28 The Reproductive

System

Learning Outcomes
These Learning Outcomes correspond by number to this chapter's sections
and indicate what you should be able to do after completing the chapter.

28-1 ■ List the basic structures of the human reproductive system,

and summarize the functions of each. p. 1102
28-2 ■ Describe the structures of the male reproductive
system, and specify the composition of semen. p. 1103
28-3 ■ Explain the differences between meiosis
and mitosis, describe the process of
spermatogenesis, and summarize the
hormonal mechanisms that regulate male
reproductive functions. p. 1111
28-4 ■ Describe the anatomy and functions
of the ovaries, the uterine tubes,
the uterus, the vagina, and the
structures of the female external
genitalia. p. 1118
28-5 ■ Describe the ovarian roles
in oogenesis, explain the
complete ovarian and uterine
cycles, and summarize
all aspects of the female
reproductive cycle. p. 1128
28-6 ■ Discuss the physiology
of sexual intercourse in males
and females. p. 1137
28-7 ■ Describe the reproductive
system changes that occur with
development and aging. p. 1139
28-8 ■ Give examples of interactions
between the reproductive
system and each of the other
organ systems. p. 1142

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 ++ CLINICAL CASE And Baby Makes Three?
1102 UNIT 6 Continuity of Life

The Millers long to have a baby. They are Her husband, Tim, rates his health as a
both 30 years old, have been married for “9” on a 10-point scale. He's a distance run-
5 years, and have been trying desperately to ner and clocks an average of 40 miles per
get pregnant. week. He says exercise helps him manage
Susan waits and watches. Each time the stress of his sales job. Neither of them
her period comes, her heart drops. Her drinks nor smokes. When should this
period has never been like clockwork. Now couple seek help for infertility? To find
she worries about the STD (sexually trans- out, turn to the Clinical Case Wrap-Up
mitted disease) she had in college. At that on p. 1147.
time, she was admitted to the infirmary with
a high fever and pelvic pain. A course of
antibiotics finally turned things around.

An Introduction to the Reproductive In both males and females, the ducts are connected to cham-
System bers and passageways that open to the outside. The structures
involved make up the reproductive tract.
Approximately 7.4 billion people currently live on the Earth. The male and female reproductive systems are functionally
This is an astonishing number given that our reproductive quite different, however. In adult males, the testes (TES-tez;
.

system is the only system that is not essential to the life of an singular, testis), or testicles, are male gonads that secrete sex
individual. Its activities do, however, affect other systems and hormones called androgens. The main androgen is testosterone,
ensure continuation of our species. In this chapter we discuss which was introduced in Chapter 18. p. 690 The testes also
how the male and female reproductive organs produce and produce the male gametes, called sperm. Males produce about
store specialized reproductive cells called gametes. We also one-half billion sperm each day. During emission, mature sperm
look at how various reproductive organs secrete hormones travel along a lengthy duct system, where they are mixed with
that play major roles in maintaining normal sexual func- the secretions of accessory glands. The mixture created is known
tion, allowing those gametes to come together to form new .

as semen (SE-men). During ejaculation, semen is expelled from

individuals. the body.
In adult females, the ovaries, or female gonads, each
28-1 Male and female reproductive month release only one immature gamete, called an oocyte
.

(O-o-sı-t). The ovaries also secrete female sex hormones, includ-

system structures produce gametes
. .

ing estrogens. The released oocyte travels along one of two short
that combine to form a new individual uterine tubes, which end in the muscular organ called the uterus
Learning Outcome List the basic structures of the human . .

(YU-ter-us). A short passageway, the vagina (va-JI-nuh), con-

reproductive system, and summarize the functions of each.
nects the uterus with the exterior.
The human reproductive system ensures the continued exis- During sexual intercourse, ejaculation introduces semen into
tence of the human species—by producing, storing, nourish- the vagina, and the sperm then ascend the female reproductive
ing, and transporting functional male and female reproductive tract. If a sperm reaches the oocyte and starts the process of
cells. These cells are called gametes (GAM-ets). fertilization, the oocyte matures into an ovum (plural, ova). The
.

The reproductive system includes the following basic uterus will then enclose and support the developing embryo as
structures: it grows into a fetus and prepares for birth (a process we cover
Gonads (GO-nadz; gone, seed), reproductive organs that in Chapter 29).
.

■■

produce gametes and hormones In this chapter we examine the anatomy of the male and
28 female reproductive systems. We consider the physiological and
■■ Ducts that receive and transport the gametes
hormonal mechanisms that regulate reproductive function.
■■ Accessory glands and organs that secrete fluids into the re- Earlier chapters introduced the anatomical reference points
productive system ducts or into other excretory ducts used in the discussions that follow. You may find it helpful
■■ Perineal structures that are collectively known as the exter- to review the figures on the pelvic girdle (Figures 8–7, 8–8,
nal genitalia (jen-ih-TA-le-uh).
.

pp. 299, 300), perineal musculature (Figure 11–13, p. 406),

.

1102  

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 that

Chapter 28 The Reproductive System   1103

pelvic innervation (Figure 13–12, p. 496), and regional blood seminal (SEM-i-nal) glands, the prostate (PROS-tat), and the
.

supply (Figures 21–24, 21–28, pp. 812, 817). bulbo-urethral (bul-bo-yu-RE-thral) gland—secrete various flu-
. .

ids into the duct system. The male external genitalia consist of the
.

Checkpoint scrotum (SKRO-tum), that encloses the testes, the urethra, and
.

the penis (PE-nis), an erectile organ. The distal portion of the

1. Define gamete.
urethra passes through the penis. This section introduces the
2. List the basic structures of the reproductive system.
anatomy and basic functions of these structures.
3. Define gonads.
See the blue Answers tab at the back of the book. The Testes and Associated Structures
Each testis is about 5 cm (2 in.) long, 3 cm (1.2 in.) wide, and
28-2 The structures of the male 2.5 cm (1 in.) thick. Each weighs 10–15 g (0.35–0.53 oz). The
testes hang within the scrotum, a fleshy pouch suspended infe-
reproductive system consist of the rior to the perineum. The scrotum is anterior to the anus, and
testes, duct system, accessory glands, posterior to the base of the penis (see Figure 28–1).
and penis
Learning Outcome Describe the structures of the male The Spermatic Cords
reproductive system, and specify the composition of semen.
The spermatic cords are paired structures extending between
Figure 28–1 shows the main structures of the male reproductive the abdominopelvic cavity and the testes (Figure 28–2). Each
system. Starting from a testis, the sperm travel within the male spermatic cord begins at the entrance to the inguinal canal
reproductive duct system, which consists of the epididymis (ep- (a passageway through the abdominal musculature). After pass-
ih-DID-ih-mus), the ductus deferens (DUK-tus DEF-eh-renz), ing through the inguinal canal, the spermatic cord descends
and the urethra before leaving the body. Accessory glands—the into the scrotum.

Figure 28–1 Sagittal Section of the Male Reproductive System. ATLAS: Plate 64

Prostatic urethra
Ureter
Pubic symphysis
Urinary bladder
Major Structures of the
Rectum
Male Reproductive System

Ducts Ejaculatory duct Accessory Glands

Membranous urethra Seminal gland
Spongy urethra Prostate
Ductus deferens Bulbo-urethral gland
Epididymis

Gonad Testis Corpus cavernosum Anus

28
External Genitalia Penis Corpus spongiosum

External urethral orifice

Scrotum

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 1104 UNIT 6 Continuity of Life

Figure 28–2 Anterior View of the Male Reproductive System. In the anatomical position, the penis is erect,
and the dorsal surface is closest to the navel. When the penis is flaccid, its dorsal surface faces anteriorly.

Testicular artery
Urinary bladder Testicular vein

Penis
Inguinal canal
Inguinal ligament
Superficial
Spermatic Cord inguinal ring
Genitofemoral nerve
Deferential artery
Ductus deferens
Pampiniform plexus Spermatic cord
Testicular artery
Scrotal septum

Epididymis Cremaster with

cremasteric fascia
Scrotal cavity
Testis (covered by visceral Layers of the Scrotum
layer of tunica vaginalis)
Parietal layer of Superficial scrotal
tunica vaginalis fascia
(inner lining of cremaster, Dartos muscle
facing scrotal cavity)
Scrotal skin (cut)
Raphe of scrotum

Each spermatic cord consists of layers of fascia and muscle A narrow space separates the inner surface of the scrotum
enclosing the ductus deferens and the blood vessels, nerves, from the outer surface of the testis. The tunica vaginalis (TU-
and lymphatic vessels that supply the testes. The blood ves- nih-ka vaj-ih-NAL-is), a serous membrane, lines the scrotal cav-
sels include the deferential artery, a testicular artery, and the ity and reduces friction between the opposing parietal (outer)
pampiniform (pam-PIN-ih-form) plexus of a testicular vein. layer and visceral (inner) layer (Figure 28–3a).
Branches of the genitofemoral nerve from the lumbar plexus The scrotum consists of a thin layer of skin and the under-
provide innervation. lying superficial fascia. The dermis contains a layer of smooth
The inguinal canals form during development as the tes- muscle, the dartos (DAR-tos) muscle. Resting muscle tone in
.

tes descend into the scrotum. At that time, these canals link the dartos muscle elevates the testes and causes wrinkling of the
the scrotal cavities with the peritoneal cavity. In normal adult scrotal surface.
males, the inguinal canals are closed, but the spermatic cords A layer of skeletal muscle, the cremaster (kre-MAS-ter),
.

create weak points in the abdominal wall that remain through- lies deep to the dermis. When the cremaster contracts during
out life. As a result, inguinal hernias—protrusions of visceral sexual arousal or in response to decreased temperature, it tenses
tissues or organs into the inguinal canal—are fairly common the scrotum and pulls the testes closer to the body. The crem-
in males. asteric reflex can be initiated by stroking the skin on the upper
thigh, causing the scrotum to move the testes closer to the body.
28 The Scrotum and the Position of the Testes For sperm to develop normally in the testes, the tem-
The scrotum is divided internally into two chambers. A raised perature must be about 1.1°C (2°F) lower than elsewhere in
thickening in the scrotal surface known as the raphe (RA-fe) of the body. The cremaster and dartos muscle relax or contract
.

scrotum divides it in two (see Figure 28–2). Each testis lies in a to move the testes away from or toward the body as needed
separate chamber, or scrotal cavity. Because the scrotal cavities to maintain acceptable testicular temperatures. When air or
are separated by a partition, infection or inflammation of one body temperature increases, these muscles relax and the testes
testis does not normally spread to the other. move away from the body. When the scrotum cools suddenly,

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 Chapter 28 The Reproductive System   1105

Figure 28–3 Anatomy of the Scrotum and the Testes.

Ductus deferens
Epididymis Mediastinum of testis

Efferent ductule
Rete
Scrotum testis
Skin
Dartos muscle Straight
Superficial tubule
scrotal fascia

Cremaster Seminiferous
tubules
Tunica vaginalis

Tunica albuginea Septa testis

Scrotal cavity

Septa testis
Lobule
Raphe of scrotum

a A transverse section of the scrotum and testes Testis LM × 26

b A section through a testis

a maze of passageways called the rete (RE-te; rete, a net) testis

.

as happens when jumping into a cold swimming pool, these

.

muscles contract to pull the testes closer to the body and keep (see Figure 28–3a,b). Fifteen to 20 efferent ductules connect
testicular temperatures from decreasing. the rete testis to the epididymis.
Because the seminiferous tubules are tightly coiled, most
Gross Anatomy of the Testes tissue slides show them in transverse section (Figure 28–4a). A
Deep to the tunica vaginalis covering the testis is the tunica delicate connective tissue capsule surrounds each tubule, and
albuginea (al-bu-JIN-e-uh), a dense layer of connective tissue
. .

areolar tissue fills the spaces between the tubules (Figure 28–4b).
rich in collagen fibers (see Figure 28–3a). These fibers are con- Within those spaces are numerous blood vessels and large
tinuous with those surrounding the adjacent epididymis and interstitial endocrine cells (Leydig cells). Interstitial endocrine
extend into the testis. There they form fibrous partitions, or cells produce androgens (androsterone and testosterone), the
septa testis, that converge toward the region nearest the entrance dominant sex hormones in males. Testosterone is the most
to the epididymis. The connective tissues in this region support important androgen.
the blood vessels and lymphatic vessels that supply and drain
the testis, and the efferent ductules, which transport sperm to the
Functional Anatomy of the Male
epididymis.
Reproductive Duct System
Histology of the Testes The testes produce sperm that are immobile and not yet capable
The septa testis subdivides the testis into a series of lobules of successfully fertilizing an oocyte. The male reproductive duct
(see Figure 28–3a). Distributed among the lobules are approxi- system is responsible for the functional maturation, nourish-
mately 800 slender, tightly coiled seminiferous (sem-ih-NIF- ment, storage, and transport of sperm.
er-us) tubules (Figure 28–3a,b). Each tubule averages about 28
80 cm (32 in.) in length. A typical testis contains nearly one- The Epididymis
half mile of seminiferous tubules. Sperm production takes These immobile sperm are transported into the epididymis by
place within these tubules. fluid currents created by the cilia that line the efferent ductules.
Each lobule contains several seminiferous tubules that The epididymis (ep-ih-DID-ih-mis; epi, on + didymos, twin)
merge into straight tubules that enter the mediastinum of the is the start of the male reproductive tract (Figure 28–5). It is a
testis. Straight tubules are extensively interconnected, forming coiled tube bound to the posterior border of each testis.

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 1106 UNIT 6 Continuity of Life

Figure 28–4 Anatomy of the Seminiferous Tubules.

Seminiferous tubule Dividing

containing late spermatocytes Nurse cell
spermatids
Interstitial
endocrine cells

Spermatogonium

Seminiferous
tubule Spermatids
containing
sperm

Lumen
Seminiferous tubule
containing early
spermatids

Heads of
maturing
sperm

Seminiferous tubules LM × 75 LM × 350

Seminiferous tubule

a A section through the seminiferous tubules. b A cross section through a single tubule.

The epididymides (ep-ih-dih-DIM-ih-dez) can be felt products of enzymatic breakdown are released into the sur-
.

through the scrotal skin. Each epididymis is almost 7 m (23 ft) rounding interstitial fluids for pickup by the epididymal
long, but it is coiled and twisted so it takes up very little space. blood vessels.
It has a head, a body, and a tail (Figure 28–5a,c). The superior ■■ It stores and protects sperm and facilitates their functional matu-
head is the largest part of the epididymis. The head receives ration. It takes up to 3 weeks for a sperm to pass through
sperm from the efferent ductules. the epididymis and complete its functional maturation.
The body is the middle part that extends inferiorly from During this time, sperm exist in a sheltered environment
the head to the tail of the epididymis on the posterior surface that is regulated by the surrounding epithelial cells.
of the testis. Near the inferior border of the testis, the number of
Transport along the epididymis involves a combination of
coils decreases, marking the start of the tail. The tail re-curves,
fluid movement and peristaltic contractions of smooth muscle
ascends, and connects to the ductus deferens. Sperm are stored
in the epididymis. After passing along the tail of the epididy-
primarily within the tail of the epididymis.
mis, the sperm enter the ductus deferens.
The epididymis has the following functions:
■■ It monitors and adjusts the composition of the fluid produced by The Ductus Deferens
28 the seminiferous tubules. The pseudostratified columnar epi-
Each ductus deferens (plural, ductus deferentia), also called
thelial lining of the epididymis has distinctive stereocilia
the vas deferens, is 40–45 cm (16–18 in.) long. It begins at the
(Figure 28–5b). These stereocilia increase the surface area
tail of the epididymis (see Figure 28–5a,c). As part of the sper-
available for absorption from, and secretion into, the fluid
matic cord, it ascends through the inguinal canal (look back at
in the tubule.
Figure 28–2). Inside the abdominal cavity, the ductus deferens
■■ It acts as a recycling center for damaged sperm. Cellular debris passes posteriorly, curving inferiorly along the lateral surface of
and damaged sperm are absorbed in the epididymis. The the urinary bladder toward the superior and posterior margin

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 Chapter 28 The Reproductive System   1107

Figure 28–5 Anatomy of the Epididymis. ATLAS: Plate 60a

Tails of sperm in
lumen of epididymis

Pseudostratified columnar
epithelium of epididymis
Stereocilia Tunica
vaginalis Spermatic Ductus
(reflected) cord deferens
Spermatic cord
Ductus deferens

Efferent ductules
Epididymis LM × 304
Straight tubule
b Sectional view of the epididymis

Rete testis
Seminiferous
Epididymis
tubule
Tunica Head of epididymis
albuginea
Body of epididymis
Testis
Tail of epididymis Testis

Scrotal
cavity

a A diagrammatic view of the c The testis and epididymis on

head, body, and tail of an gross dissection
epididymis

of the prostate (see Figure 28–6a). Just before the ductus defer- The Accessory Glands
ens reaches the prostate and seminal glands, its lumen enlarges.
The fluids secreted by the seminiferous tubules and the epididy-
This expanded portion is known as the ampulla (am-PUL-luh)
mis account for only about 5 percent of the volume of semen.
of ductus deferens.
The fluid component of semen is a mixture of secretions—
The wall of the ductus deferens contains a thick layer of
each with distinctive biochemical characteristics—from many
smooth muscle (see Figure 28–6b). Peristaltic contractions in
glands. Important glands include the seminal glands, the prostate,
this layer propel sperm and fluid along the duct, which is lined
and the bulbo-urethral glands. All of these occur only in males.
by a pseudostratified ciliated columnar epithelium. In addition
Among the major functions of these glands are (1) activat-
to transporting sperm, the ductus deferens can store sperm in
ing sperm; (2) providing the nutrients sperm need for motility;
the ampulla for several months. During this time, the sperm
(3) propelling sperm and fluids along the reproductive tract,
remain in a temporary state of inactivity with low metabolic
mainly by peristaltic contractions; and (4) producing buf-
rates.
fers that counteract the acidity of the urethral and vaginal
environments.
The Male Urethra
In males, the urethra is a passageway that extends 18–20 cm 28
(7–8 in.) from the urinary bladder to the tip of the penis. The Seminal Glands (Seminal Vesicles)
It is divided into prostatic, membranous, and spongy regions The ductus deferens on each side ends at the junction between the
(Figures 28–1, 28–7b). The male urethra is used by both the ampulla of ductus deferens and the duct that drains the seminal
urinary and reproductive systems. It conducts urine to the exte- gland (see Figure 28–6a). The seminal glands, also called the
rior and introduces semen into the female's vagina during seminal vesicles, are tubular glands embedded in connective tissue
sexual intercourse. on each side of the midline, sandwiched between the posterior

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 1108 UNIT 6 Continuity of Life

Figure 28–6 Anatomy of the Ductus Deferens and Accessory Glands.

