Saladin 7e Extended Outline

Chapter 27
The Male Reproductive System
I. Sexual Reproduction and Development (pp. 10291034)
A. The essence of sexual reproduction is that it involves two sexes whose gametes
combine to form a zygote that develops into a new individual (p. 1029)
1. The gametes must be motile so that they can come into contact, and
must be able to provide nutrients for the zygote.
2. Usually these tasks are apportioned to two different kinds of gametes: a
small motile one (sperm or spermatozoon) and a large nutrient-laden one
(egg or ovum).
3. By definition, in sexually reproducing species the individual that
produces eggs is the female, and the individual that produces sperm is
male.
4. Genetically, however, a human with a Y sex chromosome is classified
as male, and a human lacking a Y is classified as female.
5. In mammals, the female parent also carries the developing embryo
within her body.
6. The male must have a copulatory organ, the penis, for depositing sperm,
and the female must have a copulatory organ, the vagina, for receiving
sperm.
B. The male reproductive system serves to produce sperm and introduce them into
the female body, while the female reproductive system produces eggs, receives
sperm, provides for the union of the gametes, harbors the fetus, gives birth, and
nourishes the offspring. (pp. 10291030)
1. The reproductive system consists of primary and secondary sex organs.
a. The primary sex organs, or gonads, are those that produce the
gametestestes of the male and ovaries of the female.
b. The secondary sex organs are organs other than gonads that are
necessary for reproduction.
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Saladin Outline Ch.27

i. In the male, they comprise the system of ducts, glands,
and the penis, concerned with storage, survival, and
conveyance of sperm.
ii. In the female they include the uterine tubes, uterus, and
vagina, concerned with uniting the sperm and egg and
harboring the fetus.
2. According to location, the reproductive organs are classified as external
and internal genitalia. (Table 27.1)
a. The external genitalia are located in the perineum. (Fig. 27.6)
b. The internal genitalia are located mainly in the pelvic cavity,
except for the male testes and some associated ducts in the
scrotum.
3. Secondary sex characteristics are features that further distinguish the
sexes and play a role in mate attraction.
Insight 27.1 Androgen-Insensitivity Syndrome (Fig. 27.1)
a. They typically appear only as an animal approaches sexual
maturity.
b. In humans, the physical attributes that contribute to mate
attraction are culturally conditioned to a great degree.
c. In both sexes, pubic and axillary hair and their associated sex
glands, and the pitch of the voice are generally accepted as signs of
sexual maturity.
d. Other traits commonly regarded as male characteristics are facial
hair, coarse hair on the torso and limbs, and a relatively muscular
physique.
e. Other traits commonly regarded as female characteristics include
distribution of body fat, enlargement of breasts, and relatively
hairless appearance of the skin.
C. The combination of sex chromosomes received by a zygote is a determining
factor of development into male or female. (pp. 10301031)

Saladin Outline Ch.27

1. Most human cells have 23 pairs of chromosomes, of which 1 pair
constitutes the sex chromosomes.
2. The X chromosome is larger than the Y chromosome, and every egg
contains an X chromosome.
3. Of the sperm, half carry an X and the other half carry a Y.
a. Fertilization between and X egg and an X sperm yields an XX
zygote destined to become female.
b. Fertilization between an X egg and a Y sperm yields an XY
zygote destined to become male.
c. Thus, the sex of a child is determined at fertilization by the
sperm. (Fig. 27.2)
D. Sex determination does not end with fertilization, but requires an interaction
between genetics and the hormones produced by the mother and fetus. (p. 1031)
1. Up to a point, a fetus is sexually undifferentiated; its gonads begin to
develop at 5 to 6 weeks as gonadal ridges, lying alongside a primitive
kidney, the mesonephros, which later degenerates.
a. Adjacent to each gonadal ridge are two ducts, the mesonephric
(wolffian) duct, and the paramesonephric (mllerian) duct.
b. In males, the mesonephric ducts develop into the reproductive
tract, and the paramesonephric ducts degenerate; in females, the
opposite occurs. (Fig. 27.3)
2. The Y chromosome has a gene called SRY (sex-determining region of
the Y) that codes for a protein called testis-determining factor (TDF),
which interacts with genes on other chromosomes.
a. By 8 to 9 weeks, the male gonadal ridge has become a
rudimentary testis that begins to secrete testosterone.
b. Testosterone stimulates the mesonephric duct to develop into the
system of male reproductive ducts.
c. By this time, the testis also secretes a hormone, mllerianinhibiting factor (MIF), which causes atrophy of the
paramesonephric duct.
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Saladin Outline Ch.27

