Department of Zoology

B.Sc-II Semester-IV
Histology of Mammalian Organs
By- Mr. Om.S. Ingale

1.Liver

The liver is the most vital and largest internal organ in the mammalian
body, weighing approximately 1.5kg in human. Anatomically, it has
lobular structure and lies in the abdominal cavity below diaphragm in
human. It consists of four lobes; two larger ones, which are right and left,
and two smaller ones which are quadrate and caudate.
Histologically, it has a complex microscopic structure that can be viewed
from several different angles.
Physiologically, the liver also performs many essential functions ..

Functions and physiology: The liver performs several important

functions in the human body, such as given below:
− Plasma protein synthesis - albumins, lipoproteins, glycoproteins,
prothrombin, fibrinogen
− Vitamin storage and modification - vitamins A, D, and K − Iron storage
and metabolism
- transferrin, haptoglobin, hemopexin, ferritin
− Drugs and toxins degradation
− Bile production
− Carbohydrate metabolism

Histological components: The liver consists of the following major

histological components:
Parenchyma: It is represented by hepatocytes.
Stroma: It is a continuation of the surrounding capsule of Glisson. It
consists of connective tissue and contains the vessels. The capsule is also
covered by a layer of mesothelium, arising from the peritoneum covering
the liver. The connective tissue of the stroma is type III collagen
(reticulin), which forms a meshwork that provides integrity for the
hepatocytes and sinusoids.
Sinusoids: These are capillaries travelling between hepatocytes.
Spaces of Disse (perisinusoidal spaces): These are located between the
hepatocytes and the sinusoids.
 Histologically liver consists of a large number of microscopic
functional units. These functional units are hepatic lobule, portal lobule
and liver acinus.
Hepatic lobule: It It is an independent venous unit consisting of
hexagonal plates of hepatocytes stacked on top of each other. Within each
plate, the hepatocytes radiate outwards from a central vein. the
hepatocytes are arranged into strips known as hepatic lamina. Spaces
between the hepatic lamina are called hepatic lacunae, which are
occupied by hepatic sinusoids. Hepatic sinusoids travel between the strips
of hepatocytes, draining into the central vein. One portal canal is located
at each corner of the hexagonal classic lobule, making a total of six for
each lobule.

Portal lobule: Peripherally, each lobule has 3 to 6 portal areas with more
fibrous connective tissue, each of which contains interlobular structures
that comprise the portal triad. .
Hepatic regions contain liver cells and bile canaliculi that are responsible
for production, secretion and transportation of bile in the hepatic regions.
Bile canaliculi: It is formed by spaces present between the plasma
membranes of adjacent liver cells. It forms hexagonal networks around
the liver cells. The canaliculi pass to periphery of the hepatic lobules
where they form interlobular canal of Herring that finally drains into the
interlobular duct of the portal area. Bile canaliculi are intralaminar and
centrifugal in direction.

Cells of the liver

Hepatocytes: Hepatocytes constitute a major fraction of hepatic cell

population. These are cuboidal or polyhedral in shape with round central
nuclei. These cells are arranged in single-cell cords or plates. These
hepatocytes are linked together via intercellular adhesion complexes and
tight junctions. Their one side faces the persinusoidal space, while the
other faces the bile canaliculi and covered with microvilli.
Hepatocytes are responsible for most of the liver functions such as
metabolism, detoxification, synthesis, and storage of nutrients,
carbohydrates, fats, and vitamins.

Kupffer cells: Kupffer cells are resident macrophages in liver with

largest population. They are attached to the luminal surface of the
sinusoidal endothelium. These cells are essential for the phagocytosis of
foreign particles, infecting organism as well as cytokines products.
Hepatic stellate cells: These cells are located between hepatocytes.
These cells are associated with several functions such as secretion of
cytokines, storage of vitamin A and synthesis of hepatic extracellular
matrix.
Biliary epithelial cells: These cells line the bile duct in portal triads.
They are also known as the cholangiocytes. The connecting duct between
bile duct and bile canaliculi (canal of Herring) along with hepatocytes are
also lined by these biliary cells..
Endothelial cells: These cells are largest group of non-parenchymal cells
of liver and line the intrahepatic circulatory vessels of liver and provide a
large surface area for nutrient absorption.
Lymphocytes: Lymphocytes are present everywhere in the liver
parenchymal sinusoids. These lymphocytes are also a part of innate
immune system and selectively rich in NKT cells and natural killer cells
providing defense against invading pathogens.
 2. Pancreas

