albuginea on the anterior and lateral sides of the testis

(Figure 21–2).
HISTOLOGY  Having evolved in cold-blooded animals, certain molecular

JOSE MARIA COLLEGE OF MEDICINE FOUNDATION events in the process of sperm formation cannot occur at

PRIMUS EXEMPLAR the core body temperature of 37°C.

 A permissive temperature of about 34°C is maintained in
CHAPTER 21: MALE REPRODUCTIVE SYSTEM
the scrotal sac by various mechanisms.
 Each testicular artery is surrounded by a rich pampiniform
Instructor: Dr. Jed I. Enumerables
venous plexus containing cooler blood from the testis,
Textbook: Junqueira’s Basic Histology Text & Atlas (2016)
which draws heat from the arterial blood by a
countercurrent heat-exchange system. Evaporation of
M ALE RE PRO DU CTIVE SYSTE M
sweat from the scrotum also contributes to heat loss.
 Consists of the testes, genital ducts, accessory glands, and
 Relaxation or contraction of the thin dartos muscle of the
penis (Figure 21–1).
scrotum and the cremaster muscles of the spermatic cords
 Testes produce sperm but also contain endocrine cells
move the testes away from or closer to the body,
secreting hormones such as testosterone, which drives
respectively, allowing further control on testicular
male reproductive physiology.
temperature.
 Testosterone is important for spermatogenesis, sexual
differentiation during embryonic and fetal development,
and control of gonadotropin secretion in the pituitary.
 A metabolite of testosterone, dihydrotestosterone, also
begins to act on many tissues during puberty (eg, male
accessory glands and hair follicles).
 The genital ducts and accessory glands 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 (L. seed), which is introduced into the
female reproductive tract by the penis.
ME DI CA L A PPLI CAT I O N
TE STIS
An excessive accumulation of serous fluid in one or both
 Each testis (or testicle) is surrounded by a dense sides of the scrotal sac, termed a hydrocele, is the most
common cause of scrotal swelling and a condition easily
connective tissue capsule, the tunica albuginea, which
corrected surgically.
thickens on the posterior side to form the mediastinum Cryptorchidism (Gr. kryptos, hidden + orchis, testis), the
failure of one or both testes to descend from the abdomen,
testis.
occurs in about 4% of male neonates, but in most of these
 From this fibrous region, septa penetrate the organ and individuals the testes move to the scrotum during the first year.
Bilateral cryptorchidism causes infertility if not surgically
divide it into about 250 pyramidal compartments or
corrected by 2 to 3 years of age.
testicular lobules (Figures 21-2 and 21-3).
 Each lobule contains sparse connective tissue with INTE RSTITIAL TISSU E
endocrine interstitial cells (or Leydig cells) secreting  The interstitial tissue of the testis between the
testosterone, and one to four highly convoluted seminiferous tubules consists of sparse connective tissue
seminiferous tubules in which sperm production occurs. containing fibroblasts, lymphatics, and blood vessels
 The testes develop retroperitoneally in the dorsal wall of including fenestrated capillaries.
 the embryonic abdominal cavity and are moved during  During puberty interstitial cells, or Leydig cells, develop as
fetal development to become suspended in the two halves large round or polygonal cells with central nuclei and
of the scrotal sac, or scrotum, at the ends of the spermatic eosinophilic cytoplasm rich in small lipid droplets (Figures
