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Chapter 21 The Male Reproductive System


- Consists of the:
Testes (produces sperm and contains endocrine cells secreting
hormones like testosteron1)

Genital ducts x accessory glands


- both produce secretions required for sperm activity and contract to
propel spermatozoa and the secretions from the penile urethra
- These secretions provide nutrients for spermatozoa while they are
confined to the male reproductive tract.
- Spermatozoa and the secretions of the accessory glands make up
the semen.

Penis
- introduces the semen to the female reproductive tract.
Tunica Vaginalis
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Testosterone - A serous sac carried by each testis during migration from the
- Drives male reproductive physiology abdominal cavity.
Function: Important for spermatogenesis, sexual differentiation Source: Derived from the peritoneum.
during embryonic and fetal development, and control of - It consists of an outer parietal layer lining the scrotum and an inner
gonadotropin secretion in the pituitary. visceral layer covering the tunic albuginea on the anterior and lateral
sides of the testis.
Dihydrotestosterone
-metabolite of testosterone NOTE: Sperm formation cannot occur at the core body temperature
- also begins to act on may tissues during puberty (i.e. male accessory of 37oC. Thus, a temperature of about 34oC is maintained in the
glands and hair follicles). scrotal sac.

Pampiniform Venous Plexus


- Surrounds each testicular artery
- contains cooler blood from the testis, which draws heat from the
arterial blood by a countercurrent heat-exchange system.

- Evaporation of sweat from the scrotum also contributes to heat


loss.
- Relaxation or contraction of the thin dartos muscle of the scrotum
and the cremaster muscles of the spermatic cords move the testes
away from or closer to the body respectively, allowing further control
on testicular temperature.
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INTERSTITIAL TISSUE
TESTES
- The interstitial tissue of the testis between the seminiferous tubules
Tunica Albuginea
consists of sparse CT containing fibroblasts, lymphatics, and blood
- Dense CT capsule that surrounds each testis (or testicle)
vessels including fenestrated capillaries.
- thickens on the posterior side to form the mediastinum testis.
Interstitial Cells (or Leydig cells)
Testicular Lobules
- During puberty, these develop as large round or polygonal cells
- about 250 pyramidal compartments
Location of nuclei: Center
- From the tunica albuginea, septa penetrate the organ and divide it
Cytoplasm: Eosinophilic and rich in small lipid droplets.
to form these.
Function: Produces testosterone.
- Each lobule contains sparse CT with endocrine interstitial cells (or
Leydig cells)2 and one to four highly convoluted seminiferous Testosterone secretion by interstitial cells is triggered by the pituitary
tubules3 gonadotropin, luteinizing hormone (LH), which is also called
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interstitial cell-stimulating hormone (ICSH).
Leydig Cells – secrete testosterone4
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Seminiferous tubules – where sperm production occurs. Testosterone synthesis begins at puberty, when the hypothalamus
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Testosterone – a steroid hormone that promotes development of begins producing gonadotropin-releasing hormone.
the secondary male sex characteristics.
- It is synthesized by enzymes present in the SER and mitochondria. In the late embryonic testes, gonadotropin from the placenta
stimulates interstitial cells to synthesize the testosterone needed for
The testes develop retroperitoneally in the dorsal wall of the development of the ducts and glands of the male reproductive
embryonic abdominal cavity and are moved during fetal system. These fetal interstitial cells are very active during the 3rd and
development to become suspended in two halves of the scrotal sac, 4th months of pregnancy, then regress and become quiescent cells
or scrotum, at the ends of the spermatic cords. resembling fibroblasts until puberty.
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SEMINIFEROUS TUBULES Primary spermatocytes
- Where sperm are produced at a rate of about 2x108 per day in the - spherical cells with euchromatic nuclei
young adult. - these replicate their DNA, so each chromosome consists of
- Each testis has about 250-1000 tubules in its lobules, and each duplicate chromatids, and enter meiosis, during which homologous
tubule measures 150-250 um in diameter and 30-70 cm in length. chromosomes come together in synapsis, DNA recombination occurs,
and two rapid cell divisions produce haploid cells.
Straight tubule - It has 46 (44 + XY) chromosomes, the diploid number, and a DNA
- A very short, narrower segment that links each tubule into a loop to content of 4N.
the rete testis5 - The letter N denotes either the haploid number of chromosomes
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(23 in humans) or the amount of DNA in this set.
Rete Testis – a labyrinth of epithelium-lined channels embedded in - Soon after their formation, these cells enter the first meiotic
the mediastinum testis. prophase that lasts about 3 weeks.
- These are the largest cells of the spermatogenic lineage and are
Efferent Ductules – About 10-20 of these connect the rete testis to
characterized by the presence of partially condensed chromosomes
the head of the epididymis.
in various stages of synapsis and recombination.
Germinal (or Spermatogenic) Epithelium - Most spermatocytes seen in sections of testis are in this phase
- A complex, specialized stratified epithelium that lines each (prophase) of meiosis.
seminiferous tubule.
- The basement membrane of this epithelium is covered by fibrous
CT, with an innermost layer containing flattened, smooth muscle-like
myeloid cells6
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Myeloid Cells – these allow weak contractions of the tubule.