i. Even an adult male retains a tiny Y-shaped vestige of the
paramesonephric ducts in the area of the prostatic urethra.
3. The level of estrogen is always high during pregnancy; the development
of a female therefore results from the absence of androgens, not the
presence of estrogens.
E. External genitalia begin to develop from identical structures in both sexes.
(p. 1031) (Fig. 27.4)
1. By 6 weeks, the embryo has the following structures:
a. A genital tubercle, which is an anterior median bud.
b. Urogenital folds, a pair of medial tissue folds slightly posterior
to the genital tubercle.
c. Labioscrotal folds, a larger pair of tissue folds lateral to the
urogenital folds.
2. By the end of week 9, the fetus begins to show sexual differentiation,
and either male or female genitalia are distinctly formed by the end of
week 12.
a. In the female, the three structures become the clitoral glans,
labia minora, and labia majora.
b. In the male, the genital tubercle elongates to form the phallus;
the urogenital folds fuse to enclose the urethra, joining the phallus
to form the penis; and the labioscrotal folds fuse to form the
scrotum.
c. Thus these organs are homologous, and in some abnormalities of
development, they may develop into the organs of the other sex.
F. Both male and female gonads initially develop near the kidneys and then
descend into the pelvic cavity (ovaries) or scrotum (testes). (pp. 10311034)
1. In the embryo, a connective tissue cord, the gubernaculum, extends
from the gonad to the floor of the abdominopelvic cavity.
a. In males, the gubernaculum continues to grow and passes
between the internal and external abdominal oblique muscles and
into the scrotal swelling.
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Saladin Outline Ch.27

b. Independently, the peritoneum also develops a fold that extends
into the scrotum as the vaginal process.
c. These create a path through the groin called the inguinal canal,
which is the most common site of herniation in boys and men.
2. The descent of the testes begins as early as week 6. (Fig. 27.5)
a. The superior part of the embryonic gonad degenerates and the
inferior part migrates downward, guided by the gubernaculum.
b. In the seventh month, the testes abruptly pass through the
inguinal canals into the scrotum.
c. As they descend, they are accompanied by elongating testicular
arteries and veins, lymphatic vessels, nerves, spermatic ducts, and
extensions of the internal abdominal oblique muscle.
d. The vaginal process becomes separated from the peritoneal
cavity to persist as the tunica vaginalis, enfolding the anterior and
lateral sides of the testis.
e. The mechanism of descent remains obscure.
3. About 3% of boys are born with undescended testes, or cryptorchidism.
a. In most cases the testes descend with the first year of infancy,
but if they do not, testosterone injection or surgery can correct this.
b. Uncorrected cryptorchidism leads to sterility and sometimes to
testicular cancer.
4. The ovaries descend also, but to a lesser extent.
a. The gubernaculum extends from the inferior pole of the ovary to
the labioscrotal fold.
b. The ovaries eventually lodge just inferior to the brim of the
lesser pelvis.
c. The gubernaculum becomes a pair of ligaments that support the
ovary and uterus.

Saladin Outline Ch.27

II. Male Reproductive Anatomy (pp. 10341042)
A. The scrotum and penis constitute the external genitalia of the male and occupy
the perineum, a diamond-shaped area between the thighs bordered by the pubic
symphysis, ischial tuberosities, and coccyx. (pp. 10341036) (Fig. 27.6)
B. The scrotum is a pendulous pouch of skin, muscle, and fibrous connective
tissue, containing the testes. (pp. 10351036) (Fig. 27.7)
1. The skin of the scrotum has sebaceous glands, sparse hair, rich sensory
innervation, and somewhat darker pigmentation.
2. It is divided into right and left compartments by an internal median
septum.
a. This protects each testis from infections involving the other one.
b. The location of the septum is externally marked by the perineal
raphe, which extends anteriorly along the ventral side of the penis
and posteriorly as far as the margin of the anus. (Fig. 27.6)
3. The left testis is usually suspended lower than the right so that the two
are not compressed against each other between the thighs.
4. Posteriorly, the scrotum contains the spermatic cord, a bundle of fibrous
connective tissue containing the ductus deferens, blood and lymphatic
vessels, and testicular nerves.
a. The spermatic cord passes upward behind and superior to the
testis, continues along the anterior side of the pubis, and into a
4-cm-long inguinal canal, which emerges into the pelvic cavity.
b. The inferior entrance into the inguinal canal is the external
inguinal ring, and its superior exit into the pelvic cavity is the
internal inguinal ring.
5. Testes cannot produce sperm at the core body temperature of 37C and
must be held at about 35C. The scrotum has three mechanisms to regulate
temperature.
a. The cremaster muscle consists of strips of the abdominal oblique
that contract and draw the testes closer to the body when it is cold;