cords (Figure 21–2). 21–2b and 21–4).
 During migration from the abdominal cavity, each testis  These cells produce the steroid hormone testosterone,
carries with it a serous sac, the tunica vaginalis, derived which promotes development of the secondary male sex
from the peritoneum. characteristics.
 This tunic consists of an outer parietal layer lining the
scrotum and an inner visceral layer, covering the tunica
1
  Testosterone is synthesized by enzymes present in the
smooth ER and mitochondria similar to the system in
adrenal cortical cells.
 Testosterone secretion by interstitial cells is triggered by
the pituitary gonadotropin, luteinizing hormone (LH),
which is also called interstitial cell stimulating hormone
(ICSH).
 Testosterone synthesis thus begins at puberty, when the
hypothalamus begins producing gonadotropin-releasing
hormone.
 In the late embryonic testes gonadotropin from the
placenta stimulates interstitial cells to synthesize the
testosterone needed for development of the ducts and
glands of the male reproductive system.
 These fetal interstitial cells are very active during the third
and fourth months of pregnancy, then regress and become
quiescent cells resembling fibroblasts until puberty when
they resume testosterone synthesis in response to the
pituitary gonadotropin.
ME DICAL APPLICATIO N
Both interstitial cell tumors and Sertoli cell tumors
are rare.
Most (95%) testicular cancer involves germ cell
tumors, which only appear after puberty and are much more
likely to develop in men with untreated cryptorchidism.
SE M INE FE RO U S TU BU LE S
 Sperm are produced in the seminiferous tubules at a rate
of about 2 × 108 per day in the young adult.
 Each testis has from 250 to 1000 such tubules in its
lobules, and each tubule measures 150-250 μm in
diameter and 30-70 cm in length.
 The combined length of the tubules of one testis totals
about 250 m.
 Each tubule is actually a loop linked by a very short,
narrower segment, the straight tubule, to the rete testis, a
labyrinth of epithelium-lined channels embedded in the
mediastinum testis (Figures 21–2a and 21–3).
 About 10-20 efferent ductules connect the rete testis to
the head of the epididymis (Figure 21–2a).
 Each seminiferous tubule is lined with a complex,
specialized stratified epithelium called germinal or
spermatogenic epithelium (Figure 21–2b).
 The basement membrane of this epithelium is covered by
fibrous connective tissue, with an innermost layer
containing flattened, smooth muscle-like myoid cells
(Figure 21–2b), which allow weak contractions of the
tubule.
 The germinal epithelium consists of two types of cells:
1. Large nondividing Sertoli cells (Figure 21–4), which
physically and metabolically support developing sperm cell
precursors
2. Dividing cells of the spermatogenic lineage (Figure 21–5a)
 The cells of the spermatogenic lineage, comprising four or
more concentric layers of cells in the germinal epithelium,
develop from progenitor cells to fully formed sperm cells
over a period of approximately 10 weeks.
 As shown in Figure 21–5 spermatogenesis, the first part of
sperm production, involves mainly mitosis and meiosis and
is followed by spermiogenesis, the final differentiation
process occurring in the haploid male germ cells.
SPE RM ATO G ENE SIS
 Begins at puberty with proliferation of stem and
progenitor cells called spermatogonia (Gr. sperma + gone,
generation), small round cells about 12 μm in diameter.