The pancreas is both an exocrine accessory digestive organ and a

hormone secreting endocrine gland. The bulk of the pancreatic
tissue is formed by the exocrine component, which consists of
many serous pancreatic acini cells. These acini synthesize and
secrete a variety of enzymes essential to successfully “rest and
digest”.
The endocrine component is a much smaller, but equally
important, portion of the pancreas. It is composed of pancreatic
islets, which appear as islands of cells dispersed between the
pancreatic acini. These islet cells produce and secrete hormones
that regulate glucose, lipid and protein metabolism.
The pancreas is a large, mixed gland composed of five parts: the
head, uncinate process, neck, body and tail. The location of the
pancreas is mostly retroperitoneal, except for the tail. This organ
extends from the C-shaped curve of the duodenum, passes behind
the stomach and finishes at the hilum of the spleen.
Several pancreatic ducts extend throughout the pancreas,
emptying the pancreatic contents inside the duodenum.

The pancreas is covered by a thin capsule made of loose

connective tissue.
The parenchyma consists of pancreatic acini and sparsely
scattered pancreatic islets surrounded by stroma of loose
connective tissue.
Interlobular connective tissue septa project from the capsule
into the pancreatic parenchyma, organising it into lobules. The
interlobular septa house the interlobular ducts, blood vessels,
nerves, and lamellar (Pacinian) corpuscles, which are special
types of sensory receptors.
Exocrine pancreas

Secretory units
The exocrine component of the pancreas makes up about 98% of
the pancreatic tissue. It is comprised of densely packed serous
acinar (tubuloacinar) glands. These glands are called pancreatic
acini, which represent the secretory units of the pancreas. They
are formed out of simple epithelium. Each pancreatic acinus
consists of pyramidal-shaped acinar cells, which have a broad
basal portion and a narrow apical surface that surround a small
central lumen.
These acinar cells are serous secretory cells that
produce digestive enzymes.
Zymogen granules are large secretory organelles in which acinar
cells store their inactive enzymes, called zymogens or
proenzymes. Upon stimulation, the zymogens are activated and
the acinar cells release their secretions by way of exocytosis.
Duct system
 Once synthesized, the pancreatic secretions leave the acini via
the intercalated ducts. The latter are short ducts with a small
lumen that start within the acini. Centroacinar cells are continued
by simple, low cuboidal ductal cells that line the extra-acinar
portion of the intercalated ducts which extends outside the
acinus. Intercalated ducts drain into intralobular ducts, which
are lined by simple, low columnar epithelium.
The endocrine component makes up about 2% of the pancreas,
which is represented by about 1-2 million pancreatic islets (of
Langerhans). They are dispersed throughout the exocrine
component of the pancreas, most of them being located in the
tail region..
Pancreatic Islets are spherical clusters of polygonal endocrine
cells. The cells of the islets are connected to each other with
desmosomes and gap junctions, forming bands or cords of
cells. Pancreatic islets are permeated by many fenestrated
capillaries, which allow quick entry of pancreatic hormones into
the blood.
There are four main types of cells in the pancreatic islets:

• B (beta) cells - these cells secrete insulin and constitute about

70% of the islet cells. They are most commonly located in the
central part of the islet. B cells contain many secretory granules
which possess a dark center with crystallized insulin,
surrounded by a wide pale halo.
• A (alpha) cells - these cells secrete glucagon and constitute 15-
20% of the islet cells. They are usually larger than B cells and
most commonly located peripherally in the islet. Their granules
are more uniform in size, with a larger dark center surrounded
by a thinner halo compared to B cells. The granules are filled
with glucagon.
• D (delta) cells - these cells secrete somatostatin and constitute
5-10% of the islet cells. They are located diffusely throughout
the islet but most commonly in the periphery. D cells contain
larger secretory granules compared to A and B cells.
• PP (pancreatic polypeptide) cells - these cells
secrete pancreatic polypeptide and constitute <5% of the islet
cells. They are mostly located within the head of the pancreas.