Lumen of ductus deferens Smooth muscle

Ureter

Urinary
bladder
Ductus deferens Ductus deferens LM × 120

b Light micrograph showing the thick layers of

smooth muscle in the wall of the ductus deferens.
Seminal gland
Ampulla of
ductus deferens
Excretory duct of
seminal gland
Ejaculatory duct
Prostate Lumen
Prostatic urethra
Secretory
a A posterior view of the pockets
Bulbo-urethral glands
urinary bladder and
prostate, showing
subdivisions of the ductus Smooth
deferens in relation to muscle
surrounding structures. Seminal gland LM × 45

c Histology of the seminal glands. These organs

produce most of the volume of seminal fluid.

Smooth
Connective muscle
tissue and Capsule
smooth
muscle

Mucous
glands

Tubulo-alveolar
glands
Lumen

Prostate LM × 50 Bulbo-urethral gland LM × 175

d Histology of the tubulo-alveolar glands, connective tissue, e Histology of the bulbo-urethral glands, which
28 and smooth muscle within the prostate. secrete a thick mucus into the spongy urethra.

wall of the urinary bladder and the rectum. Each tubular semi- Seminal glands have an epithelial lining that contains
nal gland is about 15 cm (6 in.) in length. The body of the gland extensive folds (Figure 28–6c), reflecting that they are extremely
has many short side branches. The entire assemblage is coiled active secretory glands. The seminal glands produce the major-
and folded into a compact, tapered mass roughly 5 cm * 2.5 cm ity of the volume of semen. The secretions of the seminal
(2 in. * 1 in.). glands are discharged into the ejaculatory duct. The junction

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 Chapter 28 The Reproductive System   1109

of the duct of the seminal gland with the ampulla of ductus ionic and nutrient composition. A typical sample of semi-
deferens marks the start of the ejaculatory duct. This short nal fluid contains the combined secretions of the seminal
passageway is 2 cm (less than 1 in.). It penetrates the muscu- glands (60 percent), the prostate (30 percent), the nurse
lar wall of the prostate and empties into the prostatic urethra cells and epididymis (5 percent), and the bulbo-urethral
(see Figure 28–6a). glands (less than 5 percent). An abnormally low volume
may indicate problems with the prostate or seminal glands.
The Prostate In particular, the secretion of the seminal glands contains
The prostate is a small, muscular, rounded organ about (1) a higher concentration of fructose, which is easily me-
4 cm (1.6 in.) in diameter. The prostate encircles the proximal tabolized by sperm; (2) prostaglandins, which can stimulate
portion of the urethra as it leaves the urinary bladder (see smooth muscle contractions along the male and female
Figures 28–1, 28–6a). The glandular tissue of the prostate con- reproductive tracts; and (3) fibrinogen, which forms a tem-
sists of a cluster of 30–50 compound tubulo-alveolar glands porary semen clot within the vagina. The glandular fluid the
(Figure 28–6d). p. 171 These glands are surrounded by a seminal glands produce generally has the same osmotic con-
thick blanket of smooth muscle fibers. centration as that of blood plasma, but the compositions of
The prostate produces prostatic fluid, a solution that the two fluids are quite different. The secretions of the semi-
makes up about a quarter of the volume of semen. This slightly nal glands are slightly alkaline, helping to neutralize acids in
acidic fluid is rich in enzymes, such as acid phosphatase and the secretions of the prostate and within the vagina.
amylase, that prevent sperm coagulation in the vagina. Prostatic ■■ Enzymes. Several important enzymes are in seminal fluid,
fluid is ejected into the prostatic urethra by peristaltic contrac- including (1) a protease that may help dissolve mucus in
tions of the muscular prostate wall. the vagina; (2) seminalplasmin (sem-i-nal-PLAZ-min), a
Prostatic inflammation, or prostatitis (pros-ta-TI-tis), can
.

protein with antibiotic properties that may help prevent

occur in males at any age, but it most commonly afflicts older urinary tract infections in males; (3) a prostatic enzyme
men. Prostatitis can result from bacterial infections but also that converts fibrinogen to fibrin, coagulating the semen
occurs without pathogens. Signs and symptoms can resemble within a few minutes after ejaculation; and (4) fibrinolysin,
those of prostate cancer. Men with prostatitis may have pain which liquefies the clotted semen after 15–30 minutes.
in the lower back, perineum, or rectum. In some cases, the
A complete chemical analysis of semen appears in the
symptoms are accompanied by painful urination and mucus
Appendix.
discharge from the external urethral orifice. Antibiotics are
effective in treating most cases caused by bacterial infection.
The Penis
The Bulbo-urethral Glands The penis is a tubular organ for sexual intercourse and con-
The paired bulbo-urethral glands, or Cowper's glands, are ducting urine to the exterior. Main structures include the root,
compound tubular mucous glands located at the base of the body, and glans penis (Figure 28–7a). The root of penis is
penis (see Figure 28–7a). These glands are round, with diam- the fixed portion that attaches the penis to the body wall. It
eters nearly 10 mm (less than 0.5 in.). The duct of each gland includes left crus of penis, right crus of penis, and the bulb of
travels alongside the spongy urethra for 3–4 cm (1.2–1.6 in.) penis. This connection occurs immediately inferior to the pubic
before emptying into the urethral lumen. symphysis. The body of penis (shaft) is the tubular, movable
The bulbo-urethral glands secrete thick, alkaline mucus portion of the organ. The glans penis (head) is the expanded
(Figure 28–6e). The secretion helps neutralize any urinary acids distal end that surrounds the external urethral orifice. The neck
that may remain in the urethra, and it lubricates the tip of the of glans is the narrow portion of the penis between the body
penis. and the glans penis.
The skin overlying the penis resembles that of the scrotum.
Semen The dermis contains a layer of smooth muscle that is a con-
tinuation of the dartos muscle of the scrotum. The underlying
A typical ejaculation releases 2–5 mL of semen (ejaculate). When
areolar tissue allows the thin skin to move without distorting
sampled for analysis, semen is collected after a 36-hour period
underlying structures. The subcutaneous layer also contains 28
of sexual abstinence. Semen typically contains the following:
superficial arteries, veins, and lymphatic vessels.
Sperm. The normal sperm count ranges from 20 million to A fold of skin called the foreskin or prepuce (PRE-pus)
.

■■
.

100 million sperm per milliliter of semen. In addition, at surrounds the tip of the penis. (Figure 28–7b). The foreskin
least 60 percent of the sperm in the sample appear normal attaches to the relatively narrow neck of the penis and contin-
and will be actively swimming. ues over the glans penis. Preputial (pre-PU-she-al) glands in
.

. .

■■ Seminal Fluid. Seminal fluid, the liquid component of the skin of the neck and the inner surface of the foreskin secrete
semen, is a mixture of glandular secretions with a distinct a waxy material known as smegma (SMEG-ma). Unfortunately,

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 1110 UNIT 6 Continuity of Life

Figure 28–7 Anatomy of the Penis. ATLAS: Plate 60b

Pubic
symphysis
Membranous
urethra Ureter
Bulb of penis
Trigone of
urinary bladder
Right crus Body of
of penis Ductus deferens
penis
Ramus of
ischium Seminal Opening of
gland ejaculatory duct
Corpus spongiosum Neck Prostate
of glans Prostatic urethra

Corpora cavernosa Glans Membranous Bulbo-urethral gland

penis urethra
Crus at root of penis
Scrotum External Bulb of penis
urethral
orifice Opening from
bulbo-urethral Erectile Tissue
gland
Corpus spongiosum
a An anterior and lateral view of the penis,
Corpora cavernosa
showing positions of the erectile tissues

Spongy urethra
Dorsal artery (red), veins
(blue), and nerve (yellow)
Glans penis Foreskin
External urethral
Corpora cavernosa orifice

Dartos muscle
b A frontal section through the
Deep artery of penis
penis and associated organs
Collagenous sheath

Spongy urethra

Corpus
spongiosum

c A sectional view through the penis

? Which erectile tissue is split into two cylindrical masses that surround a central artery? Which
erectile tissue surrounds the urethra?

28 smegma can be an excellent nutrient source for bacteria. Mild urinary tract infections, HIV infection, and penile cancer.
inflammation and infections in this area are common, espe- Because it is a surgical procedure with risk of bleeding, infection,
cially if the area is not washed thoroughly and frequently. One and other complications, the practice remains controversial.
way to avoid such problems is circumcision (ser-kum-SIZH- Deep to the areolar tissue, a dense network of elastic fibers
un), the surgical removal of the foreskin. In Western societies encircles the internal structures of the penis. Most of the body of
(especially the United States), this procedure is often performed the penis consists of three cylindrical columns of erectile tissue
shortly after birth. Circumcision lowers the risks of developing (Figure 28–7b,c). Erectile tissue consists of a three-dimensional

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 Chapter 28 The Reproductive System   1111

28-3 Spermatogenesis occurs in

+ Clinical Note Circumcision the testes, and hormones from the
hypothalamus, pituitary gland, and
Circumcision, or the removal of the foreskin of the penis,
is an elective procedure for a baby boy. Certain religions
testes control male reproductive
conduct a ceremonial circumcision, as in the bris practiced functions
in the Jewish faith when the infant is 8 days old. In the brief Learning Outcome Explain the differences between meiosis and
medical process of circumcision, the penis is anesthetized mitosis, describe the process of spermatogenesis, and summarize
locally. A clamp or ring is applied to remove the prepuce. the hormonal mechanisms that regulate male reproductive
functions.
Circumcision in an adult may require a general anesthetic.
A circumcised penis is easier to keep clean. The risk of
developing a sexually transmitted disease, which can flour- Overview of Mitosis and Meiosis
ish under the foreskin, is decreased. However, these risks
Mitosis, the production of genetically identical cells, and meiosis,
can also be managed by teaching the boy to practice care-
ful personal hygiene. the production of gametes, differ significantly in terms of the
events that take place in the nucleus. Recall from Chapter 3
that human somatic cells contain 23 pairs of chromosomes,
or 46 chromosomes altogether. Each pair consists of one chro-
maze of vascular channels incompletely separated by parti- mosome provided by the father, and another provided by the
tions of elastic connective tissue and smooth muscle fibers. In mother, at the time of fertilization. The two members of each
the resting state, the arterial branches are constricted and the of the 22 pairs have similar sizes and genes and are known as
homologous (huh-MOL-o-gus) chromosomes.
.

muscular partitions are tense. This combination restricts blood

flow into the erectile tissue. During erection the penis stiffens Mitosis is part of the process of somatic cell division,
and elevates to an upright position, but the flaccid (nonerect) producing two daughter cells each containing identical num-
penis hangs inferior to the pubic symphysis and anterior to the bers and pairs of chromosomes. The pattern is illustrated in
scrotum. Figure 28–8a. Because daughter cells contain both members
The anterior surface of the flaccid penis covers two cylin- of each chromosome pair, they are called diploid (DIP-loyd;
drical masses of erectile tissue: the corpora cavernosa (KOR- diplo, double) (2n = 46) cells. (You can review the description
.

por-uh ka-ver-NO-suh; singular, corpus cavernosum). The two of mitosis and cell division in Chapter 3. pp. 150–151)
are separated by a thin septum and encircled by a dense collag- Meiosis is a specialized form of cell division that produces
enous sheath (see Figure 28–7c). The corpora cavernosa diverge only gametes. In contrast to mitosis, meiosis follows the pattern
at their bases, forming the crus of penis (see Figure 28–7a). in Figure 28–8b. It involves two cycles of cell division (meiosis I
Each crus is bound to the ramus of the ischium and pubis and meiosis II) and produces four cells, each containing 23 indi-
by tough connective tissue ligaments. The corpora cavernosa vidual chromosomes. These cells are called haploid (HAP-loyd;
extend along the length of the penis as far as its neck. The erec- haplo, single) (n = 23) cells and contain only one member of
tile tissue within each corpus cavernosum surrounds a central each homologous pair of chromosomes. Because gametes con-
artery, or deep artery of the penis (see Figure 28–7c). tain half the number of chromosomes found in somatic cells,
The penis contains the distal portion of the urethra, which the fusion of the nuclei of a male gamete and a female gamete
the relatively slender corpus spongiosum (spon-je-O-sum)
.
.

produces a cell that has the normal number of chromosomes,

surrounds (see Figure 28–7a,b). This erectile body extends to rather than twice that number. The events in the nucleus shown
the tip of the penis, where it expands to form the glans penis. in Figure 28–8b are the same for the formation of sperm in
The sheath surrounding the corpus spongiosum contains more males or oocytes in females.
elastic fibers than does that of the corpora cavernosa, and the As a cell prepares to begin meiosis, DNA replication occurs
erectile tissue contains a pair of small arteries. within the nucleus just as it does in a cell preparing for mitosis.
The duplicated chromosomes condense and become visible
with a light microscope. As in mitosis, each chromosome now
Checkpoint
28
.

consists of two duplicate chromatids (KRO-mah-tidz).

4. Name the male reproductive structures. At this point, the close similarities between meiosis and
5. On a warm day, would the cremaster be contracted or mitosis end. In meiosis, the corresponding maternal (inherited
relaxed? Why?
from the mother) and paternal (inherited from the father)
6. Trace the pathway that a sperm travels from the site of its chromosomes now come together in an event known as synap-
production to outside the body.
sis (sih-NAP-sis). Synapsis involves 23 pairs of chromosomes,
See the blue Answers tab at the back of the book. and each member of each pair consists of two chromatids.

> Go to Pearson Mastering > Study Area > Menu > Lab Tools > 3.0 > Human Cadaver and Anatomical Models > Reproductive Systems

M28_MART9867_11_GE_C28.indd 1111 23/08/17 10:29 AM

 1112 UNIT 6 Continuity of Life

Figure 28–8 A Comparison of Chromosomes in Mitosis and Meiosis.

a The fate of three homologous b The fate of three homologus

chromosome pairs during mitosis. chromosome pairs during meiosis.