These cells occupy a basal niche in the epithelial wall of
the tubules, next to the basement membrane and closely
associated with Sertoli cell surfaces (Figures 21–5, 21–6
and 21–7).
 Different stages of spermatogonia development can be
recognized by subtle changes in shape and staining
properties of their nuclei. Spermatogonia with dark, ovoid
nuclei act as stem cells, dividing infrequently and giving
rise both begins at puberty with proliferation of stem and
progenitor cells called spermatogonia (Gr. sperma + gone,
generation), small round cells about 12 μm in diameter.
 These cells occupy a basal niche in the epithelial wall of
the tubules, next to the basement membrane and closely
associated with Sertoli cell surfaces (Figures 21–5, 21–6
and 21–7).
2
 Different stages of spermatogonia development can be
recognized by subtle changes in shape and staining
properties of their nuclei. Spermatogonia with dark, ovoid
nuclei act as stem cells, dividing infrequently and giving
rise both to new stem cells and to cells with more pale-
staining, ovoid nuclei that divide more rapidly as transit
amplifying (progenitor) cells (Figure 21–7).
 These type A spermatogonia each undergo several unique
clonal divisions that leave most of the cells interconnected
as a syncytium. These become type B spermatogonia,
which have more spherical and pale nuclei.
 Each type B spermatogonium then undergoes a final
mitotic division to produce two cells that grow in size and
become primary spermatocytes, which are spherical cells
with euchromatic nuclei (Figures 21–6 and 21–7). Primary
spermatocytes replicate their DNA, so each chromosome
consists of duplicate chromatids, and enter meiosis, during
which homologous chromosomes come together in
synapsis, DNA recombination occurs, and two rapid cell
divisions produce haploid cells (see Chapter 3).
 The primary spermatocyte has 46 (44 + XY) chromosomes,
the diploid number, and a DNA content of 4N. (The letter
N denotes either the haploid number of chromosomes, 23
in humans, or the amount of DNA in this set.)
 Soon after their formation, these cells enter the first
meiotic prophase that lasts about 3 weeks. Most
spermatocytes seen in sections of testis are in this phase
of meiosis.
 The primary spermatocytes are the largest cells of the
spermatogenic lineage and are characterized by the
presence of partially condensed chromosomes in various
stages of synapsis and recombination (Figure 21–6).
 Homologous chromosomes separate in the first meiotic
division, which produces smaller cells called secondary
spermatocytes (Figures 21–5a and 21–7) with only 23
chromosomes (22 + X or 22 + Y), but each still consists of
two chromatids so the amount of DNA is 2N (see Chapter
3).
 Secondary spermatocytes are rare in testis sections
because they are very short-lived cells that remain in
interphase only briefly and quickly undergo the second
meiotic division.
 Division of each secondary spermatocyte separates the
chromatids of each chromosome and produces two
haploid cells called spermatids each of which contains 23
chromosomes (Figures 21–5a, 21–6, and 21–7).
 Because no S phase (DNA replication) occurs between the
first and second meiotic divisions, the amount of DNA per
cell is reduced by half when the chromatids separate and
the cells formed are haploid (1N).
 With fertilization, a haploid ovum and sperm produced by
meiosis unite and the normal diploid chromosome number
is restored.

