UNiT 6 THE HUMAN LiFE CYCLE

22
Reproductive
Systems
LEARNiNG OUTCOMES
After you have studied this chapter, you should be able to:

22.1 Meiosis and Sex Cell Production

Several sperm approach an egg, the winners of a race that several hundred
million sperm began (1,500×). Only one sperm cell can fertilize the egg.
THE WHOLE
HOLE PiCTURE
The male and female reproductive systems are each connected
sets of organs and glands. Some of the reproductive organs and
glands secrete hormones vital to the development and main-
tenance of secondary sex characteristics and the regulation of
reproductive functions. Reproductive organs produce and nur-
ture sex cells (gametes) and transport them to sites of fertilization.
N •
AR
SS • LE
1 Outline the process of meiosis, and explain how it mixes
up parental genes. (p. 824)
22.2 Organs of the Male Reproductive System
2 Describe the structure and function(s) of each part of
the male reproductive system. (pp. 826–835)
3 Outline the process of spermatogenesis. (p. 828)
4 Describe semen production and exit from the body.
(pp. 828–836)
5 Explain how the tissues of the penis produce an erection.
(p. 835)
22.3 Hormonal Control of Male Reproductive Functions
6 Explain how hormones control the activities of the male
reproductive organs and the development of male
secondary sex characteristics. (pp. 837–839)
22.4 Organs of the Female Reproductive System
7 Describe the the structure and function(s) of each part of
the female reproductive system. (pp. 840–848)
8 Outline the process of oogenesis. (p. 841)
22.5 Hormonal Control of Female Reproductive
Functions
9 Explain how hormones control the activities of the female
reproductive organs and the development of female
secondary sex characteristics. (p. 849)
10 Describe the major events during a female reproductive
cycle. (pp. 850–852)
22.6 Mammary Glands
11 Review the structure of the mammary glands. (pp. 852–853)
22.7 Birth Control
12 Describe several methods of birth control, including the
relative effectiveness of each method. (pp. 854–858)
22.8 Sexually Transmitted infections
13 List the general symptoms of diseases associated with
sexually transmitted infections. (p. 859)
P

Module 14: Reproductive System

RA
CTIC
E

E
823
• ASS
 U N D E R S TA N D i N G W O R D S
andr-, man: androgens—male sex hormones.
contra-, against, counter: contraception—
prevention of fertilization.
crur-, lower part: crura—diverging parts
at the base of the penis by which it
attaches to the pelvic arch.
ejacul-, to shoot forth: ejaculation—expulsion
of semen from the male reproductive tract.
fimb-, fringe: fimbriae—irregular extensions
on the margin of the infundibulum of the
uterine tube.
follic-, small bag: follicle—ovarian structure
that contains an egg.
-genesis, origin: spermatogenesis—
formation of sperm cells.
gubern-, to steer, to guide: gubernaculum—
fibrous cord that guides the descent of
a testis.
labi-, lip: labia minora—flattened, longitudi-
nal folds that extend along the margins
of the female vestibule.
mamm-, breast: mammary gland—female
accessory gland that secretes milk.
mast-, breast: mastitis—inflammation of the
mammary gland.
mens-, month: menses—monthly flow of
blood from the female reproductive tract.
mons-, an eminence: mons pubis—rounded
elevation of fatty tissue overlying the
pubic symphysis in a female.
oo-, egg: oogenesis—formation of an egg.
prim-, first: primordial follicle—ovarian fol-
licle composed of an oocyte surrounded
by a single layer of cells.
puber-, adult: puberty—the time when a
person becomes able to reproduce.
zon-, belt: zona pellucida—transparent
layer surrounding an oocyte.

22.1
| Meiosis and Sex Cell
Production
Male sex cells are sperm. Female sex cells are eggs, or oocytes
(o′o-sı̄tz), which in Latin means “egg cells.” Sex cells have one set
of genetic instructions carried on 23 chromosomes, compared to
two sets on 46 chromosomes in other body cells. When sex cells
join at fertilization, the amount of genetic information held in
46 chromosomes is restored. Sperm and oocytes are produced by a
special type of cell division called meiosis.
A GLiMPSE AHEAD | To Chapter 24
a normal fertilized human egg contains two copies of each
gene, one from the sperm and one from the egg. The different
possible combinations of variants of these genes explain why
siblings are not exactly alike.
Meiosis (mi-o′sis) includes two successive divisions, called the
first and second meiotic divisions. At the beginning of the process
each diploid cell has two sets of chromosomes (46 chromosomes)
that are in the form of 23 homologous chromosome pairs. In the first
meiotic division (meiosis I) members of each homologous pair are
separated from each other. Homologous pairs are the same, gene for
gene. They may not be identical, however, because a gene may have
variants, and the chromosome that comes from the person’s mother
may carry a different variant for the corresponding gene from the
father’s homologous chromosome. Before meiosis I, each chromo-
some is replicated, so it consists of two DNA molecules called
chromatids. Each chromatid has the complete genetic information
associated with that chromosome. The chromatids of a replicated
chromosome attach at regions called centromeres.
Each of the cells emerges from meiosis I with one member of
each homologous pair, a condition termed haploid. That is, a hap-
loid cell has one set of chromosomes (23 chromosomes, one from
each homologous pair). The second meiotic division (meiosis II)
separates the chromatids, producing cells that are still haploid, but
whose chromosomes are no longer in the replicated form. After
meiosis II, each of the chromatids is an independent chromosome.
The steps of meiosis are clearer when considered in a time
sequence (fig. 22.1). However, keep in mind that meiosis, like mito-
sis, is a continuous process. Considering it in steps simply makes it
easier to follow.
First Meiotic Division
Prophase I. Individual chromosomes appear as thin threads in
the nucleus that then shorten and thicken. Nucleoli disappear, the
nuclear membrane temporarily disassembles, and microtubules
begin to build the spindle that will separate the chromosomes.
The DNA of the chromosomes has already been replicated.
As prophase I continues, homologous chromosomes pair up
side by side and tightly intertwine. During this pairing, called syn-
apsis, the chromatids of the homologous chromosomes touch at
various points along their lengths. Often, the chromatids break in
one or more places and exchange parts, forming chromatids with
new combinations of genetic information (fig. 22.2). One chromo-
some of a homologous pair is maternal and the other is paternal.
Therefore, an exchange, or crossover, between homologous chro-
mosomes produces chromatids that contain genetic information
from both parents.
Metaphase I. During the first metaphase, chromosome pairs line
up about midway between the poles of the developing spindle, and
they are held under great tension, like two groups of people playing
tug-of-war. Each chromosome pair consists of two chromosomes,
which equals four chromatids. Each chromosome attaches to spin-
dle fibers from one pole. The chromosome alignment is random
with respect to maternal and paternal origin of the chromosomes.
Each of the 23 chromosomes contributed from the mother may be
on the left or the right, and the same is true for the paternal chromo-
somes—it is similar to the number of ways that 23 pairs of children
could line up, while maintaining the pairs. Chromosomes can line
up with respect to each other in many combinations.
Anaphase I. Homologous chromosome pairs separate, and each
replicated member moves to one end of the spindle. Each new, or
daughter, cell receives only one replicated member of a homologous
pair of chromosomes, overall halving the chromosome number.