MITOSIS Parent cell MEIOSIS I

(before chromosome replication) Site of a
Prophase crossing Prophase I
over

Duplicated Tetrad formed

chromosome by synapsis of
(two sister chromatids) homologous
2n = 6 chromosomes

Metaphase Metaphase I

Chromosomes Tetrads align at

align at the the metaphase
metaphase plate plate

Anaphase Anaphase I
Telophase Homologous Telophase I
chromosomes
separate during
anaphase I;
sister
chromatids
remain together

Daughter cells
of mitosis (2n) Sister chromatids MEIOSIS II
separate during
KEY anaphase II
Haploid
Maternal chromosomes n=3

n n n n
Paternal chromosomes
Daughter cells (gametes) of meiosis

A matched set of four chromatids is called a tetrad (TET- chromosome or the paternal chromosome. (Compare the two
rad; tetras, four) (Figure 28–8b). Some genetic material can be parts of Figure 28–8.)
exchanged between the maternal and paternal chromatids of As anaphase I proceeds, the maternal and paternal com-
a chromosome pair at this stage of meiosis. Such an exchange, ponents are randomly and independently distributed. That
called crossing over, increases genetic variation among offspring. is, as each tetrad splits, we cannot predict which daughter cell
We discuss genetics in Chapter 29. will receive copies of the maternal chromosome, and which
Meiosis includes two division cycles, referred to as meiosis I will receive copies of the paternal chromosome. As a result,
and meiosis II. We identify the stages within each cycle with I telophase I ends with the formation of two daughter cells con-
or II, for example as prophase I or metaphase II. taining unique combinations of maternal and paternal chro-
mosomes. Both cells contain 23 chromosomes.
Meiosis I The first meiotic division is called a reductional division
28 Meiosis I begins with prophase I; the nuclear envelope disap- because it reduces the number of chromosomes from the dip-
pears at the end of prophase I. As metaphase I begins, the tet- loid number (2n = 46) to the haploid number (n = 23). Each
rads line up along the metaphase plate. As anaphase I begins, of these chromosomes still consists of two chromatids. The two
the tetrads break up—the maternal and paternal chromo- chromatids will separate during meiosis II.
somes separate. This is a major difference between mitosis
and meiosis: In mitosis, each daughter cell receives a copy of Meiosis II
every chromosome, maternal and paternal, but in meiosis I, The interphase between meiosis I and meiosis II is very brief.
each daughter cell receives two copies of either the maternal No DNA is replicated during that period. Each cell then proceeds

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 Chapter 28 The Reproductive System   1113

through prophase II, metaphase II, and anaphase II. During ana- Spermatogenesis
phase II, the chromatids separate. Telophase II yields four haploid
As we have indicated, spermatogenesis (sper-ma-to-JEN-eh-
.

cells, each containing 23 chromosomes. Because the number of

sis) is the process of sperm formation (Figure 28–9). Spermato-
chromosomes is unchanged, meiosis II is an equational division.
genesis begins at puberty (sexual maturation) and continues
The chromosomes are evenly distributed among the four
until relatively late in life (past age 70).
cells, but the cytoplasm may not be. In males, in the process of
The complete process of spermatogenesis takes about 64
spermatogenesis, meiosis produces four immature gametes that
days. It involves three steps:
are identical in size. Each will develop into a functional sperm.
In females, meiosis produces one relatively huge oocyte and 1. Mitosis. Spermatogonia (sper-ma-to-GO-ne-uh) are stem
.

. .

three tiny, nonfunctional polar bodies. (If fertilization takes cells in the seminiferous tubules that undergo cell divi-
place, the oocyte completes meiosis II, yielding an ovum.) We sions throughout adult life. Some of these cells differenti-
examine the details of spermatogenesis next. We will consider ate into primary spermatocytes (sper-MA-to-sııts), which
. .

the production of female oocytes, oogenesis, in Section 28-5. prepare to begin meiosis.

Figure 28–9 The Process of Spermatogenesis. The fates of three representative chromosome pairs are shown;
for clarity, maternal and paternal chromatids are not identified.

Spermatogenesis

Mitosis of spermatogonium

Each division of a diploid spermatogonium produces two

daughter cells. One is a spermatogonium that remains in contact
with the basement membrane of the seminiferous tubule, and
the other is a primary spermatocyte that is displaced toward
the lumen. These events from spermatogonium to primary Primary
spermatocyte take 16 days. spermatocyte
(diploid, 2n)

Meiosis I DNA replication

As meiosis I begins, each primary spermatocyte contains
Primary
46 individual chromosomes. At the end of meiosis I, the
spermatocyte
daughter cells are called secondary spermatocytes.
Every secondary spermatocyte contains 23 chromo-
somes, each with a pair of duplicate chromatids. This Synapsis and
tetrad formation
phase of spermatogenesis takes about 24 days.

Secondary
spermatocytes

Meiosis II
Spermatids
The secondary spermatocytes soon enter (haploid, n)
meiosis II, which yields four haploid spermatids,
each containing 23 chromosomes. For each
primary spermatocyte that enters meiosis, four
spermatids are produced. This phase lasts only
a few hours.

Spermiogenesis (physical maturation) 28

In spermiogenesis, the last step of
spermatogenesis, each spermatid matures Sperm
into a single sperm. The process of (haploid, n)
spermiogenesis—from spermatids to
sperm—takes 24 days.

? What is the name of the cell at each stage of sperm development, from mitosis to spermiogenesis?

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 1114 UNIT 6 Continuity of Life

2. Meiosis. Primary spermatocytes then undergo meiotic divi- completed in meiosis I or meiosis II. These connections assist
sions that produce spermatids (SPER-ma-tidz), undiffer- in transferring nutrients and hormones between the cells, help-
entiated male gametes. ing ensure that the cells develop together. The bridges are not
3. Spermiogenesis. In spermiogenesis, spermatids differentiate broken until the last stages of physical maturation.
into physically mature sperm. In spermiogenesis, the last step of spermatogen-
esis, the spermatids gradually develop into mature sperm
Spermatogenesis is a continuous process, so all stages of
(Figures 28–9, 28–11). At spermiation, a sperm loses its attach-
meiosis are seen within the seminiferous tubules. Each seminif-
ment to the nurse cell and enters the lumen of the seminiferous
erous tubule contains spermatogonia, spermatocytes at various
tubule.
stages of meiosis, spermatids, and sperm. They also contain large
nurse cells, which provide a microenvironment that supports sper-
matogenesis. The sperm eventually lose contact with the nurse The Role of Nurse Cells
cells of the seminiferous tubule and enter the fluid in the lumen. Developing spermatocytes undergoing meiosis, and spermatids
undergoing spermiogenesis, are not free in the seminiferous
Mitosis and Meiosis during Spermatogenesis tubules. Instead, they are surrounded by the cytoplasm of adja-
Spermatogenesis begins at the outermost layer of cells in cent nurse cells. Nurse cells, which are also known as Sertoli
the seminiferous tubules and proceeds toward the lumen cells, play a critical supportive role in spermatogenesis.
(Figure 28–10a,b). At each step in this process, the daughter These cells have six important functions that directly or
cells move closer to the lumen. indirectly affect mitosis, meiosis, and spermiogenesis within
First, spermatogonia divide by mitosis to produce two the seminiferous tubules:
daughter cells. One daughter cell from each division remains ■■ Maintenance of the Blood Testis Barrier. A blood testis
at that location as a spermatogonium, while the other is pushed barrier isolates the seminiferous tubules from the general
toward the lumen (space) of the tubule. The displaced cell dif- circulation. This barrier is comparable in function to the
ferentiates into a primary spermatocyte. Primary spermatocytes blood brain barrier. p. 519 Nurse cells are joined by tight
then begin meiosis, giving rise to secondary spermatocytes that junctions between basal cytoplasmic processes. They form
divide and differentiate into immature spermatids. a layer that divides the seminiferous tubule into an outer
basal compartment and an inner luminal compartment.
Spermiogenesis The basal compartment contains the spermatogonia. Meiosis
The four spermatids initially remain interconnected by bridges and spermiogenesis occur in the luminal compartment (see
of cytoplasm because cytokinesis (cytoplasmic division) is not Figure 28–10a,b). Transport across the nurse cells is tightly

Figure 28–10 Spermatogenesis in a Seminiferous Tubule.

Spermatid Nurse Dividing Capillary

cell spermatocytes
Spermatids completing
spermiogenesis LUMEN Spermatids beginning
spermiogenesis
Initial spermiogenesis
Secondary
spermatocyte
Secondary spermatocyte
in meiosis II Primary
spermatocyte
Luminal compartment preparing
for meiosis I
Lumen Blood testis barrier
Nuclei of
Connective nurse cells
tissue capsule
Fibroblast
28
Interstitial il lary
Cap
endocrine cells
Spermatogonium
Spermatogonium
Sperm Basal compartment

a Nurse cells surround the stem cells of the tubule and b Stages in spermatogenesis in the
support the developing spermatocytes and spermatids. wall of a seminiferous tubule.

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 Chapter 28 The Reproductive System   1115

Figure 28–11 The Process of Spermiogenesis and Anatomy of a Sperm. ATLAS: Plate 60a

Spermatid

Mitochondria

Nucleus

Shed Tail (55 µm)

Golgi cytoplasm
apparatus
Acrosomal Sperm SEM × 780
vesicle

Fibrous sheath of flagellum

Acrosome
Middle
Nucleus Mitochondrial spiral
piece (5 µm)
Acrosome Neck (1 µm)
Spermiogenesis. The differentiation of a Centrioles
spermatid into a sperm. This process is
completed in approximately 24 days. Head (5 µm) Nucleus
Acrosome

Sperm

? In which of the four parts of a sperm are the centrioles found?

regulated, so conditions in the luminal compartment re- ■■ Support of Spermiogenesis. Spermiogenesis requires nurse
main very stable. cells. These cells surround and enfold the spermatids, pro-
●  The nurse cells produce the fluid in the lumen of a viding nutrients and chemical stimuli that promote their
seminiferous tubule. They also regulate the fluid's development. Nurse cells also phagocytize cytoplasm that
composition. This luminal fluid is very different from is shed by spermatids as they develop into sperm.
the surrounding interstitial fluid. It is high in andro- ■■ Secretion of Inhibin. Nurse cells secrete the peptide hormone
gens, estrogens, potassium ions, and amino acids. The inhibin (in-HIB-in) in response to factors released by devel-
blood testis barrier is essential to preserving the dif- oping sperm. Inhibin depresses the pituitary production
ferences between the luminal fluid and the interstitial of FSH, and perhaps the hypothalamic secretion of gonad-
fluid. otropin-releasing hormone (GnRH). The faster the rate of
●  In addition, this barrier prevents immune system cells sperm production, the more inhibin is secreted. By regulat-
from detecting and attacking the developing sperm. The ing FSH and GnRH secretion, nurse cells provide negative
plasma membranes of sperm contain sperm-specific feedback control of spermatogenesis. 28
antigens not found in somatic cell membranes, so they
■■ Secretion of Androgen-Binding Protein. Androgen-binding pro-
might be identified as “foreign.”
tein (ABP) binds androgens (primarily testosterone) in the
■■ Support of Mitosis and Meiosis. Circulating follicle-stimulat- luminal fluid of the seminiferous tubules. This protein is
ing hormone (FSH) and testosterone stimulate nurse cells. likely important for increasing androgen concentration
These stimulated nurse cells then promote the division of and stimulating spermiogenesis. FSH stimulates the pro-
spermatogonia and the meiotic divisions of spermatocytes. duction of ABP.

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 1116 UNIT 6 Continuity of Life

Maturation of Sperm Hormonal Regulation of Male

The sperm that lie within the lumen of the seminiferous tubule
Reproductive Function
have most of the physical characteristics of mature sperm, but The hormonal interactions that regulate male reproductive func-
are functionally immature and incapable of coordinated move- tion are diagrammed in Spotlight Figure 28–12. We introduced
ment or fertilization. It takes up to 2 weeks for a sperm to pass major reproductive hormones in Chapter 18. p. 690 The ante-
through the epididymis and complete its functional matura- rior lobe of the pituitary gland releases follicle-stimulating hor-
tion. During this time, sperm exist in a sheltered environment mone (FSH) and luteinizing hormone (LH). The pituitary releases
that is precisely regulated by the surrounding epithelial cells. these hormones in response to gonadotropin-releasing hormone
Sperm leaving the epididymis are mature but immobile. To (GnRH). This peptide is synthesized in the hypothalamus and
become motile (actively swimming) and fully functional, they carried to the anterior lobe by the hypophyseal portal system.
must undergo a process called capacitation. When mixed with The hormone GnRH is secreted in pulses rather than con-
the secretions of the seminal glands, sperm undergo the first tinuously. In adult males, small pulses occur at 60- to 90-minute
step in capacitation and begin beating their flagella,. Capaci- intervals. As levels of GnRH change, so do the rates of secretion
tation normally takes place in two steps: (1) Sperm become of FSH and LH (and testosterone, which is released in response to
motile when they are mixed with secretions of the seminal LH). The GnRH pulse frequency in adult males remains relatively
glands, and (2) they become capable of successful fertilization steady from hour to hour, day to day, and year to year, unlike
when exposed to conditions in the female reproductive tract. the situation in women, as we will see later in the chapter. As a
The epididymis secretes a substance (as yet unidentified) that result, plasma levels of FSH, LH, and testosterone remain within
prevents premature capacitation. a relatively narrow range until relatively late in a man's life.
Testosterone plays a major role in maintaining male
sexual function (see Spotlight Figure 28–12). Testosterone
The Anatomy of a Sperm
functions like other steroid hormones, circulating in the blood-
Sperm are among the most highly specialized cells in the body. stream while bound to one of two types of transport proteins.
Each sperm has four distinct regions: head, neck, middle piece, These proteins are (1) gonadal steroid-binding globulin (GBG),
and tail (Figure 28–11). The head is a flattened ellipse contain- which carries about two-thirds of the circulating testosterone,
ing a nucleus with densely packed chromosomes. At the tip of and (2) the albumins, which bind the remaining one-third. Tes-
the head is the acrosome (AK-ro-som), a membranous com-
. .

tosterone diffuses across the plasma membrane of target cells

partment containing enzymes essential to fertilization. During and binds to an intracellular receptor. The hormone–receptor
spermiogenesis, saccules of the spermatid's Golgi apparatus complex then binds to the DNA in the nucleus.
fuse and flatten into an acrosomal vesicle, which ultimately
forms the acrosome.
A short neck attaches the head to the middle piece.
The neck contains both centrioles of the original spermatid. + Clinical Note Dehydroepiandrosterone
The microtubules of the distal centriole are continuous (DHEA)
with those of the middle piece and tail. Mitochondria in the
middle piece are arranged in a spiral around the microtubules. Dehydroepiandrosterone, or DHEA, is the primary
Mitochondria provide the ATP required to move the tail. androgen secreted by the zona reticularis of the adrenal
The tail is the only flagellum in the human body. A flagel- cortex. p. 682 As noted in Chapter 18, these androgens,
lum is a whiplike organelle that moves a cell from one place to which are secreted in small amounts, are converted to tes-
another. Cilia beat in a predictable, wavelike fashion, but the tosterone (or estrogens) by other tissues. The significance
of this small adrenal androgen secretion in both sexes
tail of a sperm has a whip-like, corkscrew motion.
remains unclear, but DHEA is being promoted as a wonder
Unlike other, less specialized cells, a mature sperm lacks
drug for increasing vitality, strength, and muscle mass. The
an endoplasmic reticulum, a Golgi apparatus, lysosomes, per-
effects of long-term high doses of DHEA remain largely
oxisomes, inclusions, and many other organelles. The loss of unknown. However, high levels of DHEA in women have
these organelles reduces the cell's size and mass. A sperm is been linked to an increased risk of ovarian cancer as well
28 essentially a mobile carrier for chromosomes, and extra weight
as to masculinization, due to the conversion of DHEA to
would slow it down. Because a sperm lacks glycogen or other testosterone. The International Olympic Committee, NCAA,
energy reserves, it must absorb nutrients (primarily fructose) MLB, and NFL have banned the use of DHEA.
from the surrounding fluid.

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 SPOTLIGHT
Figure 28–12
Hormonal Regulation of Male Reproduction
Male reproductive function is regulated by the complex
interaction of hormones from the hypothalamus, anterior
lobe of the pituitary gland, and the testes. Negative
feedback systems keep testosterone levels within a HYPOTHALAMUS
relatively narrow range until late in life.
Negative feedback
Release of Gonadotropin-
Releasing Hormone (GnRH)
? The hypothalamus secretes the
High testosterone levels
inhibit the release of
hormone GnRH at a rate that GnRH by the hypothala-
remains relatively steady. As a mus, causing a decrease
result, blood levels of FSH, LH, in LH secretion, which
and testosterone remain within a lowers testosterone to
relatively narrow range through- normal levels.
out a man’s reproductive life.

When stimulated by GnRH, the

anterior lobe of the pituitary gland
releases luteinizing hormone (LH) and
follicle-stimulating hormone (FSH).