3
THE CLO NAL NATU RE OF M ALE G E RM CE LLS spermatozoa, which are highly specialized to deliver male
DNA to the ovum.
 The stem cells produced by mitotic divisions of
 No cell division occurs during this process, and as with
spermatogonia remain as separate cells.
spermatogenesis the cells involved remain associated with
 However, all subsequent divisions of the daughter cells
Sertoli cells.
that become transit amplifying progenitor cells have
 The haploid spermatids are small (7-8 μm in diameter)
incomplete cytokinesis after telophase and the cells
cells near the lumen of the seminiferous tubules (Figures
remain attached to one another by intercellular bridges of
21–5a and 21–6b). Spermiogenesis includes formation of
cytoplasm (Figure 21–7).
the acrosome (Gr. akron, extremity + soma, body),
 These allow free cytoplasmic communication among the
condensation and elongation of the nucleus, development
cells during their remaining mitotic and meiotic divisions.
of the flagellum (L, whip), and the loss of much of the
 Although some cells degenerate without completing
cytoplasm.
spermatogenesis and some cells may separate, clones of
 The end result is the mature spermatozoon, which is
approximately a hundred cells may remain linked through
released from the Sertoli cell surface into the tubule’s
meiosis.
lumen. Spermiogenesis is commonly divided into four
 The complete significance of this spermatogenic
phases:
syncytium is not clear, but the cytoplasmic bridges allow
1. In the Golgi phase the cytoplasm contains a prominent
the haploid cells to be supplied with products of the
Golgi apparatus near the nucleus, mitochondria, paired
complete diploid genome, including proteins and RNA
centrioles, and free ribosomes. Small proacrosomal
encoded by genes on the X or Y chromosome missing in
vesicles from the Golgi apparatus coalesce as a single
their haploid nuclei.
membrane-limited acrosomal cap close to one end of the
 The germ cells finally become separated from one another
nucleus (Figures 21–5b and 21–8). The centrioles migrate
during differentiation (Figure 21–7).
to a position farthest from the acrosomal cap and one acts
 The cellular events and changes between the final mitoses
as a basal body, organizing the axoneme of the flagellum
of spermatogonia and the formation of spermatids take
which is structurally and functionally similar to that of a
about 2 months.
cilium (see Chapter 2).
 The spermatogenic cells are not randomly distributed in
2. In the cap phase the acrosomal cap spreads over about
the spermatogenic epithelium. Cells at different stages of
half of the condensing nucleus (Figures 21–5b and 21–8).
development are typically grouped together along the
The acrosome is a specialized type of lysosome containing
tubule, with the intercellular bridges helping to coordinate
hydrolytic enzymes, mainly hyaluronidase and a trypsin-
their divisions and differentiation.
like protease called acrosin. These enzymes are released
when a spermatozoon encounters an oocyte and the
acrosomal membrane fuses with the sperm’s plasma
membrane. They dissociate cells of the corona radiate and
digest the zona pellucida, both structures that surround
the egg (see Chapter 22). This process, the acrosomal
reaction, is one of the first steps in fertilization.
3. In the acrosome phase the head of the developingsperm,
containing the acrosome and the condensing nucleus,
remains embedded in the Sertoli cell while the growing
axoneme extends into the lumen of the tubule (Figure 21–
6b). Nuclei become more elongated and very highly
condensed, with the histones of nucleosomes replaced by
small basic peptides called protamines. Flagellum growth
continues as the tail and mitochondria aggregate around
its proximal region to form a thickened middle piece
where the ATP for flagellar movements is generated
(Figure 21–5).
4. In the maturation phase of spermiogenesis, unneeded
SPE RM IO GE NE SIS cytoplasm is shed as a residual body from each

