, and foliate contain taste buds which are
responsible for mediation of the sense of taste.
1. Filiform papillae are the most numerous among the types of papilla. They are
slender, sharp-pointed structures that have a mechanical function in the
movement of food into and within the oral cavity. The shape is like a rose thorn with
its thorn directed caudad. They project above the surface of the tongue and are
108
Digestive System  Lecture Guide in Veterinary Histology

covered by a keratinized stratified squamos epithelium with a thick stratum

corneum.
2. Conical papillae occur on the root of the tongue in dog, cat and pig as well as on
the cheek and floor of the oral cavity of other species. They are larger than the
filiform type and usually are not highly keratinized.
3. Lenticular papillae are flattened lenticular-shaped (like a double convex lens)
projections of the tongue surface and occur mainly in the ruminants especially
prominent on the dorsum of the caudal one-third of the tongue. They are covered
by stratified squamos epithelium and have a core of dense irregular connective
tissue.
4. Fungiform papillae are scattered among the filiform papillae and have dome-
shaped upper surface in the horse and pig. The shape is suggestive of mushroom, a
type of fungus, and thus the name fungiform. They are covered by a nonkeratinized
stratified squamos epithelium containing one or more taste buds on the upper
surface.
5. Vallate papillae located on the dorsum and rostral to the root of the tongue, are
large, flattened structures completely surrounded by an epithelium-lined cleft or
moat. The lingual surface is covered by a stratified squamos epithelium with taste
buds on the papillary side of the moat.
6. Foliate papillae are parallel folds of the lingual mucosa located at the lateral
border just rostral to the palatoglossal arch. They are leaf-shaped separated from
each other or the associated mucous membrane by an invagination of the mucous
membrane. The folds are covered with stratified squamos epithelium, with taste
buds on the sides of the folds.

Taste buds

The taste buds are ellipsoidal clusters of sensory cells embedded in the stratified
squamos epithelium of the fungiform, vallate and foliate papilla. Substances that
stimulate the taste buds must first be dissolved in the saliva, which is normally present in
the oral cavity and increase during food intake. In addition, taste buds in the
circumvallate papillae are washed continuously by watery secretions produced by the
serous (von Ebner’s) gland located below the surface in the connective tissue of the
tongue. This continuous secretion process and washing of the taste buds allow different
chemicals in the solution to enter the taste pores in the taste buds and to stimulate their
receptor cells.
There are four primary taste sensations to which the cells in the taste buds
respond: sour, salt, bitter, and sweet. All other taste sensation is the combination of
these four. Certain regions of the tongue exhibit more sensitivity to certain taste
sensation than to others. Thus, the tip of the tongue is most sensitive to sweet and salt,
the posterior portion of the tongue to bitter, and the lateral edges to sour taste
sensation. Taste pores are small openings of the tastes buds that extend from the
basement membrane.

Three different cell types in the taste buds

1. Neuroepithelial cells are spindle-shaped, studded with microvilli (taste hairs) that
project into the taste pore, and located in the center of the bud.
 109
C.T. Reyes, DVM, MS  Digestive System

2. Supporting or sustentacular cells are spindle-shaped, located toward the periphery

and between the taste cells.
3. Basal cells located near the basement membrane.

Special Structures of the Tongue in Various Species

 In dogs, lyssa is filled with adipose tissue, striated muscle, blood vessels and nerves.
 In cat, it contains mainly fats. Lyssa of the carnivore tongue is a cordlike structure
enclosed in a dense collagenous sheath and extends lengthwise near the ventral
surface of the center of the tongue.
 In pig, it contains a similar structure.
 In horse, a middorsal fibroelastic cord with hyaline cartilage, striated muscle, and
fat is present.
 In ruminants, it has a dorsal prominence characterized by a thickened mucosa.
Flattened lenticular papilla are scattered over the surface of this area.

Teeth

The teeth are highly mineralized structures in the oral cavity that serve the
domestic mammals in procuring, cutting, and crushing food and as weapons of
offense and defense.
 The roots of the teeth are set into bony ridges called alveolar process. In these
processes are sockets (alveoli) - one for the root of each tooth.
 The teeth are suspended and held firmly in their alveoli by bundles of connective
tissue fibers known collectively as peridontal membrane or preferably as peridontal
ligament.
 Sharpey’s fibers are embedded collagenous fibers that hold the tooth in place.
They are arranged so that when pressure is exerted on the biting surface of the
tooth, the tooth being suspended by them will not be pressed further into the
narrowing socket, and at the same time the tooth is permitted some slight
movement within its alveolus.

Types of teeth in domestic animals:

1. Brachydont teeth
 Also called simple teeth
 Short and cease to grow after eruption is completed
 Divisible into a definitive crown, neck and root
 Enamel organ is intact until the time of eruption
 Ameloblasts disintegrate after eruption of the crown
 Cementum is deposited only on the dentine
2. Hypsodont teeth
 Also called complex teeth
 Much longer than the brachydont teeth and continue their growth throughout
a portion of the adult life of the animal.
 They do not have crown and neck but an elongated body with the roots and
neck forming in some species only after a delayed period. The tusks of the
boar continue to grow throughout its life and never develop roots.
 Enamel organ is ruptured prior to eruption
110
Digestive System  Lecture Guide in Veterinary Histology

 Ameloblasts do not disintegrate; they continue to secrete enamel for an

extended period beyond eruption
 Cementum is deposited on both the dentin and the enamel
 Include the cheek teeth of ruminants, all the teeth of horse, incisor teeth of
rodents and the canine teeth of the pig.

Structure of Teeth

1. Enamel is the hardest substance in the body composed of 99% mineral

(hydroxyapatite) and 1% organic matrix by weight. It covers the crown of the
brachydont teeth but lies beneath a layer of cementum in the hypsodont teeth.
Lines of Hunter-Schreger are curve lines when parallel bundles of enamel rods
change directions.
2. Dentin constitutes the major part of the tooth composed of 70% mineral (crystallites
and hydroxyapatite) and 30% organic matter. The dentin that surrounds the pulp
cavity is lined with a layer of special cells called odontoblasts whose function is
related to the production of dentin.
3. Cementum is a yellowish, mineralized layer of slightly modified bone. Bundles of
collagen fibers, called perforating cementum fibers, are embedded in the
cementum of the tooth and extend into the bony socket. These fibers form the
peridontal membrane, which anchors the tooth in the socket. They are produced by
cementoblasts.
4. Dental pulp is composed of connective tissue cells and fibers, matrix, numerous
blood vessels, and nerves.

SALIVARY GLANDS

The major salivary glands include parotid, mandibular, and sublingual glands of all
domestic animals, the zygomatic gland of carnivores, and the molar gland of cat.
These glands, located outside of the mouth, produce a watery secretion called saliva.
The minor salivary glands are small clusters of seromucous secretory units in the
submucosa of the oral cavity and are named according to their location, e.g., labial,
lingual, buccal, and palatine.

Parotid Salivary Gland

The duct is called Stensen’s duct. The parotid salivary gland in domestic animals is
the largest of the major salivary glands, predominantly serous, although occasional
isolated mucous secretory units may occur in the dog and cat. The apex of each cell is
filled with secretory granules containing digestive enzymes. The granules are referred to
as zymogen granules because they initiate the digestion of carbohydrates.

