Histology M.

Sandig

BLOOD VESSELS

Introduction:
The function of the cardiovascular system which includes the heart and blood vascular system is to
distribute oxygen, nutrients and hormones and to collect CO2 and other waste products. The blood
vascular system includes arteries, capillaries and veins. Arteries carry blood from the heart to the
capillaries (where blood and tissue exchanges occur) while veins return blood to the heart.

Arteries
The largest arteries emerge directly from the heart and branch repeatedly giving rise to a large
number of smaller arteries.

3 layers in wall of arteries:

1) Tunica Intima:
- innermost layer lining the lumen comprised of:
- endothelium (simple squamous epithelium)
- subendothelial layer of connective tissue
- internal elastic lamina
- layer of elastin, appears as wavy pink line- fenestrations permit diffusion of metabolites into
- media from lumen
2) Tunica Media:
-3-50 concentric layers of tightly packed smooth muscle cells
- Some smooth muscle cells may run longitudinally
- outer border has a sharp demarcation from a thin external elastic lamina
3) Tunica Adventitia:
- layer of connective tissue containing fibroblasts, elastic and collagenous fibres
- merges with surrounding connective tissue
- limits vessels - holds them in place

Elastic Arteries:
- largest arteries, closest to the heart - include the aorta and its main branches
- the tunica media is highly elastic
- serves as an auxiliary pump
- reduces variations in blood pressure due to pulse
Tunica Intima
- thickness varies with age, relatively thicker than in smaller arteries
- typical endothelium
- subendothelial layer of connective tissue, smooth muscle and poorly demarcated internal elastic
lamina
Tunica Media
- contains 50-60 concentric, fenestrated elastic laminae
- between laminae connective tissue containing concentric smooth muscle cells
- vasa vasorum: small blood vessels located in outer third of tunica media as well as in adventitia
- nerves are also located in the outer layer of the tunica media and adventitia (nervi vasorum)
Tunica Adventitia
- a thin layer comprised of irregularly arranged connective tissue
- small blood vessels and lymphatics present
- adventitia serves as a sheath to restrain overexpansion and prevent aneurysms

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Muscular Arteries:
- most numerous arteries which vary greatly in size
- tunica media highly muscular - muscle alters the diameter of the vessel
- adjust the volume of blood to suit the needs of the tissue being serviced
Tunica Intima
- thin endothelium often lies adjacent to the internal elastic lamina
- little connective tissue between the endothelial cells and internal elastic lamina
Tunica Media
- composed mainly of circularly arranged smooth muscle cells in connective tissue
- external elastic lamina is visible
Tunica Adventitia
- relatively thicker than in elastic arteries
- rich in elastic fibres

Atherosclerosis: inflammatory diseasev of vascular wall:

- monocytes migrate through the endothelium and enter the tunica intima
- these cells ingest lipids and become foam cells
- smooth muscle cells proliferate and lay down collagen forming a plaque in the intima.
- lipid accumulates in the thickened intima and plaque calcifies

ARTERIOLES:
- maintain and regulate blood pressure entering the capillary bed
- this is achieved by arterioles due to their narrow lumen and thick muscular wall
- arterioles are vessels with diameters less than 100µm
- the media consists of 1-5 layers of circular smooth muscle, number of layers varies with size

CAPILLARIES:
- endothelial tubes that connect the arterial and venous circulation
- arranged in the form of capillary beds
- the degree of metabolism in a region determines the closeness of the network
- the wall of capillaries consists of an endothelium and a supporting layer of connective tissue

Types of Capillaries:
1. Continuous Capillaries
- in the central nervous system, muscles and connective tissue throughout the body
- the lumen is slightly larger than the diameter of an erythrocyte
- many pinocytotic vesicles in endothelial cells permit fluid transport across the cell
- cells held together by tight and adherens junctions preventing diffusion between cells
- basement membrane underlies endothelium - exerts some control over diffusion
2. Fenestrated Capillaries
- in renal glomeruli, endocrine glands, intestine, pancreas
- similar to continuous but endothelium perforated by many pores or fenestrations
- fenestrations facilitate exchange across the endothelium
3. Discontinuous capillaries (SINUSOIDS)
- found in bone marrow, spleen and liver
- thin-walled, vascular channels with an irregular outline
- lumen up to three times larger than normal capillaries
- lie close to cells of the organ in which they are located and conform to available space

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Pericytes:
- are perivascular cells scattered around the outside of capillaries
- have long cytoplasmic processes which extend around the blood vessel
- may become phagocytic or exhibit contractile properties
- may differentiate into other cell types during growth or wound healing (muscle, fibroblasts)

ARTERIOVENOUS ANASTOMOSES:
- enable arterioles to connect with venules bypassing capillaries
- possess thick, muscular walls which are richly innervated
- contract when stimulated by mechanical or chemical stimuli
- function in regulating supply of blood to capillary bed of tissues
- numerous in skin where they bypass capillaries and thereby conserve heat

VENULES
- blood from capillaries enters postcapillary venules, collecting venules, muscular venules
- relatively large lumen and thin wall compared to an arteriole
- Media not always present and smooth muscle cells are more dispersed when present
- with increasing size, pericytes increase in number, muscle appears, adventitia thickens

VEINS:
Medium and Small Sized Veins:
- Three layers are usually less organized than arteries
- the tunica intima consists of endothelium, limited subendothelial connective tissue and
- the tunica media is much thinner than in an artery of comparable size
- contains circularly arranged smooth muscle cells like arteries, but thinner
- tunica adventitia is the thickest layer

Large Veins:
- structure highly variable

- subendothelial layer thicker and media thinner relative to smaller veins

- media contains smooth muscle cells in both circular and longitudinal bundles
- adventitia by far the thickest layer

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GASTROINTESTINAL TRACT

- long muscular tube from oral cavity to anus

- includes associated glands (liver, pancreas)
- functions in digesting food and absorbing nutrients

General Plan
(1) Mucous Membrane (Mucosa)
(a) Epithelium – protective, or absorptive-secretory
(b) Lamina Propria - loose connective tissue with lymphocytes
(c) Muscularis Mucosae - layer of smooth muscle
(2) Submucosa
(a) loose connective tissue with elastic fibres
(b) blood vessels and nerve plexus - Meissner's plexus
(3) Muscularis Externa
- 2 thick layers of smooth muscle
-inner circular
-outer longitudinal
-peristalsis
-Auerbach's plexus
(4) Serosa or Adventitia
- mesothelium or connective tissue covering

Glands
- in lamina propria, submucosa
- some lie outside GI tract, connected to it
(e.g. salivary glands, liver, pancreas)

ESOPHAGUS
- straight tube, conveys food to stomach from pharynx
- numerous longitudinal folds in lumen
(1) Mucous Membrane
(a) Epithelium - stratified squamous - non-keratinized (“wear and tear”)
(b) Lamina propria - loose connective tissue with lymphocytes, papillae, cardiac glands
(c) Muscularis mucosae - longitudinal smooth muscle
(2) Submucosa
- dense connective tissue with lymphocytes and esophageal glands
(3) Muscularis externa
- 2 concentric layers. Upper third skeletal, middle third mixed, lower third smooth muscle.