Saladin Outline Ch.27

when it is warm, the cremaster relaxes and lowers the testes farther
from the body.
b. The dartos muscle is a subcutaneous layer of smooth muscle that
also contracts when it is cold, and the scrotum becomes taut and
wrinkled, helping to hold the testes against the warm body.
c. The pampiniform plexus is an extensive network of veins from
the testis that surrounds the testicular artery in the spermatic cord,
cooling the warm arterial blood by countercurrent heat exchange
and lowering the temperature of the arterial blood by 1.5 to
2.5C. (Fig. 27.8)
C. The testes (testicles) are combined endocrine and exocrine glands that produce
sex hormones and sperm. (pp. 10361038)
1. Each testis is oval and slightly flattened, 4 cm long, 3 cm from anterior
to posterior, and 2.5 cm wide. (Fig. 27.9)
a. Its anterior and lateral surfaces are covered by the tunica
vaginalis.
b. The testis has a white fibrous capsule called the tunica
albuginea.
c. Connective tissue septa divide the testis into 250 to 300 wedgeshaped lobules, each of which contains one to three seminiferous
tubules, ducts up to 70 cm long that produce the sperm.
d. Between the seminiferous tubules are clusters of interstitial
(Leydig) cells, the source of testosterone.
2. A seminiferous tubule has a narrow lumen lined by a thick germinal
epithelium.
a. The epithelium consists of several layers of germ cells in the
process of becoming sperm, and a smaller number of tall
sustentacular (nurse or Sertoli) cells, which protect the germ cells
and promote their development.
i. Germ cells rely on sustentacular cells for nutrients, waste
removal, growth factors, etc.
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Saladin Outline Ch.27

ii. The sustentacular cells also secrete androgen-binding
protein and inhibin, both of which regulate the rate of
sperm production.
3. A sustentacular cell is shaped like a tree trunk with spreading roots
along the basement membrane.
a. This forms the boundary of the tubule; the thick trunk reaches to
the tubule lumen.
b. Tight junctions between adjacent sustentacular cells form a
bloodtestis barrier (BTB) that prevents antibodies, other large
molecules, and intercellular fluid from getting to the germ cells.
i. This barrier is important because the germ cells are
genetically different from other cells.
ii. Some cases of sterility occur when the BTB fails to form
adequately in adolescence.
c. The seminiferous tubules lead into a network called the rete
testis, embedded in the capsule on the posterior side.
i. Sperm partially mature in the rete; they are moved along
by fluid secreted by the sustentacular cells and cilia on
some rete cells.
ii. Sperm do not swim while in the male reproductive tract.
4. Each testis is supplied by a testicular artery that arises from the
abdominal aorta just below the renal artery.
a. This is a very long, slender artery that winds its way down the
posterior abdominal wall before passing through the inguinal canal
into the scrotum. (Fig. 27.7)
b. Its blood pressure is very low, and it is one of the few arteries to
have no pulse.
c. Blood flow to the testes is therefore meager and oxygen is in
short supply.

Saladin Outline Ch.27

d. In response, the sperm develop unusually large mitochondria,
which may help them survive in the hypoxic female reproductive
tract.
5. Blood leaves the testis by way of the pampiniform plexus of veins,
which converge as they pass through the inguinal canal to form the
testicular vein.
a. The right testicular vein drains into the inferior vena cava.
b. The left testicular vein drains into the left renal vein.
6. Lymphatic vessels also drain each testis, traveling through the inguinal
canal with the veins and leading to lymph nodes near the lower aorta.
a. Lymph from the penis and scrotum travels to lymph nodes
adjacent to the iliac arteries and veins and in the inguinal region.
7. Testicular nerves lead to the gonads from spinal cord segments T10 and
T11.
a. They are mixed sensory and motor nerves containing
predominantly sympathetic but also some parasympathetic fibers.
b. The sensory fibers are concerned primarily with pain, and the
autonomic fibers are predominantly vasomotor, for regulation of
blood flow.
D. After leaving the testis, sperm travel through a series of spermatic ducts to
reach the urethra. (pp. 10381040) (Fig. 27.10)
1. About 12 small efferent ductules arise from the posterior side of the
testis and carry sperm to the epididymis; they have clusters of ciliated
cells.
2. The epididymis is a site of sperm maturation and storage.
a. It consists of a clublike head, a long middle body, and a slender
tail.
b. It contains a single coiled duct about 6 m (20 ft) long, embedded
in connective tissue.
c. This duct reabsorbs about 90% of the fluid secreted by the testis.

Saladin Outline Ch.27

d. Sperm take about 20 days to reach the tail, where they are
stored.
e. The sperm remain fertile for 40 to 60 days; if not ejaculated,
they disintegrate and are reabsorbed.
3. The ductus (vas) deferens begins where the duct of the epididymis
straightens out at the tail and turns 180.
a. It is a muscular tube about 45 cm long and 2.5 mm in diameter,
passing upward through the spermatic cord and inguinal canal into
the pelvic cavity.
b. After passing between the bladder and ureter, it turns downward
behind the bladder and widens into a terminal ampulla.
c. It unites with the duct of the seminal vesicle.
d. The duct has a thick wall of smooth muscle innervated by
sympathetic nerve fibers.
4. Where the ductus deferens and duct of the seminal vesicle meet, they
form a short (2 cm) ejaculatory duct.
a. This duct passes through the prostate gland and empties into the
urethra.
b. This is the last of the spermatic ducts.
5. The male urethra is shared by the reproductive and urinary systems.
a. It is about 18 cm long and consists of three regions: the
prostatic, membranous, and spongy (penile) urethra.
b. The urethra cannot pass both urine and semen simultaneously.
E. Three sets of accessory glands are found in the male reproductive system: the
seminal vesicles, prostate gland, and bulbourethral glands. (p. 1040)
1. The seminal vesicles are a pair of glands posterior to the bladder; one is
associated with each ductus deferens.
a. A seminal vesicle is about 5 cm long with a connective tissue
capsule and underlying layer of smooth muscle.
b. The secretory portion is a very convoluted duct with numerous
branches that empties into the ejaculatory duct.
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Saladin Outline Ch.27