The parotid duct system

 The narrow lumen of the acinus opens into a short intercalated duct lined by a low
cuboidal epithelium.
 The intercalated duct joins a large striated or salivary duct lined by cuboidal or
columnar epithelium.
 The striated ducts extend to the edge of the lobule, where they join the interlobular
ducts located in the connective tissue septa between lobules lined by simple
 111
C.T. Reyes, DVM, MS  Digestive System

columnar epithelium, which changes to stratified columnar epithelium as the ducts

become larger and fuse with similar ducts draining other lobules.
 The interlobular ducts converge to form the main parotid duct where it opens into
the oral cavity, the epithelium changes from stratified columnar to stratified
squamos epithelium.

Mandibular Salivary Gland

The main duct is called the Wharton’s duct. The mandibular salivary gland is a
compound branched tubuloacinar gland. It is a mixed type, majority of secretory units
are composed of serous, and the rest are mucous with serous demilunes. It has a well-
defined capsule and fairly prominent duct system. The columnar epithelium of the
interlobular ducts gradually changes to two-layered cuboidal epithelium and finally
becomes stratified squamos epithelium before it enters the oral cavity. Goblet cells
may occur in the epithelium of the main duct.

Sublingual Salivary Gland

The sublingual salivary gland is a compound tubuloacinar gland. The main duct is
called Rivinus duct. It has an ill-defined capsule. Mixed type: majority of secretory units
are mucous; a few serous; and mucous with serous demilunes. In cow, sheep and pigs,
they are almost entirely mucous, with relatively few serous demilunes. In dogs and cats,
in addition to the typical mucous acini and demilunes, these glands contain clusters of
serous acini with well-developed intercellular canaliculi.

Zygomatic Salivary Gland

Zygomatic salivary gland is present only in carnivores named from its location
between the zygomatic arch of the temporal bone. The parenchyma is composed of
long branched tubuloacinar secretory units that are predominantly mucous secreting.
Intercalated and striated ducts are almost non-existent, but isolated patches of tall,
striated cells are present within the low cuboidal epithelium of the interlobular ducts.

Molar Salivary Gland

The molar salivary gland of cat, located in the fascia beneath the mucosa of the
lower lip near the commissure is histologically similar to zygomatic salivary gland. It is a
compound tubuloacinar gland that is predominately mucous secreting.

Minor Salivary Glands

Clusters of serous, mucous or seromucous minor salivary glands occurring
throughout the oral cavity are generally named according to their location. Mucous
tubular and acinar units frequently have serous demilunes associated with them;
however, striated ducts are not characteristics of minor salivary glands. The duct
system consists of simple cuboidal epithelium within the lobules and two-layered
cuboidal epithelium in the larger interlobular ducts.
 Lingual glands are located in the submucosa and between the intrinsic muscle
bundles of the tongue.
 Gustatory glands associated with the vallate and foliate papillas are entirely serous
and their ducts open into the furrow at the base of the papillae.
 The labial, buccal, palatine and pharyngeal glands also contribute mucous and
serous secretory products to the saliva.
112
Digestive System  Lecture Guide in Veterinary Histology

SALIVA
The saliva is a mixture of some cellular and bacterial debris and leukocytes; salt,
gases, mucus, organic material, two enzymes (ptyalin or salivary amylases and
maltase), immunoglobulins, and ionic secretions are produced by cells in different
salivary glands, although the major composition of saliva is water. The composition
varies with the type of stimulant that initiates its secretion. The sight, smell, thought,
taste, or the actual presence of food in the mouth cause an autonomic increase of
saliva secretion from the salivary glands and is release to the oral cavity.

Saliva performs numerous functions in the oral cavity

1. Aid in assimilation. It moistens the chewed foods and lubricates the bolus to facilitate
its swallowing and passage through the esophagus towards the stomach.
2. Saliva also contains numerous electrolytes (calcium, potassium, sodium, chloride,
bicarbonate ions, and others). As the saliva flows through the duct systems of the
salivary glands, its ions content is modified. Sodium and chloride ions are actively
reabsorbed from the fluid in the lumen while potassium and bicarbonate ions are
added to the salivary secretions.
3. Digestion of food. The digestive enzyme amylase, produce mainly by the serous
acini of the salivary glands, is also found in the saliva. This salivary amylase initiates
the breakdown of starch into smaller carbohydrates during the short time that the
food is present in the oral cavity.
4. Saliva also controls bacterial flora in the mouth and protects the oral cavity from
pathogens. It washes the mouth from cellular and food debris which might provide
an excellent culture medium for bacteria. The enzyme lysozyme, secreted also by
the cells of the serous glands into the saliva, hydrolyzes the cell walls of bacteria and
inhibits their growth in oral cavity. Immunoglobulins in saliva, primarily the IgA
produced by the lymphocytes and plasma cells in the connective tissue of the
glands, assist in the immunological defense against bacteria found in the mouth.

PHARYNX

The oropharynx is an extension of the buccal cavity and connects it with the
esophagus.
 The tunica mucosa is typical and contains branched, tubuloalveolar mucous
glands.
 The lining epithelium is stratified squamos with varying degrees of keratinization in
various species.
 The lamina propria is typical and contains tonsils, lymph nodules and scattered
leukocytes.
 The muscularis mucosa is absent but instead a layer of elastic fiber is present.
 Tunica muscularis is composed of striated (skeletal) muscle.
 Tunica adventitia is typical and blends with the accompanying deep fascia.

ESOPHAGUS

The esophagus is a long, soft tube that extends from the pharynx to the stomach.
Most of the esophagus is located in the thorax with a short piece extending into the
abdominal cavity. The major function of the esophagus is to convey (voluntary or
 113
C.T. Reyes, DVM, MS  Digestive System

involuntary movement) liquids and chewed food (bolus) to and from the stomach. The
lumen of the esophagus is lined by a protective, nonkeratinized stratified squamous
epithelium. The mucus secreted by the esophageal gland located below the
epithelium lubricates the lumen of the esophagus for smoother passage of the
swallowed food. The wall of the digestive tube from the esophagus to the rectum
exhibits four common layers: mucosa, submucosa, muscularis externa, and serosa.
Where the digestive tube is retroperitoneal or adheres to the abdominal wall, the
outermost layer is the adventitia. The four layers of the digestive tube, however, show
regional variations, depending on the location of the digestive tube and its specific
function in the digestive process.
 Lamina propria is typical in man and pig may contain numerous lymph nodules
and diffuse lymphatic tissue in the esophageal-gastric junction.
 Muscularis mucosae is typical; in man it is thick and complete layer; in horse,
ruminants and cat, it consist of scattered muscle bundles; in dog and pig, it is
absent in the cervical portion becomes complete near the stomach.
 Tunica submucosa is loose connective tissue and contains branched tubuloalveolar
mucosal glands. These glands are present in the entire length of the canine
esophagus; in pig, they are present in the cervical portion; in ruminants, the horse
and cat, they are present in the cervical portion.
 The tunica submucosa and tunica mucosa have longitudinal folds that permits
expansion of the esophagus.
 The tunica muscularis is entirely striated muscle in ruminants and dog; in pig the
cervical portion is striated, the thoracic is mixed and the caudal portion is smooth; in
the horse and cat, it is striated to the middle portion and becomes smooth
caudally.
 Tunica adventitia, a loose connective tissue containing blood and lymph vessels
and nerve, is typical in the cervical portion. It is replaced by tunica serosa in the
thoracic portion.