Two Types of Cells of the Germinal Epithelium


Sertoli Cells Spermatogenic lineage
Large, non-dividing cells which Dividing cells. Cells from this is
physically and metabolically comprised of four or more
support developing sperm cell concentric layers of cells in the
precursor. germinal epithelium.
These develop from progenitor
cells to fully formed sperm cells
over 10 weeks.
Spermatogenesis
- the first part of sperm production which involves mainly mitosis and
meiosis and is followed by spermiogenesis7
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Spermiogenesis
- the final differentiation process occurring in the haploid male germ
cell.
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SPERMATOGENESIS
- Begins at puberty with proliferation of stem and progenitor cells
called spermatogonia8
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Spermatogonia
- these are small round cells
- these occupy a basal niche in the epithelial wall of the tubules, next
to the basement membrane and closely associated with Sertoli cell
surfaces. Secondary spermatocytes
- Spermatogonia with dark, ovoid nuclei act as stem cells, dividing - Smaller cells produced by homologous chromosomes separate in
infrequently and giving rise both to new stem cells and to cells with the first meiotic division.
more pale-staining, ovoid nuclei that divide more rapidly as transit - Has only 23 chromosomes (22 + X or 22 + Y), but each still consists
amplifying (progenitor) cells. These… of two chromatids, so the amount of DNA is 2N.
Type A spermatogonia - These are rare in testis sections because they are very short-lived
- each undergo several unique clonal divisions that undergo several cells that remain in interphase only briefly and quickly undergo the
unique clonal divisions that leave most of the ells interconnected as a second meiotic division.
syncytium. These become… - Division of each separates the chromatids of each chromosome and
Type B spermatogonia produces two haploid cells called…
- have more spherical and pale nuclei. Spermatids
- Each type B then undergoes a final mitotic division to produce two - each of these have 23 chromosomes.
cells that grown in size and become…
Because no S phase occurs between the first and second meiotic
divisions, the amount of DNA per cell is reduced by half when the
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chromatids separate, and the cells formed are haploid (1N). With Spermiogenesis includes formation of the acrosome, condensation
fertilization, a haploid ovum and sperm produced by meiosis unit and and elongation of the nucleus, development of the flagellum, and the
the normal diploid chromosome number is restored. loss of much of the cytoplasm.
________________________________________________________ Result: Mature spermatozoon which is released from the Sertoli cell
THE CLONAL NATURE OF MALE GERM CELLS surface into the tubule’s lumen.