c. The yellowish secretion of the seminal vesicles constitutes about
60% of the semen.
2. The prostate gland surrounds the urethra and ejaculatory ducts
immediately inferior to the urinary bladder.
Insight 27.2 Prostate Diseases
a. It measures about 2 4 3 cm and is an aggregate of 30 to 50
tubuloacinar glands in a fibrous capsule.
b. The glands empty through about 20 pores in the urethral wall.
c. The stroma consists of connective tissue and smooth muscle,
like that of seminal vesicles.
d. The prostate secretion constitutes about 30% of the semen.
3. The bulbourethral (Cowper) glands are named for their position near a
dilated bulb at the inner end of the penis and their association with the
urethra.
a. They are brownish, spherical glands about 1 cm in diameter,
with a 2.5 cm duct to the urethra.
b. During sexual arousal, they produce a clear, slippery fluid that
lubricates the head of the penis.
c. The fluid also protects the sperm by neutralizing the acidity of
residual urine in the urethra.
F. The penis serves to deposit semen into the vagina. (pp. 10401041)
1. Half of the penis is an internal root, and half is the externally visible
shaft and glans. (Figs. 27.10 and 27.11)
a. The external portion is about 8 to 10 cm (34 in.) long and 3 cm
in diameter when flaccid.
b. Typical dimensions of an erect penis are 13 to 18 cm (57 in.)
long and 4 cm in diameter.
c. The glans is the expanded head at the distal end of the penis with
the external urethral orifice at its tip.
2. The directional terminology is based on that of most mammals, which
are not bipedal.
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Saladin Outline Ch.27

a. The dorsal side is the one that faces anteriorly when flaccid; the
ventral side faces posteriorly.
b. The urethra passes through the lower, more obviously ventral
half.
3. The skin is loosely attached to the penile shaft, allowing for expansion
during erection.
a. The skin continues over the glans as the prepuce, or foreskin,
which is often removed by circumcision in some cultures.
b. A ventral frenulum attaches the skin to the glans.
c. The skin of the glans is thinner and is firmly attached to the
underlying erectile tissue.
d. The glans and facing surface of the prepuce have sebaceous
glands that produce a waxy secretion called smegma.
4. The penis consists mainly of three cylindrical bodies called erectile
tissues that fill with blood during arousal and bring about enlargement and
erection.
a. A single erectile body, the corpus spongiosum, passes along the
ventral side of the penis and encloses the penile urethra; at the
distal end it expands to fill the entire glans.
b. Proximal to the glans, the dorsal side of the penis has a corpus
cavernosum on each side.
i. Each is ensheathed in a fibrous tunica albuginea, and they
are separated by a median septum, similar to the tissues of
the same name of the testes and scrotum.
5. At the body surface, the penis turns 90 posteriorly and continues
inward as the root.
a. The corpus spongiosum terminates internally as a dilated bulb,
ensheathed in the bulbospongiosus muscle and attached to the
lower surface of the muscular pelvic floor.
b. The corpora cavernosa diverge like the arms of a Y, of which
each arm is called a crus.
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Saladin Outline Ch.27

c. The crura attach the penis to the pubic arch (ischiopubic ramus)
and perineal membrane on each side.
d. Each crus is enveloped by an ischiocavernosus muscle.
III. Puberty and Climacteric (pp. 10421044)
A. Unlike any other organ system, the reproductive system remains dormant for
several years after birth. At age 10 to 12 in most boys and 8 to 10 in most girls, a
surge of pituitary gonadotropins awakens the reproductive system at the onset of
puberty. (p. 1042)
B. Adolescence is the period from the onset of gonadotropin secretion and
reproductive development until a person attains full adult height. Puberty is the
first few years of adolescence until the first menstrual period in girls or the first
ejaculation of viable sperm in boys. (p. 1042)
C. Puberty is under endocrine control. (pp. 10421043)
1. The testes secrete substantial amounts of testosterone in the first
trimester (3 months) of fetal development, but then become dormant for
the rest of infancy and childhood.
2. From puberty through adulthood, reproductive function is regulated by
hormonal links between the hypothalamus, pituitary, and gonads.
(Fig. 27.12)
3. As the hypothalamus matures, it begins to produce gonadotropinreleasing hormone (GnRH), which travels via the hypophyseal portal
system to the anterior pituitary.
a. GnRH stimulates cells called gonadotropes to secrete folliclestimulating hormone (FSH) and luteinizing hormone (LH), two
gonadotropins that act on different testicular cells.
b. LH stimulates the interstitial cells of the testes to secrete
androgens, mainly testosterone.
i. In the male, LH is sometimes called interstitial cell
stimulating hormone (ICSH).

