DIGESTIVE GLANDS

SALIVARY GLANDS
There are three pairs of the major salivary glands:
1. the parotid,
2. submandibular and
3. sublingual
and numerous minor accessory glands, scattered throughout the oral mucosa,
connecting with the oral cavity epithelium and derived from it.
The minor glands secrete continuously and are under the local control,
whereas the major glands mainly secrete in response to parasympathetic activity,
which is induced by physical, chemical and psychological stimuli.
Functions. The product of these glands is saliva – a hypotonic watery se-
cretion containing variable amount of mucus, enzymes (amylase, antibacterial en-
zyme – lysozyme), antibodies and inorganic ions and the saliva moistens and lub-
ricates the ingested food to aid swallowing.
General structure of the major salivary glands. They are compound
branched acinar or acino-tubular glands. Each looks like bunch of grapes with
twigs and berries. Each gland is covered by a connective tissue capsule and is di-
vided into numerous lobules, containing numerous secretory units (berries) and
small excretory ducts (twigs). Supporting connective tissue septa radiate between
the lobules from an outer capsule and convey blood vessels, nerves and large ex-
cretory ducts.
As it is known all epithelials and, correspondingly, glands of an anterior
compartment of the digestive system are ectodermal origin, that’s why excretory
ducts and even secretory units have 2 or more layers of cells.
 Fig. 15.14. Salivary glands

Each secretory lobule contains secretory units of one, two or three types
– serous, mucous, and mixed.
Serous acini (protein, enzyme-secreting units) are spherical containing
small pyramid-shaped cells rich in rough endoplasmic reticulum, free ribosomes,
that’s why darker stained on the slides and have the round nuclei centrally loc-
ated.
Mucous secretory unit is tubular, larger than the serous, lighter stained,
mucous cells are columnar and full of mucus; contain flattened nuclei basally
located.
Mixed acini contain both serous and mucous cells. They are the typical
mucous units, surrounded or capped by several serous cells, forming serous de-
milune.
A thin layer of the myoepithelial cells, which lie between the secretory
cells and their basal lamina, surrounds each secretory unit. They are the contract-
ile cells with processes, contraction of these cells helps to expel the secretory
product. The myoepithelial cells also underlie the cells of the proximal portion of
the duct system.
The lumen of the secretory units continuous with a duct system.
 Fig. 15.15. Structure of mixed secretory portion

System of ducts.
Secretory cells discharge their product into the narrow
 intercalated duct, that is smaller than the secretory unit, is lined
by a simple cuboidal epithelium, surrounded by the myoep-
ithelial cells. Further the secret is propels into the larger
 striated duct, which has bigger lumen and is lined by simple
cuboidal or columnar cells with the basal striations, latter are
the folds of the basal cell membrane and numerous mitochon-
dria. Cells of these ducts possess of the secretory activity,
providing the hypotonic feature of the saliva.
 Next secretory duct is called the interlobular and situated in the
interlobular connective tissue septa. It lumen is wider and the
wall is made of slightly stratified epithelium (2–3 layers of
cells).
 The wall of main excretory duct consists of the stratified epithe-
lium.
Parotid glands are chiefly serous, produce a thin watery secretion rich in
enzymes and antibodies, are the largest and have long excretory ducts. Large
amount of connective tissue, good expressed lobular structure, big amount of in-
tercalated and striated excretory ducts in lobules may be the distinguishable fea-
tures.
Submandibular glands are mixed glands that are mostly serous in human.
Some mucous units capped by serous demilune found among the predominant
serous acini. Intercalated ducts are less extensive than in parotid gland.
Sublingual glands are mixed glands but mostly mucus secreting. Some
of predominant mucous acini have serous demilunes, but purely serous acini are
present seldom. Intercalated and striated ducts are difficult to identify or may be
absent.