STOMACH
- functions as a reservoir and a digestive organ, limited absorption
- rugae - folds in mucosa
- gastric pits (foveolae) - invaginations of surface

(1) Mucous Membrane

(a) Epithelium - columnar "surface epithelial cells"
- mucus in apical portion
- no goblet cells
(b) Lamina propria - filled with gastric glands
- delicate fibres and usual ct cells plus lymphocytes

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(c) Muscularis mucosae - 2 or 3 layers of smooth muscle

(2) Submucosa - denser connective tissue containing blood and lymph vessels
(3) Muscularis Externa - 3 layers: - outer longitudinal, - middle circular, - inner oblique
(4) Serosa - thin layer of connective tissue covered with mesothelium

Gastric Glands (Fundic Glands) - produce enzymes, HCl

- 4 types of cells
1. Chief (Zymogenic) Cells
- low columnar with zymogen granules (pepsinogen and lipase)
- base of cell basophilic
2. Parietal Cells
- secrete HCl, intrinsic factor (required for Vitamin B12 absorption)
- between chief cells, very eosinophilic, large cells
- many mitochondria, no granules
- secretory canaliculus, internal microvilli
3. Mucous Neck Cells
- near gastric pit
- numerous secretory granules of mucin
4. Enteroendocrine Cells
- produce hormones: serotonin; gastrin
Cardiac glands
- around esophageal orifice
- mainly mucous cells
Pyloric glands
- short glands in pyloric region
- coiled tubules of mucous cells

SMALL INTESTINE
- duodenum, jejunum, ileum
- functions in digestion and absorption
- surface area is increased for absorption in 5 ways:

(1) length of small intestine

(2) plicae circulares: (valves of Kerckring), semicircular folds of submucosa
(3) intestinal villi: finger-like projections of epithelium
(4) microvilli: increase surface area of epithelial cells
(5) glycocalyx: Glycoproteins (enterokinases) projecting from microvilli

Intestinal Wall
(1) Mucous Membrane
(a) Epithelium simple columnar
- 2 types of cells:
(1) Columnar absorptive cells
- brush border and glycocalyx
- junctional complexes
- role in absorption and digestion
(2) Goblet cells
- apical region distended with mucigen droplets
(b) Lamina Propria
- reticular connective tissue with lymphatic elements
- central lacteal

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- smooth muscle enters villus, expresses lacteal

- crypts of Lieberkuhn (intestinal glands) invaginate from surface between villi]
- upper half - Absorptive, Goblet, Enteroendocrine cells
- lower half - Paneth cells:
secrete lysozyme, large, apical, eosinophilic granules, basal ER.
- Undifferentiated columnar stem cells:
migrate from base of crypt to villus tip - in 3 days.
Billions of cells are shed every day and replaced by upward
migration of cells from crypts
- Enteroendocrine cells:
secrete hormones such as Secretin, Cholecystokinin (CCK)
towards basal lamina
(2) Submucosa
- Brunner's glands in duodenum
- compound tubular glands, produce mucus with an alkaline, bicarbonate content
- ducts penetrate muscularis mucosae and empty into the crypts
- Peyer's patches
- groups of lymphatic nodules in ileum
- in lamina propria and submucosa
- local source of lymphocytes (no lymph filtration by these nodules)
(3) Muscularis Externa
- inner circular and outer longitudinal layers
(4) Serosa
- mesothelial cells on c.t.

LARGE INTESTINE (Colon)

- function - absorption of water and some digestion
- Intestinal wall
- no villi
- many long, straight crypts
- surface epithelium is columnar absorptive plus many goblet cells
- Paneth cells absent
- other features similar to small intestine
- muscularis externa
- outer longitudinal layer localized into 3 longitudinal bands (taeniae coli)

Appendix
- small appendage of cecum close to ileum
- small, angular lumen, often contains debris
- villi absent, epithelium as in colon
- lamina propria contains lymphoid tissue in continuous layer of nodules
- crypts not numerous, embedded in lymphoid tissue
- muscularis mucosae poorly developed, interrupted by lymphoid tissue
- submucosa contains blood vessels
- muscularis externa without taeniae coli
- typical serosa
- many lymphocytes may be present, especially in lamina propria, but not neutrophils

EXOCRINE PANCREAS
- large organ in the abdomen with its head on the duodenum and its tail touching the spleen
- covered with a thin layer of c.t. - loose septa form lobules
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- the exocrine portion produces about 1200 ml of digestive juice per day
- the endocrine portion (Islets of Langerhans) produces hormones regulating carbohydrate
metabolism

Acinar (Exocrine) Tissue

- compound acinar gland - each acinus varies from a rounded structure to a short tubule
- acini are composed of pyramidal cells with secretory capillaries between & a central lumen
- cells contain mitochondria and granular E.R. basally, secretion (zymogen) granules apically
- the base of each cell is therefore basophilic and the apex is acidophilic

Duct System
- intercalated ducts commence from central part of acinus, appear as centroacinar cells
- intercalated ducts empty into interlobular ducts which converge on main ducts
- ducts are lined with columnar epithelium,

Functions:
- production and secretion of digestive enzymes necessary for further digestion after stomach
proteases - hydrolyze proteins to amino acids
nucleases - ribonuclease, deoxyribonuclease
amylase - hydrolyzes starch and glycogen to form disaccharides
lipase - splits triglycerides into fatty acids and glycerol
- secretion is regulated according to delivery of acid contents of stomach into duodenum
- results in release of two hormones produced in duodenum

Secretin: - induces pancreatic release of a large volume of fluid that contains

bicarbonate, but few enzymes
this alkaline juice neutralizes acid in the intestine to provide appropriate pH
for optimal activity of pancreatic enzymes