Secretion of Follicle- ANTERIOR Secretion of

Stimulating Hormone
(FSH) LOBE OF THE Luteinizing
Hormone (LH)
Inhibin depresses the
pituitary production of FSH targets primarily PITUITARY LH targets the
FSH, and perhaps the
hypothalamic secretion of
the nurse cells of the
seminiferous tubules. GLAND interstitial endocrine
cells of the testes.
gonadotropin-releasing
hormone (GnRH). The
faster the rate of sperm
production, the more
inhibin is secreted. By
Interstitial Endocrine Cell Stimulation
Negative feedback

regulating FSH and GnRH

secretion, nurse cells LH induces the secretion of testosterone
provide feedback control and other androgens by the interstitial
of spermatogenesis. endocrine cells of the testes.
TESTES
Nurse Cell Stimulation Testosterone

Under FSH stimulation, and with

testosterone from the interstitial
endocrine cells, nurse cells (1) secrete
inhibin in response to factors released KEY
Inhibin by developing sperm, (2) secrete
androgen-binding protein (ABP), and Stimulation
(3) promote spermatogenesis and
Inhibition
spermiogenesis.

28

Androgen-binding protein (ABP) Nurse cell Peripheral Effects of Testosterone

binds androgens within the environment
seminiferous tubules, which facilitates both Maintains libido Stimulates Establishes and Maintains accessory
increases the local concentration spermatogenesis (sexual drive) bone and maintains male glands and organs
of androgens and stimulates the and spermiogenesis. and related muscle secondary sex of the male
physical maturation of spermatids. behaviors growth characteristics reproductive system

1117

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 1118 UNIT 6 Continuity of Life

In many target tissues, some of the arriving testosterone is

converted to dihydrotestosterone (DHT). A small amount of + Clinical Note Prostate Cancer
DHT diffuses back out of the cell and into the bloodstream, and
DHT levels are usually about 10 percent of circulating testoster- Prostate cancer is the second most common cancer
one levels. Dihydrotestosterone can also enter peripheral cells and the second most lethal cancer in males. For reasons
and bind to the same hormone receptors targeted by testoster- that are poorly understood, prostate cancer rates for Asian
one. In addition, some tissues (notably those of the external American males are relatively low compared with those of
genitalia) respond to DHT rather than to testosterone. Other either Caucasian Americans or Black Americans. For all
tissues (including the prostate) are more sensitive to DHT than ages and ethnic groups, the rate of prostate cancer is rising
to testosterone. sharply. The reason for the increase is not known. Aggres-
In adult males, negative feedback controls the level of sive diagnosis and treatment of localized prostate cancer in
elderly patients is controversial because many of these men
testosterone production. Above-normal testosterone levels
have nonmetastatic tumors (tumors that have not spread).
inhibit the release of GnRH by the hypothalamus, causing a
In addition, even if their cancer is left untreated, these men
reduction in LH secretion and lowering testosterone levels (see are more likely to die of some other disease.
Spotlight Figure 28–12).
The plasma of adult males also contains relatively small
amounts of the estrogenic hormone estradiol (2 ng/dL versus
525 ng/dL of testosterone). Seventy percent of the estradiol is
formed from circulating testosterone. Interstitial endocrine cells + Clinical Note Prostate-Specific Antigen
and nurse cells of the testes secrete the rest. An enzyme called (PSA) Testing
aromatase converts testosterone to estradiol. For unknown rea-
sons, estradiol production increases in older men. Blood tests are used to screen for prostate cancer and
to monitor the success of treatment. The most sensitive
is a blood test called prostate-specific antigen (PSA).
Checkpoint An elevated level of this antigen, which is normally pres-
ent in a low concentration, may indicate the presence of
7. What effect would a low FSH level have on sperm prostate cancer. Screening with periodic PSA tests is now
production? being recommended for some men over age 50. If prostate
See the blue Answers tab at the back of the book. cancer is detected, treatment decisions vary depending on
how rapidly the PSA level is rising.

+ Clinical Note Enlarged Prostate

28-4 The structures of the female
Enlargement of the prostate, or benign prostatic hyper- reproductive system consist of the
plasia, is so common in men over the age of 50 that it is ovaries, uterine tubes, uterus, vagina,
considered an aspect of aging. The increase in size hap-
pens as testosterone production by the interstitial endocrine
and external genitalia
cells decreases. For unknown reasons, small masses called Learning Outcome Describe the anatomy and functions of
the ovaries, the uterine tubes, the uterus, the vagina, and the
prostatic concretions may form within the glands. At the
structures of the female external genitalia.
same time, the interstitial endocrine cells begin releasing
small quantities of estrogens into the bloodstream. The A woman's reproductive system produces sex hormones and
combination of a lower testosterone level and the presence functional gametes. It must also be able to protect and sup-
of estrogens probably stimulates prostatic growth. In severe port a developing embryo and nourish a newborn infant. The
cases, prostatic swelling constricts and blocks the urethra main organs of the female reproductive system are the ovaries,
and constricts the rectum. If not corrected, the urinary the uterine tubes, the uterus, the vagina, and the components
obstruction can cause permanent kidney damage. Partial of the external genitalia—the clitoris and the labia majora and
28
surgical removal is the most effective treatment. In the pro-
labia minora (Figure 28–13). As in males, a variety of accessory
cedure known as a TURP (transurethral prostatectomy), an
glands—the urethral glands and the greater vestibular glands—
instrument pushed along the urethra cuts away the swollen
release secretions into the female reproductive tract.
prostatic tissue. Most of the prostate remains in place, and
normal function returns. The ovaries, uterine tubes, and uterus are enclosed
within an extensive mesentery known as the broad ligament

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 Chapter 28 The Reproductive System   1119

Figure 28–13 Sagittal Section of the Female Reproductive System. ATLAS: Plate 65

Major Structures
of the Female
Reproductive System
Ovarian follicle
Ovary

Uterine tube
Sigmoid
Recto-uterine colon
pouch Uterus
Vesico-uterine Perimetrium
pouch
Myometrium
Urinary bladder
Endometrium
Urethra Vaginal
fornix
Pubic symphysis Cervix

Accessory Glands Vagina

Urethral glands
Rectum

Greater vestibular
gland
Anus External Genitalia
Labium minus
Labium majus
Clitoris

(Figure 28–14a). The uterine tubes run along the superior bor- broad ligament limits side-to-side movement and rotation, and
der of the broad ligament and open into the pelvic cavity lateral the other ligaments (described in our discussion of the ovaries
to the ovaries. The broad ligament attaches to the sides and and uterus) prevent superior–inferior movement.
floor of the pelvic cavity, where it becomes continuous with the Let's look further at the structures and basic functions of
parietal peritoneum. The broad ligament subdivides this part of the female reproductive system.
the peritoneal cavity.
The pocket formed between the posterior wall of the uterus
and the anterior surface of the colon is the recto-uterine
The Ovaries
(rek-to-YU-ter-in) pouch. The pocket formed between the
.

The paired ovaries are small, lumpy, almond-shaped organs near

.

uterus and the posterior wall of the bladder is the vesico-uterine the lateral walls of the pelvic cavity (Figure 28–14b). The ova-
28
(ves-ih-ko-YU-ter-in) pouch. These subdivisions are easily seen
.

ries perform three main functions. They (1) produce immature

.

in sagittal section (see Figure 28–13). female gametes, or oocytes; (2) secrete female sex hormones,
Several other ligaments assist the broad ligament in sup- including estrogens and progesterone; and (3) secrete inhibin,
porting and stabilizing the uterus and associated reproductive involved in the feedback control of pituitary FSH production.
The mesovarium (mez-o-VA-re-um), a thickened fold of
.

organs. These ligaments lie within the mesentery sheet of the

. .

broad ligament and are connected to the ovaries or uterus. The mesentery, supports and stabilizes the position of each ovary

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 1120 UNIT 6 Continuity of Life

Figure 28–14 Relationships between the Ovaries, Uterine Tubes, and Uterus. ATLAS: Plate 67

Structures Stabilizing the Ovary

Ovarian Mesovarium Suspensory
Ovarian artery Fimbriae Uterine tube
ligament ligament Retractor
and vein

Suspensory Uterus
ligament

Infundibulum

Broad ligament
Ovary

Uterine
Ureter tube
Uterosacral ligament
Corpus
External os luteum
Medulla Cortex
Cervix Tunica
Mesenteries of
Vaginal rugae the Ovary and albuginea
Vaginal wall Uterine Tube

a A posterior view of the uterus, Mesosalpinx

uterine tubes, and ovaries
Mesovarium

Broad ligament Ovarian Egg Germinal Mature

(see Figure 28–14a). Each ovary is also stabilized by a pair of hilum nest epithelium follicle

supporting ligaments: the ovarian ligament and the suspensory b A sectional view of the ovary, uterine
(infundibulopelvic) ligament. The ovarian ligament extends tube, and associated mesenteries
from the uterus, near the attachment of the uterine tube, to the
medial surface of the ovary. The suspensory ligament extends
from the lateral surface of the ovary past the open end of the Figure 28–14b). We divide the interior tissues, or stroma, of
uterine tube to the pelvic wall. The suspensory ligament con- the ovary into a superficial cortex and a deeper medulla. Gam-
tains the major blood vessels of the ovary: the ovarian artery etes are produced in the cortex.
and ovarian vein. These vessels are connected to the ovary at
the ovarian hilum, where the ovary attaches to the mesovar- The Uterine Tubes
ium (see Figure 28–14b).
Each uterine tube (fallopian tube or oviduct) is a hollow, mus-
A typical ovary is about 5 cm long, 2.5 cm wide, and
28 8 mm thick (2 in. * 1 in. * 0.33 in.). It weighs 6–8 g (about cular cylinder measuring roughly 13 cm (5.2 in.) in length (see
Figure 28–14a).
0.25 oz). An ovary is pink or yellowish and has a nodular
consistency. The visceral peritoneum, or germinal epithelium,
covers the surface of each ovary and consists of a layer of Regions of the Uterine Tubes
columnar epithelial cells that overlies a dense connective tis- We divide each uterine tube into three regions, from lateral to
sue layer called the tunica albuginea (al-bu-JIN-e-ah) (see medial (Figure 28–15a):
. .

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 Chapter 28 The Reproductive System   1121

1. The Infundibulum. The end closest to the ovary forms an ex- Functional Anatomy of the Uterine Tube
panded funnel, or infundibulum, with numerous finger- The epithelium lining the uterine tube is composed of cili-
like projections that extend into the pelvic cavity. The pro- ated columnar epithelial cells with scattered mucin-secreting
jections are called fimbriae (FIM-bre-e) (Figure 28–16a).
. .

cells. Concentric layers of smooth muscle surround the mucosa

Fimbriae drape over the surface of the ovary, but there is no (Figure 28–15b). A combination of ciliary movement and peri-
physical connection between the two structures. The inner staltic contractions in the walls of the uterine tube transports
surfaces of the infundibulum are lined with cilia that beat oocytes toward the uterus (Figure 28–15c). In addition, the
toward the middle region of the uterine tube. uterine tube provides a nutrient-rich environment that contains
2. The Ampulla. The middle region is the called the ampulla. lipids and glycogen for both oocytes and sperm. If fertilization
The thickness of its smooth muscle layers gradually occurs, it typically takes place near the boundary between the
increases as the tube approaches the uterus. ampulla and isthmus of the uterine tube.
3. The Isthmus. The ampulla leads to the isthmus (IS-mus), a In addition to ciliated cells, the epithelium lining the uter-
short region connected to the uterine wall. ine tubes contains peg cells. The peg cells project into the lumen
of the uterine tube. They secrete a fluid that both completes the
capacitation of sperm and supplies nutrients to sperm and the
pre-embryo.
+ Clinical Note Ovarian Cancer
The Uterus
Ovarian cancer is not the most common female reproduc- The uterus protects, nourishes, and removes wastes for the
tive cancer—it is the third most common—but it is the most developing embryo (weeks 1–8) and fetus (week 9 through
dangerous, because it is so difficult to diagnose in its early delivery). In addition, contractions of the muscular uterus are
stages. A woman in the United States has a 1-in-70 chance important in expelling the fetus at birth.
of developing ovarian cancer in her lifetime. The prognosis
The uterus is a small, pear-shaped organ (Figure 28–16).
is relatively good for cancers that originate in the general
It is about 7.5 cm (3 in.) long with a maximum diameter of
ovarian tissues or from abnormal oocytes. These cancers
5 cm (2 in.). It weighs 50–100 g (1.75–3.5 oz). In its nor-
generally respond well to some combination of chemother-
apy, radiation, and surgery. However, 85 percent of ovarian mal position, the uterus bends anteriorly near its base, a con-
dition known as anteflexion (an-te-FLEK-shun) (look back at
.

cancers develop from epithelial cells, and sustained remis-

sion occurs in only about one-third of these cases. Figure 28–13). In this position, the uterus covers the superior
and posterior surfaces of the urinary bladder. If the uterus bends

Figure 28–15 Anatomy of the Uterine Tubes.

Ampulla Isthmus

Microvilli
Uterus of mucin-
secreting
Fimbriae cells
Infundibulum

Columnar
epithelium

Lamina
a Regions of the uterine
propria
tubes, posterior view
Cilia
28
Smooth
muscle

Isthmus LM × 122 Epithelial surface SEM × 4000

b A sectional view of the isthmus c A colorized SEM of the ciliated

lining of the uterine tube

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 1122 UNIT 6 Continuity of Life

Figure 28–16 Gross Anatomy of the Uterus. ATLAS: Plates 66; 67

Uterine Tube
Fimbriae Infundibulum Ampulla Isthmus

Fundus Uterine tube Suspensory

of uterus ligament of ovary
Ovarian artery
and vein

Body of Mesovarium
uterus Ovary
Ovarian ligament

Uterine
cavity Round ligament
Perimetrium
of uterus
Myometrium Broad ligament
Endometrium
See Figure 28–17

Uterine artery
and vein
Internal os of uterus
Isthmus of uterus
Cervical canal Cervix

Vaginal artery

External os
of uterus
a A posterior view with the
left portion of the uterus,
Vaginal rugae
left uterine tube, and left
ovary shown in section. See Figure 28–18 Vagina

Uterosacral POSTERIOR
ligament POSTERIOR
Cardinal Sigmoid
ligaments colon
(under broad
ligament)

Suspensory Recto-uterine
ligament pouch
of ovary
Ovary
Broad
Ovarian
ligament
ligament
Uterus
Uterine tube Uterus
Round
ligament Vesico-uterine
of uterus pouch
Urinary
bladder ANTERIOR ANTERIOR
28
c Superior view of the female pelvic cavity showing supporting
b A superior view of the ligaments that stabilize the ligaments of uterus and ovaries. In the photo, the urinary
position of the uterus in the pelvic cavity. bladder cannot be seen because it is covered by peritoneum.

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 Chapter 28 The Reproductive System   1123

backward toward the sacrum, the condition is termed retroflex- The tubular cervix projects about 1.25 cm (0.5 in.) into the
ion (re-tro-FLEK-shun). Retroflexion occurs in about 20 percent vagina. Within the vagina, the distal end of the cervix forms a
.

of adult women and has no clinical significance. (A retroflexed curving surface that surrounds the external os (os, an opening
uterus generally becomes anteflexed spontaneously during the or mouth) of the uterus. The external os leads into the cervical
third month of pregnancy.) canal, a constricted passageway that opens into the uterine
cavity of the body at the internal os.
Suspensory Ligaments of the Uterus The uterus receives blood from branches of the uterine
In addition to the broad ligament, three pairs of suspensory arteries. These vessels arise from branches of the internal iliac
ligaments stabilize the uterus and limit its range of move- arteries, and from the ovarian arteries, which arise from the
ment (Figure 28–16b). The uterosacral (yu-te-ro-SA-krul) abdominal aorta inferior to the renal arteries. The arteries to the
.