LIVER
Liver is the largest gland and largest internal organ which acts as a vast
chemical factory.
Functions. The liver has a lot of functions, which account for its com-
plex structure and specific vasculature.
1. Bile synthesis and secretion. Bile is transported from the liver to the
gallbladder, where it is stored and then expelled into duodenum. Bile neutralizes
the acid chyme and is necessary for the lipid digestion.
2. Excretion of bilirubin, which appears in the spleen as a result of the
decay of hemoglobin.
3. Protein synthesis, such as albumen, blood clotting factors (fibrinogen,
prothrombin).
4. Gluconeogenesis – lipids and amino acids are converted into glucose.
5. Storage of glycogen.
6. Storage of fat- soluble vitamins: A (retinol), D (cholecalciferol), vit-
amin K.
7. Iron is stored within the hepatocyte cytoplasm in the form of ferritin.
8. Detoxification of metabolic waste products to produce urea and break-
down some substances as alcohol.
9. Protective function due to phagocytic cells – Kupffer cells. Final
breakdown of some damaged red blood cells.
10. The liver modifies the action of hormones released by other organs:
thyroxine, growth hormone, insulin and glucagon.
11. At the embryonic life the liver works as hemopoietic organ.
Liver is the gland with the mixed secretion – exocrine and endocrine, but
parenchyma is not divided into exo- and endocrine. All of the liver functions are
carried out by the parenchymal cells and are dependent on liver vasculature.
 The liver structure

Fig. 15.16. Diagram of a classic liver lobule

Same as each gland, the liver is surrounded by the connective tissue cap-
sule and the septa of the connective tissue divide the liver into lobes and lobules.
The structural-functional unit of the liver is liver lobule.
The hepatic lobules are hexagonal in shape (a six-sided prism) and each
lobule has about 2 mm long and 1 mm in diameter. It is delimited by interlobular
connective tissue (only little, if any, visible in humans; plentiful in e.g. pigs). In
the periphery, in each of the six corners the portal triads (also known as tracts)
are presented.
The liver receives blood from two vessels, the hepatic artery and the
portal vein. The hepatic artery brings into the liver oxygenated blood (about 25
%). The portal vein brings blood from the digestive tract (absorptive material)
and spleen (about 75% of blood). They enter into the liver together and both di-
vide into branches and in angular places between the hepatic lobules take part in
the portal triad formation.
Take notes, the portal triad is the distinctive feature of the liver, consist-
ing of the branching of hepatic artery (1), branching of portal vein (2) and bile
duct (3). Portal triads are embedded in interlobular connective tissue.
In the middle of each lobule the central vein is situated. In cross sec-
tions, the lobule is filled by cords of hepatic parenchymal cells, hepatocytes,
which radiate from the central vein and are separated by vascular sinusoids. Si-
nusoids are low pressure vascular channels that receive blood from terminal
branches of the hepatic artery and portal vein at the periphery of lobules and de -
liver it into central veins. Thus the sinusoids contain the mixed blood. Sinusoids
are lined with endothelial cells and flanked by plates of hepatocytes.
 Fig. 15.17. Blood supply to the liver: the portal triad

The central veins of the neighboring lobules enter the sublobular vein, usually
big and single and are a branch of the hepatic vein, through which blood leaves
the liver.
Stellate sinusoidal macrophage, or Kupffer cell, are derived from monocytes and
line the sinusoids. Between the hepatocytes and a thin discontinuous simple
squamous epithelium, which lines the sinusoids, is a narrow perisinusoidal space

– space of Disse.

Fig. 15.18. Schematic diagram of a hepatocyte

 This is the site of exchange of materials between the hepatocytes and the
blood. In fetal life space of Disse contains blood-forming islands. The other cell
type found in the perisinusoidal space is the Ito cell (or hepatic stellate cell or
adipose cell). These cells store vitamin A.
The liver cells, called hepatocytes are polyhedral in shape, contain all
organelles in big amount, numerous mitochondria, large deposits of glycogen and
lipid droplets and many other inclusions. Big round nucleus and ability to regen-
eration are the features of hepatocytes. Good developed rER can enlarge. There
are as many as 50 Golgi units in each hepatocyte.