 The final phase of sperm production is the temperature- spermatozoon and remaining intercellular bridges are lost.

sensitive process by which spermatids differentiate into Mature but not yet functional or mobile sperm (Figure 21–
5) are released into the lumen of the tubule.

4

ME DICAL APPLICATIO N
Decreased semen quality, which is frequently
idiopathic (arising from unknown causes), is a major
cause of male infertility.
Common features of poor semen quality
include oligospermia (ejaculate volume > 2 mL), sperm
cell density less than 10-20 million/mL, abnormal sperm
morphology, and flagellar defects that impair sperm
SE RTO LI CE LLS
 The Sertoli cells, named after Enrico Sertoli (1842-1910)
who first demonstrated their physiologic significance, are
tall “columnar” epithelial cells that nourish
spermatogenic cells and divide the seminiferous tubules
into two (basal and adluminal) compartments (Figure 21–
4c-e).
 All cells of the spermatogenic lineage are closely
associated with the extended surfaces of Sertoli cells and
depend on them for metabolic and physical support.
 Sertoli cells adhere to the basal lamina and their apical
ends extend to the lumen, as shown
immunohistochemically in Figure 21–4c-e.
 In routine preparations the outlines of Sertoli cells
surrounding the spermatogenic cells are very poorly
defined (Figures 21–6 and 21–8).
 Each Sertoli cell supports 30-50 developing germ cells.
 Ultrastructurally Sertoli cells are seen to contain abundant
SER, some rough ER, well-developed Golgi complexes,
numerous mitochondria, and lysosomes (Figure 21–8).
 Their nuclei are typically ovoid or triangular, euchromatic,
and have a prominent nucleolus, features that allow
Sertoli cells to be distinguished from the neighboring germ
cells (Figure 21–6).
 Important in Sertoli cell function are elaborate tight
occluding junctions between their basolateral membranes
that form a blood-testis barrier within the seminiferous
epithelium (Figure 21–5a).
 The tightest blood-tissue barrier in mammals, this physical
barrier is one part of a system that prevents autoimmune
attacks against the unique spermatogenic cells, which first
appear after the immune system is mature and central
self-tolerance is well established.
 Spermatogonia lie in a basal compartment of the tubule,
below the tight junctions and not sealed off from the
vascularized interstitial tissue containing lymphocytes and
other immune cells.
 Newly formed primary spermatocytes temporarily
disassemble the adhesion molecules of the local occluding
junctions and move into the tubule’s adluminal
compartment while still adhering to Sertoli cells (Figure
21–5a).
 Like the spermatogonia, all spermatocytes and spermatids
lie within invaginations of the Sertoli cells surfaces.
 Adluminal migration occurs without compromising the
blood-testis barrier, which is all the more impressive when
one remembers that the germ cells remain linked by
intercellular bridges.
 Sertoli cells are also connected and coupled ionically by
gap junctions, which may help regulate the transient
changes in the occluding junctions and synchronize
activities in the spermatogenic cells.
 As the flagellar tails of the spermatids develop, they
appear as tufts extending from the apical ends of the
Sertoli cells.
 Related to their role in establishing the blood-testis
the
barrier, Sertoli cells have three general functions:
1. Support, protection, and nutrition of the developing
spermatogenic cells
- Because spermatocytes, spermatids, and developing
sperm are isolated from plasma proteins and nutrients by
the blood-testis barrier, they depend on Sertoli cells for
production or transport into the lumen of metabolites and
nutritive factors such as the iron-transport protein
transferrin.
- Thus, while protecting spermatogenic cells from
circulating immune components, Sertoli cells supply many
plasma factors needed for cell growth and differentiation.
2. Exocrine and endocrine secretion
- Sertoli cells continuously release into the seminiferous
tubules water that carries new sperm out of the testis.
Production of nutrients and androgen-binding protein
(ABP), which concentrates testosterone to a level required
for spermiogenesis, is promoted by follicle-stimulating
hormone (FSH).
- As endocrine cells, they secrete the 39-kDa glycoprotein
inhibin, which feeds back on the anterior pituitary gland to
suppress FSH synthesis and release.
- In the fetus Sertoli cells also secrete a 140-kDa
glycoprotein called müllerian-inhibiting substance (MIS)
that causes regression of the embryonic mullerian
(paramesonephric) ducts; in the absence of MIS these
ducts persist and become parts of the female reproductive
tract.
5
 3. Phagocytosis
- During spermiogenesis, excess cytoplasm shed as residual
bodies is phagocytosed and digested by Sertoli cell
lysosomes.
- No proteins from sperm normally pass back across the
blood-testis barrier.

ME DICAL APPLICATIO N

Acute or chronic inflammation of the testis, orchitis,

frequently involves the ducts connecting this organ to the
epididymis.
Common forms of orchitis are produced by infective
agents and occur secondarily to a urinary tract infection
or a sexually transmitted pathogen such as Chlamydia or
Neisseria gonorrhoeae entering the testis from the
epididymis or via the lymphatics.
Acute epididymitis is a result of sexually transmitte
infections such as gonorrhea or Chlamydia infection and
causes intrascrotal pain and tenderness.
Persistent inflammation of the epididymis, such
as that associated with gonorrhea infections, includes
massive invasion by leukocytes into the infected duct,
stimulating fibrosis that obstructs the epididymis and is a
common cause of male infertility.