Function
The hormones of the endocrine pancreas are the primary
regulators of glucose, lipid and protein metabolism.

3. Testis
Introduction:
The main functions of the male reproductive system, are to
produce spermatozoa, androgens. The male reproductive system
includes the testis, genital ducts, accessory sex glands and penis.
The testis is surrounded by a thick capsule, the tunica albuginea,
from which a conical mass of connective tissue, the mediastinum
testis, projects into the testis. The tunica albuginea is covered
externally by a serosa. From the mediastinum, delicate fibrous
septa radiate towards the tunica albuginea and divide the
parenchyma of the testis into about 300 lobuli testis, which
communicate peripherally. Each lobule contains 1- 4 convoluted
seminiferous tubules Interstitial tissue between the convoluted
tubules is continuous with a layer of loose vascular connective
tissue, the tunica vasculosa testis, which is found beneath the
tunica albuginea. Each seminiferous tubule continues near the
mediastinum into a straight tubule, a tubulus rectus. The straight
tubules continue into the rete testis, a labyrinthine system of
cavities in the mediastinum. These tubules are enclosed by a
thick basal lamina and surrounded by 3-4 layers of smooth muscle
cells (or myoid cells). The insides of the tubules are lined with
seminiferous epithelium, which consists of two general types of
cells: spermatogenic cells and Sertoli cells.
The production of sperm and eggs/ova (gametes) is a procedure
called gametogenesis (spermatogenesis and oogenesis).
Gametogenesis involves two rounds of meiotic cell division, in
which one diploid cell gives rise to 4 haploid cells. The germinal
(seminiferous epithelium) of the seminiferous tubules contains
spermatogenic cells and Sertoli cells. The spermatogenic cells
divide by mitosis, then meiosis to form gametes, which mature
into sperm by the process of spermiogenesis. Unusually, the
developing spermatogenic cells remain connected by cytoplasmic
bridges, until they have formed a mature spermatozoan.
Primary spermatocytes: lie in the cell layer luminal to the
spermatogonia. They appear larger than spermatogonia. They
immediately enter the prophase of the first meiotic division, and
results in the formation of Secondary spermatocytes
Secondary spermatocytes:are smaller than primary
spermatocytes. They rapidly enter and complete the second
meiotic division and are therefore seldom seen in histological
preparations. Their division results in the formation of
Spermatids.
Spermatids: lie in the luminal part of the seminiferous
epithelium. They are small (about 10 µm in diameter) with an
initially very light (often eccentric) nucleus. The chromatin
condenses during the maturation of the spermatids into
spermatozoa, and the nucleus becomes smaller and stains darker.
The terminal phase of spermatogenesis is called
spermiogenesis and consists of the differentiation of the newly
formed spermatids into Spermatozoa
Spermatozoa:The mature human spermatozoon is about 60 µm
long and actively motile. It is divided into head, neck and tail.The
head (flattened, about 5 µm long and 3 µm wide) chiefly consists
of the nucleus (greatly condensed chromatin!). The anterior 2/3
of the nucleus is covered by the acrosome, which contains
enzymes important in the process of fertilisation. The posterior
parts of the nuclear membrane forms the so-called basal plate.
The neck is short (about 1 µm) and attached to the basal plate. A
transversely oriented centriole is located immediately behind the
basal plate. The neck also contains nine segmented columns of
fibrous material, which continue as the outer dense fibres into the
tail. The tail is further divided into a middle piece, a principal
piece and an end piece. In the middle piece (about 5 µm long),
the axonema and dense fibres are surrounded by a sheath of
mitochondria. The middle piece is terminated by a dense ring, the
annulus. The principal piece is about 45 µm long. It contains a
fibrous sheath, which consists of dorsal and ventral longitudinal
columns interconnected by regularly spaced circumferential