STOMACH

The stomach is a dilated, hollow organ of the digestive tube that is situated
between the esophagus and the small intestine. The luminal surface of the stomach is
lined by a single layer of mucus-secreting, columnar epithelium, whose secretion forms
a thick layer of mucous that lines and protects the stomach surface from the corrosive
action of the gastric juices. The surface epithelium of the stomach invaginates the
underlying lamina propria of the mucosa and forms numerous gastric pits. The tubular
glands open into these pits.
The main functions of the stomach are to receive, store, mixed, and digest the
ingested food products. Some of these stomach functions are performed by
mechanical and chemical actions, which reduce the ingested food material to a
semi-liquid mass called chyme. The mechanical reduction of food products in the
stomach is produced by strong, muscular contractions of its thick wall in the form of
peristalsis. This peristaltic activity churns and mixes the food with gastric juices produce
by the gastric glands when the food enters and distends the stomach wall. Nerve cell
and nerve fibers located in the submucosal nerve plexus and myenteric nerve plexus
in the stomach regulate the peristaltic activity of the stomach musculature. The
114
Digestive System  Lecture Guide in Veterinary Histology

stomach also has some absorptive functions; however, these are limited to the
absorption of water, alcohol, salts, and certain drugs.
The chemical reduction of food in the stomach is the result of the watery gastric
secretions produced by different cells in the gastric glands, especially those in the
fundus and body region of the stomach. The main components of the gastric
secretions are pepsin, hydrochloric acid, mucus intrinsic factor, water, and different
electrolytes.

Anatomically, the stomach is divided into:

1. Cardia is the most superior region in the stomach, surrounds the entrance of
esophagus in the stomach.
2. Fundus (body or corpus) form the major portion of the stomach; their mucosa
contains deep gastric glands that produce most of the gastric secretions or juices
for digestion of food.
3. Pylorus is the most inferior organ of the stomach, ends at the border of the
duodenum of the small intestine.

THE NON-GLANDULAR REGION

The non-glandular region is absent in carnivores and small in pigs. In horse, it

extends a considerable distance and ends abruptly at the margo plicatus. In
ruminants, it reaches the greatest development and is subdivided into rumen,
reticulum and omasum.
 The lining epithelium is non-secretory and may be non-keratinized stratified
squamos, depending on species and diet.
 Lamina propria is composed of irregularly arranged collagen, elastic and reticular
fibers.
 Lamina muscularis is distinct
 The junction between the glandular and non-glandular stomach is abrupt, with the
epithelium changing from stratified squamos to tall columnar type.

THE GLANDULAR STOMACH

The glandular mucosa has extensive folds called rugae, which flattens as the
stomach fills. Permanent elevations, called gastric areas, subdivide the mucosa and
bulge into the lumen. The surface is studded with small depressions or invaginations
called gastric pits, continuous with gastric glands and receive their secretory products.
 Tunica mucosa and part of the underlying tunica submucosa are thrown into a
tortuous folds plica gastricae (gastric folds) by numerous small grooves.
 Lamina propria is typical and consists of loose connective tissue, presents gastric
glands thus the connective tissue is scanty because it is interdigitated between the
extensive gastric glands.
 Lamina propria may contain a distinct zone, the lamina subglandularis at the
junction of the lamina propria and lamina muscularis.
 Lamina muscularis is relatively thick, usually comprising three layers: an inner and
outer circular layer and a middle longitudinal layer.
 Subglandular layer (lamina subglandularis) in carnivores is interposed between the
base of the glands and the lamina muscularis. It is composed of the stratum
 115
C.T. Reyes, DVM, MS  Digestive System

granulosum, an inner layer containing many fibroblasts, and the stratum

compactum, an outer layer that is sheet of collagen fibers.

Distinct regions and glands of the stomach:

1. Esophageal region is the non-glandular portion of the stomach
 Lined by stratified squamos epithelium, keratinized or non-keratinized.
 Limited in carnivores, man and swine; in ruminants, it is extensive and is
subdivided into rumen, reticulum and omasum.
2. Cardiac gland region are branched, coiled tubular glands that opens into the
gastric pits.
 The beginning is marked by a transition called the margo plicatus in the horse.
 It occupies a narrow strip at the junction of the glandular and non-glandular
mucosae in all domestic mammals except the pig, where it covers nearly half of
the stomach including the diverticulum ventriculi.
 The neck and upper body are lined by cuboidal cells, which are mucus
secreting. The remaining cells of the gland are columnar, mucus secreting cells.
Some parietal cells may be present in the canine cardiac gland while some
chief cells may be present in porcine cardiac glands. Argentaffin cells
(enteroendocrine cells) are located between the lining cells of the glands. The
argentaffin cells secrete hormones, which control the secretory and muscular
activities of the gastrointestinal organs.
3. Pyloric gland region are branched, coiled and relatively short compared to other
gastric glands
 The gastric pits are deeper than in other regions of the stomach
 Predominant cell type is mucus secreting cell similar to that in the cardiac gland
 Has a well-developed inner circular layer of the tunica muscularis that forms the
pyloric sphincter at the gastroduodenal junction. This encircles the caudal
portion of the stomach and causes the submucosa and mucosa to bulge into
the lumen. In ruminants and pigs, this protuberance called the pyloric tortus is
especially prominent. At the pyloric-duodenal junction, the submucosal glands
are found in the submucosa of the pyloric region.
 Occupies approximately half of the stomach in carnivores but only one-third in
the horse stomach and in the ruminant abomasum. In the pig, the pyloric region
is small, representing about one-fourth of the stomach.
4. Fundic gland region are straight, branched tubular glands that extends to the
lamina muscularis.
 Well-developed in all species
 Composed of four regions: base, body, neck and isthmus. The isthmus or opening
of the gland into the gastric pit is continuous with a constricted part of the gland,
the neck. The body continuous from the neck and terminates as a slightly dilated
and bend adenomere, the base.

Species Differences
 In carnivores, the cardiac zone is relatively narrow area, with the fundic and pyloric
regions occupying the remainder of the stomach. In dog, the fundic region is
divided into two zones:
 Light zone has thinner mucosa with thinner gastric pits and has short tortuous
glands that appear in groups and do not reach the muscularis mucosae.
116
Digestive System  Lecture Guide in Veterinary Histology

 Dark zone is adjacent to the pyloric region and has thicker mucosae, shallow
gastric pits, and the fundic glands that resemble those of other species.
 In pig, the stomach has a large cardiac zone, and the parietal cells in the fundic
region tend to occur in clusters.
 In horse, the stomach has an extensive nonglandular region that terminates
abruptly, forming the margo plicatus. The cardiac region is almost non-existent,
whereas the pyloric and fundic zones follow the normal pattern.