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Saladin Outline Ch.27

c. FSH stimulates the sustentacular cells to secrete a protein called
androgen-binding protein (ABP) into the lumen of the
seminiferous tubules.
i. ABP raises testosterone levels in the tubules and
epididymis by binding and accumulating the hormone.
ii. Without FSH and ABP, testosterone has no effect on the
testes.
iii. Germ cells have no androgen receptors and do not
respond to it.
4. Many of the physiological processes of puberty are not noticeable at
first, but in time, androgens have many conspicuous effects.
a. They stimulate growth of the sex organs.
i. Testes and scrotum enlarge around age 13.
ii. The penis continues to grow for about 2 more years.
iii. Internally, the ducts and accessory glands also enlarge.
b. Testosterone stimulates a burst of generalized body growth.
i. Limb bones elongate rapidly, and more muscle mass
develops.
ii. The larynx enlarges, deepening the voice and making the
thyroid cartilage more prominent.
iii. Erythropoiesis accelerates, resulting in a higher RBC
count.
iv. The basal metabolic rate increases, accompanied by an
increase in appetite.
c. Dihydrotestosterone (DHT) stimulates development of pubic
hair, axillary hair, and facial hair.
i. The skin becomes darker and thicker and secretes more
sebum.
ii. The apocrine scent glands of the perineal, axillary, and
beard areas develop in conjunction with the hair in those
regions.
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d. Accumulation of testosterone by ABP leads to the onset and
rising rate of sperm production.
i. If testosterone secretion ceases, the sperm count and
semen volume decline rapidly, and a male becomes sterile.
e. Testosterone inhibits GnRH secretion by the hypothalamus and
GnRH sensitivity of the pituitary.
i. FSH and LH secretions are thus held in check.
ii. Over the course of puberty, however, the pituitary
becomes less sensitive to negative feedback and secretes
increasing amounts of gonadotropins, resulting in rising
androgens.
f. Testosterone also stimulates the brain and awakens the libido
(sex drive), although the neurons convert the testosterone to
estrogen, which is what directly affects the behavior.
i. Increasing libido and sensitivity causes erections to occur
frequently.
ii. Ejaculation often occurs during sleep (nocturnal
emissions).
5. FSH secretion and sperm production can be modulated without having
to reduce LH and testosterone via the hormone inhibin.
a. Inhibin is secreted by the sustentacular cells and suppresses FSH
output from the pituitary.
b. When the sperm count drops below 20 million sperm/mL,
inhibin secretion drops and FSH secretion rises.
D. During aging, testosterone secretion declines; at age 20, it is about 7 mg/day,
but as little as one-fifth of this level by age 80. (pp. 10431044)
1. There is a corresponding decline in the number and activity of
interstitial cells and sustentacular cells.
2. FSH and LH levels rise significantly after age 50 and may produce
changes associated with the controversial concept of the male climacteric
or andropause.
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3. Most men experience little or no effects, but in some cases may
experience mood changes, hot flashes, and illusions of suffocation.
4. About 20% of men in their 60s and 50% of men in their 80s experience
erectile dysfunction (impotence or ED).
a. ED is the frequent inability to produce or maintain an erection
sufficiently for intercourse (Table 27.2)
b. ED and declining sexual activity can have an impact on older
peoples perception of the quality of life.
c. Over 90% of men with ED, however, remain able to ejaculate.
IV. Sperm and Semen (pp. 10441049)
A. Spermatogenesis is the process of sperm production; it occurs in the
seminiferous tubules and involves three principal events: (p. 1044)
1. Remodeling of a relatively large germ cell into small, mobile cells with
flagella.
2. Reduction of the chromosome number by one-half.
3. A shuffling of the genes so that each chromosome of the sperm carries
new gene combinations.
B. Meiosis is the process of cell division in which recombination (shuffling) of
genes and reduction in chromosome number are achieved. (pp. 10441046)
1. In nearly all living organisms except bacteria, there are two forms of
cell division, mitosis and meiosis.
a. Mitosis is the basis for division of the single-celled fertilized
egg, growth of an embryo, and all postnatal growth and tissue
repair.
b. It is essentially the splitting of a cell with a distribution of
chromosomes resulting in two genetically identical daughter cells.
c. Mitosis has four stages: prophase, metaphase, anaphase, and
telophase.
2. Mitosis and meiosis have important similarities and differences.