Fig. 15.19. Schematic diagram of a plate of hepatocytes

interposed between hepatic sinusoids

Hepatocytes usually are described as having six surfaces. Two of cells

surfaces appear facing the sinusoids – vascular surface. Four other surfaces face
neighboring cells and form bile canaliculi – biliary surface. So, the bile can-
aliculi haven’t own wall – they are formed by appose grooves in the surfaces of
the adjacent hepatocytes, while the wall of the interlobular bile duct consists of a
layer cuboidal shaped epithelial cells with big round nuclei. Microvilli are
present abundantly on the sinusoidal face and project sparsely into bile canaliculi.
Each hepatic cell expels the component of bile into the tiny bile can-
aliculus, and the bile canaliculi, in turn, expel the bile into the interlobular bile
duct.
In addition, there are several points of view of the liver lobules. They are
called: classical lobule, portal lobule and liver acinus.
The classical lobule has the central vein at the center, and the portal
canals at the peripheral angles of the liver.
The portal lobule has a portal canal (triad) at the center and central veins
at the peripheral angles of the lobule.
The liver acinus is designated by the way of connecting of two central veins and
two portal canals.
Epithelial parenchyma of the liver has good ability for regeneration.
The interlobular bile ducts form anastomosing network surrounding the branches
of the portal vein. Towards the porta, the lumen of the ducts gradually becomes
larger, the epithelium becomes taller and surrounding them connective tissue be-
comes thicker.
The main ducts from the lobes of the liver fuse to form the common hepatic
duct.

Fig. 15.20. Electron micrograph showing plate of hepatocytes (H),

space of Disse (D), sinusoid (E), bile canaliculi (arrow)

The common hepatic duct is about 3 cm long and has all alimentary canal mem -
branes except a muscularis mucosae. Leaving liver hepatic duct through the
cystic duct enters the gallbladder. Then common bile duct (ductus choledochus)
enters the duodenum. Near the duodenum the smooth muscle layer becomes
more prominent and functions as a sphincter in regulating the flow of bile.

The gallbladder is a pear-shaped sac, which stores bile and concentrates

it up to 10 times. About 1 liter bile a day is produced. The wall of the gallbladder
consists of columnar epithelium, lamina propria, layer of smooth muscles, per-
imuscular connective tissue and the serous layer, covering a part of the organ.
The mucosa forms folds, the epithelial cells have short and irregular microvilli, in
the connective tissue the simple tubuloalveolar mucus producing glands are loc-
ated.
Gallbladder muscles contract under the hormones of enteroendocrine
cells of the small intestine in response to the presence of fat in the duodenum and
bile is discharged.
Liver development occurs through a progressive series of interactions
between the embryonic endoderm and mesoderm. The hepatocytes and biliary
epithelial cells are derived from the endoderm and the stroma, stellate cells,
Kupffer cells, sinusoidal endothelial cells and other vascular structures are de-
rived from mesoderm. The first morphological sign of embryonic liver develop-
ment is the formation of the hepatic diverticulum, an outgrowth of the ventral
foregut epithelium. The hepatic diverticulum invades the neighboring septum
transversum, which is the mesodermal plate that separates the embryonic thoracic
and abdominal cavities. Hepatoblasts from the endoderm proliferate within the
septum transversum and become organized in cords around the portal veins,
which are in the process of developing from mesoderm.

Pancreas
Pancreas is the gland with the mixed secretion – exocrine and endocrine.
The pancreatic functions:
1. As the predominantly exocrine gland, the pancreas is a typical serous
gland that synthesizes and secretes into the duodenum enzymes that are essential
for digestion in the intestine. Enzymes are as follows:
1) trypsinogen, pepsinogen, peptidase for the digestion of the proteins;
2) Amylase digests carbohydrates;
3) Lipase digests lipids;
4) deoxyribonuclease and ribonuclease digest nucleic acids.
Notes, they become active only after they reach the lumen of the intest-
ine under the influence of the enterokinases which are situated in the intestinal
absorptive cell membrane.
2. The endocrine component synthesizes and secretes hormones into the
blood.