Cell Types in the Stomach

1. Chief cells (or zymogen cells) are pyramidal-shaped cells with a basally positioned
nucleus and apically positioned secretory granules. They are the predominant cells
of the fundic glands.
 The basophilic, cuboidal cells distributed predominantly in the lower region of the
gastric glands in the body of the stomach are chiefly of the zymogenic cells. The
cytoplasm of these cells exhibits numerous secretory granules that contain
pepsinogen, an active form of pepsin.
 The basal portion contains numerous rER and free ribosomes, which impart a
distinct basophilia.
 Synthesize and secrete gastric enzymes in inactive forms such as pepsinogen
and prorenin. Pepsinogen is activated by hydrochloric acid within the lumen of
the stomach. Prorenin activated by pepsin into renin. Renin, which is only present
in young animals, converts soluble milk proteins (casein) into an insoluble form
(paracasein). Paracasein is then proteolysed by pepsin, the active form of
pepsinogen. It also secretes gastric lipase.
2. Parietal cells (or oxyntic cells) are large, polygonal, distinctive eosinophilic cells
located primarily in the upper half of the gastric glands between other gland cells.
 These cells secrete hydrochloric acid, a major component of gastric juice. In
humans, the parietal cells produce gastric intrinsic factor, a glycoprotein that is
necessary for vitamin B12 absorption in the small intestine. Vitamin B12 is necessary
for red blood cell production in the red bone marrow; deficiency of this vitamin
leads to anemia.
3. Mucous neck cells are located in the upper region of the gastric gland near the
gastric pits.
 The product of this cells along with that of the cells in the surface epithelium, is
mucus; it covers the stomach lining with its protective layer.
 Mucus secretion is less viscous than that secreted by the surface lining cells. It
may be capable of differentiating into surface lining and/or glandular lining
cells.
 The secretion may protect the fundic gland from the proteolytic and hydrolytic
activities of the proteases and HCl, respectively.
4. Transitional cell is described at times in the isthmus and may be responsible for the
replacement of the lining and glandular cells.
5. Enteroendocrine (APUD-amine precursor uptake decarboxylation) cells secrete a
variety of polypeptides and proteins with hormonal activity that influences various
functions of the digestive tract. APUD cells are not confined to the gastrointestinal
tract but are also found in the respiratory organs and other organs of the body.
 117
C.T. Reyes, DVM, MS  Digestive System

THE RUMINANT STOMACH

The stomach of ruminants has four structurally distinct parts. The first three parts
(rumen, reticulum and omasum) are collectively called the forestomach, lined with
non-glandular mucous membrane having a stratified squamos epithelium. The
forestomach breaks down ingesta through mechanical and chemical processes. The
rumen serves as a large fermentation vat in which microorganisms break down the
ingested food and produce volatile fatty acids (VFA) particularly butyric, acetic and
propionic acids, which are then absorbed through the mucosa in the blood. The
reticulum and omasum exert a mechanical action on the ingesta reducing the mass of
fine particles. Absorption of metabolites occurs in the epithelium of the forestomach.
The fourth part, the abomasum, is the glandular portion similar to the simple stomach of
other species. The enzymatic digestive processes of the abomasum further degrade
the ingesta to such substances as glucose and amino acids in similar manner to the
glandular portion of simple stomach.

1. Rumen or paunch is characterized by the presence of small tongue-shaped conical

papillae that project into the lumen from the tunica mucosa.
 Devoid of lymphatic nodules and glands
 Tunica mucosa - the lining epithelium is keratinized stratified squamous type and
performs at least three important functions: protection, metabolism and absorption
 Lamina propia consists of a dense feltwork of collagen, elastic and reticular
fibers
 Tela submucosa extends into the papilla, composed of loose connective
tissue and blends into the lamina propria without any distinct line of
demarcation. The region may be similar to the stratum compactum of the
lamina subglandularis and is sometimes mistaken for muscularis mucosae
 Muscularis mucosa is absent.
 Tunica muscularis is composed of two layers of smooth muscle, a circularly
arranged inner layer and a longitudinally oriented outer layer.
 Tunica serosa is composed of collagen and elastic connective tissue and is
covered by a mesothelium.

2. Reticulum or honeycomb has a mucosa with permanent anastomosing folds, giving

it the appearance of a honeycomb.
 The reticular or esophageal sulcus begins at the cardia and passes ventrally on the
medial wall of the reticulum to end at the reticulo-omasal orifice. The entire sulcus
is lined by stratified squamos epithelium. Contraction of the sulcus closes off a
channel that directs ingested fluids from the esophagus to the abomasum, thus
bypassing the rumen and reticulum.
 Same morphology as the rumen except that:
 Tunica mucosa presents numerous anastomotic primary folds, which are
oriented on the surface view as a reticulum or honeycomb. From the primary
folds, numerous secondary and tertiary papillae project.
 At the tips of the primary folds the lamina propia presents an isolated mass of
smooth muscle of the muscularis mucosae
 Tunica submucosa, tunica muscularis and tunica serosa are all present and
typical.
118
Digestive System  Lecture Guide in Veterinary Histology

3. Omasum or book is characterized by the presence of numerous rounded, horny

papillae as well as smaller papillae that stud the surface of the mucosa.
 lining epithelium is keratinized stratified squamos and the agrandular lamina
propria is composed of dense subepithelial capillary network
 lamina muscularis mucosae is present and continuous and forms a double layer of
smooth muscle that follows the contour of the papillae as well as that of the
unraised surface lining. Another layer of smooth muscle is interdigitated between
the smooth muscle of the lamina muscularis mucosae within the laminae. It is
continuous with the inner layer of the tunica muscularis.
 other layers are all present and typical.

4. Abomasum or true stomach is the glandular portion of the ruminant stomach and
similar in structure to the simple or glandular stomach.
 junction of the omasum with the abomasum is marked by a mucosal fold, the
omaso-abomasal fold, where the epithelium changes abruptly from stratified
squamos to simple columnar epithelium. The lamina propria becomes less dense
on the abomasal side of the fold, and frequently there is a lymphatic nodule
beneath the epithelial junction.

SMALL INTESTINE

The small intestine consists of three parts: the duodenum, jejunum, and ileum. It is
a long tube that extends from the junction with the stomach to the junction with the
large intestine. This region is highly modified for the secretion and absorption facilitated
by several specialized structures.

The efficiency of secretory and absorptive functions is enhanced by three structural

features that increase the surface area exposed to the intestinal contents:

1. The tunica mucosa is thrown into circularly or spirally disposed folds called plicae
ciculares (valves of Kerking). These folds also contain portions of the tunica
submucosa and disappear in the lower and/or middle of the ileum. The plicae
circulares are permanent, spiral folds of mucosa (with submucosal core) that
extend into the intestinal lumen; they are most prominent in the proximal portion of
the small intestine, where most of the absorption takes place, and decrease in
prominence closer to the ileum.
2. The surface of the tunica mucosa over the folds is studded with tiny finger-like
projections called villi. The villi cover the surface of the intestine, extend into the
lumen and are also prominent in the proximal portion of the small intestine. The villi
are broader in the duodenum and have no muscularis mucosa and tunica
submucosa. In the upper jejunum, the villi are tongue-like and lower down they are
finger-shaped.
3. Presence of microvilli on the free or luminal surfaces of the absorptive columnar
cells. Microvilli are cytoplasmic extensions that cover the apices of the intestinal
absorptive cells. They are visible under a light microscope as a striated (brush)
border.
 119
C.T. Reyes, DVM, MS  Digestive System

Important Digestive Functions of the Small Intestine:

1. Completing digestion (initiated in the stomach) of food products (chyme) by
chemicals and enzymes produced in the liver and pancreas, and by cells in its own
mucosa;
2. Selective absorption of nutrient into the blood and lymph capillaries;
3. Transportation of chyme and digestive waste material to the large intestine
4. Release of hormones that regulates the digestive process.