824 UNiT 6 | ThE humaN LifE CyCLE


First meiotic
division
(23 chromosomes, each
with 2 chromatids)
Paired homologous
chromosomes
(46 chromosomes,
each with 2 chromatids)
(23 chromosomes, each
with 2 chromatids)
FiGURE 22.1 Sex cells are formed by a special type of cell division, meiosis. This illustration follows two representative pairs of
homologous chromosomes.
Telophase I. The original cell divides in two. Nuclear mem-
branes form around the chromosomes, nucleoli reappear, and the
spindle fibers disassemble into their constituent microtubules.
Second Meiotic Division
After telophase I, the second meiotic division begins. Meiosis II
is similar to a mitotic division (see fig. 22.1). During prophase II,
chromosomes condense and reappear, still replicated. They move
into positions midway between the poles of the developing spin-
dle. In metaphase II, the replicated chromosomes attach to spindle
fibers. In anaphase II, centromeres separate, freeing the chroma-
tids to move to opposite poles of the spindles. The former chroma-
tids are now considered to be chromosomes. In telophase II, each
of the two cells resulting from meiosis I divides to form two cells.
Therefore, each cell undergoing meiosis has the potential to pro-
duce four gametes. In males, the gametes mature into four sperm
cells. In females, three of the products of meiosis are “cast aside”
as polar bodies, and one cell becomes the egg.
Meiosis generates astounding genetic variety. Any one of a per-
son’s more than 8 million possible combinations of 23 chromosomes
can combine with any one of the more than 8 million combinations
of his or her mate, raising the potential variability to more than
70 trillion genetically unique individuals! Crossing over contributes
even more genetic variability. Figure 22.3 illustrates in a simplified
manner how maternal and paternal traits reassort during meiosis.
Second meiotic
division
(23 chromosomes,
each chromatid now an
independent chromosome)
CAREER CORNER
Nurse-Midwife
a couple chooses a certified nurse-midwife (CNm) to deliver
their third child because they prefer a home birth. Their first
two children had been delivered in a hospital. The children
are healthy and the births were uncomplicated.
The nurse-midwife meets with the couple throughout
the pregnancy, for at least an hour each time, and is present
for the entire labor and birth. She uses soothing words and
gentle massage to ease the baby into the world. The nurse-
midwife places the newborn on the mother’s chest while she
completes the initial examination.
a CNm provides pregnancy care, but can also provide
primary care, gynecologic care, family planning, treatment
for sexually transmitted infections, and newborn care for the
first month of life. She or he can do physical exams, prescribe
medications, and order and interpret medical tests to provide
a diagnosis.
a nurse-midwife is a registered nurse who has a graduate
degree in midwifery from an accredited program and has
passed a certification or licensing exam. many nurse-midwives
deliver babies in patients’ homes, but some are affiliated with
hospitals. Nurse-midwives work with physicians who can help
in case of emergency.
CHAPTER 22 | Reproductive Systems 825

(a) (b) (c)
FiGURE 22.2 Crossing over mixes up genetic traits.
(a) homologous chromosome pair, (b) chromatids cross over, (c) crossing
over recombines genes. The different colors indicate that one of the
homologous chromosomes comes from the individual’s father and one
from the mother.
PRACTiCE
1 What are the male and female sex cells called?
2 Describe the major events of meiosis.
3 how does meiosis provide genetic variability?
22.2
| Organs of the Male
Reproductive System
Organs of the male reproductive system are specialized to pro-
duce and maintain the male sex cells, or sperm cells; transport
these cells and supporting fluids to the outside; and secrete male
sex hormones.
The primary sex organs (gonads) of this system are the two
testes in which the sperm cells (spermatozoa) and the male sex
hormones are formed. The other structures of the male repro-
ductive system are termed accessory sex organs (secondary sex
organs). They include the internal reproductive organs and the
external reproductive organs (fig. 22.4; reference plates 3 and 4,
pp. 40–41).
Testes
The testes (tes′teˉz; sing., testis) are ovoid structures about 5 cen-
timeters in length and 3 centimeters in diameter. Both testes, each
suspended by a spermatic cord, are within the cavity of the saclike
scrotum (fig. 22.4 and reference plate 12, p. 48).
Descent of the Testes
In a male fetus, the testes originate from masses of tissue posterior
to the parietal peritoneum, near the developing kidneys. Usually a
month or two before birth, the testes descend to the lower abdomi-
nal cavity and pass through the abdominal wall into the scrotum.
The male sex hormone testosterone, which the developing tes-
tes secrete, stimulates the testes to descend. A fibrous cord called
the gubernaculum (goo″ber′-nak′u-lum) is attached to each devel-
oping testis and extends into the inguinal region of the abdominal
cavity. The gubernaculum passes through the abdominal wall and
is fastened to the skin of the scrotum. As the body grows the testis
826 UNiT 6 | ThE humaN LifE CyCLE
Parent cell Paternal
chromatids
Gene for
blood type
Gene for
Maternal eye color
chromatids
Gene for
hair color
Result of
crossing
over
FiGURE 22.3 as a result of crossing over, the genetic information
in sex cells varies from cell to cell. Colors represent parent of origin.
although only one eye color gene pair is illustrated, eye color is polygenic
(involving more than one gene pair).
remains anchored to the gubernaculum, and guided by the guber-
naculum, the testis descends through the inguinal canal (ing′gwı̆-
nal kah-nal′) of the abdominal wall and enters the scrotum, where
it remains. Each testis carries a developing ductus (vas) deferens,
blood vessels, and nerves. These structures later in development
form parts of the spermatic cord that suspends the testis in the
scrotum (fig. 22.5).
If the testes do not descend into the scrotum, they cannot
produce sperm cells because the temperature in the abdominal
cavity is too high. If this condition, called cryptorchidism, is left
untreated, the cells that normally produce sperm cells degener-
ate, causing infertility.
PRACTiCE
4 What are the primary sex organs of the male reproductive system?
5 Describe the descent of the testes.
as a testis descends, a pouch of peritoneum, called the vaginal
process, moves through the inguinal canal and into the scro-
tum. in about 25% of males, this pouch stays open, providing
a potential passageway through which a loop of intestine may
be forced by great abdominal pressure, producing an indirect
inguinal hernia. if the protruding intestinal loop is so tightly
constricted within the inguinal canal that its blood supply stops,
the condition is called a strangulated hernia. Without prompt
treatment, the blood-deprived tissues may die.
 Ureter