. .

ligaments extend from the lateral surfaces of the uterus to the uterus are extensively interconnected, ensuring a reliable flow
anterior face of the sacrum, keeping the body of the uterus from of blood to the organ despite changes in its position and shape
moving inferiorly and anteriorly. The round ligaments arise during pregnancy. Numerous veins and lymphatic vessels also
on the lateral margins of the uterus just posterior and inferior drain each portion of the uterus.
to the attachments of the uterine tubes. These ligaments extend The organ is innervated by autonomic fibers from the
through the inguinal canal and end in the connective tissues hypogastric plexus (sympathetic) and from sacral segments S3
of the external genitalia. (Unlike the inguinal canals in males, and S4 (parasympathetic). Sensory information reaches the cen-
these canals in females are very small, containing only the tral nervous system (CNS) within the posterior roots of spinal
round ligaments and the ilioinguinal nerves; this makes inguinal nerves T11 and T12. The most delicate anesthetic procedures used
hernias in women very rare.) The round ligaments restrict pos- during labor and delivery, known as segmental blocks, target only
terior movement of the uterus. The cardinal ligaments extend spinal nerves T10–L1.
from the base of the uterus and vagina to the lateral walls of
the pelvis. These ligaments tend to prevent inferior movement The Uterine Wall
of the uterus. The muscles and fascia of the pelvic floor provide The dimensions of the uterus are highly variable. In women of
additional mechanical support. reproductive age who have not given birth, the uterine wall is
about 1.5 cm (0.6 in.) thick.
Gross Anatomy of the Uterus
Layers of the Uterine Wall. The uterine wall is made up
We divide the uterus into four main anatomical regions
of three main layers. The fundus and the posterior surface of
(Figure 28–16a). The uterine body is the largest portion of the
the uterine body and isthmus are covered by an outer serous
uterus. The fundus is the rounded portion of the body supe-
membrane that is continuous with the peritoneal lining. This
rior to the attachment of the uterine tubes. The body ends at
incomplete serosa is called the perimetrium. The wall has
a constriction known as the isthmus of the uterus. The cervix
a thick, middle, muscular myometrium (mı-o-ME-tre-um;
.

. . .

(SER-viks) is the inferior portion of the uterus that extends

myo-, muscle + metra, uterus) and a thin, inner, glandular
from the isthmus to the vagina.
endometrium (en-do-ME-tre-um) (Figure 28–17a).
.

. .

The myometrium is the thickest portion of the uterine

+ Clinical Note Pap Smear wall. It makes up almost 90 percent of the mass of the uterus.
Smooth muscle in the myometrium is arranged into longitudi-
nal, circular, and oblique layers. The smooth muscle tissue of
For many young women, the Pap smear is the reason for the myometrium provides much of the force needed to move a
a regular visit to the doctor. This important screening test fetus out of the uterus and into the vagina.
evaluates the health of the cervix, the inferior portion of the
The endometrium makes up about 10 percent of the mass
uterus that projects into the vaginal canal. Early detection of
of the uterus. The glandular and vascular tissues of the endo-
cervical cancer, the most common reproductive cancer in
metrium support the physiological demands of the growing
women ages 15–34, is the main goal of the test. Developed
by Dr. Georgios Papanikolaou (hence the name of the test), fetus. Vast numbers of uterine glands open onto the endometrial
the simple, quick, and inexpensive Pap smear is a form of surface and extend deep into the lamina propria, almost to the 28
exfoliative cytology. The practitioner opens the patient's myometrium. Under the influence of estrogens, the uterine
vagina with a speculum, and gently scrapes the cervix glands, blood vessels, and epithelium change with the phases
with a swab to collect epithelial cells. These cells are then of the monthly uterine cycle.
studied in the pathology lab for any evidence of cancerous
changes. Histology of the Uterine Wall. The endometrium is divid-
ed into a functional layer—the layer closest to the uterine

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 1124 UNIT 6 Continuity of Life

Figure 28–17 Histology of the Uterine Wall.

Endometrium

Straight artery Simple Uterine Functional Basilar

columnar glands layer layer Myometrium
Perimetrium epithelium
Myometrium
Endometrium
Uterine glands
Uterine cavity
Spiral artery

Arcuate arteries
Uterine
Radial artery cavity

Uterine artery

Uterine wall LM × 32

a A diagrammatic sectional view of the uterine b The basic histological structure of the
wall, showing the endometrial regions and the uterine wall
blood supply to the endometrium

cavity—and a basal layer, adjacent to the myometrium Gross Anatomy of the Vagina
(Figure 28–17b). The functional layer contains most of the The vagina extends between the cervix and the body exterior
uterine glands and contributes most of the endometrial thick- (Figure 28–18a). At the proximal end of the vagina, the cervix
ness. This layer undergoes dramatic changes in thickness and projects into the vaginal canal. The shallow recess surround-
structure during the uterine cycle. The basal layer attaches the ing the cervical protrusion is known as the vaginal fornix
endometrium to the myometrium and contains the terminal (FOR-niks). The vagina is typically 7.5–9 cm (3–3.6 in.) long.
branches of the tubular uterine glands. The structure of the basal Its diameter varies because it is highly distensible.
layer remains fairly constant over time, but the functional layer The two bulbospongiosus muscles extend along either side
undergoes cyclical changes in response to sex hormone levels. of the vaginal entrance. Contractions of the bulbospongiosus
Within the myometrium, branches of the uterine arter- muscles constrict the vagina. p. 407 These muscles cover
ies form arcuate arteries, which encircle the endometrium the bulb of vestibule, a mass of erectile tissue on each side of
(see Figure 28–17a). Radial arteries supply straight arteries, the vagina (Figure 28–19). The bulb of vestibule has the same
which deliver blood to the basal layer of the endometrium, and embryonic origins as the corpus spongiosum of the penis in
spiral arteries, which supply the functional layer. males.
The vagina opens into the vestibule, a space bordered by
28
The Vagina the female external genitalia (see Figure 28–18a). The hymen
.

The vagina is an elastic, muscular tube that has three major (HI-men) is an elastic epithelial fold of variable size that par-
functions. It (1) is a passageway for the elimination of men- tially blocks the entrance to the vagina. An intact hymen is typi-
strual fluids; (2) receives the penis during sexual intercourse, cally stretched or torn during first sexual intercourse, tampon
and holds sperm prior to their passage into the uterus; and use, pelvic examination, or physical activity.
(3) forms the inferior portion of the birth canal, through which The vagina lies parallel to the rectum, and the two are in
the fetus passes during delivery. close contact posteriorly. Anteriorly, the urethra extends along

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 Chapter 28 The Reproductive System   1125

Figure 28–18 Anatomy of the Vagina.

Stratified
Lumen of squamous
vaginal epithelium Lamina Blood Smooth
canal (nonkeratinized) propria vessels muscle fibers

Vaginal fornix

Vaginal artery
Vaginal vein

Rugae

Vaginal
canal

Hymen
Greater
vestibular gland

Labium minus Vestibule

The vaginal wall LM × 25

a Vaginal structures b Histology of vagina

the superior wall of the vagina from the urinary bladder to the
external urethral orifice, which opens into the vestibule.
Figure 28–19 Anatomy of the Female External Genitalia. The primary blood supply of the vagina is by the vaginal
branches of the internal iliac (or uterine) arteries and veins.
External Innervation is from the hypogastric plexus, sacral nerves S2–S4,
Genitalia and branches of the pudendal nerve. pp. 497, 810, 817
Mons pubis

Clitoris Histology of the Vagina

Prepuce In sectional view, the lumen of the vagina appears constricted,
Glans forming a rough H-shape. The vaginal walls contain a network
Vestibule
of blood vessels and layers of smooth muscle (Figure 28–18b).
Urethral opening The lining is moistened by secretions of the cervical glands and
Labia
minora
Bulb of vestibule by the movement of water across the permeable epithelium.
Vaginal entrance The vaginal lumen is lined by a nonkeratinized stratified
Hymen (torn)
squamous epithelium (see Figure 28–18b). In the relaxed state,
Labia Greater vestibular .

this epithelium forms folds called rugae (RU-ge). The underly-

.

majora gland
ing lamina propria is thick and elastic, and it contains small
blood vessels, nerves, and lymph nodes. The vaginal mucosa
is surrounded by an elastic muscular layer consisting of layers 28
Anus of smooth muscle fibers arranged in circular and longitudinal
bundles continuous with the uterine myometrium. The portion
of the vagina adjacent to the uterus has a serosal covering that
is continuous with the pelvic peritoneum. Along the rest of the
vagina, the muscular layer is surrounded by an adventitial layer
of fibrous connective tissue.

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 1126 UNIT 6 Continuity of Life

The vagina contains a population of resident bacteria, usu- labia majora and the mons pubis are covered with coarse hair,
ally harmless, supported by nutrients in the cervical mucus. but the inner surfaces of the labia majora are hairless. Seba-
The metabolic activity of these bacteria creates an acidic ceous glands and scattered apocrine sweat glands secrete onto
environment, which restricts the growth of many pathogens. the inner surface of the labia majora, moistening and lubricat-
However, this acidic environment also inhibits the motility of ing them.
sperm.
Vaginitis (vaj-ih-NI-tis), an inflammation of the vagina, is The Breasts
.

caused by fungal, bacterial, or parasitic infections. In addition

Female breasts are two projections anterior to the pectoral
to any discomfort that may result, the condition may affect the
muscles that include a mammary gland, a variable amount
survival of sperm and thereby reduce fertility.
of fat, a nipple, and an areola. Breasts are rudimentary in the
The hormonal changes associated with the ovarian cycle, the
male. A newborn infant cannot fend for itself, and several of its
monthly cycle that produces an oocyte, also affect the vaginal
key systems have yet to complete development. Over the initial
epithelium. By examining a vaginal smear—a sample of epi-
period of adjustment to an independent existence, the infant is
thelial cells shed at the surface of the vagina—a clinician can
nourished from the milk secreted by the maternal mammary
estimate the corresponding stages in the ovarian and uterine
glands. Milk production, or lactation (lak-TA-shun), takes
.

cycles. This diagnostic procedure is an example of exfoliative

place in these glands. In females, mammary glands are special-
cytology. p. 172
ized organs of the integumentary system that are controlled
mainly by hormones of the reproductive system and by the
The Female External Genitalia placenta, a temporary structure that provides the embryo and
The area containing the female external genitalia is the vulva fetus with nutrients.
(VUL-vuh), or pudendum (pu-DEN-dum; see Figure 28–19). The On each side, a mammary gland lies in the subcutaneous
.

vagina opens into the vestibule, a central space surrounded by tissue of the pectoral adipose tissue deep to the skin of the
small folds known as the labia minora (LA-be-uh mi-NOR- chest (Figure 28–20a). Each breast has a nipple, a small conical
.

uh; singular, labium minus). The labia minora are covered with projection where the ducts of the underlying mammary gland
smooth, hairless skin. The urethra opens into the vestibule just open onto the body surface. The reddish-brown skin around
anterior to the vaginal entrance. The urethral glands discharge each nipple is the areola (a-RE-o-luh). Large sebaceous glands
.

into the urethra near the external urethral opening. deep to the areolar surface give it a grainy texture.
Anterior to this opening, the clitoris (KLIT-o-ris) projects The glandular tissue of a mammary gland consists of
.

into the vestibule. A small, rounded tissue projection, the separate lobes, each containing several secretory lobules.
clitoris is derived from the same embryonic structures as the Ducts leaving the lobules converge, giving rise to a single
penis in males, and has a major role in female sexual response. lactiferous (lak-TIF-er-us) duct in each lobe. Near the
Internally, it contains erectile tissue called corpus caverno- nipple, that lactiferous duct enlarges, forming an expanded
sum of clitoris (comparable to those structures of the penis) chamber called a lactiferous sinus. Typically, 15–20 lactif-
that form its body. A small erectile glans sits atop the body. erous sinuses open onto the surface of each nipple. Dense
The bulb of vestibule along the sides of the vestibule are connective tissue surrounds the duct system and forms parti-
comparable to the male corpus spongiosum. These erectile tions that extend between the lobes and the lobules. These
tissues engorge with blood during sexual arousal. Extensions bands of connective tissue, the suspensory ligaments of the
of the labia minora encircle the body of the clitoris, forming breast, originate in the dermis of the overlying skin. A layer
its prepuce. of areolar tissue separates the mammary gland complex from
A variable number of small lesser vestibular glands dis- the underlying pectoralis muscles. Branches of the internal
charge their secretions onto the exposed surface of the vestibule thoracic artery supply blood to each mammary gland (look
between the orifices of the vagina and urethra. During sexual back at Figure 21–20, p. 806).
arousal, the greater vestibular glands, located on each side of Figure 28–20b,c compares the histology of inactive and
the distal portion of the vagina, secrete into the vestibule. These active mammary glands. An inactive, or resting, mammary gland
mucous glands keep the area moist and lubricated. The ves- is dominated by a duct system rather than by active glandu-
28
tibular glands have the same embryonic origins as the bulbo- lar cells. The size of the mammary glands in a nonpregnant
urethral glands in males. woman reflects primarily the amount of adipose tissue present,
The outer margins of the vulva are formed by the mons not the amount of glandular tissue. The secretory apparatus
pubis and the labia majora. The mons pubis is a pad of adi- normally does not complete its development unless pregnancy
pose tissue covering the pubic symphysis. Adipose tissue also occurs. An active mammary gland is a tubulo-alveolar gland,
accumulates within the labia majora (singular, labium majus), consisting of multiple glandular tubes that end in secretory
prominent folds of skin that encircle and partially conceal the alveoli. We will discuss the hormonal mechanisms involved in
labia minora and adjacent structures. The outer margins of the lactation in Chapter 29.

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 Chapter 28 The Reproductive System   1127

Figure 28–20 Anatomy of the Breast. ATLAS: Plate 28

Secretory alveoli

Pectoralis major Lactiferous duct

Connective
tissue
Adipose tissue
Resting mammary gland LM × 100
Suspensory
ligaments b An inactive mammary gland of a
nonpregnant woman

Lobules of two
lobes of the
mammary gland

Lactiferous duct
Secretory alveoli
Areola
Nipple

Lactiferous duct
Lactiferous sinus
Milk

Active mammary gland LM × 131

a The mammary gland of the left breast c An active mammary gland of a nursing woman

+ Clinical Note Fibrocystic Disease and Breast Cancer

The uterine cycle also influences cyclical changes in the mam- Notable risk factors include (1) a family history of breast cancer,
mary glands. Occasional discomfort or even inflammation of (2) a first pregnancy after age 30, and (3) early menarche (first
mammary gland tissues can occur late in the cycle. If inflamed menstruation) or late menopause (end of menstruation).
lobules become walled off by scar tissue, cysts are created. Despite repeated studies, no links have been proven
Clusters of cysts can be felt in the breast as discrete masses, a between breast cancer and birth control pills, fat consump-
condition known as fibrocystic disease. Because its symptoms tion, or alcohol use. It appears likely that multiple factors are
are similar to those of breast cancer, biopsies may be needed to involved. In some families an inherited genetic variation has 28
distinguish between this benign condition and breast cancer. been linked to higher-than-normal risk of developing the dis-
Breast cancer is a malignant, metastasizing tumor of ease. However, most women never develop breast cancer—
the mammary gland. It is the leading cause of death in women even women in families with a history of the disease. Mothers
between ages 35 and 45, but it is most common in women over who breast-fed (nursed) their babies have a 20 percent lower
age 50. An estimated 12.4 percent of U.S. women will develop incidence of breast cancer after menopause than do mothers
breast cancer at some point in their lifetime. The incidence is who did not breast-feed. The reason is not known. Adding to
highest among Caucasian Americans, somewhat lower in Black the mystery, nursing does not appear to affect the incidence of
Americans, and lowest in Asian Americans and Native Americans. premenopausal breast cancer.

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 1128 UNIT 6 Continuity of Life

28-5 Oogenesis occurs in the ovaries,

+ Clinical Note Laparoscopy and hormones from the hypothalamus,
pituitary gland, and ovaries control
Laparoscopy is a minimally invasive procedure that allows
a direct look into the abdominopelvic cavity. The technique
female reproductive functions
is often employed to assess the health of female pelvic Learning Outcome Describe the ovarian roles in oogenesis,
structures when another diagnostic test, such as an x-ray, explain the complete ovarian and uterine cycles, and summarize all
aspects of the female reproductive cycle.
is inconclusive. The surgeon makes a tiny incision through
the abdominal wall when the patient is under general anes- Three interrelated processes help to ensure the continuation of
thesia. Through this opening, carbon dioxide gas is injected our species from the female perspective: oogenesis, the ovar-
into the abdomen in order to spread out the organs and ian cycle, and the uterine cycle. Oogenesis is the production
increase visibility. Then the laparoscope is inserted. The of female gametes called oocytes. Oogenesis takes place in
scope is a flexible tube bearing a tiny light and an equally the ovaries within structures called follicles. The ovarian cycle
tiny camera. The operator can look around, take photos,
is the monthly series of events associated with the maturation
and collect biopsies. Laparoscopy can determine, for
of an oocyte. The uterine cycle involves a series of events that
example, the presence of a blocked uterine tube or a cyst
prepares the uterus for implantation of a fertilized oocyte. This
on an ovary.
section describes these events and how they are influenced by
hormones.

+ Clinical Note Mammoplasty Oogenesis

Ovum production, or oogenesis (o-o-JEN-eh-sis; oon, egg),
. .