Regions of the Small Intestine

1. Duodenum - is the fixed portion of the small intestine. It presents regularly shaped,
blunt and wide villi. Duodenal mucosa forms the highly corrosive action of the
gastric secretions. When acidic chyme enters the duodenum from the stomach, the
glands secrete an alkaline fluid rich in bicarbonate ions. This alkaline fluid then
neutralizes the acidic chyme from the stomach and protects the duodenal surfaces
from digestion. The alkaline secretion also provides a more favorable environment
for the continued action of the digestive enzymes.
2. Jejunum - is the mesenteric region of the small intestine. They are essentially similar
to the duodenum. The villi are slender, smaller and fewer than in the duodenum.
The remaining mural elements are typical.
3. Ileum - is similar to jejunum except for the following:
a) More goblet cells
b) Presence of Peyer’s patches which are very prominent especially in pigs
c) Villi are club-shaped
d) Plica circulares are no longer present.

The Mural Elements

1. Tunica mucosa. The lumen is lined by simple columnar epithelium.
The lining epithelium consists of several types of cells:
a) Columnar cells. Most cells in the intestinal epithelium are tall, columnar
absorptive cell with a prominent striated (brush) border (microvilli) covered by a
thick glycocalyx coat. The outer glycocalyx coat not only protects the intestinal
surface from digestion, but also contains the enzyme required for the terminal
digestion of carbohydrates and proteins and the carriers for transporting the
products of digestion into the cell. Intestinal cells absorb amino acids, glucose,
and fatty acids - the end products of protein, carbohydrates, and fat digestion,
respectively. Amino acids and glucose are transported through intestinal cells to
the blood capillaries. Most of the fatty acids, however, enter the lymphatic
vessels called the lacteals that are located in the lamina propria of the villi.
b) Goblet cells. Interspersed among the columnar absorptive cells are the goblet
cells that increase in number toward the distal region of the small intestine
(ileum). The goblet cells secrete mucus that lubricates, coats, and protects the
intestinal surface from the corrosive action of digestive chemicals and enzymes.
c) Enteroendocrine or APUD (amine precursor uptake decarboxylation) cells are
also found in the epithelium of the villi and intestinal glands. These cells secrete
numerous regulatory hormones of the intestine, including gastric inhibitory
peptide, secretin, and cholecystokinin (pancreozymin). These intestinal hormones
control the release of gastric and pancreatic secretions, intestinal motility, and
contractions of the gall bladder.
120
Digestive System  Lecture Guide in Veterinary Histology

d) Paneth cells are specialized pyramidal cells that contain deep-staining

eosinophilic granules and have basally displaced nucleus, located at the base
of the intestinal glands. Their exact function is not completely known however,
these cells produce lysozyme, an antibacterial enzyme that digest the cell walls
of some bacteria and appears to control the microbial flora of the small
intestine. Paneth cells are present in ruminants, horse and man but are absent in
other domestic species.
e) Additional cells found in the crypts are:
 Oligomucous cells (young goblet cells),
 Midcrypt columnar cells (young columnar cells)
 Undifferentiated cells at the base of the crypt
 Migrant cells such as leukocytes and lymphocytes can also be found.

Lamina propria is generally areolar or loose connective tissue that forms the core of
the villi and surrounds the intestinal glands.
 Numerous aggregations of lymphoid nodules, called Peyer’s patches, are found
in the ileum. Present in the Peyer’s patches are B-lymphocytes, T lymphocytes,
and macrophages. Overlying the lymphoid nodules are M cells (membranous
epithelial cells). M cells continually sample the antigens of the intestinal lumen,
ingest the antigens, and transport them to the underlying lymphocyte, where the
specific antibody to the foreign antigen is developed.
 The crypts of Lieberkuhn are located in the intestinal mucosa and open into the
intestinal lumen at the base of the villi.
 Lacteal is single lymphatic capillary located in the center of the lamina propria
of the villus.
Lamina subglandularis is present in the initial part of the intestine of carnivores.
Lamina muscularis mucosa is composed of inner circular and outer longitudinal
layers of smooth muscle.

2. Tela submucosa is a connective tissue layer of collagen and elastic bundles located
between the lamina muscularis and tunica muscularis.
 Submucosal glands (Brunner’s gland or duodenal glands) are simple branched
tubuloalveolar that open into the base of the crypts. They may be mucous in
ruminants and dog; serous in pig and horse and seromucous (mixed) in cat. The
serous or mucous secretory product from these glands lubricates the surface
epithelium and provides protection from the acid gastric chyme. The lumina of
glands are largest in goat and smallest in the pig. In carnivores, man and small
ruminants, they are confined to the initial or middle portion of the duodenum. In
horse, pig and large ruminants, they extend into the duodenum.
 Solitary lymphatic nodules are present throughout the small intestine.
 Large masses of aggregated lymphatic nodules occur in all three segments of the
small intestine, but most prominent in the ileum.
3. Tunica muscularis in all species consists of inner circular and outer longitudinal
smooth muscle layers. It is thickest in horse, in which the two layers are nearly equal in
thickness. The connective tissue between the two layers contains myenteric plexus
(Auerbach’s plexus).
 121
C.T. Reyes, DVM, MS  Digestive System

4. Tunica serosa is similar to that of the stomach, is loose connective tissue continuous
with that of the mesentery. Both are invested by the mesothelium of the peritoneum.

LARGE INTESTINE

The large intestine is the caudal continuation of the alimentary canal, which
begins from the ileo-caecal junction and ends in the anus. The principal functions of
the large intestine are to absorb water and minerals from the residual contents and
form feces. Consistent with these functions, the epithelium of the large intestine
contains columnar absorptive cells (similar to those in the epithelium of the small
intestine) and mucus-secreting goblet cells. Unabsorbed and undigested food residues
in the small intestine are forced into the large intestine by peristaltic action of the
muscles in muscularis externa of the small intestine. The residues that enter the large
intestine are in the semi-fluid state; by the time they have reached the terminal portion
of the large intestine, however, these residues have acquired the semisolid consistency
of feces.

Characteristics Common to all Segments of the Large Intestine

1. Absence of villi.
2. Absence of Paneth cells
3. Increase in the number of lymphatic nodules
4. Longer, straighter, more compact intestinal glands. The intestinal glands of the large
intestine are deeper in the small intestine, and contain different enteroendocrine
(APUD) cells. The cells of the large intestine produce no digestive enzymes than in
the small intestine.
5. Gradual increase of goblet cells caudally. Goblet cells appear more numerous in
the large intestine than in the small intestine. They produce mucus for lubricating
the lumen of the large intestine to facilitate passage of the hardened feces.
6. Absence of plica circulares but longitudinal mucosal folds are present
7. Diffuse lymphatic tissue and lymph nodules are very conspicuous

The Regions of the Large Intestine

1. Caecum
 presents taeniae ceci, the thickened, longitudinally oriented bands of smooth
muscle and elastic fibers from the outer longitudinal layer of the tunica
muscularis.
 In herbivores with simple stomach (e.g. horse) the relatively large caecum is an
important bacterial fermentation reservoir, but in carnivores, it is small.
 It has a substantial number of lymphatic nodules scattered throughout its length.
In dog, pig and ruminants, it is located around the ileocaecal opening; while in
horse and cat, they are concentrated near the blind end of the caecal sac.