Four Phases of Spermiogenesis


[1] Golgi phase [2] Cap Phase
Cytoplasm contains a The acrosomal cap spreads over
prominent Golgi apparatus near about half of the condensing
the nucleus, mitochondria, nucleus. The acrosome is a
paired centrioles, and free specialized type of lysosome
polyribosomes. containing hydrolytic enzymes,
Small proacrosomal vesicles mainly hyaluronidase and a
from the Golgi coalesce as a trypsin-like protease called
single membrane-limited ACROSIN. These enzymes are
acrosomal cap close to one end released when a spermatozoon
of the nucleus. The centrioles encounters an oocyte and the
migrate to a position farthest acrosomal membrane fuses
from the acrosomal cap and with the sperm’s plasma
one acts as a basal body, membrane. They dissociate
organizing the axoneme of the cells of the corona radiata and
flagellum (structurally and digest the zona pellucida (Both
functionally similar to cilium) surrounds the egg). This
The stem cells produced by mitotic divisions of spermatogonia process, the ACROSOMAL
remain as separate cells. However, all subsequent divisions of the REACTION, is one of the first
daughter cells, which become transit amplifying progenitor cells, steps of fertilization.
have incomplete cytokinesis after telophase and the cells remain [3] Acrosome phase [4] Maturation phase
attached to one another by intercellular bridges of cytoplasm. The head of the developing Unneeded cytoplasm is shed as
sperm containing the acrosome a residual body from each
These allow free cytoplasmic communication among the cells during
and the condensing nucleus spermatozoon and remaining
their remaining mitotic and meiotic divisions. remains embedded in the intercellular bridges are lost.
Sertoli cell while the growing Fully formed, but not yet
Although some cells degenerate without completing
axoneme extends into the functional or mobile, sperm are
spermatogenesis and some cells may separate, clones of
lumen of the tubule. Nuclei released into the lumen of the
approximately a hundred cells may remain linked through meiosis.
become more elongated and seminiferous tubule.
The complete significance of this spermatogenic syncytium is not very highly condensed, with the
clear, but the cytoplasmic bridges allow the haploid cells to be histones of nucleosomes
supplied with products of the complete diploid genome, including replaced by small basic
proteins and RNA encoded by genes on the X or Y chromosome peptides called protamines.
missing in their haploid nuclei. The germ cells finally become Flagellum growth continues
separated from one another during differentiation. distally in the tail and
mitochondria aggregate around
The cellular events and changes between the final mitoses or it proximally to form a
spermatogonia and the formation of spermatids take about 2 thickened middle piece where
months. The spermatogenic cells are not randomly distributed in the the ATP for flagellar
spermatogenic epithelium. Cells at different stages of development movements is generated.
are typically grouped together along the tubule, with the intercellular ________________________________________________________
bridges helping to coordinate their divisions and differentiations. SERTOLI CELLS
________________________________________________________ - Enrico Sertoli (first demonstrated their physiologic significance)
SPERMIOGENESIS Shape: tall “columnar” epithelial cells
- The final phase of sperm production, is the temperature-sensitive Function:
process by which spermatids differentiate into… - nourish the spermatogenic cells and divide the seminiferous tubules
Spermatozoa into two (basal and adluminal) compartments.
- highly specialized to deliver male DNA to the ovum. - Each supports 30-50 developing germ cells.
Nuclei: Ovoid or triangular; Euchromatic with a prominent nucleolus
- No cell division occurs during this process, and as with
spermatogenesis the cells involved remain associated with Sertoli All cells of the spermatogenic lineage are closely associated with the
cells. extended surfaces of the Sertoli cells and depend on them for
metabolic and physical support.
The haploid spermatids are small cells near the lumen of the
seminiferous tubules.
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Contains abundant SER, some RER, well-developed Golgi complexes, Müllerian-inhibiting Substance
numerous mitochondria, and lysosomes. - a glycoprotein that causes regression of the embryonic Müllerian
(paramesonephric) ducts
Sertoli cells adhere to the basal lamina and their apical ends extend ________________________________________________________
to the lumen. INTRATESTICULAR DUCTS
- These are the straight tubules (or tubuli recti), the rete testis, and
Tight Occluding Junctions the efferent ductules
- Important in Sertoli cell function Function: carry spermatozoa and liquid from the seminiferous
- between basolateral membranes which form a blood-testis barrier tubules to the duct of the epididymis.
within the seminiferous epithelium.
- Tightest blood-tissue barrier in mammals Loops of seminiferous tubules
- one part of a system preventing autoimmune attacks against the - join the rete testis by the short straight tubules which are lined
unique spermatogenic cells initially only by Sertoli cells. These empty into the rete testis, an
- First appear after the immune system is mature and central self- interconnected network of channels lined with cuboidal epithelium
tolerance is well established. and supported by CT of the mediastinum.
Basal Compartment (of the tubule) Rete testis
- where spermatogonia lie - drains into about 20 efferent ductules lined by an unusual
- below the tight junctions and not sealed off from the vascularized epithelium in which groups of non-ciliated cuboidal cells alternate
interstitial tissue containing lymphocytes and other immune cells. with groups of taller ciliated cells and give the tissue a scalloped
appearance.
Adluminal Comparment (of the tubule) - The non-ciliated cells absorb some of the fluid secreted by the
- newly formed primary spermatocytes temporarily disassemble the Sertoli cells of seminiferous tubules.
adhesion molecules of the local occluding junctions and move here - This absorption and the ciliary activity create a fluid flow which