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a. In mitosis, each daughter cell has 46 chromosomes (23 pairs); in
meiosis, the chromosome number of daughter cells is reduced by
half (a reduction division).
i. In mitosis, the parent cell is diploid (2n), and so are the
daughter cells.
ii. In meiosis, the parent cell is diploid, but the daughter
cells are haploid (n) with 23 unpaired chromosomes.
b. In mitosis, the chromosomes do not change their genetic
makeup; in meiosis, the chromosomes of each homologous pair
join and exchange portions of their DNA, creating new gene
combinations.
c. In mitosis, each parent cell produces two daughter cells; in
meiosis, each parent cell produces four daughter cells.
i. Each male germ cell gives rise to four sperm.
ii. Female germ cell division is somewhat different.
3. Because sexual reproduction is biparental, meiosis is necessary so that
the zygote produced by union of two gametes has the correct chromosome
number.
4. The stages of meiosis are fundamentally the same in both sexes,
consisting of two cell divisions in succession with the following phases:
prophase I, metaphase I, anaphase I, telophase I, interkinesis, prophase II,
metaphase II, anaphase II, and telophase II. (Fig. 27.13)
a. In prophase I, each pair of homologous chromosomes line up
side by side and form a tetrad (four chromatids).
i. One chromosome of each tetrad is from the individuals
father and the other from the mother.
ii. The paternal and maternal chromosomes exchange
segments of DNA in a process called crossing-over.
b. After crossing-over, the chromosomes line up at the midline of
the cell in metaphase I.
c. They separate at anaphase I.
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d. The cell divides into two cells at telophase I.
e. The centromeres do not divide, and the chromatids do not
separate from each other at anaphase I. Each chromosome is still
double-stranded, but each daughter cell has only 23 chromosomes.
f. Meiosis II is like mitosis; the chromosomes line up at the equator
during metaphase II, the centromeres divide, and each
chromosome separates into two chromatids.
g. These chromatids are drawn to opposite poles of the cell during
anaphase II.
h. At the end of meiosis II, there are four haploid cells.
5. Fertilization combines the 23 chromosomes from the male parent with
23 chromosomes from the female parent to reestablish the diploid number
of 46 in the zygote.
C. Spermatogenesis involves meiosis. (pp. 10461047) (Fig. 27.14)
1. The first stem cells destined to become sperm are primordial germ cells.
a. These form in the yolk sac, a membrane associated with the
developing embryo.
b. In the fifth to sixth week of development, they crawl into the
embryo and colonize the gonadal ridges, where they become
spermatogonia.
2. Spermatogonia remain dormant throughout childhood, lying along the
periphery of the seminiferous tubule near the basement membrane, outside
the bloodtestis barrier (BTB).
3. At puberty, testosterone secretion reactivates the spermatogonia,
bringing on spermatogenesis.
4. The essential steps of spermatogenesis are as follows. (Fig. 27.14)
a. Spermatogonia divide by mitosis.
i. One daughter cell remains near the tubule wall as a stem
cell, a type A spermatogonium. Type A cells serve as a
lifetime supply of stem cells.

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Saladin Outline Ch.27

ii. The other daughter cell, a type B spermatogonium,
migrates slightly away from the wall.
b. The type B spermatogonium enlarges and becomes a primary
spermatocyte.
i. The tight junction between two sustentacular cells ahead
of the primary spermatocyte is broken down.
ii. A new tight junction forms behind the spermatocyte, so
that it is now protected by the BTB.
c. Now safely isolated from blood-borne antibodies, the primary
spermatocyte undergoes meiosis I, which gives rise to two haploid
and genetically unique secondary spermatocytes.
d. Each secondary spermatocyte undergoes meiosis II, dividing
into two spermatids.
e. A spermatid divides no further, but undergoes spermiogenesis,
in which it differentiates into a spermatozoon. (Fig. 27.15)
i. It sprouts a tail (flagellum) and discards most of its
cytoplasm.
ii. It will not move under its own power until ejaculation.
5. Each stage is a little closer to the lumen than the earlier stages.
a. Throughout meiotic divisions, the daughter cells remain
connected to each other by cytoplasmic bridges.
b. Eventually the mature sperm separate, depart from their
supportive sustentacular cells, and are washed down the tubule by
a slow flow of fluid.
c. It takes about 70 days for a type B spermatogonium to become a
mature spermatozoa.
d. 300,000 mature sperm are produced every minute, or 400
million per day, in a young male.
D. The spermatozoon has two parts: a pear-shaped head and a long tail. (p. 1047)
(Fig. 27.16)

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1. The head, about 4 to 5 m long and 3 m wide, has three structures: a
nucleus, an acrosome, and a flagellar basal body.
a. The nucleus is the most important of these, as it contains the
haploid set of chromosomes.
b. The acrosome is a lysosome in the form of a thin cap covering
the apical half of the nucleus; its enzymes later penetrate the egg.
c. The basal body of the tail flagellum is nestled in an indentation
at the basal end of the nucleus.
2. The tail is divided into three regions.
a. The midpiece is a cylinder about 5 to 9 m long and half as wide
as the head and contains numerous large mitochondria
i. These mitochondria coil tightly around the axoneme of
the flagellum.
ii. They produce ATP for the beating of the tail when the
sperm migrates up the female reproductive tract.
b. The principal piece, 40 to 45 m long, constitutes most of the
tail and consists of the axoneme surrounded by a sheath of
supportive fibers.
c. The endpiece, 4 to 5 m long, consists of the axoneme only and
is the narrowest part.
E. Semen or seminal fluid is the fluid expelled during orgasm. (pp. 10481049)
1. A typical ejaculation discharges 2 to 5 mL of semen, composed of about
10% sperm and spermatic duct secretions, 30% prostatic fluid, 60%
seminal vesicle fluid, and a trace of bulbourethral fluid.
a. Most of the sperm emerge in the first one or two jets of semen.
2. Semen usually has a sperm count of 50 to 120 million sperm/mL.
a. A sperm count any lower than 20 to 25 million sperm/mL is
usually associated with infertility. (Table 27.2)
3. The prostate and seminal vesicles contribute the following constituents
to semen.