INTRATE STICU LAR DU CTS

 The intratesticular ducts are the straight tubules (or tubuli

recti), the rete testis, and the efferent ductules (Figure
21–2), all of which carry spermatozoa and liquid from the E X CRE TO RY G ENITAL DU CTS
seminiferous tubules to the duct of the epididymis (Table
 The excretory genital ducts are those of the epididymis,
21–1).
the ductus (or vas) deferens, and the urethra.
 The loops of seminiferous tubules join the rete testis by
 They transport sperm from the scrotum to the penis
the short straight tubules, which are lined initially only by
during ejaculation.
Sertoli cells (Figure 21–9).
 These empty into the rete testis, an interconnected
E PIDIDYM IS
network of channels lined with cuboidal epithelium and
 The long, coiled duct of the epididymis, surrounded by
supported by connective tissue of the mediastinum (Figure
connective tissue, lies in the scrotum along the superior
21–9).
and posterior sides of each testis (Figure 21–2).
 The rete testis drains into about 20 efferent ductules lined
 About 4-5 m in length, it includes a head region where the
by an unusual epithelium in which groups of nonciliated
efferent ductules enter, a body, and a tail opening into the
cuboidal cells alternate with groups of taller ciliated cells
ductus deferens.
and give the tissue a characteristic scalloped appearance
 Passage of sperm through the duct of the epididymis
(Figure 21–10).
normally takes 2-4 weeks, during which spermatozoa
 The nonciliated cells absorb some of the fluid secreted by
undergo maturation and acquire the ability to fertilize.
the Sertoli cells of seminiferous tubules.
Important changes within sperm while passing through the
 This absorption and the ciliary activity create a fluid flow
epididymis include:
that carries sperm passively out of the testis toward the
1. Development of the competence for independent forward
epididymis.
motility,
 A thin layer of circularly oriented smooth muscle cells in
2. Maturation of the acrosome, and
the walls of efferent ductules aids the movement of sperm
3. Biochemical and organizational changes within the cell
into the duct of the epididymis.
membrane.
 Fluid within the epididymis contains glycolipid
“decapacitation factors” that bind the plasma membranes
of sperm and block acrosomal reactions and fertilizing
ability until these factors are removed as part of the
capacitation process in the female reproductive tract.

6
  The epididymal duct is lined with pseudostratified
columnar epithelium consisting of columnar principal
cells, with characteristic long stereocilia, and small round
stem cells (Figure 21–11).
 The principal cells secrete glycolipids and glycoproteins,
but also absorb most of the remaining water and remove
residual bodies or other debris not removed earlier by
Sertoli cells.
 The duct epithelium is surrounded by a few layers of
smooth muscle cells, arranged as inner and outer
longitudinal layers as well as a circular layer in the tail of
the epididymis.
 At ejaculation peristaltic contractions of this muscle move
the sperm rapidly along the duct and empty the
epididymal tail and distal body regions.
ACCE SSO RY GLANDS
DU CTU S OR VAS DE FE RE NS
 From the epididymis the ductus (or vas) deferens, a long
straight tube with a thick, muscular wall and a relatively
small lumen, continues toward the prostatic urethra
where it empties (Figure 21–1).
 As shown in Figure 21–12, its mucosa is slightly folded
longitudinally, the lamina propria contains many elastic
fibers, and the epithelial lining is pseudostratified with
some cells having sparse stereocilia.
 The very thick muscularis consists of longitudinal inner and
outer layers and a middle circular layer. The muscles
produce strong peristaltic contractions during ejaculation,
which rapidly move sperm along this duct from the
epididymis.
 The ductus (vas) deferens forms part of the spermatic
cord, which also includes the testicular artery, the
pampiniform plexus, and nerves (Figure 21–2).
 Following the general path along which the embryonic
testes descend, each ductus passes over the urinary
bladder where it enlarges as a ampulla (L. a small bottle)
where the epithelium is thicker and more extensively
folded (Figure 21–13).
 Within the prostate gland, the ends of the two ampullae
merge with the ducts of the two seminal vesicles, joining
these ducts to form the ejaculatory ducts which open into
the prostatic urethra.
 The histology of the intratesticular and excretory ducts is
summarized in Table 21–1.
M E DICAL APPLICATIO N
The accessibility of the ductus (vas) deferens in
the spermatic cords allows for the most common surgical
method of male contraception: vasectomy.
In this procedure a very small incision is made
through the scrotal skin near the two ducts and each vas
is exposed, cut, and the two ends (or only the end
leading to the abdomen) are cauterized and tied.
After vasectomy sperm are still produced, but
they degenerate and are removed by macrophages in the
epididymis (and in the scrotal sac if the short portion of
the vas is left open-ended.)
Inflammatory and other changes occur in the
mucosa of the epididymis, but serious adverse effects of
vasectomy are usually minimal.
A vasectomy may be reversed by surgically
reconnecting the two ends of each ductus deferens.
However, even successful surgery very often
fails to restore fertility, due to incomplete sperm
maturation in the epididymis changed by postvasectomy
inflammation.
 The accessory glands of the male reproductive tract
produce secretions that are mixed with sperm during
ejaculation to produce semen and that are essential for
reproduction.
 The accessory genital glands are the seminal vesicles (or
glands), the prostate gland, and the bulbourethral glands
(Figure 21–13).
7
 PRO STATE G LAND