while still adhering to Sertoli cells. carries sperm passively out of the testis toward the epididymis.
- Adluminal migration occurs without compromising the blood-testis - A thin layer of circulatory oriented smooth muscle cells in the walls
barrier even if germ cells remain linked by intercellular bridges. of efferent ductules aids the movement of sperm into the duct of
epididymis.
- Sertoli cells are also connected by coupled ionically by gap junctions
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(which may help regulate the transient changes in the occluding
EXCRETORY GENITAL DUCTS
junctions and synchronize activities in the spermatogenic cells)
- These are those of the epididymis, the ductus (or vas) deferens,
- As the flagellar tails of the spermatids develop, they appear as tufts and the urethra.
extending from the apical ends of the Sertoli cells. Function: Transport sperm from the scrotum to the penis during
ejaculation.
Three General Functions of Sertoli Cells
[1] Support, protection, and nutrition of the developing [1] EPIDIDYMIS
spermatogenic cells - has a long highly-coiled duct surrounded by CT and lies in the
Spermatocytes, spermatids, and developing sperm depend on scrotum along the superior and posterior sides of each testis.
Sertoli cells for production or transport into the lumen of - Regions of the epididymis:
metabolites and nutritive factors such as the iron-transport
Head Body Tail
protein transferrin (because they are isolated from plasma
Where the efferent Where sperm cells Where sperm is
proteins and nutrients by the blood-testis barrier). Thus, while
ductules enter. undergo further stored until
protecting spermatogenic cells, Sertoli cells supply many plasma
subtle modifications ejaculation.
factors needed for cell growth and differentiation.
Epithelium: Its duct is lined with pseudostratified columnar
[2] Exocrine and endocrine secretion [3] Phagocytosis epithelium12 consisting of columnar principal cells13 with long
These release (into the seminiferous During spermiogenesis, stereocilia and small round stem cells.
tubules) water that carries new sperm excess cytoplasm shed - Its duct is surrounded by a thin, circular layer of smooth muscle
out of the testis. Production of as residual bodies is cells, supplemented in the tail region with the addition of inner and
nutrients and androgen-binding phagocytosed and outer longitudinal layers.
protein (ABP)9 is promoted by follicle- digested by Sertoli cell
stimulating hormone (FSH). As lysosomes. No proteins At ejaculation, peristaltic contractions of these muscle layers rapidly
endocrine cells, they secrete the from sperm normally empty stored sperm from this tail region (which is continuous with
glycoprotein inhibin10. In the fetus, pass back across the the ductus deferens)
Sertoli cells also secrete Müllerian- blood-testis barrier.
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inhibiting substance (MIS)11. In the Principal Cells – these remove most of the water and residual
absence of MIS, these ducts persist bodies entering the epididymis with the sperm and secrete various
and become parts of the female products, greatly changing the fluid in which sperm is suspended.
reproductive tract
9 - These epithelial cells12 release abundant exosomes and other
Androgen-binding protein
vesicles from multivesicular bodies by apocrine secretion.
- concentrates testosterone to a level required for spermiogenesis
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Inhibin (39-kDa) – feeds back on the anterior pituitary gland to
suppress FSH synthesis and release.
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- Passage of sperm through the duct of the epididymis normally takes A] SEMINAL VESICLES
2-4 weeks, during which time the spermatozoa cell membranes bind - Two
and integrate many components secreted by the principal cells and - Exocrine glands
released from their exosomes. - Its production of viscid, yellowish secretion depends on
testosterone.
In general, changes within spermatozoa while passing through the - Consist of highly tortous tubes, each enclosed by a CT capsule.
epididymis include: Epithelium: Its folds are lined with simple or pseudostratified
[a] Development of competence for forward motility. columnar epithelial cells rich in secretory granules.
[b] Final modifications within the acrosome - The lamina propria contains elastic fibers and is surrounded by
[c] Reorganization of the cell membrane surrounding the sperm smooth muscle with inner circular and outer longitudinal layers
head, including the addition of cholesterol and other “decapacitation which empty the gland during ejaculation.
factors”, which block the acrosomal reaction (a key event in - Fluid from seminal vesicles make up about 70% of the ejaculate and
fertilization). The fertilizing ability of spermatozoa is inhibited until its components are:
their cell membranes are modified again as part of the capacitation
process, which occurs in the female reproductive tract. Fructose Prostaglandins Fibrinogen
Major energy Stimulate activity in Allows semen to
[2] DUCTUS OR VAS DEFERENS source for sperm, as the female coagulate after
- Each has a long straight tube with a thick, muscular wall and a well as inositol, reproductive tract. ejaculation.
relatively small lumen. citrate, and other
- This tube leaves the scrotum and continues toward the prostatic metabolites.
urethra where it empties.
- Its mucosa is slightly folded longitudinally, the lamina propria B] PROSTATE GLAND
contains many elastic fibers. - dense organ surrounding the urethra below the bladder.
Epithelium: pseudostratified with some cells having sparse - Consists of a collection of 30-50 tubuloacinar glands embedded in a
stereocilia. dense fibromuscular stroma in which smooth muscle contracts at
- The very thick muscularis consists of longitudinal inner and outer ejaculation. Ducts from individual glands may converge but all empty
layers and a middle circular layer. directly into the prostatic urethra, which runs through the center of
- The muscles produce strong peristaltic contractions during the prostate.
ejaculation (which rapidly move sperm along this duct from the
epididymis).