Urinary bladder
Large intestine

Superior pubic
ramus (cut)
Seminal vesicle
Ductus
(vas) deferens Ejaculatory duct

Prostate gland

Bulbourethral
Urethra gland

Corpus cavernosum Urogenital

Corpus spongiosum diaphragm

Penis Anus

Epididymis
Glans penis
Testis
Prepuce
Scrotum

(a)

Ureter

Urinary bladder
Ampulla
Seminal vesicle

Ejaculatory duct
Prostate gland
Bulbourethral
gland
Bulb of
penis Ductus (vas) deferens
Root of
Crus of penis
penis

Epididymis

Testis

Penis

Urethra Body of
penis
Glans penis

(b)

FiGURE 22.4 male reproductive organs. (a) Sagittal view and (b) posterior view. The paired testes are the primary sex organs, and the other
reproductive structures, both internal and external, are accessory sex organs.

CHAPTER 22 | Reproductive Systems 827

 The seminiferous tubules are lined with a specialized strati-
Abdominal
wall fied epithelium, which includes the spermatogenic cells that give
rise to the sperm cells. Other specialized cells, called interstitial
Testis (in″ter-stish′al) cells (cells of Leydig), lie between the seminifer-
Lower
ous tubules (fig. 22.6 and fig. 22.7). Interstitial cells produce and
Rectum secrete male sex hormones.
abdominal
cavity
Gubernaculum

Developing Pubic
penis symphysis The epithelial cells of the seminiferous tubules can give rise
to testicular cancer, a common cancer in young men. in most
(a) cases, the first sign is a painless testis enlargement or a small
lump or area of hardness on the testis. if a biopsy (tissue sam-
Peritoneum ple) reveals cancer cells, surgery is performed to remove the
affected testis (orchiectomy). Radiation and/or chemotherapy
in many cases prevents the cancer from recurring.

Testis
Vaginal Inguinal canal PRACTiCE
process
(cavity) Gubernaculum 6 Where in the testes are sperm cells produced?
7 Which cells produce male sex hormones?
(b)

Formation of Sperm Cells

Ductus deferens
Tunica Spermatic cord
vaginalis
Testis
Scrotum
Gubernaculum
(c)
FiGURE 22.5 During fetal development, each testis develops near
a kidney and then descends through an inguinal canal and enters the
scrotum, completing the journey by the eighth gestational month (a–c).
Structure of the Testes
A tough, white, fibrous capsule called the tunica albuginea
encloses each testis. Along the capsule’s posterior border, the con-
nective tissue thickens and extends into the organ, forming a mass
called the mediastinum testis. From this structure, thin layers of
connective tissue, called septa, pass into the testis and subdivide it
into about 250 lobules.
A lobule contains one to four highly coiled, convoluted semi-
niferous tubules (sem″ ı̆-nif′er-us tu′būlz), each approximately 70
centimeters long when uncoiled. These tubules course posteriorly
and unite to form a complex network of channels called the rete
testis (re′te tes′tis). The rete testis is in the mediastinum testis and
gives rise to several ducts that join a tube called the epididymis.
The epididymis, in turn, is coiled on the outer surface of the testis
and continues to become the ductus deferens (figs. 22.6).
The epithelium of the seminiferous tubules consists of supporting
cells called sustentacular cells (Sertoli cells) and spermatogenic
cells (fig. 22.7). The sustentacular cells are columnar and extend
the full thickness of the epithelium, from its base to the lumen of
the seminiferous tubule. The sustentacular cells support, nourish,
and regulate the spermatogenic cells, which give rise to sperm
cells (spermatozoa).
In the male embryo, undifferentiated spermatogenic cells
are called spermatogonia (fig. 22.7). Each spermatogonium has
46 chromosomes (23 pairs) in its nucleus, the number for most
human body (somatic) cells. Hormones stimulate the spermato-
gonia to become active. Some of the cells undergo mitosis (see
chapter 3, pp. 108–110). Mitotic cell division gives rise to two
new cells, with each of these new cells containing 46 chromo-
somes. One cell (type A) maintains the supply of undifferentiated
cells, the other cell (type B) differentiates to become a primary
spermatocyte. Sperm production arrests at this stage.
At puberty, mitosis resumes, and new spermatogonia form.
Testosterone secretion increases, and the primary spermatocytes
divide by meiosis, each forming two secondary spermatocytes.
Each of these cells divides to form two spermatids, which mature
into sperm cells. Meiosis halves the number of chromosomes
in each cell (fig. 22.8). Spermiogenesis is the further develop-
ment of spermatids into sperm (see fig. 22.10). The combined
processes of meiosis and spermiogenesis constitute spermato-
genesis (sper″mah-to-jen′ĕ-sis), taking about 65 to 75 days from
start to finish.
The spermatogonia are located within the seminiferous
tubules, adjacent to the inside surface of the surrounding basement
membrane. As spermatogenesis proceeds, cells in more advanced
stages are pushed along the sides of sustentacular cells toward the
lumen of the seminiferous tubule (fig. 22.9).

828 UNiT 6 | ThE humaN LifE CyCLE

 Epididymis Ductus deferens

Rete testis

Tunica vaginalis

Plane of section

Lumen of
(a) seminiferous
Tunica tubule
albuginea
Testis Basement
membrane
Seminiferous tubules
Sperm cells
Interstitial cells
Spermatogonia

FIGURE 22.6 Structure of the testis. (a) Sagittal

section of a testis. (b) Cross section of a seminiferous tubule. (b)