Cholesystokinin: - causes pancreatic secretion of large amounts of digestive enzymes

also activates contraction of the gallbladder to add bile to the intestine
this bile release occurs in response to fats in the intestine

LIVER
- largest gland in body
- exocrine function (secretes bile into duodenum)
- endocrine function (synthesizes and releases substances into blood)
- interposed between blood vessels of intestinal tract and general circulation
- receives venous blood from intestine via portal vein
- receives arterial blood via hepatic artery
- drains into inferior vena cava via hepatic veins
- the liver therefore receives (in the portal blood) the material absorbed from the intestine

Functions:
(1) absorbed products of digestion are metabolized or transformed & returned to the blood
(2) toxic substances in the blood are degraded or detoxified
(3) bile is produced for release into the intestine where it plays a role in digestion of lipids
(4) the protein components of blood plasma is produced
(5) carbohydrates are stored as glycogen & released as glucose - maintains blood glucose

Organization:
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- hepatocytes form plates radially surrounding terminal branches of the hepatic veins (central veins)
- plates of cells are exposed to blood flowing in parallel vascular channels (hepatic sinusoids)

- Classical Liver Lobules

- hexagonal areas with a portal area at each corner and a central vein at the centre
- each portal area (portal tract/space) consists of:
(1) a branch of the portal vein
(2) a branch of the hepatic artery
(3) a bile duct
(4) sometimes a lymphatic vessel is also seen

Blood Supply and Bile Ducts:

- the portal vein, hepatic artery and bile duct enter at the porta hepatis (on inferior surface)
- the hepatic artery carries much less blood than the portal vein
- all three vessels branch repeatedly until they reach a portal area
- they are accompanied throughout by a network of lymphatics
- terminal branches of artery, vein and bile duct leave the portal area and feed sinusoids
- blood leaves the sinusoids at the central vein
- the central vein passes to a collecting vein -- to the hepatic veins -- to the vena cava
- they control blood flow through the liver which serves as an important reservoir

Sinusoids:
- irregular shape
- 2 types of lining cells:
(1) Endothelial cells
- large fenestrations in cells
- discontinuities (spaces) between cells
- provides little or no filtration barrier except to cells
(2) Kupffer cells
- situated on or between endothelial cells
- highly phagocytic, lysosomal activity (destroy bacteria, erythrocytes)
- surface folds and processes
- support for these cells is by fine reticular fibres

Perisinusoidal Space (Space of Disse):

- perivascular space between endothelial cells and hepatic parenchymal cells (hepatocytes)
- numerous microvilli on hepatocytes project into space - increase surface area for absorption
- mainly plasma in space - some reticular fibres
- forms a space comparable to a lymph capillary

Hepatic Parenchymal Cells (Hepatocytes):

- rough and smooth E.R., numerous mitochondria
- Golgi complex lie adjacent to canaliculi
- glycogen and lipid deposits

Bile Canaliculi:
- minute canals between adjacent hepatocytes
- wall is formed from hepatic cell plasmalemma
- lumen is an expansion the intercellular space
- microvilli project into the lumen
- tight junctions occur at each margin of a canaliculus
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Bile Ducts:
- from bile canaliculi
- drain into interlobular bile ducts
- main ducts fuse to form hepatic duct
- receives cystic duct, continues to the duodenum as the common bile duct
- extrahepatic ducts show many folds in the mucosa lined with tall columnar epithelium
- incomplete smooth muscle layer

GALLBLADDER
- pear-shaped, hollow organ
- fundus, body, neck, continues into cystic duct
- wall consists of
- mucosa (simple columnar ep. with microvilli) thrown into many folds
- lamina propria
- contains folds of surface epithelium
- fibromuscular layer
- serosa
- functions in concentration and storage of bile by pumping NaCl from the lumen across the
epithelium. Water passively follows.
- Cholecystokinin from intestinal mucosa induces contraction of muscle layer and release of bile

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URINARY SYSTEM

- functions: to clear the blood of the waste products of metabolism

to regulate the concentrations of many constituents of the body fluids
- consists of 2 kidneys and excretory passages to store and conduct excreted materials

KIDNEYS:
- bean shaped, lie in retroperitoneal space on the posterior aspect of the abdominal cavity
- surrounded by capsule, divided into cortex and medulla
- nephrons - functional units in which blood undergoes ultrafiltration & reabsorption

Anatomy:
Surface features
- hilus - a concavity on the medial border
- ureter - a large excretory duct which passes from the hilus to the bladder
- pelvis - the upper expanded end of the ureter which fills the hilus
- papilla - a conical protrusion of renal substance enveloped by a minor calyx
- collecting ducts - 10 to 25 ducts which perforate each papilla
Hemisected view
- medullary pyramids - 8-12 pyramids in the medulla with tips projecting into calyx
- renal columns - cortical tissue extending into the medulla between pyramids
- medullary rays - medullary material extends into the cortex as fine radial rays
- lobe - a pyramid with its associated overlying cortex
- lobule - all nephrons draining into a collecting duct
- has a medullary ray at its centre
- interlobular arteries lie between the lobules

Uriniferous Tubule:
- one uriniferous tubule includes a nephron & the collecting duct into which it empties

Nephron
- 1-2 million per kidney
- each is a long tube which begins blindly and ends by joining a collecting duct
- the tube has several segments, each with a specific structure, function & location
the segments are as follows:
1) Renal corpuscle
2) Proximal convoluted tubule
3) Loop of Henle
- thin segment
- thick segment
4) Distal convoluted tubule

1) Renal Corpuscle:
(a) Bowman's capsule:
- a thin-walled expansion at the end of the tube
- a double-walled cup composed of squamous epithelium
- it has a visceral layer which is applied to the glomerular capillaries
- it also has a parietal layer which forms the outside wall of the cup
- between the two layers is a narrow cavity called urinary (Bowman's) space
- each corpuscle has a vascular pole where the afferent and efferent arterioles enter and leave
- each corpuscle has a urinary pole where Bowman's space is continuous with the lumen of the

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proximal convoluted tubule

(b) Glomerulus:
- a globular tuft of capillary loops, each covered by a thin epithelial layer
- the endothelial cells are fenestrated (i.e. have many circular pores)
- the capillary endothelium possesses a very thick basement membrane
- the parietal layer remains a typical squamous epithelium
- cells of the visceral layer become highly modified to form podocytes
- these cells derive their name from their foot-like processes
- podocytes have processes called major or primary processes
- each major process extends from the nuclear region
- minor processes arise from the major processes as side branches
- they attach to the outer surface of the capillary basement membrane
- minor processes interdigitate with those of adjoining podocytes
- thin gaps between the minor processes are called filtration slits
- filtration slits are covered by a thin slit diaphragm composed of nephrin

