John W. Bigbee, Ph.

D
November 11, 2016
john.bigbee@vcuhealth.org

HISTOLOGY OF THE
MALE REPRODUCTIVE SYSTEM

I. General concepts
A.

Components
1.

Testis - paired organs

a.
b.

2.

Genital ducts
a.
b.
c.
d.

3.

Seminiferous tubules
Rete testis

Epididymis paired organs

Ductus deferens paired ducts
Ejaculatory duct paired ducts
Urethra

Major genital glands

a.
b.
c.

Seminal vesicles - paired glands

Prostate - single gland
Bulbourethral glands - paired glands

4. Penis
B. Functions of the male reproductive system components
1.
2.
3.
4.
5.

Produce sperm
Produce male sex hormones
Produce seminal fluid
Propel sperm and seminal fluid (semen) to exterior
Copulation

TESTIS
I.

General organization
A.

Paired, ovoid organ - serves both to produce sperm and secrete testosterone

B.

Connective tissue stroma

1.

Tunica albuginea - Connective tissue capsule

2.

Septa connective tissue partitions that separate the testis into lobules

3.

Mediastinum testis
a.

Thickening of tunica albuginea, projecting into the testis from its posterior
surface.

b.

Contains the rete testis, a network of ducts that receive the seminiferous
tubules and connect with the efferent ducts which carry sperm out of the

testis.

C.

Parenchyma
1.

Consists of the seminiferous tubules and interstitial cells of Leydig

2.

Seminiferous tubules Produce sperm

mediastinum.
3.
II.

a.

1-4 hairpin-shaped, tubules per lobule

b.

Convoluted portion - lined by the seminiferous epithelium where

sperm production occurs

c,

Straight portion (tubuli recti) - non-coiled, terminal ends of each

seminiferous tubule; connect with the rete testis in the
Lined by non-sperm forming cuboidal epithelium.

Interstitial cells of Leydig - produce testosterone

Interstitial cells of Leydig

A.

Clusters of in the connective tissue between the seminiferous tubules

B.

Steroid homone-secreting cytology

1.

Abundant SER, mitochondria with tubular cristae and lipid droplets in the
cytoplasm

2.

III.

Secrete testosterone, regulated by luteinizing hormone (LH), from the

gonadotrope cells in the adenohypophysis. LH in males is called interstitial cell
stimulating hormone (ICSH).

Seminifererous tubules lined by the seminiferous epithelium composed of Sertoli cells and
spermatogenic cells
A.

Sertoli cells (supporting cells)

1.

Tall, columnar cells that rest on the basement membrane and extend to the lumen.

2.

Nucleus is euchromatic, ovoid and infolded; long axis oriented perpendicular to

the basement membrane.

3.

Numerous lateral processes surround and invest maturing spermatogenic cells.

4.

Secrete androgen-binding protein to concentrate testosterone

5.

Regulated by follicle stimulating hormone (FSH) from the gonadotrope cells in

the adenohypophysis and testosterone from interstitial cells.

6.

Form the blood-testis barrier

a.

Formed by tight junctions between lateral processes of adjacent Sertoli

cells; divide the seminiferous epithelium into basal and luminal
compartments

b.

Basal compartment Contains cells in the early stages of

spermatogenesis; direct contact with tissue fluid

c.

Luminal compartment - Contains cells in the later stages of

spermatogenesis; tight junctions prevent free diffusion of tissue fluid
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contents, especially antibodies, thereby protecting cells from immunologic

attack.
B.

Meiosis
1.

Cell division process restricted to the germinal cells in the gonads,

ovary and testis, for the production of gametes, egg and sperm.

2.

Results in reduction of the chromosome number by one-half

(haploid). Fusion of the gametes to form a zygote reconstitutes the
normal diploid number of chromosomes present in somatic cells.

3.

Results in four daughter cells

4.

Meiosis occurs in two stages

a.

b.

C.

Meiosis I (reductional division)

i.

At anaphase, duplicated chromosomes of each homologous pair

migrate to the spindle pole.

ii.

Each daughter cell contains one-half the number of

chromosomes. Each chromosome still consists of two sister
chromatids.

Meiosis II (equational division)

i.

Begins soon after Meiosis I

ii.

Same phases as mitosis but involves segregation of sister

chromatids to daughter cells by the same phases seen during
mitosis.

Overview of spermatogenesis
5

1.

Spermatogonia are progenitor cells which divide by mitosis to produce more

spermatogonia (self renewal) and primary spermatocytes

2.

Primary spermatocytes contain 23 paired, replicated chromosomes (chromatids)

3.

Primary spermatocytes divide by Meiosis I which segregates one member of each

pair to the daughter cells (secondary spermatocytes), thereby reducing
chromosome number from 2N to 1N (haploid) (reductional

4.