Basement
Spermatogenesis happens continually in a male, starting at
membrane puberty. The resulting sperm cells collect in the lumen of each
Sustentacular seminiferous tubule, then pass through the rete testis to the epi-
cells didymis, where they accumulate and mature.
(Sertoli cells)
Seminiferous Spermatogonia Structure of a Sperm Cell
tubule Sperm cells A mature sperm cell is a tiny, tadpole-shaped structure about
0.06 millimeter long. It consists of a flattened head, a cylindrical
Interstitial cells midpiece (body), and an elongated tail (flagellum).
(cells of Leydig) The oval head of a sperm cell is primarily composed of a
nucleus and contains highly compacted chromatin consisting of 23
chromosomes. A small caplike covering over the head, called the
FIGURE 22.7 Light micrograph of a seminiferous tubule (250×). acrosome, contains enzymes that aid the sperm cell in penetrating
the layers surrounding the oocyte during fertilization (fig. 22.10). An
enzyme on the sperm cell membrane also contributes to this process.
Near the base of the epithelium, tight junctions fuse the The midpiece of a sperm has a central, filamentous core and
membranous processes of adjacent sustentacular cells (fig. 22.9). many mitochondria organized in a spiral. The tail consists of sev-
These tightly packed cells and their attachments form the blood- eral microtubules enclosed in an extension of the cell membrane.
testis barrier, which prevents some substances from reaching the The mitochondria provide ATP for the lashing movement of the
developing sperm. The blood-testis barrier helps maintain a favor- tail that propels the sperm cell. The micrograph in figure 22.11
able environment by isolating developing sperm from the male’s shows a few mature sperm cells.
immune system, which might otherwise react to the sperm as
abnormal, nonself cells.
Many toxic chemicals that affect sperm hamper the tail’s abil-
ity to propel them, preventing the sperm from reaching an
Anton van Leeuwenhoek was the first to view human sperm egg and exposing it to the chemical. An exception is cocaine,
under a microscope in 1678, concluding that they were para- which attaches to thousands of binding sites on a human sperm
sites in semen. By 1685, he had modified his view, writing that cell, without apparently harming the cell or impeding its abil-
sperm contain a preformed human being and are seeds requiring ity to ferry cocaine to an egg. However, it is not known what
nurturing in a female to start a new life. Although his interpre- harm, if any, the drug causes. We do know that fetuses exposed
tation was incorrect, he did identify sperm as playing a role in to cocaine in the uterus may suffer a stroke, or, as infants, be
human reproduction. unable to react normally to their surroundings.

CHAPTER 22 | Reproductive Systems 829

 Secondary Second meiotic
spermatocyte division

Spermatids Sperm cells

First meiotic
Primary division
spermatocyte (23 chromosomes, each
with 2 chromatids)

Paired homologous
chromosomes

(46 chromosomes,
each with 2 chromatids)

(23 chromosomes, each

with 2 chromatids) (23 chromosomes,
each chromatid now an
independent chromosome)
FIGURE 22.8 During spermatogenesis four sperm cells result from meiosis of a primary spermatocyte.
Two representative homologous chromosome pairs are shown.

Q Why is it important that a sperm possess only 23 chromosomes?

Answer can be found in Appendix G.

PRACTICE The inner lining of the epididymis is composed of pseu-

8 Review the events of spermatogenesis.
9 Explain the function of the sustentacular cells in the
seminiferous tubules.
10 Describe the structure of a sperm cell.
Male Internal Accessory Reproductive
Organs
The internal accessory organs of the male reproductive system are
specialized to nurture and transport sperm cells. These structures
include the two epididymides, two ductus deferentia, two ejacula-
tory ducts, the urethra, as well as the two seminal vesicles, prostate
gland, and two bulbourethral glands.
Epididymides
The epididymides (ep″ı̆-di-dy′mides; sing., epididymis) are tightly
coiled, threadlike tubes about 6 meters long (see fig. 22.4 and refer-
ence plate 12, p. 48). Each epididymis is connected to ducts in a tes-
tis. It emerges from the top of the testis, descends along its posterior
surface, and then courses upward to become the ductus deferens.
dostratified columnar cells that bear nonmotile cilia (fig. 22.12).
These cells secrete glycogen and other substances that support
stored sperm cells and promote their maturation.
When immature sperm cells reach the epididymis, they are
nonmotile. However, as they travel through the epididymis as a
result of rhythmic peristaltic contractions, they mature. Following
this aging process, the sperm cells can move independently and
fertilize egg cells. However, they usually do not move indepen-
dently until after ejaculation.
Ductus Deferentia
The ductus deferentia (duk′tus def′er-en′sha; sing., ductus def-
erens), also called vasa deferentia, are muscular tubes about
45 centimeters long lined with pseudostratified columnar epithe-
lium (fig. 22.13). Each ductus deferens originates at the lower end
of the epididymis and passes upward along the medial side of a
testis to become part of the spermatic cord. The ductus deferens
passes through the inguinal canal, enters the abdominal cavity
outside the parietal peritoneum, and courses over the pelvic brim.
From there, it extends backward and medially into the pelvic cavity,
where it ends behind the urinary bladder.

830 UNIT 6 | THE HUMAN LIFE CYCLE

 Lumen of seminiferous
tubule

Changes in
chromosome
structure

Maturing spermatid
(23 chromosomes,
1 chromatid per chromosome)

Sustentacular cells

Nucleus of
sustentacular cell

Spermatid

Developmental sequence
(23 chromosomes, 1
chromatid per chromosome)

Meiosis II
Secondary spermatocyte
(23 chromosomes, 2
chromatids per chromosome)
Wall of seminiferous

Primary spermatocyte
Meiosis I (46 chromosomes, 2
tubule

chromatids per chromosome)

Tight junction
between sustentacular
cells (blood-testis barrier)

Daughter cell in late

interphase (Type B
spermatogonium,
46 chromosomes, 2
chromatids per chromosome)
Spermatogonium
mitosis Daughter cell in late
interphase (New type
A spermatogonium,
46 chromosomes, 2
chromatids per chromosome)

Basement
membrane

FiGURE 22.9 mitosis in spermatogonia results in type a spermatogonia that continue the germ cell line and type B spermatogonia that give
rise to primary spermatocytes. The primary spermatocytes, in turn, give rise to sperm cells by meiosis. Changes in chromosome number and structure are
represented by a single pair of chromosomes. Note that as the cells approach the lumen they mature.

CHAPTER 22 | Reproductive Systems 831

 Flagellum
Nucleus

Mitochondria

Golgi apparatus

Excess
cytoplasm

Excess cytoplasm
and most organelles lost FiGURE 22.11 human sperm cells (1,400×).

Tail

Epithelial
cells
Mitochondria
Midpiece
Centriole
Nonmotile
cilia

Head
Sperm cells

Acrosome
(a)

Acrosome
Head
Nucleus

Midpiece
(with mitochondria)

Tail

FiGURE 22.12 Cross section of a human epididymis (200×).

(b)

FiGURE 22.10 Spermiogenesis—sperm cell maturation. (a) The head of

the sperm develops largely from the nucleus of the formative cell. (b) Parts of
a mature sperm cell.