Filtration Barrier:
- the filtration barrier between the blood in the glomerular capillaries and Bowman's space consists of:
1) fenestrated endothelium of the glomerular capillaries
2) basement membrane of the glomerular capillaries
3) slit diaphragm in the filtration slits
- ultrafiltration from the glomerular capillaries into Bowman's space occurs because:
1) the efferent arteriole is smaller than the afferent one, thereby increasing blood pressure
in the glomerular capillaries
2) large surface for filtration due to numerous glomerular capillaries
3) the barrier separating the blood from the capsule is very thin

Mesangium:
- the area where the capillary loops are attached to the vascular pole
- mesangial cells lie between the adjacent capillaries
- function in turn-over of glomerular basement membrane and in regulating filtration rates through
their contractility
(c) Juxtaglomerular Apparatus:
1) Macula densa
- a region in the distal tubule where the cells are narrow & closely packed
- cells function in sampling electrolyte conc. of the urinary ultrafiltrate
2) Juxtaglomerular cells
- smooth muscle cells in the tunica media of the afferent arteriole
- large, pale staining epithelioid cells with conspicuous granules of renin
- cells sensitive to stretch i.e. perceive increased blood pressure
- juxtagl. cells are intimately related to the base of the macula densa cells
- therefore also alerted to increased Na+ levels in ultrafiltrate by macula ddensa cells

2) Proximal Tubule:
- commences at the urinary pole of a renal corpuscle
- constitutes the bulk of the renal cortex
- comprised of two parts, a convoluted portion and a straight portion
- lined with a single layer of cuboidal cells with a brush border (microvilli)
- the basal plasmalemma has numerous infoldings with mitochondria between
- indicative of active transport (sodium pump)
- numerous apical canaliculi and small vesicles - suggests absorption of protein

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- cells actively transport sodium - chloride and water follow

- cells resorb glucose, amino acids, protein
3) Loop of Henle:
- consists of: - a straight part of the proximal tubule in the descending limb
- a thin segment in the descending and ascending limbs
- a straight part of the distal tubule in the ascending limb
(a) Thick Segment - Descending Limb
- the proximal convoluted tubule straightens out and forms the descending limb
- its structureand function in the descending limb are typical of proximal tubule
(b) Thin Segment - Descending and Ascending Limbs
- a transition occurs very abruptly from cuboidal to squamous
- the diameter of the thin segment is much reduced
- the cells have a thin cytoplasm so their nuclei bulge into the lumen
- the wall is freely permeable to sodium and water
(c) Thick Segment - Ascending Limb
- the cells change from squamous to cuboidal
- ascends from medulla to cortex to reach glomerulus of origin
- forms a macula densa at the vascular pole
- cells have basal infoldings with mitochondria but no brush border
- actively transport sodium from the urine to the interstitium

4) Distal Convoluted Tubule:

- continues from macula densa
- follows a short convoluted course
- continues into a collecting tubule in a medullary ray

Collecting Ducts
- not really part of the nephron
- they pass in medullary rays down into the medulla
- several distal tubules empy into each collecting duct
- in the medulla several collecting ducts join to form larger papillary ducts (of Bellini)
- papillary ducts open directly onto the apex of a papilla
- more water passes out to the medulla and the urine becomes increasingly concentrated

Blood Vessels:
- renal arteries arise from the abdominal aorta
- each renal artery divides into three main branches, each branch to 3 or 4 pyramids
- branches divide in the hilum into interlobular arteries which ascend between the pyramids
- at the corticomedullary junction several arcuate arteries are formed
- arcuates run parallel to the kidney surface
- arcuates give rise to interlobular arteries which ascend into the cortex between the rays
- side branches form intralobular arteries which branch to form afferent glomerular arterioles
- efferent arterioles arise from the glomeruli and break up into a capillary net or vasa recta
- venous drainage has a similar arrangement to the arterial supply

Ureter
- continuous with its upper expanded end, the papilla
- lined with pseudostratified transitional epithelium that changes thickness when stretched
- lumen has a stellate outline in transverse section due to longitudinal folds
- thick muscularis in two layers - performs slow peristaltic movements
- surrounded by adventitia
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Urinary Bladder
- similar to ureter, transitional epithelium thicker, binucleate cells present
- surface cells contain angular vesicles which represent a reservoir of surface membrane
- they allow for expansion of the surface during distension
- the mucosa is normally folded in the relaxed state
- three layers in the muscularis, prominent middle layer

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FEMALE REPRODUCTIVE SYSTEM

OVARY
- the ovaries are paired organs which are variable in size
- the anterior wall of each ovary (the hilum) is attached to the broad ligament of the uterus
- cortex - a cellular connective tissue stroma containing follicles
- covered by germinal epithelium which is simple cuboidal in type
- tunica albuginea - a fibrous ct stroma with abundant intercellular substance
- the cortex contains ovarian follicles at various stages of deevelopment
- the follicles contain oocytes
- medulla - the medulla contains loose connective tissue in which elastic fibers, smooth muscle
fibres, blood vessels, lymphatics and nerves, are located

Development
- ovaries develop from the genital ridges
- mesoderm at the surface forms the germinal epithelium
- oogonia develop in the yolk sac, migrate to the ovary and divide many times
- oogonia enlarge to form primary oocytes (fourth month)
- they become enclosed by epithelial cells to form primordial follicles (7th month)
- they enter meiosis and rest in prophase until after puberty
- 2 million oocytes remain at birth
- 400,000 oocytes remain at puberty, 400-500 will mature fully

Follicular Maturation
- FSH stimulates growth - the oocyte enlarges
- follicular cells form several layers which make up the zona granulosa
- a zona pellucida forms around the oocyte - provides nutrients via the granulosa cells
- an antrum forms in the granulosa
- the follicle becomes ovoid with an eccentric ovum
- a theca folliculi forms:
theca interna - a vascular layer with secretory cells
theca externa - a connective tissue layer
- 10-14 days are required for maturation of the follicle
- estrogen is produced by cells in the theca, granulosa and stroma

Ovarian Steroidogenesis
- androgens (e.g. testosterone) are produced by cells in the theca interna & stroma (LH)
- the androgens pass across the follicular basement membrane to the granulosa cells
- the granulosa cells use the androgens as a substrate for estrogen production
- estrogen production in granulosa cells predominates in the proliferative phase (FSH)
- progesterone production in lutein cells predominates in the secretory phase (LH),