hoops. The fibrous sheath and the dense fibres do not extend to
the tip of the tail.
Sertoli cells:
Like all epithelial cells, the Sertoli cells are avascular. Sertoli
cells suport the germ cell progenitors and help to transfer
nutrients from the nearby capillaries. The developing
spermatogonia rely on the Sertoli cells for all of their nourishment.
Leydig cells: are 'interstitial' cells (as they lie between the
tubules). They have pale cytoplasm because they contain many
cholesterol-lipid droplets. The Leydig cells make and secrete
testosterone, in response to lutenising hormone from the pituitary.
Testosterone promotes production of spermatozoa, secretion
from the accessory sex glands, and acquisition of male secondary
characteristics. This process does not start until puberty when LH
stimulates the Leydig cells to produce testosterone.
Sperm-
 Ovary
THE OVARY -The ovary is a rounded body approx. 3 x 1.5 cm
long and 1 cm thick. It is encapsulated by the tunica albuginea, a
dense layer of connective tissue which is covered by the germinal
epithelium (Ovarian surface epithelium), a layer of simple
squamous or cuboidal epithelium. The ovarian follicles, which
enclose the oocytes, are primarily embedded in the cortical region
of the ovary while the medullary portion contains a rich vascular
bed. The ovarian stroma consists of characteristic spindle shaped
fibroblasts that respond to hormonal stimuli, reticular fibers and
ground substance.
PRIMORDIAL FOLLICLE- An ovarian follicle progresses
through several distinct phases before it releases its ovum. During
the first five months of development, a finite number of
primordial follicles form in the fetal ovary. These follicles consist
of oocytes surrounded by a single layer of squamous follicular
cells. At puberty, they begin to develop further and become
primary follicles.
EARLY PRIMARY FOLLICLE- At the start of each menstrual
cycle a limited number of primordial follicles are triggered to
develop. The first apparent histological stage is the early primary
follicle that consists of a central oocyte surrounded by a single
layer of follicular cells which have become cuboidal. The zona
pellucida is a thin band of glycoproteins that separates the oocyte
and follicular cells. Proteins on the surface of sperm will bind to
specific glycoproteins in the zona pellucida.
LATE PRIMARY FOLLICLE- The late primary follicle stage
is reached when the follicular cells proliferate into a stratified
epithelium known as the zona granulosa..
SECONDARY FOLLICLE- The characteristic feature that
distinguishes secondary from primary follicles is the appearance
of a follicular antrum within the granulosa layer. The antrum
contains fluid which is rich in hyaluronan and proteoglycans.
estrogens.
GRAAFIAN FOLLICLE- The Graafian follicle is the stage
after the first meiotic division has completed but before ovulation.
The follicle is characterized by a large follicular antrum that
makes up most of the follicle. It is surrounded by the zona
pellucida and a layer of several cells known as the corona radiata.
When released from the Graafian follicle and into the oviduct, the
ovum will consist of three structures: oocyte, zona pellucida and
corona radiata.
CORPUS LUTEUM After release of the ovum, the remaining
cells of the granulosa and theca interna form the corpus luteum.
The center contains the remains of the blood clot that formed after
ovulation. Surrounding the clot are glanulosa lutein cells and on
the outside theca lutein cells. These cells produce progesterone
and to a lesser extent cholesterol. CORPUS ALBICANS If
fertilization does not occur, the cells of the corpus luteum remain
active for roughly 14 days until the levels of LH fall and the
corpus luteum involutes to form the corpus albicans. The
secretory cells of the corpus luteum degenerate, are phagocytosed
by macrophages and replaced by fibrous material.
Stomach
The stomach is a key part of the gastrointestinal (GI) tract, sitting between
the esophagus and duodenum. Its functions are to mix food with stomach
acid and break food down into smaller particles using chemical and
mechanical digestion.