832 UNiT 6 | ThE humaN LifE CyCLE


Lumen
Epithelium
Smooth
muscle
(a)
Sperm in lumen of
ductus deferens
Pseudostratified
columnar epithelium
Smooth muscle
(b)
FiGURE 22.13 Ductus (vas) deferens. (a) micrograph of a
cross section of the ductus deferens (40×). (b) Light micrograph of the wall
of the ductus deferens (400×).
Near its termination, the ductus deferens dilates into a portion
called the ampulla. Just outside the prostate gland, the tube becomes
slender again and unites with the duct of a seminal vesicle. The
fusion of these two ducts forms an ejaculatory duct, which passes
through the prostate gland and empties into the urethra through a slit-
like opening (see fig. 22.4).
Seminal Vesicles
The seminal vesicles (or seminal glands) are convoluted, saclike
structures about 5 centimeters long, each attached to the ductus
deferens on the posterior surface and near the base of the urinary
bladder (see fig. 22.4). The glandular tissue lining the inner wall
of the seminal vesicle secretes a slightly alkaline fluid. This fluid
helps regulate the pH of the tubular contents as sperm cells travel
to the outside. Additionally, seminal vesicle fluid neutralizes the
acidic secretions of the vagina, helping to sustain the sperm cells
that enter the female reproductive tract. Seminal vesicle secretions
also include fructose, a monosaccharide that provides energy to
the sperm cells, and prostaglandins, which stimulate muscular
contractions of the female reproductive organs, aiding the move-
ment of sperm cells toward the egg cell.
As sperm move through the ductus deferens into the ejacu-
latory duct, the contents of the seminal vesicles empty into the
ejaculatory ducts. This greatly increases the volume of the fluid
discharged from the ductus deferens.
PRACTiCE
11 Describe the structure of the epididymis.
12 Trace the path of the ductus deferens.
13 What is the function of a seminal vesicle?
Prostate Gland
The prostate (pros′tāt) gland (see figs. 22.4 and 22.14) is a
chestnut-shaped structure about 4 centimeters across and 3 cen-
timeters thick that surrounds the proximal portion of the ure-
thra, just inferior to the urinary bladder. It is composed of many
branched tubular glands enclosed in connective tissue. Septa of
connective tissue and smooth muscle extend inward from the cap-
sule, separating the tubular glands. The ducts of these glands open
into the urethra. The prostate gland releases its secretions into the
urethra as smooth muscle contracts in its capsular wall. As this
release occurs, the contents of the ductus deferens and the seminal
vesicles enter the urethra, which increases the volume of the fluid.
The prostate gland secretes a thin, milky fluid. This slightly
acidic secretion also contains citrate, a nutrient for sperm, and
prostate-specific antigen (PSA), an enzyme which helps liquefy
semen following ejaculation. Clinical Application 22.1 discusses
prostate cancer.
Bulbourethral Glands
The bulbourethral (bul″bo-u-re′thral) glands (Cowper’s glands)
are two small structures, each about a centimeter in diameter.
They are inferior to the prostate gland lateral to the intermediate
part of the urethra and are enclosed by muscle fibers of the uro-
genital diaphragm (see fig. 22.4).
The bulbourethral glands are composed of many tubes whose
epithelial linings secrete a mucuslike fluid. This fluid is released in
response to sexual stimulation and lubricates the end of the penis
in preparation for sexual intercourse (coitus). However, females
secrete most of the lubricating fluid for intercourse.
Secretory cells of the
prostate gland
Smooth muscle
Lumen of urethra
FiGURE 22.14 Light micrograph of a cross section through the
prostate gland (10×).
CHAPTER 22 | Reproductive Systems 833
 C L i N i C A L A P P L i C AT i O N 22.1
Prostate Cancer
Each year in the united States about 240,000 men are diagnosed
with prostate cancer, and about 30,000 die of the disease. how-
ever, many prostate tumors grow so slowly that they are not likely
to affect survival and do not require treatment.
The diagnostic process typically begins with a digital rectal
exam, in which a physician inserts a finger into the patient’s rec-
tum to feel for an enlargement of the prostate gland. This exam is
typically coupled with a blood test to detect elevated levels of a
biomarker called prostate-specific antigen (PSa). Normally, secre-
tory epithelium in the prostate gland releases PSa, which liquefies
the ejaculate. When cancer cells accumulate, more PSa is pro-
duced, and it enters capillaries in the prostate.
health-care organizations’ recommendations for PSa screen-
ing change often, and range from screening all men to screening
none, although any man with symptoms of an enlarged prostate
(frequent and slowed urination) should be tested. The contro-
versy is that the value of saving lives following screening must be
balanced against the risk that high levels of PSa in the absence of
cancer can lead to unnecessary biopsies.
if the physician feels an enlargement of the patient’s prostate,
or if the PSa level remains high or rises on tests repeated a few
months later, the next step is a biopsy procedure to sample cells
Semen
The fluid the urethra conveys to the outside during ejaculation is
called semen (se′men). It consists of sperm cells from the testes
and secretions of the seminal vesicles, prostate gland, and bulbo-
urethral glands. Semen is slightly alkaline (pH about 7.5), and it
includes prostaglandins and nutrients.
The volume of semen released at one time varies from 2 to 5
milliliters. The average number of sperm cells in the fluid is about
120 million per milliliter.
Sperm cells remain nonmotile while they are in the ducts of the
testis and epididymis, but begin to swim as they mix with the secre-
tions of accessory glands. However, sperm cells cannot fertilize an
egg cell until they enter the female reproductive tract, because here
they undergo capacitation, which weakens the acrosomal mem-
branes. When sperm cells are placed with egg cells in a laboratory
dish to achieve fertilization—a technique called in vitro fertilization,
discussed in From Science to Technology 23.1 (pp. 870–871)—
chemicals are added to simulate capacitation.
Although sperm cells can live for many weeks in the ducts of the
male reproductive tract, they usually survive only up to six days after
being expelled to the outside, even when they are maintained at body
temperature. The ability of a sperm cell to fertilize an oocyte gener-
ally lasts only twenty-four to forty-eight hours after the sperm enter
the female reproductive tract. On the other hand, sperm cells can be
stored and kept viable for years if they are frozen at a temperature
below –100°C. Clinical Application 22.2 describes some causes of
male infertility.
834 UNiT 6 | ThE humaN LifE CyCLE
from several sites in the gland. a cancer detected with a biopsy is
assigned a two-digit number, called a Gleason score, which indi-
cates how specialized the cancer cells are. The less specialized
the cells, the more aggressive the disease. imaging technologies
can then assess whether the cancer has spread beyond the pros-
tate capsule.
Treatment of prostate cancer may be necessary if the tumor
has a high Gleason score or fits the genetic profile of being likely
to spread. Treatments include surgery to remove the prostate
gland, radiation, and hormones. adverse effects of treatment
include urinary incontinence and erectile dysfunction, which may
improve with time. fortunately, for many men “active surveil-
lance” to regularly monitor the disease is sufficient. This means
PSa tests twice a year and a biopsy every one to two years, with
treatment only if the condition worsens.
New tests measure the expression of specific genes that
change activity dramatically when the disease spreads. in one large
study that evaluated a gene expression test, doctors switched
37% of men being treated to active surveillance and 23% on active
surveillance to treatment based on test results. it will take a few
years to determine how well gene expression profiling improves
selection of patients for treatment.
PRACTiCE
14 Where is the prostate gland located?
15 What is the function of the bulbourethral glands?
16 What are the components of semen?
Male External Reproductive Organs
The male external reproductive organs are the scrotum, which encloses
two testes, and the penis. The urethra passes through the penis.
Scrotum
The scrotum is a pouch of skin and subcutaneous tissue that hangs
from the lower abdominal region posterior to the penis. The sub-
cutaneous tissue of the scrotal wall lacks fat but contains a layer of
smooth muscle fibers that constitute the dartos muscle. Exposure
to cold stimulates these muscle fibers to contract, the scrotal
skin to wrinkle, and the testes to move closer to the body, where
they can absorb heat. In warmer temperatures, the dartos muscle
relaxes and the scrotum hangs loosely. The testes move away from
the body into an environment 3°C (about 5°F) below body tem-
perature, more conducive to sperm production and survival.
A medial septum divides the scrotum into two chambers,
each of which encloses a testis. Each chamber also contains a
serous membrane, which covers the front and sides of the testis
and the epididymis, helping to ensure that the testis and epididy-
mis move smoothly within the scrotum (see fig. 22.4).
Penis
The penis is a cylindrical organ that conveys urine and semen through
the urethra to the outside. It is also specialized to enlarge and stiffen,
which enables it to enter the vagina during sexual intercourse.
The body, or shaft, of the penis is composed of three columns
of erectile tissue, which include a pair of dorsally located corpora
cavernosa and a single, ventral corpus spongiosum. A tough cap-
sule of white dense connective tissue called a tunica albuginea
(too′nı̆-kah al″bu-jin′e-ah) surrounds each column. Skin, a thin
layer of subcutaneous tissue, and a layer of connective tissue
enclose the penis (fig. 22.15).
The corpus spongiosum, surrounding the urethra, enlarges at
its distal end to form a sensitive, cone-shaped glans penis. The
glans covers the ends of the corpora cavernosa and bears the ure-
thral opening—the external urethral orifice. The skin of the glans
is very thin, hairless, and contains sensory receptors for sexual
stimulation. A loose fold of skin called the prepuce (foreskin)
originates just posterior to the glans and extends anteriorly to
cover the glans as a sheath. The prepuce can be removed by a sur-
gical procedure called circumcision.
At the root of the penis, the columns of erectile tissue sepa-
rate. The corpora cavernosa diverge laterally in the perineum and
are firmly attached to the inferior surface of the pubic arch by con-
nective tissue. These diverging parts form the crura (sing., crus) of
the penis. The single corpus spongiosum is enlarged between the
crura as the bulb of the penis, which is attached to membranes of
the perineum (see fig. 22.4b).
PRACTiCE
17 Describe the structure of the penis.
18 What is circumcision?
Erection, Orgasm, and Ejaculation
During sexual stimulation, parasympathetic impulses from the
sacral portion of the spinal cord release the vasodilator nitric
oxide, which dilates the arteries leading into the penis, increasing
Skin
Subcutaneous
tissue
Connective tissue
(fascia)
External urethral orifice
(a)
blood flow into erectile tissues. At the same time, the increas-
ing pressure of arterial blood entering the vascular spaces of the
erectile tissue compresses the veins of the penis, reducing flow
of venous blood away from the penis. Consequently, blood accu-
mulates in the erectile tissues, and the penis swells and elongates,
producing an erection (fig. 22.16).
in erectile dysfunction (impotence), the penis cannot become
erect or sustain an erection. Causes of erectile dysfunction
include underlying disease such as diabetes mellitus; paraly-
sis; treatments such as prostate surgery or certain drugs;
smoking cigarettes; and drinking alcohol. Development of
drugs to treat erectile dysfunction grew out of understand-
ing the physiology of erection. The first drug, Viagra (silde-
nafil), blocks the enzyme that breaks down cyclic guanosine
monophosphate in the erectile tissues, which is necessary for
an erection to persist.
The culmination of sexual stimulation is orgasm (or′gazm),
a pleasurable feeling of physiological and psychological release.
Orgasm in the male is accompanied by emission and ejaculation.
Emission (e-mish′un) is the movement of sperm cells from
the testes and secretions from the prostate gland and seminal ves-
icles into the urethra, where they mix to form semen. Emission is
a response to sympathetic impulses from the spinal cord, which
stimulate peristaltic contractions in smooth muscle in the walls of
the testicular ducts, epididymides, ductus deferentia, and ejacula-
tory ducts. Other sympathetic impulses stimulate rhythmic con-
tractions of the seminal vesicles and prostate gland.
As the urethra fills with semen, sensory impulses are stimu-
lated and pass into the sacral part of the spinal cord. In response,
motor impulses are conducted from the spinal cord to certain
skeletal muscles at the base of the erectile columns of the penis,
rhythmically contracting them. This movement increases the pres-
sure in the erectile tissues and aids in forcing the semen through
Superficial dorsal vein
Deep dorsal vein
Dorsal nerve
Dorsal artery
Deep artery
Corpora cavernosa
Tunica albuginea
Urethra
Corpus spongiosum
Prepuce
Glans penis FiGURE 22.15 Structure
of the penis. (a) interior and (b) cross
(b) section of the penis.
CHAPTER 22 | Reproductive Systems 835
 C L i N i C A L A P P L i C AT i O N 22.2
Male infertility