Graafian Follicle
- granulosa cells around the antrum form the membrana granulosa
- follicular cells form the cumulus oophorus and the corona radiata
- an LH surge triggers the resumption of meiosis
- the first meiotic division occurs before ovulation
- the secondary oocyte and first polar body are formed
- the second meiotic division is entered and arrested at metaphase until fertilization
- the mature ovum and second polar body are formed

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Ovulation
- the Graafian follicle bulges on the surface of the ovary
- the surface layers of the ovary become weakened and rupture
- the ovum is released
- the ovum is drawn into the oviduct by its fimbriae
- fertilization occurs in the ampulla of the oviduct and meiosis II is completed

Corpus Luteum
- after ovulation, the follicle collapses
- the membrana granulosa is thrown into folds
- the granulosa cells differentiate into granulosa lutein cells
- the theca interna cells form theca lutein cells
- the follicle is now called the corpus luteum
- bleeding into the cavity forms a clot which contains much fibrin
- the lutein cells are invaded by capillaries and connective tissue
- the corpus luteum secretes progesterone and some estrogen
- if the ovum is not fertiized a corpus luteum of menstruation forms
- it lasts 14 days then degenerates to form a corpus albicans
- if the ovum is fertilized a corpus luteum of pregnancy forms
- it lasts for 6 months then gradually declines to form a corpus albicans after birth

Atretic Follicles
- several follicles develop each cycle
- one matures and the remainder undergo follicular atresia
- the ovum and follicular cells are resorbed & replaced by connective tissue to form a scar

FALLOPIAN TUBES (oviducts, uterine tubes)

- muscular tubes, 12 cm. long

- 4 segments: infundibulum, ampulla, isthmus, intramural part
- 3 layers: - mucosa: columnar epithelium with some cells ciliated, others secretory
- - muscle: inner circular, outer longitudinal layers
- - serosa: a fold of peritoneum

UTERUS

- a single, pear-shaped organ with a muscular wall

- 2 parts - the body and the cervix

Body
Myometrium:
- 3 layers of smooth muscle, thick circular middle layer
- the muscle functions in expulsion of the fetus at birth
- contractility of the muscle is increased by oxytocin
Endometrium:
- simple columnar epithelium, two cell types - ciliated and secretory
- tubular glands with ciliated and secretory cells produce a glycogen rich secretion
- the stroma resembles mesenchyme with reticular fibers, fibroblasts, ground substance.
- two types of arteries called regular and coiled
- functional and basal layers of the endometrium are identified

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- changes during the menstrual cycle:

Proliferative phase (follicular, estrogenic)

- mitosis occurs in the epithelium and stroma
- the endometrium increases in thickness
- the glands are straight and narrow
- the coiled arteries do not extend beyond the lower half of functional layer
- the ground substance in the stroma is abundant
- occurs from day 4 to approximately day 14 of the cycle

Secretory phase (luteal, progestational)

- this phase lasts 12 - 14 days
- glycogen accumulates in the epithelial cells
- secretion accumulates in the glands
- the glands are tortuous and sacculated
- coiled arteries extend into the superficial portion of the endometrium
- edema occurs in the stroma
- ischemia
- occurs for 2 - 3 days at the end of the secretory phase
- the coiled arteries become constricted for prolonged periods
- blanching of the superficial zone of the endometrium occurs
- the constricted arteries dilate
- blood & tissue fluid in the stroma lead to lifting of the endometrium

Menstrual phase
- days 1 - 4: the functional layer and blood are lost
- on day 4 the epithelial cells glide out from the glands to restore the surface epithelium

Cervix

Endocervix
- the lowest segment of the uterus, adjoining the vagina
- the wall is fibromuscular, i.e mainly dense c.t. with about 15% smooth muscle
- the lining is a simple columnar ep. with mucus secreting and some ciliated cells
- many tubular, highly branched glands
- their lining is similar to the surface i.e. mucus producing cells
- cyclic changes occur in secretory activity of the glands - 10X increase at ovulation
- the secretory activity changes with the circulating levels of ovarian hormones

Exocervix
- covered with stratified squamous epithelium - similar to vagina
- susceptible to cervical erosions - patches of columnar epithelium, inflammation

VAGINA
- a distensible muscular tube lined with non-keratinized stratified squamous epithel.
- cells in the more superficial layers accumulate large amounts of glycogen
- the glycogen may serve as a nutrient for the sperm while in the vagina
- there are no glands in the vagina - it is lubricated by mucoid cervical secretions

MAMMARY GLAND
- specialized accessory glands of the skin

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Histology M. Sandig

- resemble sweat glands in their structure and mode of development

- consist of 15-20 compound (branched) tubular glands, one per lobe
- each gland has its own duct opening at the nipple - called a lactiferous duct
- marked changes in the amount and activity of the glandular tissue occur during
pregnancy and lactation

Resting Mammary Gland

- at puberty the glands enlarge rapidly in the female due to estrogen and prolactin
- enlargement due to development of c.t., particularly adipose tissue
- there is also some development of duct tissue
- true secretory units do not develop until pregnancy
- each lobe is subdivided into lobules which contain alveolar ducts
- alveolar ducts are covered by small, saccular evaginations called alveoli
- alveolar ducts and alveoli represent the secretory portion of the gland
- ducts are the principal epithelial tissue seen in the resting gland
- ducts are lined with cuboidal or low columnar secretory cells
- myoepithelial cells form a basket-like network around the secretory portion
- each lobe is surrounded by interlobar C.T. containing many fat cells
- interlobular c.t. is dense with some fat, intralobular C.T. is loose with little fat
- intralobular ducts drain into interlobular ducts which join to form a single excretory duct from each l
obe - the lactiferous duct
- lactiferous ducts from the different lobes converge to enter the base of the nipple
- they become expanded and the widened segments are termed lactiferous sinuses
- these sinuses are believed to act a little reservoirs for milk in the lactating breast

Nipple
- the nipple and the areola (skin surrounding the nipple) are covered with thin skin
- the epidermis is invaded by unusually long dermal papillae
- brings capillaries close to the surface which imparts a pink colour to this area
- smooth muscle: - disposed longitudinally along the lactiferous ducts
- disposed circumferentially within the nipple and around its base
- the muscle enables the nipple to become erect

Changes During Pregnancy and Lactation

During the First Half of Pregnancy

- rapid growth & branching from terminal portion of duct system - new alveoli form
- interlobular adipose tissue regresses with the growth of the glandular tissue
- remaining c.t. contains extensive capillary networks