Mammoplasty includes three types of surgical altera- begins before a woman's birth, accelerates at puberty, and ends
tion of the breast. Breast augmentation is most frequently at menopause (end of menstruation). Between puberty and
desired. In this procedure, the size or contour of the mam- menopause, oogenesis occurs on a monthly basis as part of the
mary gland is modified by the placement of fluid-filled ovarian cycle.
implants under the skin, and either superficial or deep to Oogenesis is summarized in Figure 28–21. This process
the pectoral muscles. The surgeon may be able to insert begins during fetal development, when female reproductive
the implant through an incision along the margin of the
stem cells called oogonia (o-o-GO-ne-uh; singular, oogo-
.

. . .

areola of the nipple. A second approach is to enter from

nium) undergo mitosis, producing diploid primary oocytes
the axilla. A third option is to make a small incision directly
underneath each breast. Breast reduction can be per- (2n). After 5 months of development, fetal ovaries contain
formed when a woman desires that cosmetic result, or an estimated 7 million primary oocytes. Not all survive.
when she is suffering back and neck pain due to excessive At birth the ovaries have approximately 2 million primary
breast size. Finally, women who have undergone surgery oocytes. However, most of these primary oocytes do not
for breast cancer may seek breast reconstruction. As a survive until puberty, degenerating in a process called atresia
case in point, actor and activist Angelina Jolie went pub-
.

(ah-TRE-ze-uh). By puberty, their number has dropped to

.

lic in 2013 with her prophylactic bilateral mastectomy, or about 400,000.

removal of the breasts in response to a high risk of breast The remaining primary oocytes begin meiosis I, proceed
cancer. as far as prophase I, and then remain in that state until the
female reaches puberty. At that point, some are stimulated to
finish meiosis I, producing haploid secondary oocytes (n).
Note that the cytoplasm of the primary oocyte is unevenly dis-
Checkpoint
tributed during the two meiotic divisions (see Figure 28–21).
8. Name the structures of the female reproductive system. Oogenesis produces one secondary oocyte, which contains
9. What effect would blockage of both uterine tubes by scar most of the original cytoplasm, and two or three polar bodies,
tissue (resulting from an infection such as gonorrhea) nonfunctional cells that later disintegrate.
28
have on a woman's ability to conceive?
The secondary oocyte then begins meiosis II, but is sus-
10. What benefit does the acidic pH of the vagina provide?
pended at metaphase II. In the process of ovulation, the ovary
11. Would the blockage of a single lactiferous sinus interfere releases such a secondary oocyte. This secondary oocyte will
with the delivery of milk to the nipple? Explain.
not complete meiosis II and become a mature ovum unless
See the blue Answers tab at the back of the book. fertilization occurs.

M28_MART9867_11_GE_C28.indd 1128 23/08/17 10:29 AM

 Chapter 28 The Reproductive System   1129

Figure 28–21 The Process of Oogenesis. For clarity, maternal and paternal chromatids are 1 Primordial Ovarian Follicles in Egg
not identified. Nest. Primary oocytes are located
in the outer portion of the ovarian
cortex, near the tunica albuginea,
Oogenesis
in clusters called egg nests. A single
squamous layer of follicle cells sur-
Mitosis of oogonium
Oogonium rounds each primary oocyte within
Unlike spermatogonia, the oogonia
(stem cell) an egg nest. The primary oocyte and
(ō-ō-GŌ-nē-uh), or female reproduc-
tive stem cells, complete their mitotic its follicle cells form a primordial
divisions before birth. ovarian follicle. At puberty, each
Oogonium Primary
oocyte ovary contains about 200,000 pri-
(diploid, 2n) mordial follicles. Forty years later,
few if any primary ovarian follicles
Meiosis I DNA replication remain, although only about 500
Between the third and seventh months of fetal secondary oocytes will have been
development, the daughter cells, or primary Primary ovulated.
oocytes (Ō-ō-sīts), prepare to undergo meiosis. oocyte
Tetrad Beginning at puberty, primor-
They proceed as far as the prophase of meiosis I,
but then the process comes to a halt. dial ovarian follicles are continu-
ously activated to join other follicles
First polar
The primary oocytes remain in a state of body
already in develop­ ment. The acti-
suspended development until puberty, vating mechanism is unknown, but
when rising levels of FSH trigger the start
of the ovarian cycle. Each month after the
local hormones or growth factors
ovarian cycle begins, some of the Secondary within the ovary may be involved.
primary oocytes are stimulated to oocyte The activated primordial ovarian
undergo further development. Meiosis I
First polar body follicle will either eventually mature
is then completed, yielding a first polar may not complete
body and a secondary oocyte. and release a secondary oocyte or de-
meiosis II
generate through follicular atresia.
2 Formation of Primary Ovarian Fol-
Meiosis II Secondary oocyte
released (ovulation) in licle. The preliminary steps in fol-
Each month after the ovarian cycle metaphase of meiosis II licle development vary in length but
begins, usually one secondary
oocyte leaves the ovary suspended may take almost a year to complete.
in metaphase of meiosis II. Follicle development begins with
the activation of primordial ovarian
At the time of fertilization, a second
Second
Sperm follicles into primary ovarian folli-
polar
polar body forms and the fertilized body
(n) cles. The follicular cells enlarge, di-
secondary oocyte is then called a vide, and form several layers of cells
mature ovum. (A cell in any of the Nucleus
of oocyte around the growing primary oocyte.
preceding steps in oogenesis is
sometimes called an immature (n) The cells begin to produce sex hor-
ovum.) mones called estrogens. Microvilli
from the surrounding follicle cells
intermingle with microvilli originat-
Fertilization ing at the surface of the oocyte. This
(see Figure 29–1)
region is called the zona pellucida
. .

(ZO-na pe-LU-sid-uh; pellucidus,

translucent). The microvilli increase
28
Follicle Development the surface area available for the transfer of materials from the
Specialized structures in the cortex of the ovaries called ovarian follicular cells to the growing oocyte.
follicles (o-VAR-e-an FOL-i-klz) are the sites of both oocyte
. .

3 Formation of Secondary Ovarian Follicle. Many primordial

growth and meiosis I of oogenesis. Follicle development and ovarian follicles develop into primary ovarian follicles, but
the ovarian cycle are shown in Figure 28–22. only a few mature further in the next four months or so. This

M28_MART9867_11_GE_C28.indd 1129 23/08/17 10:29 AM

 1130 UNIT 6 Continuity of Life

Figure 28–22 Follicle Development and the Ovarian Cycle. The photomicrographs show the changes in an ovarian follicle as it devel-
ops. It enters the 28-day ovarian cycle as a tertiary ovarian follicle. The drawing of the ovary illustrates the sequence and relative sizes of the various
stages in the development, ovulation, and degeneration of an ovarian follicle. Follicles do not physically move around the periphery of the ovary.

1 Primordial ovarian follicles 2 Formation of primary

ovarian follicle
3 Formation of secondary
ovarian follicle
in egg nest

Granulosa Thecal cells

cells
Zona
Primary Primary pellucida
oocyte oocytes
Nucleus
Follicle of primary
cells Zona oocyte
pellucida Granulosa
Thecal cells
cells
LM × 270 LM × 270 LM × 245

Primordial Primary Secondary Tertiary

ovarian follicles ovarian follicle ovarian follicle ovarian follicle
4 Formation of
tertiary ovarian follicle

Antrum
containing
Released follicular
secondary fluid
oocyte Granulosa
cells

Corona
radiata

Secondary
oocyte
Corona radiata
LM × 80
Corpus albicans Corpus luteum

7 Formation of 6 Formation of 5 Ovulation

corpus albicans corpus luteum

Follicular
fluid

Secondary
oocyte within
corona radiata
Ruptured
28 follicle wall
Outer
surface
of ovary
LM × 170
LM × 25 LM × 25

? At what stage of development is the oocyte during ovulation?

M28_MART9867_11_GE_C28.indd 1130 23/08/17 10:29 AM

 Chapter 28 The Reproductive System   1131

process is apparently under the control of a growth factor pro- oocyte begins meiosis II but stops short of dividing. Meiosis II
duced by the oocyte. The transformation begins as the wall of will not be completed unless fertilization occurs.
the follicle thickens. At this stage, the complex is known as a Generally, on day 14 of a 28-day ovarian cycle, the
secondary ovarian follicle. The primary oocyte continues to secondary oocyte and its surrounding follicular cells lose their
grow slowly. connections with the follicular wall and drift free within the
4 Formation of Tertiary Ovarian Follicle. During the next 2 to antrum. The granulosa cells still associated with the secondary
3 months, the follicle wall continues to grow and the deeper oocyte form a protective layer known as the corona radiata
.

(ko-RO-nuh ra-de-AH-tuh).
. . .

follicular cells begin secreting follicular fluid. This fluid accumu-

lates in small pockets that gradually expand between the inner 5 Ovulation. At ovulation (ov-u-LA-shun), the tertiary fol-
.

. .

and outer cellular layers of the follicle. The secondary ovarian licle releases the secondary oocyte. The distended follicular
follicle as a whole has doubled in size, and is now called a ter- wall then ruptures, releasing the follicular contents, including
tiary ovarian follicle. Tertiary ovarian follicles, also called vesicular the secondary oocyte, into the pelvic cavity. The sticky fol-
follicles, continue to grow and accumulate follicular fluid. licular fluid keeps the corona radiata attached to the surface of
the ovary near the ruptured wall of the follicle. The oocyte is
The Ovarian Cycle then moved into the uterine tube by contact with the fimbriae
that extend from the tube's funnel-like opening (look back at
The tertiary follicles are then ready to complete their matura-
Figure 28–14a), or by fluid currents produced by the cilia that
tion as part of the 28-day ovarian cycle. The monthly process of
line the tube.
the maturation, ovulation, and degeneration of a tertiary ovar-
Usually only a single oocyte is released into the pelvic cav-
ian follicle is known as the ovarian cycle (see Figure 28–22).
ity at ovulation, although many primordial ovarian follicles
We can divide the ovarian cycle into a follicular phase (follicu-
may have developed into primary ovarian follicles, and several
logenesis) and a luteal phase (luteogenesis), each lasting about
primary ovarian follicles may have been converted to secondary
14 days. The luteal phase involves the follicle after ovulation,
ovarian follicles. These follicles undergo follicular atresia.
when it becomes an endocrine structure called a corpus luteum
.

(LU-te-um) that secretes a sex hormone.

.

The Luteal Phase

The Follicular Phase After ovulation occurs, the luteal phase, or postovulatory phase,
begins. This phase ends with the degeneration of the corpus
In the follicular phase, or preovulatory phase, a tertiary ovarian
luteum.
follicle is ready to complete its maturation. Ovulation marks
the end of this phase. 6 Formation of Corpus Luteum. The empty tertiary ovarian follicle
At the start of each ovarian cycle, an ovary contains only a initially collapses, and ruptured vessels bleed into the antrum.
few tertiary follicles destined for further development. Stimu- Under the stimulation of LH, the remaining granulosa cells then
lated by FSH, one of these follicles becomes dominant by day invade the area and proliferate to create the corpus luteum.
5 of the cycle and it rapidly increases in size. This remaining
The corpus luteum is so named (lutea, yellow) because
tertiary ovarian follicle, also called a mature graafian (GRAF-
this is the color of the cholesterol it contains. This cholesterol
e-an) follicle, is roughly 15–20 mm (0.6–0.8 in) in diameter. It
.

is used to manufacture the steroid hormone progesterone

is formed by days 10–14 of the ovarian cycle. Its large size cre-
(pro-JES-ter-on). Progesterone is the main hormone during
. .

ates a prominent bulge in the surface of the ovary. The oocyte

the luteal phase. Progesterone's primary function is to prepare
and its covering of follicular cells projects into the expanded
the uterus for pregnancy by stimulating the maturation of the
central chamber of the follicle, the antrum (AN-trum). The
uterine lining and the secretions of uterine glands. The corpus
antrum is surrounded by a mass of granulosa cells. As the
luteum also secretes moderate amounts of estrogens, but levels
granulosa cells enlarge and multiply, adjacent cells in the
are not as high as they were at ovulation.
ovarian stroma form a layer of thecal endocrine cells (theca,
a box) around the follicle. Thecal endocrine cells and granu- 7 Formation of Corpus Albicans. The corpus luteum begins to
losa cells work together to produce the sex hormones called degenerate about 12 days after ovulation (unless fertilization
estrogens. takes place). Progesterone and estrogen levels then decline 28
Until this time, the primary oocyte has been suspended markedly. Fibroblasts invade the nonfunctional corpus luteum,
in prophase of meiosis I. As the development of the remain- producing a knot of pale scar tissue called a corpus albicans
ing tertiary ovarian follicle ends, rising LH levels prompt the (AL-bi-kanz).
primary oocyte to complete meiosis I. The completion of the The disintegration of the corpus luteum marks the end of
first meiotic division produces a secondary oocyte and a small, the ovarian cycle. A new ovarian cycle then begins with another
nonfunctional polar body (see Figure 28–21). The secondary group of tertiary ovarian follicles.

M28_MART9867_11_GE_C28.indd 1131 23/08/17 10:29 AM

 1132 UNIT 6 Continuity of Life

The Uterine (Menstrual) Cycle age 45–55. Over the interim, the regular appearance of uterine
cycles is interrupted only by circumstances such as illness, stress,
The uterine cycle, or menstrual (MEN-stru-ul) cycle, is a repeating starvation, or pregnancy.
.

series of changes in the structure of the endometrium (Figure 28–23). The uterine cycle is divided into three phases: (1) the men-
The uterine cycle averages 28 days in length, but it can range from strual phase, (2) the proliferative phase, and (3) the secretory phase.
21 to 35 days in healthy women of reproductive age. The phases take place in response to hormones associated with
The uterine cycle begins at puberty. The first cycle, known the regulation of the ovarian cycle. The menstrual and prolif-
as menarche (me-NAR-ke; men, month + arche, beginning), erative phases occur during the follicular phase of the ovarian
.

typically occurs at age 11–12. The cycles continue until meno- cycle. The secretory phase corresponds to the luteal phase of the
pause (MEN-o-pawz), the termination of the uterine cycle, at ovarian cycle.
.

Figure 28–23 A Comparison of the Structure of the Endometrium during the Phases of the Uterine Cycle.

Uterine
glands Uterine cavity
Perimetrium

Endometrium
Uterine
cavity

Myometrium

Detail of LM × 150
Cervix uterine glands

Uterine Uterine cavity Uterine Uterine cavity

glands glands

Basal layer Functional layer Functional layer

of endometrium

ENDOMETRIUM

Basal layer

MYOMETRIUM MYOMETRIUM
28
Menstrual phase LM × 63 Proliferative phase LM × 66 Secretory phase LM × 52

a The endometrium during the b The endometrium during the c The endometrium during the
menstrual phase of the uterine cycle proliferative phase of the uterine cycle secretory phase of the uterine cycle

? Which layer of the endometrium is affected most during the uterine cycle?

M28_MART9867_11_GE_C28.indd 1132 23/08/17 10:29 AM

 Chapter 28 The Reproductive System   1133

The Secretory Phase

Tips & Tools
During the secretory phase of the uterine cycle, the uterine
Progesterone literally means a steroid (-one) that favors glands enlarge, accelerating their rates of secretion. The arteries
(pro-) gestation (-gest). that supply the uterine wall elongate and spiral through the
tissues of the functional layer (see Figure 28–23c). This activity
Menstrual Phase occurs under the combined stimulatory effects of progesterone
The uterine cycle begins with the menstrual phase. This phase and estrogens from the corpus luteum. The secretory phase
is marked by the degeneration and sloughing (shedding) of begins at the time of ovulation and persists as long as the corpus
the endometrial functional layer, leading to menstruation luteum remains intact.
(men-stru-A-shun), also called menses (MEN-sez) (see
.

Secretory activities peak about 12 days after ovulation.

. .