male infertility—the inability of sperm cells to fertilize an egg man’s sperm output that are the lowest for which he is still likely to
cell—has several causes. if, during fetal development, the tes- be fertile. it is clear that the male body manufactures many more
tes do not descend into the scrotum, the higher temperature of sperm than are necessary to fertilize an egg.
the abdominal cavity or inguinal canal impedes development of
sperm cells in the seminiferous tubules. Certain diseases, such as
mumps, may inflame the testes (orchitis), impairing fertility.
The quality and quantity of sperm cells are essential factors in
FiGURE 22A Curvilinear
Computer analysis path VCL
the ability of a man to father a child. if a sperm head is misshapen,
improves the
if the acrosome is too tough to burst and release enzymes, or if consistency Average P
too few sperm cells reach the well-protected egg, fertilization may and accuracy of path ALH VA
not happen. The structure of the sperm tail is particularly impor- describing sperm
tant. male infertility can result from sperm tails that are irregularly density, motility, BCF
shaped, coiled, bent, shortened, or absent. and morphology, VSL
Computer-aided sperm analysis (CaSa) is a technique used each important
to evaluate a man’s fertility. it can analyze the pathways of up to in diagnosing
200 moving sperm in a few seconds. CaSa assesses the number male infertility.
of cells per milliliter of seminal fluid (density), sperm motility, and Computer monitor/
the size and shape of sperm cell parts (morphology). data display Digital image
in a sperm analysis, a man abstains from intercourse for two to
three days, then provides a sperm sample. This may be done either
in a clinical setting or at home using a kit ordered from the internet, Digital
camera
with the sample mailed to a lab. The man provides information
about his reproductive history and possible exposure to toxins. The
CaSa system captures images with a digital camera and analyzes
and integrates information on sperm density, motility, and morphol-
ogy (fig. 22a). a computer may also be used to track sperm prog-
ress, in a woman’s body, toward fertilizing an egg cell (fig. 22B).
Devices are being developed that will enable a man to esti-
mate his sperm count at home. They indicate whether a man’s Computer
sperm count is above or below the World health Organization’s
designation of 20 million sperm per milliliter of ejaculate as the
lower criterion for normal fertility. if several tests performed days
apart fall below this level, the man should consult a fertility spe- Temperature Microscope Keyboard Printer
cialist for further testing. Table 22a indicates characteristics of a control