2. Colon
 Presents taenia coli. The tunica mucosa is substantially thicker, owing to the
increased length of the intestinal glands as compared to the small intestine. In
pig and horse, the outer longitudinal layer of the tunica muscularis of the colon
(similar to caecum) forms a large, flat muscle bands containing numerous elastic
fibers, the taenia coli.
122
Digestive System  Lecture Guide in Veterinary Histology

 Diameter is generally greater than that of the small intestine

 Lamina muscularis is interrupted such that intestinal glands may extend into the
submucosa
 Submucosa often becomes distended by the lymphatic tissue.

3. Rectum
 Visible grossly because of pitlike depressions, rectal pits, in the mucosa overlying
these lymphatic nodules.
 In ruminants, the rectal mucosa is thrown into longitudinal folds, the rectal
columns (columnae rectales) near the junction with the anus
 In dogs, about 100 solitary lymphatic nodules are prominent feature of the
rectum
 It has similar structure to other regions of the large intestine except:
 Taenia coli are absent
 Tunica muscularis is thicker
 More goblet cells
 Tunica adventitia replaces the tunica serosa
 Lamina propria may present erectile tissue

4. Anus
 A mucocutaneous junction marked by a transition of the lining epithelium from
simple columnar to nonkeratinized stratified squamos at the anorectal line.
 At the level of the recto-anal junction, the lamina muscularis mucosa and the
outer layer of the tunica muscularis end. The inner layer of the tunica muscularis
continues and terminates as the internal anal sphincter.
 Tunica adventitia is present and blends with the surrounding connective tissue
may be associated with the anal glands and the circumanal glands in certain
species.

Three distinct zones of the anal canal (In pig and carnivores)
a) Columnar zone (zona columnaris ani) contains longitudinal columns or anal
columns, between which are anal sinuses.
b) Intermediate zone (zona intermedia) is relatively narrow. The mucosa is lined
with stratified squamos epithelium and anal glands occupy the submucosa. In
carnivores, the ducts from the anal sacs open at the junction of the
intermediate and cutaneous zones.
c) Cutaneous zone (zona cutanea) is lined by keratinized stratified squamos
epithelium. In the outermost part of the cutaneous zone, near the junction with
the skin, the mucosa of the dog contains large modified sebaceous glands,
the circumanal glands.

ACCESSORY DIGESTIVE ORGANS

The liver, gallbladder, and pancreas are the accessory digestive organs that are
connected to the small intestine by excretory ducts. The cells on the liver and
pancreas secrete numerous chemicals and enzymes for digestion; these secretions are
then delivered into the duodenum by a duct common to both organs. The gallbladder
serves primarily as reservoir for bile secreted by the liver.
 123
C.T. Reyes, DVM, MS  Digestive System

LIVER

The liver is a single, largest gland in the body and accounts for 2-5% of the total
body weight. It opens into the duodenum via the common bile duct. It has a highly
strategic location and is characterized by a multiplicity of complex functions.

Functions of the Liver

1. Excretion (waste products). Bile salts solubilize cholesterol and facilitate its excretion
from the body.
2. Secretion (bile). The main exocrine function of the hepatocytes is the secretion of
the bile, which enters tiny bile canaliculi located between adjacent hepatocytes. In
the liver the bile flows toward the bile duct located at the periphery of the liver
lobule and exactly in the opposite direction from the flow of blood.
3. Storage (lipids, vitamins A and B, glycogen). Liver also stores glucose (as glycogen),
fats and various vitamins. When the cells of the body need glucose, glycogen
stored in the liver is converted back into glucose and released into the blood
stream.
4. Synthesis (fibrinogen, globulins, albumin, prothrombin). The endocrine functions of
the liver are related to the synthesis of the numerous plasma proteins, including
albumin and blood clotting factor prothrombin and fibrinogen.
5. Phagocytosis (foreign particulate matter). Kupffer cells, specialized phagocytic cells
derived from blood monocytes and located in the hepatic sinusoids, phagocytize
particulate material and cellular debris and remove it from the blood.
6. Detoxification (lipid-soluble drugs). Liver cells detoxify various drugs and harmful
chemicals.
7. Conjugation (toxic substances, steroid hormones)
8. Esterification (free fatty acids to triglycerides). The bile salts that are present in the
bile are necessary to emulsify the fats that enter the small intestine (duodenum)
from the stomach. This action of the bile promotes easier digestion of the fats by the
fat-digesting enzymes, the pancreatic lipase produce by the pancreas. The cells in
the small intestine subsequently absorb the digested fats.
9. Metabolism (protein, carbohydrates, fats, hemoglobin, drugs)
10. Hemopoiesis (in embryo and potentially in the adult). In fetus, the liver cells
perform hemopoiesis- an important function in the blood cell production. Thus, the
liver is an essential organ for life.

Cell Types in the Liver

1. Hepatocytes. These are the primary structural and functional subunits of the liver.
The hepatocytes are highly versatile cells that perform numerous vital functions.
Since the hepatocytes synthesize and release a secretory product (bile) into a
system of ducts, they are exocrine cells. The hepatocytes are also endocrine cells
because they release many of their products directly into blood stream as the
blood flows through the sinusoids and pass the hepatocytes. Thus, the liver cells
perform both important endocrine and exocrine functions.
124
Digestive System  Lecture Guide in Veterinary Histology

2. Kupffer’s cells are interspersed among the sinusoidal endothelial cells and on their
luminal surfaces.

3. Fat-storing cells are stellate cells that are associated with sinusoids but lie in the
space of Disse. They accumulate fat and store vitamin A in the form of retinyl esters.

Blood Supply of the Liver

All venous blood that returns from the digestive organs and spleen by way of the
portal vein first percolates through the liver. Since venous blood is poor in oxygen, the
hepatic artery from the aorta also supplies the liver. Thus, a liver has a dual blood
supply.

1. Hepatic portal vein. This large vein is formed by the junction of mesenteric and
splenic veins. Mesenteric veins deliver oxygen-poor, nutrient-rich blood from
capillaries in the intestinal walls. The splenic vein delivers by-products of RBC
destruction from the splenic sinusoids. The hepatic portal vein, which supplies about
75% of the liver’s total blood volume, enters through the hilum on the liver’s inferior
surface. It branches repeatedly to form the portal venules that penetrate the liver
parenchyma and empty their blood into the hepatic sinusoids.
2. Hepatic artery is a smaller vessel, a branch of the celiac artery and enters the liver
alongside the portal vein. It follows the latter’s branching pattern and empties
oxygen-rich blood into the same sinusoids. It supplies about 25% of the liver’s blood
volume.
3. Hepatic sinusoids. Serving as the liver’s capillaries, these lies between the radially
oriented hepatocyte plates and receive blood from branches of both the portal
vein and the hepatic artery. The mixing of venous and arterial blood occurs only in
the hepatic sinusoids of the liver. The blood then flows towards the central vein that
is found in the liver lobules. The liver sinusoids are dilated blood channels lined by a
discontinuous layer of the fenestrated endothelial cells and Kupffer’s cells, which
are separated from the underlying liver cells, the hepatocytes, by a perisinusoidal
space called the space of Disse. As a result of this arrangement, the material
carried in the mixed blood percolates through the discontinuous endothelial wall
and comes in direct contact with the hepatocytes. This allows for a more efficient
exchange of materials found in the blood with those of the hepatocytes and vice
versa.
4. Central veins lies at the center of the classic liver lobule that receive blood from the
sinusoids and deliver it to larger sublobular veins, which merge to form even larger
hepatic veins.
5. Hepatic veins collect oxygen- and nutrient-poor blood from the sublobular veins.
They converge to form large veins that exit the liver’s upper surface and empty into
the posterior vena cava.