During the Second half of Pregnancy

- growth slows down, alveoli enlarge due to secretion of colostrum
- colostrum is a cloudy, watery fluid that is the first secretion after birth occurs
- it is believed to contain antibodies that provide the newborn with some immunity
- termination of pregnancy and expulsion of the placenta leads secondarily to a surge in production
of lactogenic hormone (prolactin) - induces milk production
- the cells of the alveoli have two distinct secretory products formed and released by different
mechanisms:
Protein - elaborated in RER, passes to Golgi for packaging and concentrating
- transported to surface in vesicles and released
Lipids - arise as lipid droplets free in the cytoplasm

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Histology M. Sandig

- increase in size, pass to the apical surface

- project into the lumen - covered with plasmalemma and released

- milk accumulates in the lumen of alveoli and ducts

- when a baby begins to nurse, the hypothalamus is stimulated to produce oxytocin
- causes the myoepithelial cells to contract and milk is squeezed up into the nipple
- called the milk ejection reflex

Regression of Mammary Gland

- after lactation ceases, the gland regresses and returns to a resting state
- the secretion is resorbed, alveoli decrease in size and some cells degenerate
- the connective tissue and fat become abundant again

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Histology M. Sandig

MALE REPRODUCTIVE SYSTEM

TESTIS

Capsule -
- layers:
(1) tunica vaginalis - mesothelial cells
(2) tunica albuginea - dense c.t. and smooth muscle - help to move sperm
(3) tunica vasculosa - network of blood vessels
- tunica albuginea forms mediastinum testis at the posterior pole - region of entrance and exit
- radial septa from the mediastinum form 250 lobules which contain the seminiferous tubules

Seminiferous tubules
- each testis contains about 800 highly convoluted seminiferous tubules, 1-4 in each lobule
- each tubule is 30 to 100 cm long and about 200 μm in diameter
- they are lined by germinal (seminiferous) epithelium
- they are surrounded by c.t. and contractile myoid cells (boundary or peritubular tissue)

Seminiferous epithelium
- a modified statified cuboidal epithelium
- cells at the periphery of the tubule rest on a basement membrane
- there are 2 categories of cells:
Sertoli cells
- tall cells with irregular, indistinct outlines
- they have a central, pale nucleus and a prominent nucleolus which aids in identification
- they have abundant smooth E.R. and lysosomes
Functions:
(1) support and nutrition of the germ cells (developing sperm)
(2) blood-testis barrier
(3) release of spermatozoa
(4) removal of residual sperm cytoplasm
(5) produce androgen binding protein (ABP) which maintains a sufficient concentration of
testosterone in the tubules for spermatogenesis to occur

Spermatogenic cells
- located within the seminiferous epithelium amongst sertoli cells(4-8 cells deep)
-
Spermatogenesis:
the entire sequence of events by which spermatogonia are transformed into spermatozoa
1) Spermatogonia -
- located adjacent to the basement membrane - 2 types:
Type A: - fine chromatin, 1-2 nucleoli against envelope
- division yields an A and a B type
Type B: - coarser chromatin, 1 central nucleolus
- successive division yields primary spermatocytes
2) Primary Spermatocytes -
- large, spherical cells with coarse chromatin
- they are located in the middle zone of the epithelium
- primary spermatocytes undergo the first meiotic division
3) Secondary Spermatocytes -
- they are smaller cells and are nearer the lumen
- their lifespan is of short duration so few are seen
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Histology M. Sandig

- they undergo the second meiotic division

4) Spermatids -
- they are even smaller and are located next to the lumen
- they are spherical cells with round nuclei
- cell division is incomplete
- therefore they remain attached by intercellular bridges to form a syncytium

Spermiogenesis:
the sequence of events by which spermatids are transformed into spermatozoa
1. the nucleus becomes elongated, then fusiform
2. an acrosome forms a head-cap over the nucleus
3. centrioles pass to opposite poles of the cell and a flagellum forms
4. mitochondria regroup around the base of the flagellum
5. surplus cytoplasm is cast off as a residual body

Spermatozoa:
1) Head
- the nucleus is streamlined to enhance motion and condensed to reduce size
- the acrosome contains hyaluronidase which is released at the ovum
- it helps the sperm to pass through the corona radiata prior to fertilization
2) Neck
- contains two centrioles
3) Mid-Piece
- contains an axoneme consisting of 9+2 microtubules in the core
- nine outer dense fibers are present
4) Principal Piece
- mitochondria are absent but the microtubules continue
- outer dense fibers continue and a fibrous sheath appears
5) End Piece
- the microtubules continue
- the outer dense fibers and the fibrous sheath are absent

Leydig Cells (interstitial) cells -

- they are large, acidophilic cells that occur in clusters in spaces between the tubules
- extensive smooth E.R. and a well developed Golgi
- they produce testosterone

DUCT SYSTEM

Tubuli Recti -
- they are formed by merging of the seminiferous tubules
- they are located at the apex of each lobule
- Sertoli cells but no spermatogenic cells are present within them
Rete Testis -
- it serves to combine the spermatozoa from different tubules
- it comprises a network (rete) of anastomosing channels in the mediastinum
- it is lined with simple cuboidal or squamous epithelium
- few sperms are seen within it because they traverse the rete quickly
Ductuli Efferentes -
- comprise several spiral ducts which emerge on the surface of the testis
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Histology M. Sandig

- they are lined with a simple columnar epithelium with ciliated & some glandular cells
- smooth muscle and c.t. underlies the epithelium
Ductus Epididymis -
- a single, long, coiled tube
- lined with pseudostratified columnar ep. with stereocilia and secretory characteristics
- a circular smooth muscle layer aids in movement of sperm
- it serves in accumulation, storage and maturation of sperm
- 90% of the fluid leaving the testis is absorbed in the ductus epididymis and ductuli
efferentes
- it creates an environment favourable for continued maturation of the sperm
Vas Deferens (Ductus Deferens)
- the vas deferens passes from the epididymis to the seminal vesicle
- it has a thick, muscular wall and a narrow stellate lumen
- it is lined with pseudostratified epithelium with stereocilia, like the ductus epididymis
- it has a thick muscular coat (three layers) that plays a role in moving the sperm