Sexual stimulation the urethra to the outside—a process called ejaculation (e-jak″u-
la′shun). During ejaculation, the posterior pituitary gland releases
a burst of oxytocin, which stimulates contractions of the epididy-
Parasympathetic Veins are mides, seminiferous tubules, and prostate gland, aiding the move-
neurons release compressed, ment of sperm.
nitric oxide, causing reducing blood The sequence of events during emission and ejaculation is coor-
dilation of small flow away from
arteries to penis penis dinated so that the fluid from the bulbourethral glands is expelled
first. This is followed by the release of fluid from the prostate gland,
the passage of the sperm cells, and finally, the ejection of fluid
Blood accumulates in the vascular from the seminal vesicles into the urethra. Ejaculation forcefully
spaces within erectile tissues of penis expels the semen from the body (fig. 22.17).
Immediately after ejaculation, sympathetic impulses constrict
the arteries that supply the erectile tissue, reducing the inflow of
Penis swells and becomes erect blood. Smooth muscle in the walls of the vascular spaces partially
contracts, and the veins of the penis carry excess blood out of these
FiGURE 22.16 mechanism of penile erection in the male. spaces. The penis gradually returns to its flaccid state, and usually

836 UNiT 6 | ThE humaN LifE CyCLE

 FiGURE 22B a computer tracks
sperm cell movements. in semen,
sperm cells swim in a straight line
(a), but as they are activated by
biochemicals in the woman’s body,
their trajectories widen (b). The
sperm cells in (c) are in the mucus
of a woman’s cervix, and the sperm
cells in (d) are attempting to digest
through the structures surrounding
an egg cell.

(a) (b) (c) (d)

TABLE 22A Ranges of Semen and Sperm Characteristics in Healthy Men

Characteristic Range
Volume/ejaculate 1.5–5.0 mL

Number of sperm 20–150 million/mL

Concentration of sperm 12–16 million/mL

Sperm vitality 55–63%

Sperm motility 38–42%

morphologically normal 3–4%

mL = milliliter

another erection and ejaculation cannot be triggered for ten to thirty
minutes or longer. Table 22.1 summarizes the functions of the male
reproductive organs.
Spontaneous emission and ejaculation are common in sleeping
adolescent males. Changes in hormonal concentrations that
accompany adolescent development and sexual maturation
cause these events.
22.3
| Hormonal Control of Male
Reproductive Functions
Hormones secreted by the hypothalamus, the anterior pituitary
gland, and the testes control male reproductive functions. These
hormones initiate and maintain sperm cell production and oversee
the development and maintenance of male sex characteristics.

Hypothalamic and Pituitary Hormones

PRACTiCE
19 What controls blood flow into penile erectile tissues?
20 Distinguish among orgasm, emission, and ejaculation.
21 Review the events associated with emission and ejaculation.
The male body before ten years of age is reproductively imma-
ture with undifferentiated spermatogenic cells in the testes. Then
a series of changes leads to development of a reproductively func-
tional adult. The hypothalamus controls many of these changes.
Recall from chapter 13 (pp. 498–499) that the hypothala-
mus secretes gonadotropin-releasing hormone (GnRH), which

CHAPTER 22 | Reproductive Systems 837

FiGURE 22.17 Culmination of intense
mechanism of emission sexual stimulation
TABLE 22.1 Functions of the Male Reproductive
and ejaculation in the Organs
male.
Organ Function
Sympathetic impulses contract smooth muscle
Testis

Seminiferous tubules Produce sperm cells

Peristaltic Rhythmic Rhythmic interstitial cells Produce and secrete male sex hormones
contractions contractions contractions
in testicular ducts, in erectile in bulbourethral Epididymis Promotes sperm cell maturation; stores
epididymides, columns of glands, prostate sperm cells; conveys sperm cells to
ductus deferentia, penis gland, and ductus deferens
and ejaculatory seminal vesicles
ducts Ductus deferens Conveys sperm cells to ejaculatory duct

Seminal vesicle Secretes an alkaline fluid containing

nutrients and prostaglandins that helps
regulate the ph of semen
Emission—semen moves into urethra
Prostate gland Secretes a slightly acidic fluid that
contains citrate, a nutrient for sperm
Ejaculation—semen Bulbourethral gland Secretes fluid that lubricates end of
is forcefully the penis
expelled from urethra
Scrotum Encloses, protects, and regulates
temperature of testes