Liver Lobules (Models of Liver Subdivision)

1. Classic liver lobule is based on the direction of blood flow. The liver substructure
exhibits a pattern of interlocking hexagons; each of these is a classic lobule.
 125
C.T. Reyes, DVM, MS  Digestive System

 Portal triad. One triad occupies a potential portal space at each of the six
corners of the lobule. Each triad contains three main elements surrounded by a
connective tissue: a portal venule (branch of portal vein), a hepatic arteriole
(branch of hepatic artery) and a bile ductule (tributary of the larger bile ducts).
 Central vein is a single vein that marks the center of each lobule. It can be
distinguished from the portal triad by its larger opening and lack of a connective
tissue investment.
 Hepatocyte plates and hepatic sinusoids. Many such plates radiate from the
central vein toward the lobule periphery (like the spokes of a wheel). The plates
are separated by hepatic sinusoids, which receive blood from vessels in the triad,
converging on the lobule center to empty directly into the central vein.

2. Portal lobule is based mainly on the direction of bile flow, which is opposite to that
of blood. The liver parenchyma is divided into interlocking triangles, each of which
has a portal triad at the corner and a central vein at each of its three corners. Bile
produced by the hepatocytes, enters the membrane-bound canaliculi between
them and flows within the hepatocyte plates toward the bile duct in the portal
triad. Liver lymph in the spaces of Disse flow in the same direction as bile., toward
the lymphatic vessels in the triad.

3. Hepatic acinus (acinus of Rappaport) is based on changes in oxygen, nutrient, and

toxin content as blood flowing through the sinusoids is acted on by hepatocytes.
Each diamond shaped acinus contains two central veins and two portal triads that
define its four corners. The diamond is divided into two triangles by a line
connecting the portal triads. Along this line run terminal branches of the portal and
hepatic vessels that deliver blood to the sinusoids.

Each triangle can be divided into three zones according to their distance from the
terminal distributing vessels:

a) Zone I - (peripheral zone) closer to the vessels, has the best blood supply. Blood
in zone I sinusoids has higher oxygen, nutrient, and toxin concentrations than in
the other zones. As the blood flows toward the central vein, these substances are
gradually removed by the hepatocytes. Thus, hepatocytes in this zone have a
higher metabolic rate and larger glycogen and lipid stores. They are also more
susceptible to damage by blood-borne toxins, and their energy stores are the first
to be depleted during fasting.
b) Zone II - (intermediate zone)
c) Zone III - (centrolobular zone) closer to the central vein, has the poorest blood
supply.

Biliary System

1. Bile is consists of bile acids, phospholipids, cholesterol, bilirubin, water and

electrolytes. About 90% of the bile comes from recycled substances that were
added to the intestinal contents in the duodenum and then reabsorbed into the
portal circulation by the epithelial lining of the distal part of the intestine.
Hepatocytes merely reabsorbed them from the sinusoids and transport them back
126
Digestive System  Lecture Guide in Veterinary Histology

to the bile canaliculi. About 10% of the bile is synthesized de novo in the
hepatocyte’s sER.

 Bile acids. Cholic acid is synthesized from cholesterol and conjugated with
glycine or taurine to form glucocholic and taurocholic acid, respectively.
 Bilirubin is water-insoluble byproduct of the hemoglobin catabolism that
accompanies the disposal of worn erythrocytes by the cells of the mononuclear
phagocytic system in the spleen, liver and bone marrow. It is carried by the
blood to the hepatocytes, which conjugate it with glucoronic acid to form
bilirubin gluconide. This water-soluble substance is secreted with other bile
components, into the bile canaliculi.

2. Biliary tract. Bile in the canaliculi flows towards the portal triads (opposite to the
blood which flows in the sinusoids). At the lobule periphery, the canaliculi empty
into short, narrow bile ductules (also called cholangioles or Hering’s canal), which
are lined by the cuboidal cells with clear cytoplasm. The ductules deliver the bile to
the bile ducts in the portal triads. The bile duct empty into successively larger ducts,
ending in a single hepatic duct that joins the cystic duct from the gall bladder to
form the common bile duct (ductus choledochus). This empties the bile into the
duodenum. Where the common bile duct penetrates the duodenal wall, it is
encircled by a thick layer of smooth muscle, the sphincter of Oddi. Although the
liver produces bile continuously, the sphincter opens fully only when a particular
fatty meal enters the duodenum. When the sphincter is closed, bile backs up the
common duct, through the cystic duct, and into the gallbladder.

The hepatic and common bile duct are similar in structure to that of the
gallbladder. Both circular and longitudinal muscle layers are present. It is thickest in
bovine and thinnest in carnivores. In other domestic animals, the muscle layer is
continuous.

 In horse, the hepatic duct, carries unconcentrated bile, and the major
pancreatic duct opens into the diverticulum duodeni.
 In pig, the bile duct opens onto the duodenal papilla.
 In dog, the bile duct has 2.5 to 3.0 cm intramural section in the duodenal wall
before opening with a minor pancreatic duct at the major duodenal papilla.
 In cat, the bile duct opens into the duodenum similarly but is accompanied
by a major pancreatic duct.

GALLBLADDER

The gallbladder is a small, hollow organ attached to the inferior surface of the
liver. It receives, stores, and concentrates bile by absorbing its water. In response to the
entrance of dietary fats into the small intestine, a hormone, cholecystokinin, is released
into the blood stream by the enteroendocrine cells located in the intestinal mucosa.
Cholecystokinin is carried in the blood to the gallbladder and causes rhythmic
contractions of the smooth muscle in the wall of the gallbladder. This action forces the
bile to enter the duodenum by way of the common bile duct.
 127
C.T. Reyes, DVM, MS  Digestive System

 Lining epithelium is simple columnar

 No muscularis mucosae.
 Lamina propria- tunica submucosa consists of loose connective tissue.
 Tunica mucosa is extensively folded in the dog and cat and the invaginations
are often misinterpreted as simple glands (Rokitansky-Aschoof sinuses). In ox, it
may present mucosal glands.
 Tunica muscularis - smooth muscle fibers generally course in a circular direction.
 Adventitia and serosa, the outer layer consists of both the adventitia that
attaches it to the liver and serosa that covers its free (peritoneal) surface.

PANCREAS

The pancreas, the main accessory digestive organ, is an encapsulated,

lobulated, compound tuboloacinar gland containing both exocrine and endocrine
secretory units. It produces an alkaline fluid with numerous digestive enzymes that
breakdown proteins, fats, and carbohydrates into smaller molecules for absorption in
the small intestine. The stroma consists of a thin capsule that gives rise to delicate
connective tissue septa separating the parenchyma into distinct lobules.
 Very similar to the parotid salivary glands histologically except that the striated
intralobular ducts, pancreatic islets and centroacinar cells are present.
 The secretory unit is called pancreatic acinus.
 The acinar cells are conical or pyramidal with basophilic basal regions due to the
accumulation of numerous mitochondria and rER. The pancreatic acinus opens
into the intercalated duct lined by simple squamos or cuboidal epithelium.
Larger ducts have simple columnar lining epithelium, may contain goblet cells
and simple branched tubuloalveolar mucous glands.
 The lining cells at the beginning of the duct overlap with the secretory acinar
cells and are therefore called centro-acinar cells.