The stomach can perform these roles due to the layers of the stomach wall.
These are the gastric mucosa, submucosa, muscularis externa and
serosa. The outer layer of the stomach wall is smooth, continuous with the
parietal peritoneum. The inner wall (mucosa and submucosa layers) is
thrown into folds known as rugae, or gastric folds, The various tissue
layers of the stomach wall then combine their functions to digest the bolus
into a viscous, pulpy fluid called chyme.

Mucosa- The various tissue layers of the stomach wall then combine their
functions to digest the bolus into a viscous, pulpy fluid called chime . The
innermost layer of the stomach wall is the gastric mucosa. It is formed by a
layer of surface epithelium and an underlying lamina propria and
muscularis mucosae. The surface epithelium is a simple columnar
epithelium. It lines the inside of the stomach as surface mucous cells and
forms numerous tiny invaginations, or gastric pits, which appear as
millions of holes all throughout the stomach lining. These gastric pits are
important as they are connected to the various glands of the stomach.

There are 3 types of glands found in the stomach; cardiac, gastric and
pyloric, named after the region in which they are found. These glands
produce the digestive enzymes and mucous secretions of the stomach.

Surface mucous cells-- The surface mucous cells, also known as foveolar
epithelium, are the simple columnar epithelium lining the lumen of the
stomach. They secrete alkaline, highly viscous mucus,

Gastric pits

Gastric pits are formed by invaginations of the surface epithelium. Gastric

pits connect to gastric glands and thus allow the glandular products to be
delivered into the stomach lumen.

Gastric glands-

Gastric glands open into the base of gastric pits. They are found throughout
the entire inner surface of the stomach and are divided into 3 types
depending on the region in which they are found. Gastric glands
proper (principal glands) are found in the fundus/body of the stomach. The
cells of these glands produce around two litres of gastric juice a day. The
mucus secreting pyloric glands are only associated with the pyloric antrum
and cardiac glands are located only within the cardia of the stomach.

Gastric pits and gastric glands are made up of the same 5 cell types:
mucous neck cells, stem cells, parietal (oxyntic) cells, chief (zymogenic)
cells and enteroendocrine cells.

Lamina propria and muscularis mucosae

The lamina propria is the layer of connective tissue located just deep to the
surface epithelium. It contains blood and lymphatic vessels, lymphoid
tissue and surrounds the gastric glands.

The muscularis mucosae layer consists of two thin layers of smooth

muscle. It separates the lamina propria from the underlying submucosa.
The inner layer of muscularis mucosae consists of circular fibres while the
outer layer fibres are arranged longitudinally. Its function is to help expel
the secretions of the gastric glands into the stomach lumen.

Submucosa-Deep to the mucosa is a thick layer of connective tissue

known as the gastric submucosa. Its arrangement means that it is durable,
yet flexible and mobile. Aside from rich vasculature and lymphatics, this
layer also holds the submucosal (Meissner’s) plexus.

Muscularis externa-The gastric muscularis externa, also known as tunica

muscularis, is the smooth muscle located deep to the submucosa. It is made
up of 3 layers: inner oblique, middle circular and outer longitudinal. The
muscularis externa layer produces churning movements required for
mechanical digestion. When these layers contract, they throw the mucosa
and submucosa into rugae.