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a. The prostate produces a thin, milky white fluid containing
calcium, citrate, and phosphate ions, a clotting enzyme, and a
protein-hydrolyzing enzyme called serine protease (prostatespecific antigen, PSA).
b. The seminal vesicles contribute a viscous yellowish fluid.
i. This is the last component of the semen to emerge.
ii. It contains fructose and other carbohydrates, citrate,
prostaglandins, and a protein called proseminogelin.
4. Semens stickiness promotes fertilization, and it arises when the clotting
enzyme activates proseminogelin, converting it to a sticky fibrinlike
protein, semenogelin.
a. Semenogelin entangles the sperm and sticks to the walls of the
inner vagina and cervix, ensuring that it does not simply drain back
out.
b. It may also promote the uptake of sperm-laden clots of semen
into the uterus.
5. Twenty to 30 minutes after ejaculation, serine protease in the prostatic
fluid breaks down semenogelin and liquefies the semen.
6. Two requirements must be met for sperm motility: an elevated pH and
an energy source.
a. The pH of the vagina is about 3.5 to 4.0, but the prostatic fluid
buffers the vaginal and seminal acidity, raising the pH to about 7.5.
b. Sperm use fructose and other sugars contributed by the seminal
vesicles to synthesize ATP for movement.
Insight 27.3 Reproductive Effects of Pollution
7. The sperm now become very active, thrashing their tails and crawling
up the mucosa of the vagina and uterus.
a. The prostaglandins of the semen may thin the mucus in the
cervical canal of the female and also may stimulate peristaltic
waves in the uterus and uterine tubes, helping to spread semen.

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V. Male Sexual Response (pp. 10491053)
A. Scientific studies of the physiology of sexual intercourse (coitus, coition, or
copulation) were hampered historically by repressive attitudes. Researchers
Masters and Johnson divided intercourse into four phases: excitement, plateau,
orgasm, and resolution. (p. 1049)
B. Part of the anatomical foundation of sexual response in the male has to do with
blood circulation and nerve supply to the penis. (p. 1049)
1. Each internal iliac artery gives rise to an internal pudendal (penile)
artery, which enters the root of the penis and divides in two.
a. The dorsal artery travels dorsally along the penis not far beneath
the skin, supplying skin, fascia, and corpus spongiosum.
(Fig. 27.11)
b. The deep artery travels through the core of the corpus
cavernosum and gives off smaller helicine arteries, which penetrate
the trabeculae and empty into the lacunae.
i. When the deep artery dilates, the lacunae fill with blood
and the penis becomes erect.
ii. When the penis is flaccid, most of its blood supply
comes from the dorsal arteries.
c. Numerous anastomoses are present between the two arteries.
d. A median deep dorsal vein drains blood from the penis. It runs
between the two arteries and empties into a plexus of prostatic
veins.
3. The penis is richly innervated by sensory and motor nerve fibers.
a. The glans has an abundance of tactile, pressure, and temperature
receptors, especially on its proximal margin and frenulum.
b. These receptors lead by way of a pair of dorsal nerves to the
internal pudendal nerves, then via the sacral plexus to segments S2
to S4 of the spinal cord.
c. Sensory fibers of the shaft, scrotum, perineum, and elsewhere
are also highly important to stimulation.
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4. Both autonomic and somatic motor fibers carry impulses from
integrating centers in the spinal cord to the penis and other pelvic organs.
a. Sympathetic fibers arise from T12 to L2, pass through the
hypogastric and pelvic plexuses, and innervate the penile arteries,
trabecular muscle, spermatic ducts, and accessory glands.
i. They dilate the penile arteries and can induce erection
even when the sacral region of the spinal cord is damaged.
ii. They also initiate erection in response to input to the
special senses and to sexual thoughts.
b. Parasympathetic fibers extend from S2 to S4 through the
pudendal nerves to the arteries of the penis. They are involved in
an autonomic reflex arc that causes erection in response to direct
stimulation of the penis and perineal region.
C. The excitement phase and plateau phases precede orgasm. (p. 1051)
1. The excitement phase is characterized by vasocongestion, myotonia,
and increases in heart rate, blood pressure, and pulmonary ventilation.
(Fig. 27.17)
a. The bulbourethral glands secrete fluid during this phase.
b. The excitement phase can be initiated by a broad spectrum of
erotic stimuli. Conversely, emotions can inhibit sexual response
when a person is anxious, stressed, or preoccupied.
c. The most obvious manifestation of male sexual excitement is
erection of the penis.
i. It is an autonomic reflex mediated predominantly by
parasympathetic nerve fibers that travel alongside the deep
and helicine arteries.
ii. These fibers trigger secretion of nitric oxide (NO), which
leads to relaxation of the deep arteries and lacunae.
iii. It is debated whether this relaxation causes erection
itself or also blocks outflow of blood from the penis.