 The prostate gland is a dense organ that surrounds the

urethra below the bladder.
 It is approximately 2 cm × 3 cm × 4 cm in size and weighs
about 20 g.
 The prostate is a collection of 30-50 tubuloacinar glands
embedded in a dense fibromuscular stroma in which
smooth muscle contracts at ejaculation (Figure 21–13b).
 Ducts from individual glands may converge but all empty
directly into the prostatic urethra, which runs through the
center of the prostate.
 As shown in Figure 21–15, the glands are arranged in three
major zones around the urethra:
1. Transition zone occupies only about 5% of the prostate
volume, surrounds the superior portion of glands.

SE M INAL VE SICLE S 2. Central zone comprises 25% of the gland’s tissue and
contains the periurethral submucosal glands with longer
 The two seminal vesicles consist of highly tortuous tubes,
ducts.
each about 15-cm long, enclosed by a connective tissue
3. peripheral zone, with about 70% of the organ’s tissue,
capsule.
contains the prostate’s main glands with still longer ducts
 The unusual mucosa of the tube displays a great number
(Figure 21–16).
of thin, complex folds that fill much of the lumen (Figure
 The tubuloacinar glands of the prostate are all lined by a
21–14).
simple or pseudostratified columnar epithelium and
 The folds are lined with simple or pseudostratified
produce fluid that contains various glycoproteins,
columnar epithelial cells rich in secretory granules.
enzymes, and small molecules such as prostaglandins and
 The lamina propria contains elastic fibers and is
is stored until ejaculation.
surrounded by smooth muscle with inner circular and
 A clinically important product of the prostate is prostate-
outer longitudinal layers that empty the gland during
specific antigen (PSA), a 34-kDa serine protease that helps
ejaculation.
liquefy coagulated semen for the slow release of sperm
 The seminal vesicles are exocrine glands in which
after ejaculation.
production of their viscid, yellowish secretion depends on
 Small amounts of PSA also leak normally into the prostatic
testosterone.
vasculature; elevated levels of circulating PSA indicate
 Fluid from seminal vesicles typically makes up about 70%
abnormal glandular mucosa typically due to prostatic
of the ejaculate and its components include the following:
carcinoma or inflammation.
1. Fructose, a major energy source for sperm, as well as
 Small spherical concretions, 0.2-2 mm in diameter and
inositol, citrate, and other metabolites;
often partially calcified, are normally present in the lumens
2. Prostaglandins, which stimulate activity in the female
of many prostatic tubuloacinar glands (Figure 21–16).
reproductive tract; and
 These concretions, called corpora amylacea, containing
3. Fibrinogen, which allows semen to coagulate after
primarily deposited glycoproteins and keratan sulfate, may
ejaculation.
become more numerous with age but seem to have no
physiologic or clinical significance.
 The prostate is surrounded by a fibroelastic capsule, from
which septa extend and divide the gland into indistinct
lobes.
 Like the seminal vesicles, the prostate’s structure and
function depend on the level of testosterone.