Serosa-

Gastric serosa is the outermost layer of the stomach wall. It consists of a

layer of simple squamous epithelium, known as mesothelium, and a thin
layer of underlying connective tissue. The mesothelium produces serous
fluid, which lubricates the outer wall of the stomach and ensures its smooth
movement in the abdominal cavity. The serosa is continuous with the
parietal peritoneum. It is absent at the attachment sites of the greater and
lesser omenta to the stomach, as well as over a small superoposterior area
near the cardiac orifice where the stomach is attached to the diaphragm via
gastrophrenic and gastropancreatic folds.
HISTOLOGY OF MAMMALIAN KIDNEY
The kidneys are paired retroperitoneal organs of the urinary system. Its
main function is to filter blood and produce urine. Each kidney consists
of a cortex, medulla and calyces. Nephrons are the main functional units
of the kidney that removes metabolic wastes and excess water from the
blood and adjust water, salt and pH to maintain the homeostatic balance
of tissue fluids. It is central organs or homeostasis in the organisms. In
human, around 180 liters of blood are filtered per day by the kidneys,
accounting for around 20% of cardiac output. It is also known to regulate
blood pressure through the renin-angiotensin-aldosterone system,
erythrocyte production through production of erythropoietin, and
circulating calcium and phosphate levels, in part through the activation of
vitamin D.
Structure of kidneys- The kidney is usually a bean shaped organ with a
convex lateral surface, concave medial surface and superior and inferior
poles. The medial surface features the hilum of the kidney, which is the
passageway for the renal vessels and the ureter. A connective tissue
capsule, called as renal capsule and a layer of perinephric (perirenal) fat
protect and cushion the kidney. The capsule contains a layer of contractile
cells called myofibroblasts, which make the capsule able to adapt to the
constant pressure changes within the kidney.
The kidney parenchyma consists of two layers; an outer cortex and inner
medulla. There are about one million nephrons ensheathed in the renal
capsule. Urine is collected into a system of renal calyces, which is a
series of distinctive chambers within a kidney. Calyces gradually increase
in size, starting with the minor calyces, which open into larger major
calyces that empty into the renal pelvis. From the renal pelvis, the urine
passes into the ureter. The portion of the kidney which contains the
calyces, renal pelvis, ureter and renal vessels is called the renal sinus.
Nephron –
The nephron is the functional unit of the kidney. It produces concentrated
urine by creating an ultrafiltrate from blood. A nephron consists of two
main parts: a renal corpuscle and its associated renal tubule system. Renal
corpuscles are located in the renal cortex, while their tubular systems
extend into the medulla. Depending on their distribution and morphology,
there are two main types of nephrons in the kidney; cortical and
juxtamedullary. Cortical nephrons have their corpuscles close to the
kidney capsule. Their tubules are very short, extending only into the
upper medulla andare known as proximal tubule.
Renal corpuscles -The renal corpuscle is the filtration apparatus of the
nephron. Each corpuscle consists of two main elements; the glomerulus
and glomerular (Bowman's) capsule. The glomerulus is a network of
capillaries formed by branches of the renal artery (afferent and efferent
arterioles). It surrounds the glomerulus and hence also known as
glomerular capsule. It consists of two layers (parietal and visceral), which
bound a cavity called the glomerular capsular space (Bowman’s / urinary
space). The inner visceral layer is made of special cells called podocytes.
Podocytes cover the walls of glomerular capillaries, interdigitating with
each other and forming narrow slits between their projections. The outer
parietal layer is made of simple squamous epithelium and is continuous
with the nephron tubules. The afferent and efferent arterioles enter the
renal corpuscle at the vascular pole, while the site where the glomerular
capsule narrows and continues as the proximal thick segment of the
nephron is called the urinary pole. Renal tubular system The tubule
system is the part of the nephron which processes glomerular ultrafiltrate
into urine by reabsorbing necessary molecules and secreting the
unnecessary and waste substances. It consists of three parts; proximal
tubule, nephron loop, and distal tubule.
Proximal tubule: The proximal tubule is the first part of the tubular
system. It consists of convoluted and straight segments. The proximal
convoluted tubule is located within the renal cortex and is continuous
with the capsular space. The straight proximal tubule (or thick descending
limb) extends down into the medulla. Both parts are composed of simple
cuboidal epithelium, rich in mitochondria and microvilli (brush border).
Medullary loop or nephron loop: The nephron loop is the U-shaped
bend of a nephron which extends through the medulla of the kidney.
Histologically, it consists of two parts; thin descending and thin
ascending limbs. Both limbs are composed of simple squamous
epithelium. The cells have few organelles, little to no microvilli and low
secretion abilities. The two limbs work in parallel, with the surrounding
vasa recta capillaries, to adjust the filtrate’s salt (e.g. sodium, chloride,
potassium) and water levels. More specifically, the descending limb is
highly permeable to water, less permeable to solutes, while the ascending
limb is the opposite.
Distal tubule: The distal tubule also consists of straight and convoluted
segments. The straight distal tubule (thick ascending limb) continues on
from the thin ascending limb of the nephron loop at the level between the
inner and outer medulla. The convoluted distal tubule projects into the
cortex. Both parts of the distal tubule are composed of simple cuboidal
epithelium, similar in morphology to the proximal tubule.
Juxtaglomerular apparatus (JGA): It is a collection of cells that lies
into the vascular pole of the nephron. It is formed by 3 types of cells;
macula densa, juxtaglomerular granular (JG) cells and extraglomerular
mesangial (Lacis) cells. It regulates glomerular blood flow and filtration
rate, and systemic blood pressure.
Collecting system or collecting tubules
The collecting system of the kidney is a series of tubes that moves urine
from the nephrons into the minor calyces. Several distal convoluted
tubules from neighbouring nephrons drain into a collecting duct via
connecting/collecting tubules. Collecting ducts then travel through the
kidney medulla, converging at the apex of each renal pyramid. Here,
several ducts merge to form a single large papillary duct (of Bellini),
which opens into the minor calyx.