Pancreas structure
The exocrine pancreas occupies about 98–99 % of the whole volume of
organ and is a compound branched tubuloacinar serous gland that resembles the
parotid gland, with which it can be confused.
Like each gland, the pancreas is covered with connective tissue capsule.
Many connective tissue septa divide gland into distinctive lobes and lobules. In-
terlobular ducts, blood vessels, nerves are arranged in the interlobular septa. Lob-
ules consist of the secretory units and short intralobular ducts.
 Fig. 15.21. Pancreatic acinar cell

The cells that synthesize and secrete digestive enzymes are arranged in
grape-like clusters called acini − secretory units, very similar to what is seen in
salivary glands. The pancreocytes are characterized by a distinct basophilia in the
basal part of the cell and by the presence of acidophilic zymogen granules in the
apical cytoplasm. Zymogen granules contain a variety of digestive enzymes, rich
alkaline fluid in an inactive form. The basal zone contains a lot of endoplasmic
reticulum, elongated mitochondria and a well-developed Golgi complex.
Like any exocrine gland pancreas has not only secretory units, but also a
system of excretory ducts. Duct cells secrete a watery, bicarbonate-rich fluid
which flush the enzymes through the ducts and play main pivotal role in neutral-
izing acid within the small intestine. Pancreatic ducts are classified into four
types:
The (1) intercalated duct begins actually within the acinus and the duct
cells and is lined with cuboidal epithelium that extends up into the lumen of the
acinus to form what are called centroacinar cells. These cells secrete a fluid
reach in sodium and becarbonate. The intercalated duct is very short and drain to
short (2) intralobular collecting duct, wall of which is formed with the cuboidal
epithelium. The network of intralobular ducts leads to the larger (3) interlobular
ducts, which are lined with a columnar epithelium in which enteroendocrine and
occasional goblet cells may be found. Notes, the big interlobular and (4) main
pancreatic duct except epithelium have a layer of connective tissue with some
amount of smooth muscle cells – so called mucosa. That’s why look at the slides
a little folded.
The most distinctive feature of the pancreas are islets of Langerhans –
the endocrine component of the pancreas is scattered throughout of organ. They
constitute about 1-2 % of the pancreatic volume.
Islets may contain a few cells or many hundreds of cells. Each islet con-
sists of polygonal pale-stained cells with a network of fenestrated capillaries
 Absence of an inadequate amount of insulin leads to elevated blood
glucose level and the presence of the glucose in the urine, a condition known as
diabetes mellitus.
2. A-cells constitute about 15-20% and are located peripherally in the is-
lets. They secrete glucagon, which has metabolic effects oppose the action of in-
sulin. It stimulates the breakdown of glycogen in the liver, and release of glucose
into blood stream.
3. D-cells constitute about 5-10%, are also located peripherally. They
secrete somatostatin is identical with the hypothalamic hormone, that regulates
somatotropin of pituitary gland and may inhibit insulin and glucagon secretion.
The minor islet cells constitute about 5%. They are:
PP-cells responsible for secretion of pancreatic polipeptide (PP), which
stimulates gastric chief cells, inhibits bile secretion, intestinal motility and pan-
creatic enzymes;
D1-cells responsible for secretion of VIP (vasoactive intestinal poly-
peptide, which stimulates secretory activity and motility in the gut;
EC-cells responsible for secretion of secretin (increases pancreatic and
gallbladder activity), and motilin (inhibits gastric secretion and motility).
At the foregut/midgut junction the septum transversum generates 2 pan-
creatic buds (dorsal and ventral endoderm) which will fuse to form the pancreas.
The dorsal bud arises first and generates most of the pancreas. The ventral bud
arises beside the bile duct and forms only part of the head and uncinate process
of the pancreas.