- The Vas Deferens forms part of the spermatic cord, which also
includes the testicular artery, the pampiniform plexus, and nerves.
- Each ductus passes over the urinary bladder where it enlarges as an
AMPULLA where the epithelium is thicker and more extensively
folded.
-Within the prostate gland, the ends of two ampullae merge with the
Three Major Zones of Glands around the Urethra
ducts of the two seminal vesicles, joining these ducts to form
Transition Zone Central Zone Peripheral Zone
ejaculatory ducts which open into the prostatic urethra.
- Occupies 5% of the - comprises 25% of - 70% of the organ’s
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prostate volume. the gland’s tissue. tissue.
ACCESSORY GLANDS
- Surrounds the - contains the - Contains the
- these produce secretions, which become mixed with sperm during
superior portion of periurethral prostate’s main
ejaculation to produce semen and are essential for reproduction.
the urethra submucosal glands glands with still
- These include seminal vesicles (or glands), prostate gland, and the
- contains the with longer ducts. longer ducts.
bulbourethral glands.
periurethral
mucosal glands

Epithelium: lined by simple or pseudostratified columnar epithelium


Function: Produce a fluid that contains a complex mixture of
exosomes, various glycoproteins, enzymes, and small molecules such
as prostaglandins and is stored until ejaculation.

The prostate is surrounded by a fibroelastic capsule, where septa


extend and divide the gland into indistinct lobes.
The prostate’s structure and function depend on the level of
testosterone (like in seminal vesicles)

Prostate-specific Antigen (PSA)


- clinically important product of prostate
- 34-kDa serine protease
- helps liquefy coagulated semen for the slow release of sperm after
ejaculation.
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- Elevated levels of circulating PSA indicate abnormal glandular Helicine Arteries
mucosa typically due to prostatic carcinoma or inflammation. - Nutritive arterioles and small coiling that are formed by branching
central arteries in the corpora cavernosa.
Corpora Amylacea - This leads to the cavernous vascular spaces of erectile tissue.
- small, spherical concretions that are often partially calcified
- normally present in the lumens of many prostatic tubuloacinar Penile erection
glands. - involves blood filling the cavernous spaces in the three masses of
- Contains primarily deposited glycoproteins and keratan sulfate erectile tissue.
- may become more numerous with age. - Triggered by external stimuli to the CNS, erection is controlled by
autonomic nerves in these vascular walls.
C] BULBOURETHRAL GLAND (or Cowper glands) - Parasympathetic stimulation relaxes the trabecular smooth muscle
- Paired, round glands and dilates the helicine arteries, allowing increase blood flow and
Location: Urogenital diaphragm filling of the cavernous spaces.
- these empty into the proximal part of the penile urethra. - This enlarges the corpora cavernosa and causes them to compress
- Each glands has several lobules with tubuloacinar secretory units the dorsal veins against the dense tunica albuginea, which blocks the
surrounded by smooth muscle cells. venous outflow and produces tumescence and rigidity in the erectile
Epithelium: Lined by a mucus-secreting simple columnar epithelium tissue.
which is also testosterone-dependent. - Beginning at ejaculation, sympathetic stimulation constricts the
Function: During erection, these glands as well as urethral glands, helicine arteries and trabecular muscle, decreasing blood flow into
along the penile urethra, release a clear mucus-like secretion which the spaces, lowering the pressure there, and allowing the veins to
coats and lubricates the urethra in preparation for the imminent drain most blood from the erectile tissue.
passage of sperm.
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PENIS
- Consists of three cylindrical masses of erectile tissue, plus the penile
urethra, surrounded by skin.
Epithelium:
Most of the penile urethra is lined by pseudostratified columnar
epithelium.

Corpus Cavernosum14 Corpus Spongiosum


Two dorsal erectile masses Ventral and it surrounds the urethra

Glans
- formed when the corpus spongiosum expands at its ends.
> Epithelium: Stratified squamous epithelium continuous with
that of the thin epidermis covering of the glans
surface.
Urethral Glands
- small, mucous-secreting glands
Location: Found along the length of the penile urethra.

Prepuce (or Foreskin)


- covers the glans in uncircumcised men.
- a retractable fold of thin skin with sebaceous glands on the internal
surface.
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Corpora Cavernosa
- Each surrounded by a dense fibroelastic layer (tunica albuginea).

All three erectile tissues consist of many venous cavernous spaces


lined with endothelium and separated by trabeculae with smooth
muscle and CT continuous with the surrounding tunic.

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