Figure 28–23a). Endometrial degeneration occurs in patches. Over the next day or two, the glands become less active. The
It is caused by constriction of the spiral arteries, which reduces uterine cycle ends as the corpus luteum stops producing stimu-
blood flow to areas of the endometrium. Deprived of oxygen latory hormones. A new uterine cycle then begins with the onset
and nutrients, the secretory glands and other tissues in the of menstruation and the disintegration of the functional layer.
functional layer begin to deteriorate. Eventually, the weakened The secretory phase generally lasts 14 days. For this reason,
arterial walls rupture, and blood pours into the connective tis- you can identify the date of ovulation by counting backward
sues of the functional layer. Blood cells and degenerating tissues 14 days from the first day of menstruation.
then break away and enter the uterine cavity, to be shed as they
pass through the external os and into the vagina. Only the func- Hormonal Coordination of the Ovarian and
tional layer is affected, because the deeper, basal layer receives Uterine Cycles
blood from the straight arteries, which remain unconstricted.
The female reproductive tract is under hormonal control
The sloughing of tissue is gradual, and repairs begin almost
that involves interplay between secretions of both the pitu-
immediately at each site. Nevertheless, before menstruation has
itary gland and the gonads. Circulating hormones control the
ended, the entire functional layer has been lost. The menstrual
female reproductive cycle, coordinating the ovarian and uter-
phase generally lasts from 1 to 7 days. During this time about
ine cycles for proper reproductive function. If the two cycles are
35–50 mL (1.2–1.7 oz) of blood are lost.
not properly coordinated, infertility, the inability to conceive or
The process can be relatively painless. However, painful
carry a pregnancy to term, results.
menstruation, or dysmenorrhea, can result from myometrial
As in males, GnRH from the hypothalamus regulates repro-
contractions (“cramps”), uterine inflammation, or from condi-
ductive function in females. However, in females, the GnRH
tions involving adjacent pelvic structures.
pulse frequency and amplitude (amount secreted per pulse)
change throughout the ovarian cycle. If the hypothalamus were
The Proliferative Phase a radio station, the pulse frequency would correspond to the
The basal layer, including the basal parts of the uterine glands, radio frequency it is transmitting on, and the amplitude would
survives menstruation intact. In the days after menstruation, be the volume. We will consider changes in pulse frequency,
the epithelial cells of the uterine glands multiply and spread because their effects are both dramatic and reasonably well
across the endometrial surface, restoring the uterine epithelium understood. Circulating levels of estrogens and progesterone
(see Figure 28–23b). Further growth and vascularization com- primarily control changes in GnRH pulse frequency. Estro-
pletely restore the functional layer. gens increase the GnRH pulse frequency, and progesterone
During this reorganization, the endometrium is in the decreases it.
proliferative phase. Restoration takes place at the same time The endocrine cells of the anterior lobe of the pituitary
as the tertiary ovarian follicles enlarge in the ovary. Estrogens gland respond as if each group of endocrine cells is monitoring
secreted by the developing ovarian follicles stimulate and sus- different frequencies. As a result, each group of cells is sensitive
tain the proliferative phase. to some GnRH pulse frequencies and insensitive to others. For
By the time ovulation occurs, the functional layer is sev- example, consider the gonadotropes, the cells that produce FSH
eral millimeters thick. Prominent mucous glands extend to its and LH. At one pulse frequency, the gonadotropes respond 28
border with the basal layer. At this time, the uterine glands are preferentially and secrete FSH, but at another frequency, these
manufacturing mucus that is rich in glycogen. This specialized cells release primarily LH. FSH and LH production also occurs
mucus appears to be essential for the survival of the fertilized in pulses that follow the rhythm of GnRH pulses. If GnRH is
ovum through its earliest developmental stages. (We consider absent or is supplied at a constant rate (without pulses), FSH
these stages in Chapter 29.) The entire functional layer is highly and LH secretion stops in a matter of hours.
vascularized, with small arteries spiraling toward the endome- Spotlight Figure 28–24 shows the changes in circulating
trial surface from larger arteries in the myometrium. hormone levels that accompany the ovarian and uterine cycles.

M28_MART9867_11_GE_C28.indd 1133 23/08/17 10:29 AM

 SPOTLIGHT
Figure 28–24
Hormonal Regulation of Female Reproduction

The ovarian and uterine cycles must operate in synchrony to ensure proper
reproductive function. If the two cycles are not properly coordinated,
infertility results. A female who doesn’t ovulate cannot conceive, even if her
uterus is perfectly normal. A female who ovulates normally, but whose
uterus is not ready to support an embryo, will also be infertile.

As in males, GnRH from

HYPOTHALAMUS
the hypothalamus 1
regulates reproductive Release of Gonadotropin-Releasing
Hormone (GnRH)
function in females.
However, in females, The cycle begins with the release of GnRH,
GnRH levels change which stimulates the production and
throughout the course secretion of FSH and the production—
but not the secretion—of LH.
of the ovarian cycle.

Release of GnRH

3
Luteal Phase of
2
Follicular Phase of ANTERIOR the Ovarian Cycle
the Ovarian Cycle LOBE OF The combination of increased
The follicular phase begins when Production
and secretion
PITUITARY Production
of LH
GnRH pulse frequency and
FSH stimulates growth and
development of a group of tertiary
of FSH GLAND elevated estrogen levels
stimulates LH secretion.
ovarian follicles. Usually only one Secretion
follicle becomes dominant. of LH On or around day 14, a massive
surge in LH level triggers (1) the
As tertiary ovarian follicles completion of meiosis I by the
develop, FSH levels decline due primary oocyte, (2) the forceful
to the negative feedback effects rupture of the follicular wall, (3)
of inhibin. Negative ovulation, roughly 9 hours after the
feedback LH peak, and (4) formation of the
Developing ovarian follicles also corpus luteum.
After
secrete estrogens, especially
estradiol, the dominant hormone
OVARY day 10
The corpus luteum secretes
prior to ovulation. Before
day 10 • Meisois I progesterone, which stimulates and
completion sustains endometrial development.
In low concentrations, estrogens • Ovarian follicle • Ovulation
inhibit LH secretion. This development • Corpus luteum After ovulation, progesterone
inhibition gradually decreases formation levels rise and estrogen levels fall.
• Secretion of
as estrogen levels increase. inhibin This suppresses GnRH secretion.
If pregnancy does not occur, the
• Secretion of
estrogens corpus luteum will degenerate
Secretion of
after 12 days, and as progesterone
progesterone
levels decrease, GnRH secretion
increases, and a new cycle begins.

Effects Stimulation Establishment and Maintenance Stimulation of KEY

on CNS of bone and maintenance of of accessory endometrial Stimulation
muscle growth female secondary glands and growth and
sex characteristics organs secretion Inhibition

1134

M28_MART9867_11_GE_C28.indd 1134 23/08/17 10:29 AM

 This illustration combines the key events in the ovarian and uterine cycles. The monthly
hormonal fluctuations cause physiological changes that affect core body temperature.
During the follicular phase—when estrogens are the dominant hormones—the basal body
temperature, or the resting body temperature measured upon awakening in the morning, is
about 0.3°C (0.5°F) lower than it is during the luteal phase, when progesterone dominates.

FOLLICULAR PHASE OF OVARIAN CYCLE LUTEAL PHASE OF OVARIAN CYCLE

50

40 LH
Gonadotropic
hormone GnRH pulse frequency (pulses/day)
levels 30
(FSH and LH)
20 FSH

10

Follicle stages
during the
ovarian cycle
Corpus Mature Corpus
Tertiary ovarian follicle development Ovulation luteum corpus albicans
formation luteum

Progesterone

Ovarian
hormone
levels Estrogens
Inhibin

Destruction of Repair and Secretion by

functional zone regeneration of uterine glands
functional zone
Endometrial
changes
during the
uterine cycle

Phases of the MENSTRUAL PHASE PROLIFERATIVE SECRETORY PHASE

uterine cycle PHASE

36.5ºC (97.7ºF)
Basal body
temperature
36.2ºC (97.2ºF)

28/0 7 14 21 28/0
Days
1135

M28_MART9867_11_GE_C28.indd 1135 23/08/17 10:29 AM

 1136 UNIT 6 Continuity of Life

Early in the follicular phase of the ovarian cycle and prior to tissues. It is the dominant hormone prior to ovulation. In
day 10, the levels of estrogens are low and the GnRH pulse estradiol synthesis, androstenedione is first converted to tes-
frequency is 16–24 per day (one pulse every 60–90 minutes). tosterone, which the enzyme aromatase converts to estradiol
At this frequency, FSH is the dominant hormone released by (Figure 28–25). The synthesis of both estrone and estriol pro-
the anterior lobe of the pituitary gland. The estrogens released ceeds directly from androstenedione.
by developing follicles inhibit LH secretion. As tertiary ovarian Estrogens have multiple functions that affect the activi-
follicles develop, the FSH level decreases due to the negative ties of many tissues and organs throughout the body. Among
feedback effects of inhibin. Follicular development and matu- the important general functions of estrogens are (1) stimulat-
ration continue, however, supported by the combination of ing bone and muscle growth; (2) maintaining female second-
estrogens, FSH, and LH. ary sex characteristics, such as body hair distribution and the
As one of the tertiary ovarian follicles becomes dominant, location of adipose tissue deposits; (3) affecting CNS activity,
the concentration of circulating estrogens rises steeply. As a especially in the hypothalamus, where estrogens increase the
result, the GnRH pulse frequency increases to about 36 per sexual drive; (4) maintaining functional accessory reproductive
day (one pulse every 30–60 minutes). The increased pulse fre- glands and organs; and (5) initiating the repair and growth of
quency stimulates LH secretion. In addition, around day 10 of the endometrium.
the cycle, the effect of estrogen on LH secretion changes from
Regulation of the Luteal Phase. The high LH level that trig-
inhibition to stimulation. The switchover takes place only after
gers ovulation also promotes progesterone secretion and the
the levels of estrogens have risen above a specific threshold
formation of the corpus luteum. As the progesterone level
value for about 36 hours. (The threshold value and the time
rises and the levels of estrogens fall, the GnRH pulse frequency
required vary among individuals.) High levels of estrogens also
decreases sharply, soon reaching 1–4 pulses per day. This
increase gonadotrope sensitivity to GnRH. At about day 14,
frequency of GnRH pulses stimulates LH secretion more than
the estrogen levels have peaked, the gonadotropes are at maxi-
it does FSH secretion, and the LH maintains the structure and
mum sensitivity, and the GnRH pulses are arriving about every
secretory function of the corpus luteum.
30 minutes. The result is a massive release of LH from the
The corpus luteum secretes moderate amounts of estrogens,
anterior lobe of the pituitary gland. This sudden surge in LH
but progesterone is the main hormone of the luteal phase. Its
concentration triggers (1) the completion of meiosis I by the
primary function is to continue the preparation of the uterus for
primary oocyte, (2) the forceful rupture of the follicular wall,
pregnancy. Progesterone enhances the blood supply to the func-
and (3) ovulation. Typically, ovulation occurs 34–38 hours
tional layer and stimulates the secretion of uterine glands. The
after the LH surge begins, roughly 9 hours after the LH peak.
progesterone level remains high for the next week, but unless
pregnancy occurs, the corpus luteum begins to degenerate.
Hormonal Regulation of the Ovarian Cycle
Approximately 12 days after ovulation, the corpus luteum
The phases of the ovarian cycle are regulated by various hor-
becomes nonfunctional, and progesterone and estrogen lev-
mones. Follicle-stimulating hormone (FSH) and luteinizing
els fall markedly. The blood supply to the functional layer is
hormone (LH) are the main hormones influencing the follicu-
restricted, and the endometrial tissues begin to deteriorate.
lar phases. Progesterone is the dominant hormone directing
As progesterone and estrogen levels drop, the GnRH pulse
the luteal phase. If implantation does not occur, the menstrual
frequency increases, stimulating FSH secretion by the anterior
cycle begins. This section describes the hormonal regulation of
lobe of the pituitary gland, and the ovarian cycle begins again.
the ovarian cycle.
The hormonal changes involved with the ovarian cycle in
Regulation of the Follicular Phase. Follicular development turn affect the activities of other reproductive tissues and organs.
begins under FSH stimulation. Each month some of the ter- At the uterus, the hormonal changes maintain the uterine cycle.
tiary ovarian follicles begin to grow but only one will become
dominant. As the follicles enlarge, thecal endocrine cells start Hormonal Regulation of the Uterine Cycle
producing androstenedione, a steroid hormone that is a key in- Spotlight Figure 28–24 also shows the changes in the endome-
termediate in the synthesis of estrogens and androgens. The trium during a single uterine cycle. Progesterone and estrogen
28 granulosa cells absorb androstenedione and convert it to estro- levels decrease as the corpus luteum degenerates, resulting in
gens. In addition, thecal endocrine cells scattered throughout menstruation. The shedding of endometrial tissue continues
the ovarian stroma secrete small quantities of estrogens. Circu- for several days, until rising levels of estrogens stimulate the
lating estrogens are bound primarily to albumins, with lesser repair and regeneration of the functional layer of the endo-
amounts carried by gonadal steroid-binding globulin (GBG). metrium. The proliferative phase then continues until a rising
Of the three estrogens circulating in the bloodstream—
.
progesterone level marks the arrival of the secretory phase. The
estradiol, estrone, and estriol—estradiol (es-tra-DI-ol) is the combination of estrogen and progesterone then causes the uter-
most abundant and has the most pronounced effects on target ine glands to enlarge and increase their secretions.

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 Chapter 28 The Reproductive System   1137

Figure 28–25 Pathways of Steroid Hormone Synthesis in Males and Females. All gonadal steroids
are derived from cholesterol. In men, the pathway ends with the synthesis of testosterone, which may subsequently
be converted to dihydrotestosterone. In women, an additional step after testosterone synthesis leads to estradiol
synthesis. The synthesis of progesterone and estrogens other than estradiol involves alternative pathways.

Aromatase
Cholesterol Androstenedione Other Estrogens
Estrone

Progesterone Androgens Estriol

CH3 OH OH
C=O In
some
tissues

O O H
Progesterone
O Testosterone Dihydrotestosterone

Aromatase

Estradiol KEY
= Common pathways
OH
= Primary pathways
in females
= Primary pathways
in males
HO Estradiol

? What is the precursor for all steroid hormones in both males and females?

Checkpoint Sexual intercourse can lead to several consequences, includ-

ing pregnancy (both wanted and unintended) and sexually
12. Which layer of the uterus is sloughed off, or shed, during
menstruation? transmitted infections. We also discuss these subjects in this
13. What changes would you expect to observe in the ovarian section.
cycle if the LH surge did not occur?
14. What effect would a blockage of progesterone receptors Human Sexual Function
in the uterus have on the endometrium? Human sexual function refers to how the body reacts during
15. What event in the uterine cycle occurs when the levels of the sexual response.
estrogens and progesterone decrease?
See the blue Answers tab at the back of the book. Male Sexual Function
Complex neural reflexes coordinate sexual function in males.
28-6 The autonomic nervous system The reflex pathways use the sympathetic and parasympathetic
divisions of the autonomic nervous system (ANS). During sex-
influences male and female sexual ual arousal, erotic thoughts, the stimulation of sensory nerves 28
function in the genital region, or both lead to an increase in parasym-
Learning Outcome Discuss the physiology of sexual intercourse pathetic outflow over the pelvic nerves. This outflow in turn
in males and females.
leads to erection of the penis. The parasympathetic innervation
Sexual intercourse introduces semen into the female repro- of the penile arteries involves neurons that release nitric oxide
ductive tract. The process, in which the autonomic nervous at their axon terminals. The smooth muscles in the arterial
system plays a critical part, affects both the male and female walls relax when nitric oxide is released. At that time, the vessels
reproductive systems. dilate, blood flow increases, and the vascular channels become