Penis Conveys urine and semen to outside of

body; inserted into the vagina during
enters the blood vessels leading to the anterior pituitary gland. In sexual intercourse; the glans penis
response, the anterior pituitary gland secretes the gonadotropins is richly supplied with sensory nerve
endings associated with feelings of
(go-nad″o-trōp′inz) called luteinizing hormone (LH) and follicle- pleasure during sexual stimulation
stimulating hormone (FSH). LH, which in males has been referred
to as interstitial cell-stimulating hormone (ICSH), promotes devel-
opment of the interstitial cells of the testes. The interstitial cells
secrete male sex hormones, mainly testosterone. FSH stimulates
during childhood. Between the ages of thirteen and fifteen, a
the sustentacular cells of the seminiferous tubules to proliferate,
young man’s androgen production usually increases rapidly. This
grow, mature, and respond to the effects of the male sex hormone
phase in development, when an individual becomes reproductively
testosterone. Then, in the presence of FSH and testosterone, sus-
functional, is puberty (pu′ber-te). After puberty, testosterone
tentacular cells stimulate the spermatogenic cells to undergo
secretion continues throughout the life of a male.
spermatogenesis, giving rise to sperm cells (fig. 22.18). The sus-
tentacular cells also secrete a hormone called inhibin, which inhib-
its the anterior pituitary gland by negative feedback. This action
in a group of disorders called male pseudohermaphroditism, tes-
prevents oversecretion of FSH.
tes are present but a block in testosterone synthesis prevents the
genetically male fetus from developing male structures. The child
Male Sex Hormones appears to be a female, but at puberty, the adrenal glands begin
Male sex hormones are termed androgens (an′dro-jenz). Interstitial to produce testosterone (as they normally do in both sexes) and
masculinization ensues. The voice deepens as muscles build up
cells of the testes produce most male sex hormones, but the adrenal
into a masculine physique. Breasts do not develop, nor does
cortex also synthesizes small amounts (see chapter 13, p. 512).
menstruation occur. The clitoris may enlarge so greatly under the
The hormone testosterone (tes-tos′tĕ-roˉn) is the most important
adrenal testosterone surge that it looks like a penis.
androgen. It is secreted by the testes and transported in the blood,
loosely attached to plasma proteins. Like other steroid hormones,
testosterone binds receptor molecules, which are usually in the
nuclei of its target cells (see chapter 13, p. 490). However, in many
target cells, such as those in the prostate gland, seminal vesicles, Actions of Testosterone
and male external accessory organs, testosterone is first converted to Cells of the embryonic testes first produce testosterone after about
another androgen called dihydrotestosterone (di-hi″dro-tes-tos′tĕ- eight weeks of development. This hormone stimulates the forma-
rōn), which stimulates the cells of these organs. Most androgen tion of the male reproductive organs, including the penis, scrotum,
molecules that do not reach receptors in target cells are changed by prostate gland, seminal vesicles, and ducts. Later in development,
the liver into forms that can be excreted in bile or urine. testosterone causes the testes to descend into the scrotum.
Testosterone secretion begins during fetal development and During puberty, testosterone stimulates enlargement of the tes-
continues for several weeks following birth; then it nearly ceases tes (the primary male sex characteristic) and accessory organs of the

838 UNiT 6 | ThE humaN LifE CyCLE

FIGURE 22.18 ​The hypothalamus Hypothalamus
controls maturation of sperm cells and GnRH
development of male secondary sex + – Androgens prevent
characteristics. Negative feedback oversecretion of GnRH
among the hypothalamus, the anterior Androgens prevent over-
lobe of the pituitary gland, and the – secretion of LH
testes controls the concentration of
Pituitary
testosterone in the male body. gland Inhibin prevents
oversecretion of FSH

FSH LH Increased Thickening and

muscular growth strengthening of bones
and thickening of the skin

Enlargement of
Bloodstream accessory
reproductive organs

Enlargement of the
FSH stimulates meiosis Increased growth of larynx and thickening
in primary spermatocytes body hair on face, chest, of the vocal folds
to form immature sperm Inhibin axillary and pubic regions
cells; FSH stimulates
secretion of inhibin by
supporting cells Testosterone and
other androgens Release into
+ bloodstream
LH stimulates
interstitial cells to Stimulation
secrete androgens +
(primarily testosterone) Inhibition
Testes

reproductive system, as well as development of male secondary sex
characteristics, which are special features associated with the adult
male body. Secondary sex characteristics in the male include:
1. increased growth of body hair, particularly on the face, chest,
axillary region, and pubic region; growth of hair on the scalp
may slow
2. enlargement of the larynx and thickening of the vocal folds,
with lowering of the pitch of the voice
3. thickening of the skin
4. increased muscular growth, broadening of the shoulders, and
narrowing of the waist
5. thickening and strengthening of bones
Testosterone also increases the rate of cellular metabolism
and red blood cell production. For this reason, the average num-
ber of red blood cells in a microliter of blood is usually greater in
males than in females. Testosterone stimulates sexual activity by
affecting certain parts of the brain.
Regulation of Male Sex Hormones
The extent to which male secondary sex characteristics develop
is directly related to the amount of testosterone that the intersti-
tial cells secrete. The hypothalamus regulates testosterone output
through negative feedback (fig. 22.18).
As the concentration of testosterone in the blood increases,
the hypothalamus becomes inhibited, decreasing its stimulation of
the anterior pituitary gland by GnRH. As the pituitary’s secretion
of LH falls in response, the amount of testosterone the interstitial
cells release decreases.
As the blood testosterone concentration drops, the hypothalamus
becomes less inhibited, and it once again stimulates the anterior pitu-
itary gland to release LH. The increasing secretion of LH causes the
interstitial cells to release more testosterone, and blood testosterone
concentration increases. Testosterone level decreases somewhat dur-
ing and after the male climacteric, which is a decline in sexual function
that accompanies aging. At any given age, the testosterone concentra-
tion in the male body is regulated to remain relatively constant.
  Practice
22 Which hormone initiates the changes associated with male
sexual maturity?
23 Describe several male secondary sex characteristics.
24 Explain how the secretion of male sex hormones is regulated.
22.4
| Organs of the Female
Reproductive System
The organs of the female reproductive system are specialized
to produce and maintain the female sex cells, the egg cells (or
oocytes); transport these cells to the site of fertilization; provide
a favorable environment for a developing offspring; move the off-
spring to the outside; and produce female sex hormones.
The primary sex organs (gonads) of this system are the two
ovaries, which produce the female sex cells and sex hormones.
The accessory sex organs (secondary sex organs) of the female
reproductive system are the internal and external reproductive
organs (fig. 22.19; reference plates 5 and 6, pp. 42–43).

CHAPTER 22 | Reproductive Systems 839