GLANDS
Seminal Vesicles -
- the seminal vesicle is a coiled tube with saccular outpouchings
- lined with a pseudostratified epithelium whose cells contain secretion granules
- cells produce a yellowish, viscous liquid containimg fructose to provide energy for the sperm
Prostate Gland -
- the prostate is comprised of 30-50 small compound glands that empty into the urethra
- it is characterized by large cavities and branching tubules
- the tubules are lined with cuboidal or pseudostratified ep containing secretion granules
- stroma is abundant and contains muscle fibers
- secretion serves as a vehicle for transport of the sperm
- secretion is granular in appearance and becomes condensed to form lamellated bodies
called prostatic concretions which may become calcified

PENIS
- the penis is a common outlet for urine and seminal fluid

Erectile Tissue:
- the erectile tissue in the penis is enclosed by a fibrous layer, the tunica albuginea.
- it forms 2 corpora cavernosa and a single corpus spongiosum which surrounds the urethra
- erectile tissue contains many vascular spaces supported by collagenous trabeculae that
connect with the tunica albuginea.
- the erectile tissue contains vascular spaces (blood sinuses) called cavernous spaces
- helicine arteries arise from the vessels that run deep in the erectile tissue, and empty directly into
the vascular spaces.

Blood Supply:
internal pudendal arteries
- deep arteries and dorsal arteries
⎥
- nutritive and helicene arteries ⎦ AVA's ⎦ deep dorsal vein
⎥ ⎥
trabeculae cavernous spaces
Flaccid:
- nutritive arteries to trabeculae open
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Histology M. Sandig

- AVA's open
- helicene arteries close
Erect:
- parasympathetic vasodilator impulses occur
- vessels and cavernous smooth muscle relax
- sympathetic vasoconstrictor impulses inhibited
- AVA's close
- blood flow in deep artery increases
- helicene arteries open
- cavernous spaces fill
- nutritive artery becomes compressed

Two chemicals control erection: nitric oxide and phosphodiesterase.

Sexual stimulation causes nerve impulses to be transported from the brain down the spinal cord to
nerves in the penis which produce nitric oxide. Nitric oxide molecules spread rapidly across gap
junctions of smooth muscle cells surrounding the cavernous spaces. Within the smooth muscle cells,
nitric oxide molecules activate guanylate cyclase to produce cyclic GMP from GTP. cGMP relaxes
the smooth muscle cell wall surrounding the sinusoids by inducing the sequestration of Ca++. The
lowered concentrations of Ca++ lead to relaxation of the smooth muscle cells, which leads to the
accumulation of arterial blood in the sinusoids. Sinusoids engorged with blood compress the small
veins that drain blood from the penis and the penis becomes erect. The enzyme phosphodiesterase
(PDE) is produced to destroy cGMP and terminate erection. By blocking PDE activity, cGMP levels
remain elevated and the penis remains erect. Sildenafil (Viagra) is a potent blocker of PDE activity.

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Histology M. Sandig

ENDOCRINE GLANDS

- glands can be divided into exocrines and endocrines

- exocrines are attached by a duct to the surface onto which their product is emptied
- endocrines are ductless glands and secrete their product into the blood stream
- organs that have a purely endocrine functions are:
adrenals, thyroid, parathyroid, pituitary and pineal
- organs that have endocrine and other functions are:
testes, ovaries and pancreas
- endocrine glands secrete hormones

- Definition - "hormone" - a chemical product that is transported by the blood or body fluids and
has a specific regulatory effect on cells remote from its origin.
THYROID GLAND
- consists of a lateral lobe on each side of the trachea
- lobes connected by a narrow isthmus
- gland surrounded by a fibro-elastic capsule
- capsule continues inward as septa which divide gland into lobules

Follicle
- the structural unit of the thyroid
- a closed, single-layered epithelial sac
- follicular cells
- height varies with activity
- contain colloid droplets, lysosomes
- parafollicular cells
- found in the c.t. between the follicles and beween the follicular cells and the b.m.
- produce calcitonin - a hormone which lowers blood calcium
- colloid
- a gel-like material that fills the follicular cavity
- represents storage of reserve secretion

Synthesis, Storage, Uptake and Secretion in the Thyroid

(1) Synthesis of thyroglobulin:
- uptake of amino acids from the blood, synthesis of protein in the rER, addition of
carbohydrate in the Golgi complex, release from formed vesicles at the apical surface of the
cell into the lumen of the follicle
(2) Uptake of circulating iodide:
- accomplished by a mechanism of active transport utilizing an iodide pump located basally
within the cell
(3) Oxidation of Iodide:
- iodide is oxidized by thyroid peroxidase to an intermediate which combines in the colloid with
the tyrosine residues of thyroglobulin to form triiodothyronine (T3) and tetraiodothyronine (T4
- thyroxine).
(4) Liberation of T3 and T4:
- when stimulated by TSH, thyroid follicular cells take up colloid by endocytosis.
- peptide bonds between the iodinated residues and the thyroglobulin molecule are broken by
proteases in lysosomes and the T3 and T4 are liberated into the cytoplasm, cross the basal
plasmalemma and are discharged into the capillaries

Function:

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Histology M. Sandig

- thyroid hormone regulates metabolic rate

- also involved in development, differentiation and growth
- hypothyroidism
- if present at birth: causes cretinism - dwarfing and mental retardation
- if present in the adult: causes myxedema
- hyperthyroidism
- metabolic processes increase, body weight diminishes
- thyrotropic hormone
- produced by pituitary gland
- stimulates the release of thyroid hormone

PARATHYROID GLAND -
- two glands on each side of the neck behind each thyroid lobe
- each gland is small and ovoid - about the size of an apple seed
- a fibroelastic capsule gives rise to septa which divide the gland into lobules
- cells appear as a dense mass arranged in anastomosing cords
- many capillaries are located between the cords
- cell types -
(1) Chief (principal) cell
- most numerous type, pale staining
- produces parathyroid hormone
(2) Oxyphil cell
- large, acidophilic cells - characteristic of humans
- function unknown

- parathyroid hormone regulates calcium concentration in bone and body fluids

- calcitonin from the thyroid counterbalances parathyroid hormone
- parathyroid hormone increases blood calcium levels by:
(1) increasing resorptive activity of osteoclasts in bone
(2) increasing calcium resorption in the kidney
(3) increasing calcium absorption in the gut

ADRENAL GLANDS
- triangular bodies at the cranial pole of each kidney
- consist of a cortex and medulla which are different in their origin, structure and function
- cortex develops from peritoneal mesothelium
- medulla is derived from primitive nervous tissue
- capsule. trabeculae and hilus