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iv. Erection is aided by the bulbospongiosus and
ischiocavernosus muscles, which compress the root of the
penis, and the tunica albuginea, which is a tight, fibrous
sleeve of tissue.
d. As the corpora cavernosa expand, the penis becomes enlarged,
rigid, and elevated to an angle conducive to entry (intromission).
e. Once intromission is achieved, tactile and pressure sensations
further accentuate the erection reflex.
f. The corpus spongiosum swells and becomes visible as a cordlike
ridge along the ventral surface of the penis, but it does not become
as engorged and hardened as the corpora cavernosa.
g. The testes also become as much as 50% larger during
excitement.
2. In the plateau phase, respiratory rate, heart rate, and blood pressure are
sustained at a high level for a few seconds to a few minutes before
orgasm.
Insight 27.4 Treading Erectile Dysfunction (Fig. 27.18)
D. The orgasm, or climax, is a short but intense reaction that lasts 3 to 15 seconds
and is usually marked by the discharge of semen. (p. 1052)
1. Ejaculation is the most significant aspect from the standpoint of
producing offspring; it occurs in two stages, called emission and
expulsion.
2. In emission, the sympathetic nervous system stimulates peristalsis in the
smooth muscle of the ductus deferens, which propels sperm from the tail
of the epididymis, along the ductus, and into the ampulla.
a. Contractions of the ampulla propel the sperm into the prostatic
urethra.
b. Contractions of smooth muscle in the prostate gland force
prostatic fluid into the urethra.
c. Secretions of the seminal vesicles join the semen soon after the
prostatic secretion.
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Saladin Outline Ch.27

d. The contractions and seminal flow of this phase create an urgent
sensation that ejaculation is inevitable.
3. Semen in the urethra activates somatic and sympathetic reflexes that
result in expulsion.
a. Sensory signals travel to the spinal cord via the internal
pudendal nerve and reach an integrating center in the upper lumbar
region.
b. From here, sympathetic nerve fibers carry motor signals to the
prostate gland and seminal vesicles, causing smooth muscle
contraction to express more fluid into the urethra.
i. The sympathetic reflex also constricts the internal
urethral sphincter so urine cannot enter the urethra and
semen cannot enter the bladder.
c. Somatic motor signals leave the third and fourth sacral segments
of the cord and travel to the bulbospongiosus, ischiocavernosus,
and levator ani muscles.
i. The bulbospongiosus undergoes five or six strong,
spasmodic contractions that compress the urethra and
forcibly expel the semen.
ii. Most sperm are ejected in the first mL of semen, mixed
primarily with prostatic fluid.
iii. Seminal vesicle secretion follows and flushes most
remaining sperm from the ejaculatory ducts and urethra.
4. Some sperm may seep from the penis prior to ejaculation, and
pregnancy can therefore result from genital contact even without male
orgasm.
5. Orgasm is accompanied by an intense feeling of release from tension.
6. Ejaculation and orgasm are not the same, although they usually occur
together.
a. It is possible to have all of the sensations of orgasm without
ejaculation.
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Saladin Outline Ch.27

b. Ejaculation occasionally occurs with little or no sensation of
orgasm.
E. Immediately following orgasm is the resolution phase (p. 1052)
1. Discharge of the sympathetic nervous system constricts the internal
pudendal artery and reduces the flow of blood into the penis.
2. It also causes contraction of the trabecular muscles, which squeeze
blood from the lacunae.
3. The penis may remain semierect long enough to continue intercourse,
which may be important to the females attainment of climax.
4. Gradually, the penis undergoes detumescence, becoming soft and
flaccid again.
5. Cardiovascular and respiratory functions return to normal.
6. Resolution in men is followed by a refractory period, lasting anywhere
from 10 minutes to a few hours, during which it is usually impossible to
attain another erection and orgasm.
7. Two of the most common concerns related to sex are sexually
transmitted diseases (STDs) and contraception.
8. Reproductive disorders specific to males are described in Table 27.2.
Insight 27.5 Sexually Transmitted Diseases

Cross Reference
Additional information on topics mentioned in Chapter 27 can be found in the chapters
listed below.
Chapter 4: Mitosis
Chapter 10: Inguinal hernia
Chapter 17: Action of hormones
Chapter 28: Female reproductive system
Chapter 28: Oogenesis
Chapter 28: Contraception

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