8
 ME DICAL APPLICATIO N

The prostate gland is prone to three common problems:

(1) Chronic prostatitis, usually involving bacteria or other
infectious agents;
(2) Nodular hyperplasia or benign prostatic
hypertrophy, occurring mainly in the periurethral
mucosal glands where it often leads to compression of
the urethra and problems with urination; and
(3) Prostate cancer (adenocarcinoma), the most common
cancer in nonsmoking men, occurring mainly in glands of

BU LBO U RE THRAL G LANDS

 The paired round bulbourethral glands (or Cowper

glands), 3-5 mm in diameter, are located in the urogenital
diaphragm (Figure 21–13) and empty into the proximal
part of the penile urethra.
 Each gland has several lobules with tubuloacinar secretory
units surrounded by smooth muscle cells and lined by a
mucus-secreting simple columnar epithelium that is also
testosterone-dependent (Figure 21–13d).
 During erection the bulbourethral glands, as well as
numerous, very small, and histologically similar urethral
glands along the penile urethra, release a clear mucus-like
secretion that coats and lubricates the urethra in
preparation for the imminent passage of sperm.

PE NIS

 The penis consists of three cylindrical masses of erectile

tissue, plus the penile urethra, surrounded by skin (see
Figure 21–1).
 Two of the erectile masses—the corpora cavernosa—are
dorsal; the ventral corpus spongiosum surrounds the
urethra (Figure 21–17).
 At its end the corpus spongiosum expands, forming the
glans (Figure 21–13a).
 Most of the penile urethra is lined with pseudostratified
columnar epithelium. In the glans, it becomes stratified
squamous epithelium continuous with that of the thin
epidermis covering the glans surface. Small mucus-
secreting urethral glands are found along the length of the
penile urethra.

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 In uncircumcised men the glans is covered by the prepuce
or foreskin, a retractable fold of thin skin with sebaceous
glands on the internal surface.
 The corpora cavernosa are each surrounded by a dense
fibroelastic layer, the tunica albuginea (Figures 21–17 and
21–18).
 All three erectile tissues consist of many venous cavernous
spaces lined with endothelium and separated by
trabeculae with smooth muscle and connective tissue
continuous with the surrounding tunic (Figure 21–19).
 Central arteries in the corpora cavernosa branch to form
nutritive arterioles and small coiling helicine arteries,
which lead to the cavernous vascular spaces of erectile
tissue.
 Arteriovenous shunts are present between the central
arteries and the dorsal veins.
 Penile erection involves blood filling the cavernous spaces
in the three masses of erectile tissue.
 Triggered by external stimuli to the CNS, erection is
controlled by autonomic nerves in these vascular walls.
Parasympathetic stimulation relaxes the trabecular
smooth muscle and dilates the helicine arteries, allowing
increased blood flow and filling the cavernous spaces.
 This enlarges the corpora cavernosa and causes them to
compress the dorsal veins against the dense tunica
albuginea, which blocks the venous outflow and produces
tumescence and rigidity in the erectile tissue.
 Beginning at ejaculation, sympathetic stimulation
constricts the helicine arteries and trabecular muscle,
decreasing blood flow into the spaces, lowering the
pressure there, and allowing the veins to drain most blood
from the erectile tissue.

ME DI CA L A PPLI CA T IO N

At the beginning of an erection acetylcholine from

parasympathetic nerves causes the vascular endothelial cells of
the helicine arteries and cavernous tissue to release nitric oxide
(NO).
NO diffuses into the surrounding smooth muscle cells
and activates guanylate cyclase to produce cyclic GMP, which
causes these cells to relax and promotes blood flow for the
erection.
Erectile dysfunction, or impotence, can result from
diabetes, anxiety, vascular disease, or nerve damage during
prostatectomy.
The drug sildenafil may alleviate the problem by
inhibiting the phosphodiesterase that degrades cyclic GMP in
the smooth muscle cells of helicine arteries and erectile tissue.
The subsequent higher level of cGMP promotes
relaxation of these cells and enhances the neural effect to
produce or maintain an erection.

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