Secondary spermatocytes divide by Meiosis II (equational division) which

segregates one chromatid of each replicated chromosome to the daughter
(spermatids)

division)

cells
D.

Spermatogenic cells - form a stratified epithelium (seminiferous epithelium)

1.

2.

3.

Spermatogonia
a.

Diploid cells resting on the basement membrane.

b.

Two types, one divides mitotically for self-renewal, the other type divides
mitotically to form primary spermatocytes.

Primary spermatocytes
a.

Form in the basal compartment, then probably migrate through the

tight junctions between Sertoli cell processes and enter the luminal
compartment.

b.

Remain in prophase about one-third of the spermatogenic cycle, so

many are seen.

c.

Largest germ cell; nuclei contain highly condensed chromosomes.

d.

Complete meiosis I (reductional division) to form haploid secondary

spermatocytes.

Secondary spermatocytes
a.

Haploid cells; stage lasts only eight hours of the entire 74-day
spermatogenic cycle.

c.

Divide by meiosis II (equational division) to form spermatids.

4.

Spermatids
a.

Located near the lumen of the seminiferous tubules

b.

No further division; undergo cyto-differentiation to form

spermatozoa by a process called spermiogenesis.
i.

Intercellular bridges break down.

ii.

Nucleus condenses and elongates

iii.

Acrosome forms. An acrosome is a large vesicle that

contains hydrolytic enzymes which aid the sperm in
the zona pellucida surrounding the oocyte.

iv.

Flagellum forms

penetrating

c.
5.

IV.

Haploid cells

Spermatozoa
a.

Released from Sertoli cells into the lumen of the seminiferous

tubules by a process called spermiation

b.

Anatomically mature, but incapable of fertilization at this time.

Courseofspermwithintestis
A.

Seminiferoustubules(convolutedportion),spermproduction

B.

Seminiferoustubules,(straightportion;tubulirecti),connects convoluted portion of

seminiferous tubules with rete testis

C.

Rete testis
1.

Meshwork of channels within mediastinum of testis, lined by simple cuboidal

epithelium

2.

Connects straight portion of seminiferous tubules with efferent ducts leading to

the head of the epididymis
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V.

Efferent ducts
A.

Connect rete testis the with the duct of epididymis

B.

Lined with a simple epithelium with scalloped border composed of alternating taller,
ciliated cells and shorter cuboidal cells. Function to propel spermatozoa and
testicular fluid

C.

As the efferent ducts exit the testis, they become highly coiled and form conical
structures, the coni vasculosi (sing. conus vasculosus), whose broad bases form
the head of the epididymis.

resorb

part of

GENITAL DUCTS ENTIRELY EXTERNAL TO THE TESTIS

I.

Epididymis
A.

A comma-shaped organ lying posterior to the testis that is divided into head, body and
tail subdivisions.

B.

Head region composition

1.

Efferent ducts

2.

Duct of epididymis - a single duct formed by fusion of efferent ducts

a.

Structure

b.

C.

II.

i.

Highly coiled, single tube (5 m long)

ii.

Lined by tall pseudostratified columnar epithelium with stereocilia.

Function
i.

Storage site for sperm

ii.

Absorption of excess testicular fluid

iii.

Movement of sperm toward ductus deferens

Body and tail regions of the epididymis contain the continuation of the duct of the
epididymis.

Ductus (vas) deferens

A.

Thick muscular tube conveying and propelling sperm from duct of epididymis to the
ejaculatory duct in the prostate.

B.

Histology
1.

Pseudostratified columnar epithelium with stereocilia surrounds a narrow lumen

with longitudinal folds

C.

2.

Thick smooth muscle layer - inner and outer longitudinal, middle circular layers

3.

Located in spermatic cord along with the spermatic artery, pampiniform venous
plexus of veins and a nerve plexus.

Continuous with the ejaculatory duct which traverses the prostate to join the urethra

GENITAL GLANDS
I.

II.

Seminal vesicles
A.

Paired glands lying posterior to urinary

bladder

B.

Each is composed of a single, highly

tortuous tube lined by pseudostratified
columnar epithelium; luminal surface has
complex folds.

C.

Provides the bulk of the seminal fluid;

contains fructose, citrate, proteins and
prostaglandins

Prostate
A.

Single, midline gland; largest of the genital glands; surrounds the prostatic urethra

B.
Composed of 30-50 glands which open into the prostatic urethra; consist ofthree groups,
central, transitional and peripheral; peripheral glands constitute 70% of the prostate.

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D.

C.

Histology

1.

Lined by pseudostratified columnar epithelium

2.

Prostatic concretions - lamellated, spherical bodies that are condensations of

secretory products. The number of concretions increases with age.

Function - contributes a thin, milky fluid to semen, that is rich in citric acid and acid
phosphatase. Contains hydrolytic enzymes including Prostate Specific Antigen (PSA) a
serine protease and fibinolysin.