Adrenal Cortex
- three layers
(1) Zona Glomerulosa
- cells arranged in ovoid groups
(2) Zona Fasciculata
- cells in parallel cords separated by sinusoids
- spongiocytes
(3) Zona Reticularis
- cells in anastomosing cords
- signs of degeneration
- more than forty steroid hormones produced in adrenal cortex

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Histology M. Sandig

- divided into mineralocorticoids, glucocorticoids and sex hormones

Adrenal Medulla
- small in man
- vascular supply: arterial plexus in capsule supplies both:
(1) sinusoids in cortex, and
(2) capillary plexus in medulla
- blood leaves via collecting veins at cortico-medullary junction
- cells are polarized and arranged around the vessels
- called "chromaffin cells" due to staining properties
- contain catecholamines: epinephrine or norepinephrine
- augments the action of sympathetic nerve endings

ENDOCRINE PANCREAS
- Islets of Langerhans -
- large, pale staining areas scattered throughout the pancreas
- supported by reticular tissue
- irregular, anastomosing cords and clumps of cells separated by capillaries
- special methods reveal at least 3 cell types:
alpha (A) cells
beta (B) cells
delta (D) cells
- beta cells produce insulin which enables the body to utilize glucose
- alpha cells produce glucagon which offsets the influence of insulin
- delta cells produce somatostatin which inhibits glucagon release and decreases pancreatic
exocrine secretion

PITUITARY GLAND (Hypophysis)

- endocrine glands are ductless glands of internal secretion

- endocrines elaborate secretory products called "hormones"
- hormones are chemical products that are transported by the blood or body fluids and have a
specific regulatory effect on cells remote from their origin
- the pituitary gland is structurally and functionally the most complex of the endocrines
- it is attached to the base of the brain
- it occupies a key position in the interplay of the nervous and endocrine systems
- it produces several hormones, including tropic hormones - i.e. hormones required specifically
to activate other endocrine glands
- it has 2 major subdivisions
- neurohypophysis - derived from the brain and resembles nervous tissue
- adenohypophysis - derived from oral ectoderm and resembles endocrine tissue
Development
- the infundibulum evaginates from the floor of the brain
- Rathke's pouch forms from the oral ectoderm
- the Infundibulum forms the infundibular stem, then the infundibular process (pars nervosa)
- Rathke's pouch forms the pars distalis, pars intermedia and pars tuberalis

Divisions
- neurohypophysis: median eminence, infundibular stem, pars nervosa (posterior pituitary)
- adenohypophysis: pars tuberalis, pars intermedia, pars distalis (anterior pituitary)

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Histology M. Sandig

Blood Supply
- derives from two groups of blood vessels that come from the internal carotid artery
- the inferior hypophyseal arteries mainly supply the neurohypophysis
- the superior hypophyseal arteries feed a primary capillary plexus in the pars tuberalis
- capillary loops pass from the plexus into the median eminence and infundibular stem
- a primary capillary plexus drains into veins that pass to sinusoidal capillaries in the pars distalis
which form a secondary capillary plexus
- this is the hypophyseal portal system
- nerve fibers from the hypothalamus end around the capillaries in the infundibular stem and empty
releasing and inhibitory factors into them
- the activity of the pars distalis is thus partly under nervous control
- releasing factors stimulate cells in the pars distalis to release hormones
- inhibitory factors inhibit hormone release

Adenohypophysis -
(a) Pars Distalis
- the largest division of the pituitary
- the cells are of two types:
(1) chromophobes - 50% - do not stain intensely
- include undifferentiated, degranulated, and supporting (follicular) cells
(2) chromophils - 50%: (acidophils - 40%, basophils - 10%)
- surgical removal of the pars distalis causes:
- cessation of growth
- atrophy of other endocrine organs
- metabolic disturbances
- these disturbances are due to elimination of hormones produced by the pars distalis:
(1) LTH (luteotropic hormone, prolactin, lactogenic hormone)
- promotes the secretion of progesterone by the corpus luteum of the ovary
- promotes secretion of milk
- promotes development of mammary glands during pregnancy
(2) STH (somatotropic hormone, growth hormone)
- enhances body growth
- acts on the liver and kidney to produce somatomedins
- somatomedins increase the rate of mitosis in the epiphyseal plate
- imbalance causes dwarfism, gigantism, acromegaly
(3) TSH (thyroid stimulating hormone, thyrotropic hormone)
- stimulates the thyroid
(4) FSH (follicle stimulating hormone)
- stimulates the growth of ovarian follicles and estrogen secretion
- activates spermatogenic eipthelium of the testes and stimulates Sertoli cells
(5) LH (luteinizing hormone)
- promotes ovulation, corpus luteum formation, progesterone secretion
- stimulates Leydig cells of the testes to secrete testosterone
(6) ACTH (adrenocorticotropic hormone, corticotropin)
- stimulates secretion in the adrenal cortex

- acidophils: - somatotrophs (STH), mammotrophs (LTH)

- basophils: - gonadotrophs (FSH, LH), thyrotrophs (TSH), corticotrophs (ACTH)

- hormone production and release in the pars distalis is under the influence of
- releasing factors (hormones) from the hypothalamus (TSH-RF, etc.)

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Histology M. Sandig

- inhibitory factors (hormones) from the hypothalamus (Somatostatin, PIH)

- the direct effect of hormones from target organs on median eminence and distalis

(b) Pars Tuberalis -

- forms a collar around the infundibular stem
- cells occur as groups, cords, vesicles
- produce gonadotropins (FSH and LH)

(c) Pars Intermedia -

- poorly developed: cells, vesicles and colloid
- function poorly understood
- produces melanocyte stimulating hormone (MSH) in some species
- MSH is similar to ACTH

Neurohypophysis -
- pars nervosa and infundibular stem (stalk)
- 2 components:

1) pituicytes - may be only supportive

2) axons of cells from hypothalamus
- hypothalamo-hypophyseal tract - 100,000 unmyelinated axons
- arise from supraoptic and paraventricular nuclei
- contain Herring bodies:
- accumulations of stored neurosecretory material (hormones) in axons:
- Oxytocin (mainly paraventricular)
- induces contraction of smooth muscle in the uterus during childbirth
- induces contraction of myoepithelial cells in the mammaries during nursing (milk
ejection reflex)
- Antidiuretic hormone (ADH) also called vasopressin (mainly supraoptic)
- induces water re-absorption in the kidney
- promotes contraction of smooth muscle of blood vessels - raises blood pressure
- Neurophysin (binding protein)

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