Exocrine Pancreas

Most of the pancreas consists of the exocrine secretory units or acinar cells, which
secrete the pancreatic enzymes. Pancreatic secretions are regulated both by
hormones and vagal stimulation. Two intestinal hormones, secretin and
cholecystokinin, secreted by the enteroendocrine cells of the duodenal mucosa into
the blood stream, regulate the pancreatic secretions.
In response to the presence of acidic chyme in the small intestine, secretin
release stimulates the pancreatic cells to secrete large amount of watery fluid rich in
sodium bicarbonate ions. This fluid, which has little or no enzymatic activity, is produced
primarily by the epithelial cells that line the smaller pancreatic ducts. The function of
this fluid is to neutralize the acidic chyme and create optimal environment for the
activity of the pancreatic enzymes.
In response to the presence of fats and proteins in the small intestine,
cholecystokinin release stimulates the acinar cells in the pancreas to secrete large
amounts of different digestive enzymes. The pancreatic enzymes that are produced in
the acinar cells enter the duodenum in an inactive form and are then activated by a
hormone secreted by the intestinal mucosa.
128
Digestive System  Lecture Guide in Veterinary Histology

Endocrine Pancreas

Scattered throughout the secretory acini in the pancreas are the pale-staining,
spherical units of the cells called pancreatic islets (islets of Langerhan’s); these islets
constitute the endocrine portion of the pancreas. These islet cells produce important
hormones, insulin and glucagons.

Three main types of cells identified in the pancreatic islets

1. The alpha cells produce the hormone glucagons, whose main physiological
function is to increase the levels of glucose in the blood. This function is primarily
accomplished by accelerating the conversion of glycogen, amino acids, and fatty
acids in the liver into glucose; this conversion elevates the sugar levels in the blood.
2. The beta cells on the pancreatic islets produce the hormone insulin, whose release
is stimulated by the elevation of the glucose after meal. The main physiological
function of insulin is to lower the glucose level of the blood by accelerating
membrane transport of glucose into cells, especially into the cells of liver, muscle,
and fat. Insulin also accelerates the conversion of glucose into glycogen in the liver.
The effects on insulin in the levels of blood glucose are just the opposite to that of
the glucagons.
3. The delta cells secrete the hormone somastostatin, which decreases and inhibits the
secretion of both insulin and glucagons through local action within the pancreatic
islets.

THE AVIAN DIGESTIVE SYSTEM

Regions of the Digestive System

1. Buccal Cavity - differs from that of mammals in the following:

 Lips and cheeks are absent
 Presence of cornified beak appended to the upper and lower jaws
 Tongue is narrow, pointed, devoid of taste buds and contains a bone, the
entoglossal bone
 Teeth are absent although rudimentary tooth buds are present in some birds
 Definitive salivary glands are absent, instead simple branched tubulo mucous
glands are present
 Lining epithelium is keratinized stratified squamos.

2. Esophagus and Crop - presents the following features:

 Lining epithelium is keratinized stratified squamos
 Lamina propria is loose connective tissue with diffuse lymphatic tissue and
lymphatic nodules
 Tunica mucosa and submucosa are arranged in longitudinal folds
 Muscularis mucosa is undulating
 Lamina propria presents simple branched tubuloalveolar mucous glands

The crop or ingluvies is an esophageal diverticulum. The lining epithelium is

thicker than that of the esophagus. Simple branched tubuloalveolar mucous glands
 129
C.T. Reyes, DVM, MS  Digestive System

are present in anseriform birds (ducks and geese) but absent in galliform (chicken
and turkey) and columbiform (pigeon) birds. In columbiform birds, the superficial
cells of the lining epithelium undergo a fatty change to a substance called crop
milk. The crop moistens the food through its mucoid secretions as well as macerates
the materials through the muscular contractions of the tunica muscularis. The other
elements of the wall are similar to the esophagus.

3. Proventriculus - the glandular stomach of birds. The structure of the wall conforms to
the general description of the GI tract except for the following:
 Tunica mucosa is extensively folded into flattened ridges separated by grooves.
Mucosal or rugosal glands, which are simple branched tubular glands, open into
the base of the groove. Elevations in the tunica mucosa (papillae) contain the
opening of the excretory duct of the submucosal or subrugosal glands
 Lining epithelium is simple columnar cells
 Lamina propria is typical but with numerous diffuse and nodular lymphatic tissue
 Muscularis mucosa is interrupted with strands interdigitating between the
mucosal glands
 Tunica submucosa presents large and numerous submucosal or subrugosal
glands which are compound branched tubular glands. The adenomeres radiate
360 degrees around a central excretory duct lined by tall columnar epithelium,
simple or pseudostratified
 Tunica muscularis has three layers: inner longitudinal, middle circular and outer
longitudinal
 Tunica serosa is typical

4. Ventriculus or gizzard or muscular stomach - is connected to the proventriculus by a

narrow isthmus, which is devoid of submucosal glands.
 Lining epithelium is low columnar cells with a cornified secretory product
 Lamina propria presents mucosal glands which secrete cornified substance
covering the lining epithelium
 No muscularis mucosa
 Tunica submucosa is typical and blends with the lamina propria
 Tunica muscularis is present and consists of smooth muscle and dense white
fibrous connective tissue
 Tunica serosa is present but thin

5. Lower Digestive Tract

a) Small intestine is not divisible histologically into different regions. It is similar to the
small intestine of mammals except for the following:
 No muscularis mucosa
 Lamina propria - tunica submucosa contain large quantities of diffuse and
nodular lymphatic nodules
 Tunica muscularis may contain a third layer of circular muscle in the inner
portion.

b) Ceca - are two blind sacs appended to the junction of the small and large
intestines.
130
Digestive System  Lecture Guide in Veterinary Histology

Ceca present the following features:

 Villi are present at the orifice but diminish and are lost at the end of the organ
 Muscularis mucosa is absent
 Lamina propria - tunica submucosa contain numerous diffuse and nodular
lymphatic tissue. The accumulation of nodules at the caecal orifice is called
caecal tonsil.
 Sphincter from the inner circular layer of tunica muscularis is formed at the
junction of the caeca to the small and large intestines. Water absorption
occurs in the caeca and cellulose digestion has also been reported.

c. Rectum - has short and thick villi and an increased number of goblet cells. Other
than this feature, the region is similar to the small intestine.

d. Cloaca - a common orifice for digestive, excretory and reproductive systems. It is

divisible into three regions: coprodeum, urodeum and proctodeum.
 Lining epithelium is simple columnar
 Tunica mucosa is extensively folded
 Lymphatic tissue is present in the associated connective tissue
 Proctodeum presents the bursa of Fabricius.

e. Anus - presents the following features:

 Lining epithelium is keratinized stratified squamos
 Tunica mucosa is highly folded
 Muscularis mucosa is absent
 Striated muscle of the tunica muscularis forms the anal sphincter.