PENIS
I.

Composition
A.

B.
II.

Three cylindrical masses of erectile tissue surround by tunica albuginea composed of

dense connective tissue
1.

Corpora cavernosa - paired dorsal cylinders

2.

Corpus spongiosum (corpus cavernosum urethrae); single, ventral cylinder that

surrounds the penile urethra

3.

Composed of sponge-like cavernous spaces (venous spaces) lined by

endothelium; separated by CT septa

Covered by thin skin

Blood supply and the process of erection

A.

Each corpus cavernosum contains a deep artery which distributes blood to nutritive
arteries and helicine arteries. Helicine arteries distribute blood to the cavernous

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spaces

III.

B.

In the flaccid state, there is minimal blood flow to the corpora, mostly through the
nutritive arteries.

C.

Erection
1.

Parasympathetic activity causes relaxation of smooth muscle (vasodilation) of the

deep and helicine arteries.

2.

The subsequent filling of the cavernous spaces expands the tissue which pushes
against the tunica albuginea, causing the penis to become erect and turgid.

3.

Corpus spongiosum does not become as erect as the other cavernous bodies
because the tunica albuginea is thinner here. Lumen of the urethra remains open
for the passage of sperm during ejaculation.

4.

Return to flaccid state occurs with decline of parasympathetic activity.

Penile urethra
A.

Located within corpus spongiosum

B.

Histology
1.

Epithelium: pseudostratified columnar epithelium becomes stratified squamous

moist near it opening

2.

Glands of Littre - mucous-secreting glands, provides lubrication for the ejaculate

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MALE REPRODUCTIVE SYSTEM

LABORATORY
TESTIS
Immature testis:

Slide #166, Testis, immature

This slide contains a section through the immature human testis in the scrotum. One the left side of the
section, it also includes a portion of the epididymis.
The outermost layer in this section is the scrotal wall which is separated from the testis by a space.
Directly covering the surface of the tesits is the tunica albuginea, which consists of an eosin-stained,
dense layer of connective tissue. The connective tissue septa which separate the testis into lobules
connect with the tunica albuginea. Notice on the left side of the testis, the tunica albuginea extends into
the center forming the mediastinum. The mediastinum contains a network of anastomosing channels,
the rete testis, which are lined with simple cuboidal epithelium. The rete testis connects with the
straight portions (tubuli recti) of the seminiferous tubules. In this section, the best examples of tubuli
recti are visible on the right end of the mediastinum.
Interstitial cells (of Leydig):

Slide #163, Testis

Examine the slide at low magnification and note the thick tunica albuginea at the bottom with its large
blood vessels. The remainder of the section contains the convoluted portions of seminiferous tubules.
Zoom in on this section and note the wide variety of profiles which reflects the highly coiled structure of
the tubules. Zoom in further to an area of connective tissue between the tubules and identify the
interstitial cells. These steroid secreting cells are present in small clusters and have a highly vacuolated
cytoplasm reflecting the large number of lipid droplets.
Seminiferous tubules:

Slide #163, Testis

This slide also shows the detailed structure of seminiferous tubules and the stages of spermatogenesis in
their epithelium. The seminiferous epithelium is composed of spermatogenic, germ cells and
supporting Sertoli cells. The epithelium is supported by a basement membrane and a prominent
connective tissue capsule. First, locate the Sertoli cells. Each cell is columnar shaped and the nucleus is
typically located in the basal one-third of the cell. The nucleus is ovoid rather than round, is often
indented and is usually oriented perpendicular to the basement membrane. The cell extends to the
luminal surface however the outline of the cell cannot be distinguished. Spermatogenic cells at all
stages of development are surrounded by cellular processes of the Sertoli cell.
Next review the details of the germ cells, always keeping in mind how each cell divides and the number
of chromosomes each possesses.
Spermatogonia:
These cells are located on the basement membrane. They are roughly cuboidal in
shape and their spherical nuclei will be used as a reference for identifying the nuclei of the other cell
types. A and B spermatogonia cannot be distinguished. These cells are diploid and divide by mitosis.

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Primary spermatocytes:
These cells are mostly located in the middle third of the epithelium in the
luminal compartment. Many spermatocytes will be visible and their nuclei contain large, condensed
chromosomes. These cells are arrested in prophase and their nuclei measure about 150% of the size of
the spermatogonia nuclei. They are the largest cells in the spermatogenic series. They are diploid and
are undergoing Meiosis I.
Secondary spermatocytes:
You are not required to identify these cells as they are very short lived and
hence very few are present. These cells are located closer to the lumen and have nuclei which are about
the same size as that of spermatogonia. These cells are formed when primary spermatocytes complete
Meiosis I during which the chromosome number is reduced by one-half. Hence these cells are haploid.
Secondary spermatocytes divided by Meiosis II to produce haploid spermatids.
Spermatids: Spermatids lie closer to the lumen and show a variety of cell shapes. Once formed,
spermatids do not divide. They first resemble the spherical secondary spermatocytes from which they
were derived and are often referred to early spermatids. Gradually, each begins to elongate, loses a
portion of its cytoplasm, develops a flagellum and are often referred to late spermatids. The process is
called spermiogenesis and all the while, the cells are associated with Sertoli cells.
Spermatozoa:
Mature spermatozoa are released by Sertoli cells into the lumen by the process
called spermiation. Only those cells which are free in the lumen are spermatozoa.
GENITAL DUCTS
Efferent ducts:

Slide #167, Testis, mediastinum,

The tissue in the lower two-thirds of this section contains the continuation of the rete testis out of the
testis (above) as it connects with the efferent ducts. In this lower area, locate a tubule of the rete testis.
These tubules have a low, simple cuboidal epithelium and in some cases you can see a second, taller
epithelium which is that of the efferent ducts. As the rete testis connects with the efferent ducts, these
dual epithelium tubules indicate areas of junction. Next locate one of the efferent ducts, which lined
entirely by this taller epithelium. Note that this epithelium has an irregular, scalloped margin and
contains both columnar and cuboidal cells. Each of the efferent ducts, together with the surrounding
connective tissue and blood vessels, coil into a cone called the conus vasculosus. On this slide, there
are three of these structures as indicated by the double arrows in the image below. The distal ends of the
efferent ducts anastomose to form the single duct of the epididymis, but this junction is not visible in
this section.
Epididymis: Slide #171, Epididymis, upper
This section is taken from the body of the epididymis. The epididymis is surrounded by a dense
connective tissue covering which enclosed the highly coiled, single duct of the epididymis are. Zoom
in one section of the duct and notes the pseudostratified columnar epithelium with stereocilia.
Beneath the epithelium is a thick muscle layer which helps move the sperm along the highly coiled duct.
The spermatozoa are not motile at this time. In the center of the duct is an accumulation of
spermatozoa.

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Spermatic cord: Slide #168, Spermatic cord Masson

This section of the spermatic cord has been stained with the Masson stain which highlights the
connective tissue in green. The large, thick walled structure to the left is the ductus deferens, the large
artery in the mass of tissue on the right is the testicular artery. The testicular artery is surrounded by a
system of veins called the pampinoform plexus which establishes a countercurrent of blood flow to aid
in regulating the temperature of the testis.
GENITAL GLANDS
Seminal vesicle:

Slide #174, Seminal vesicle

The seminal vesicle is a single tubular gland which is highly coiled and saccular. It joins the ductus
deferens just before it enters the prostate as the ejaculatory duct. Each of the circular/oval profiles you
see are sections of the same, coiled tube. The mucosa of the seminal vesicle is folded and complex,
which subdivides the lumen into numerous crypts. This complex mucosa is diagnostic for the seminal
vesicle. The epithelium is pseudostratified columnar.
Prostate gland:
Slide #173, Prostate
Unlike the seminal vesicle, the prostate gland is composed of many individual glands which empty
independently into the urethra. Each gland is embedded in an abundant stroma of dense irregular
connective tissue and the mucosa is much simpler than the seminal vesicle. The epithelium is
pseudostratified columnar. Diagnostic for the prostate is the presence of eosinophilic prostatic
concretions in the lumen of some of the glands. These increase with age, but are mostly asymptomatic.
PENIS
Penis: Slide #194, Penis
This cross section through the penis is inverted such that the ventral surface of the organ is oriented
toward the top of the slide. Locate the paired, dorsally located corpora cavernosa and the single
ventral corpus spongiosum. These structures are surrounded by a tunica albuginea, which is much
thicker around the corpora cavernosa. Note that the septum between the two corpora is incomplete.
Examine the erectile tissue, and identify the cavernous spaces and the intervening trabeculae. Within
each corpora cavernosa is a deep artery, located near the midline. The corpus spongiosum encloses the
penile urethra.

15

Structure to identify
Immature Testis
Mediastinum
Rete testis
Seminiferous tubules
Convoluted portions
Straight portions (tubuli recti)
Tunica albuginea

Genital ducts
Duct of the epididymis
Ductus deferens
Efferent ducts
Rete testis
Tubuli recti

Testis
Interstitial cells of Leydig

Spermatic cord
Ductus deferens
Pampiniform plexus of veins
Testicular artery

Mediastinum
Primary spermatocytes

Genital glands
Prostate
Prostatic concretions
Seminal vesicle

Sertoli cells
Spermatogonia
Spermatids (early and late)

Penis
Corpora cavernosa
Corpus spongiosum
Deep arteries
Glands of Littre
Penile urethra
Tunica albuginea

Spermatozoa
Tunica albuginea

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