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 1138 UNIT 6 Continuity of Life

engorged with blood. The skin covering the glans penis contains Any physical or psychological factor that affects a single
numerous sensory receptors, and erection tenses the skin and component of the system can result in male sexual dysfunc-
increases sensitivity. Subsequent stimulation can initiate the tion. Erectile dysfunction (ED), or impotence, is an inability to
secretion of the bulbo-urethral glands, providing lubrication achieve or maintain an erection. Various physical causes may be
for the spongy urethra and the surface of the glans penis. responsible for ED, because erection involves vascular changes
During intercourse, the sensory receptors of the penis are as well as neural commands. For example, low blood pressure
rhythmically stimulated. This stimulation eventually results in in the arteries supplying the penis, due to a circulatory blockage
the coordinated processes of emission and ejaculation. Emission such as a plaque, will reduce the ability to attain an erection.
occurs under sympathetic stimulation. The process begins when Drugs, alcohol, trauma, or illnesses that affect the ANS or the
the peristaltic contractions of the ampullae of the ductus defer- CNS can have the same effect. But male sexual performance
entia push seminal fluid and sperm into the prostatic urethra. can also be strongly affected by the psychological state of the
The seminal glands then begin contracting, and the contractions individual. Temporary periods of erectile dysfunction are fairly
increase in force and duration over the next few seconds. Peri- common in healthy individuals who are experiencing severe
staltic contractions also appear in the walls of the prostate. The stresses or emotional problems. Depression, anxiety, and fear of
combination moves the seminal mixture into the membranous ED are examples of emotional factors that can result in sexual
and penile portions of the urethra. As the contractions proceed, dysfunction. Prescription drugs, which enhance and prolong
sympathetic commands also cause the contraction of the uri- the effects of nitric oxide on the erectile tissue of the penis, have
nary bladder and the internal urethral sphincter. The combina- proven useful in treating many cases of erectile dysfunction.
tion of elevated pressure inside the bladder and the contraction
of the sphincter effectively prevent semen from passing into Female Sexual Function
the bladder. The events in female sexual function are largely comparable to
Ejaculation occurs as powerful, rhythmic contractions those of male sexual function. During sexual arousal, parasym-
appear in the ischiocavernosus and bulbospongiosus, two pairs of pathetic activation leads to engorgement of the erectile tissues
superficial skeletal muscles of the pelvic floor. The ischiocaver- of the clitoris and vestibular bulbs, and increased secretion
nosus muscles insert along the sides of the penis. Their contrac- from cervical mucous glands and greater vestibular glands.
tions serve primarily to stiffen that organ. The bulbospongiosus Clitoral erection increases the receptors' sensitivity to stimula-
muscles wrap around the base of the penis. The contraction of tion, and the cervical and vestibular glands lubricate the vaginal
these muscles pushes semen toward the external urethral open- walls. A network of blood vessels in the vaginal walls becomes
ing. The contractions of both muscles are controlled by somatic filled with blood at this time, and the vaginal surfaces are also
motor neurons in the inferior lumbar and superior sacral seg- moistened by fluid that moves across the epithelium from
ments of the spinal cord. (The positions of these muscles are underlying connective tissues. (This process accelerates during
shown in Figure 11–13b, p. 406.) Contraction of the smooth intercourse as the result of mechanical stimulation.) Parasym-
muscle within the prostate acts to pinch off the urethra, pre- pathetic stimulation also causes contraction of subcutaneous
venting urine from passing through the erect penis. smooth muscle of the nipples, making them more sensitive to
Ejaculation is associated with intensely pleasurable sensa- touch and pressure.
tions associated with perineal muscle contraction, an experience During sexual intercourse, rhythmic contact of the penis
known as male orgasm (OR-gazm). Several other noteworthy with the clitoris and vaginal walls—reinforced by touch sen-
physiological changes take place at this time, including pro- sations from the breasts and other stimuli (visual, olfactory,
nounced but temporary increases in heart rate and blood pres- and auditory)—provides stimulation that can lead to orgasm.
sure. After orgasm, a resolution phase occurs as heart rate and Female orgasm is accompanied by peristaltic contractions of
blood pressure decrease and blood begins to leave the erectile the uterine and vaginal walls and, through impulses traveling
tissue and the erection begins to subside. This process of losing over the pudendal nerves, rhythmic contractions of the bulbo-
an erection, called detumescence (de-tu-MES-ens), is medi-
. .

spongiosus and ischiocavernosus muscles. The latter contrac-

ated by the sympathetic nervous system. A refractory period tions give rise to the intensely pleasurable sensations of orgasm.
occurs during the initial period of resolution in which it is Unlike resolution and the refractory period in men, women
28 generally impossible for another orgasm to occur. may have little delay in achieving further orgasms.
In sum, arousal, erection, emission, ejaculation, orgasm,
and resolution are controlled by a complex interplay between
the sympathetic and parasympathetic divisions of the ANS.
Contraception and Infertility
Higher centers, including the cerebral cortex, can facilitate Many different methods can be used to promote or avoid
or inhibit many of the important reflexes, modifying sexual pregnancy. For example, at the time of ovulation, the basal
function. body temperature (BBT) decreases noticeably, making the rise

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 Chapter 28 The Reproductive System   1139

in temperature over the next day even more noticeable (see transmitted infections (STIs), are transferred from individ-
Spotlight Figure 28–24). Urine tests that detect LH are avail- ual to individual, primarily or exclusively by sexual intercourse.
able, and testing daily for several days before expected ovu- At least two dozen bacterial, viral, and fungal infections are cur-
lation can detect the LH surge more reliably than the BBT rently recognized as STIs. The bacterium Chlamydia can cause
changes. Because fertilization typically occurs within a day of pelvic inflammatory disease (PID) and infertility. The Zika
ovulation, this information can be used to time intercourse virus (ZIKV) is a cause of fetal brain defects, including micro-
according to the wishes of the couple. Using this to avoid con- cephaly. Recent evidence indicates it can be spread through
ception is called the rhythm method. sexual contact. AIDS, caused by a virus, is deadly.
Other forms of contraception (methods to prevent The incidence of STDs is a significant health challenge in
pregnancy) include abstinence, barrier methods, intrauter- the United States. According to the Centers for Disease Control
ine devices, chemical methods such as “the pill,” and surgical and Prevention (CDC), an estimated 20 million new cases
methods. occur each year, almost 50 percent in people aged 15–24.
Some couples find they have the opposite problem, and
they are unable to conceive. Infertility is defined as the inabil- Checkpoint
ity to conceive or carry a pregnancy to term after 1 year of
16. What happens when the arteries within the penis dilate?
unprotected intercourse. Infertility, which affects about 10 per-
cent of the U.S. population, can happen in both males and
17. List the physiological events of sexual intercourse in both
sexes, and indicate those that occur in males but not in
females. females.
For men, the most common issues are low sperm count
18. An inability to contract the ischiocavernosus and
and motility. Most males with lower sperm counts are infer- bulbospongiosus muscles would interfere with which
tile, because too few sperm survive the ascent of the female part of the male sex act?
reproductive tract to perform fertilization. A low sperm count 19. What changes occur in females during sexual arousal as
may reflect inflammation of the epididymis, ductus deferens, the result of increased parasympathetic stimulation?
or prostate. Common abnormalities in sperm are malformed See the blue Answers tab at the back of the book.
heads and “twin” sperm that did not separate at the time of
spermiation.
For women, all of their reproductive organs and hormonal 28-7 Changes in levels of reproductive
regulation must work properly to conceive and carry a preg- hormones cause functional changes
nancy to term. If menarche does not appear by age 16, or if throughout the life span
the normal uterine cycle of an adult woman is interrupted for Learning Outcome Describe the reproductive system changes
6 months or more, the condition of amenorrhea (a-men-o- that occur with development and aging.
. .

RE-uh) exists. Primary amenorrhea is the failure to initiate men-

Sex hormones have widespread effects on the body and begin
struation. This condition may indicate developmental abnor-
early in development. They affect brain development and
malities, such as nonfunctional ovaries, the absence of a uterus,
behavioral drives, muscle mass, bone mass and density, body
or an endocrine or genetic disorder. It can also result from
proportions, and the patterns of hair and body fat distribution.
malnutrition: Puberty is delayed if the leptin level is too low.
As noted earlier in the chapter, these systems become fully func-
p. 692
tional at puberty. At puberty, secretion of sex hormones in both
Severe physical or emotional stresses can cause transient
sexes accelerates markedly, initiating sexual maturation and the
secondary amenorrhea. In effect, the reproductive system gets
appearance of secondary sex characteristics. Later on, the aging
“switched off.” Factors associated with amenorrhea include
process affects all body systems, including the reproductive
drastic weight loss, anorexia nervosa, and severe depression or
systems of both men and women.
grief. Amenorrhea has also been observed in marathon runners
and other women engaged in training programs that require
Development of the Genitalia
sustained high levels of exertion, which severely reduce body
lipid reserves. Male and female reproductive systems develop from an embry-
onic aggregation of cells, termed primordial cells. Primordial 28
cells form the first trace of an organ. Both reproductive systems
Sexually Transmitted Diseases (STDs) follow a similar pattern of development from the same primor-
Sexual activity carries with it the risk of infection with a vari- dial tissues. They become distinct from each other because of
ety of microorganisms. The consequences of such an infec- the influence of sex hormones.
tion may range from merely inconvenient to potentially lethal. Around weeks 5–6 of embryonic development, an embryo
Sexually transmitted diseases (STDs), also called sexually is said to be sexually indifferent, because the gonads and other

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 1140 UNIT 6 Continuity of Life

internal and external reproductive structures are similar in both both testes are cryptorchid, the male will be sterile (infertile) and
sexes. Whether male or female gonads develop depends on the unable to father children. If the testes cannot be moved into the
embryo's genetic makeup. (We discuss the basis of sex deter- scrotum, they will usually be removed. This surgical procedure
mination in Chapter 29.) The male gonads begin to form dur- to remove the testes is called an orchiectomy (or-ke-EK-to-me).
. . .

ing week 7. Female gonads begin forming about a week later. About 10 percent of males with uncorrected cryptorchid testes
Once the testes have formed, they begin to secrete testosterone eventually develop testicular cancer.
continuously throughout fetal development and until a few As each testis moves through the body wall, the ductus
days after birth. Testosterone stimulates the differentiation and deferens and the testicular blood vessels, nerves, and lymphatic
development of male reproductive structures. Without testos- vessels accompany it. Together, these structures form the body
terone, the sexually indifferent embryo will develop female of the spermatic cord (see Figure 28–2, p. 1104).
reproductive structures.
The development of male and female genitalia from a Effects of Aging
sexually indifferent stage is shown in Figure 28–26. At week 4
As aging occurs, reductions in sex hormone levels affect appear-
of development, a temporary cloacal opening marks the site
ance, strength, and a variety of physiological functions. The
of the future genitalia (Figure 28–26a). A cloaca is a cavity or
most striking age-related changes in the female reproductive
chamber at the end of the digestive tract open to both excre-
system occur at menopause. Comparable age-related changes
tory and reproductive products. By week 6, the cloacal opening
in the male reproductive system occur more gradually and over
becomes subdivided, thus isolating the digestive tract from
a longer period of time.
future urogenital structures. An anterior urogenital membrane
will eventually form the urethra and urinary bladder. The geni-
The Male Climacteric
tal tubercle present in the sexually indifferent stage will give
Changes in the male reproductive system take place more grad-
rise to the penis in the male and clitoris in the female (see
ually than do those in the female reproductive system. The
Figure 28–26b,c).
period of declining reproductive function, which corresponds
During fetal development, the testes form inside the body
to perimenopause in women, is known as the male climacteric
cavity adjacent to the kidneys. A bundle of connective tis-
or andropause. The level of circulating testosterone begins to
sue fibers—called the gubernaculum testis (gu-bur-NAK-u-lum
. .

decrease between the ages of 50 and 60, and levels of circulating

TES-tis; plural, gubernacula)—extends from each testis to the
FSH and LH increase. Although sperm production continues
posterior wall of a small anterior and inferior pocket of the peri-
(men well into their 80s can father children), older men expe-
toneum. As the fetus grows, the gubernacula do not get any lon-
rience a gradual reduction in sexual activity. This decrease may
ger, so they lock the testes in position. As a result, the position
be linked to the declining testosterone level. Some clinicians
of each testis changes as the body enlarges. The testis gradually
suggest the use of testosterone replacement therapy to enhance
moves inferiorly and anteriorly toward the anterior abdominal
the libido (sexual drive) of elderly men, but this may increase
wall. During the seventh developmental month, fetal growth
the risk of prostate disease.
continues rapidly, and circulating hormones stimulate a con-
traction of the gubernaculum testis. During this time, each
testis moves through the abdominal musculature, along with Menopause
small pockets of the peritoneal cavity. This process is called Menopause is usually defined as the time when ovulation and
the descent of the testes into the scrotum (see Figure 28–26b). menstruation cease. Menopause typically occurs at age 45–55,
Descent of the female gonads is much less than that in males. but in the years immediately preceding it, an interval called
The ovaries come to lie below the rim of the true pelvis. They perimenopause, the ovarian and uterine cycles become irregular.
are held in place by the suspensory ligament, ovarian ligament, A shortage of primordial ovarian follicles is the underlying
and round ligament of the uterus (see Figure 28–16). cause of the irregular cycles. With the arrival of perimenopause,
In cryptorchidism (krip-TOR-ki-dizm; crypto, hidden + orchis, the number of primordial ovarian follicles responding each
testis), one or both of the testes have not descended into the month begins to decrease markedly. As their numbers decrease,
scrotum by the time of birth. Typically, the cryptorchid (abdom- the levels of estrogens decrease and may not increase enough to
28 inal) testes are lodged in the abdominal cavity or within the trigger ovulation. By age 50, there are often no primordial ovar-
inguinal canal. Cryptorchidism occurs in about 3 percent of ian follicles left to respond to FSH. In premature menopause,
full-term deliveries and in about 30 percent of premature births. this depletion occurs before age 40.
In most instances, normal descent takes place a few weeks later. Menopause is accompanied by a decrease in circulating
The condition can be surgically corrected if it persists. Cor- concentrations of estrogens and progesterone, and a sharp and
rective measures should be taken before puberty (sexual matu- sustained increase in the production of GnRH, FSH, and LH.
ration), because a cryptorchid testis will not produce sperm. If The decrease in the levels of estrogens leads to reductions in

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 Chapter 28 The Reproductive System   1141

Figure 28–26 The Development of Male and Female Genitalia.

a Sexually Indifferent Stage

Genital tubercle
By 6 weeks, the cloacal opening has been
subdivided into a posterior anal membrane and
an anterior urogenital membrane. Urethral folds Urethral fold
develop and prominent genital swellings form Urogenital membrane
lateral to each urethral fold. The genital tubercle
Genital swelling Cloacal
will develop into either the penis in males, or
clitoris in females, depending on the presence opening
or absence of testosterone.
Anal fold
4 WEEKS

6 WEEKS

b Development of the Male Genitalia c Development of the Female Genitalia

Genital tubercle

Genital swelling
Urethral folds Urethral fold

Scrotal swelling Urogenital

membrane

Anus Anus

External
Spongy urethral
orifice
urethra
Glans of Clitoris
penis Labia minora In the female, the urethral
folds do not fuse; they
Line of Urethra develop into the labia
At 10 weeks, the tips of the fusion minora. The genital
Labia majora
urethral folds are moving swellings will form the
together to form the spongy Opening labia majora. The genital
urethra, and paired scrotal Scrotum to vagina tubercle develops into the
swellings have developed clitoris. The urethra opens
Hymen
from the genital swellings. At to the exterior immediately
birth, the urethral folds have posterior to the clitoris.
closed and testes have The hymen remains as an
descended into the scrotum. elaboration of the
BIRTH BIRTH
urogenital membrane.

COMPARISON OF MALE AND FEMALE GENITALIA

MALE FEMALE
Penis Clitoris
Corpora cavernosa Erectile tissue
Corpus spongiosum Bulbs of vestibule 28
Proximal shaft of penis Labia minora
Spongy urethra Vestibule

Bulbo-urethral glands Greater vestibular glands

Scrotum Labia majora

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 1142 UNIT 6 Continuity of Life

the size of the uterus and breasts, accompanied by a thinning

Checkpoint
of the urethral and vaginal epithelia. The decreased concentra-
tions of estrogens have also been linked to the development of 20. What structure in a sexually indifferent embryo may
develop into a penis or clitoris and what hormone is
osteoporosis, presumably because bone deposition proceeds
involved in that development?
at a slower rate. A variety of neural effects are reported as well,
21. What is the male climacteric?
including “hot flashes,” anxiety, and depression. Hot flashes
22. Define menopause.
typically begin while the levels of estrogens are decreasing, and
cease when the levels of estrogens reach minimal values. These 23. Why does the level of FSH increase and remain high
during menopause?
intervals of increased body temperature are associated with
surges in LH production. The hormonal mechanisms involved See the blue Answers tab at the back of the book.
in other CNS effects of menopause are poorly understood. In
addition, the risks of atherosclerosis and other forms of cardio-
28-8 The reproductive system secretes
vascular disease increase after menopause.
Some women experience only mild symptoms, but some
hormones affecting growth and
experience acutely unpleasant symptoms in perimenopause metabolism of all body systems
or during or after menopause. For most of those women, hor- Learning Outcome Give examples of interactions between the
reproductive system and each of the other organ systems.
mone replacement therapy (HRT) involving a combination of
estrogens and progesterone can control the unpleasant neural Normal human reproduction is a complex process that involves
and vascular changes associated with menopause. The hor- multiple systems. The hormones discussed in this chapter play
mones are administered as pills, by injection, or by transdermal a major role in coordinating reproductive events (Table 28–1).
“estrogen patches.” However, recent studies suggest that taking Physical factors also play a role. The man's sperm count must
estrogen replacement therapy for more than 5 years increases be adequate, the semen must have the correct pH and nutrients,
the risk of heart disease, breast cancer, Alzheimer's disease, and erection and ejaculation must occur in the proper sequence.
blood clots, and stroke. HRT should be used only after a full The woman's ovarian and uterine cycles must be properly coor-
discussion and assessment of the potential risks and benefits, dinated; ovulation and oocyte transport must occur normally.
and taken for as short a time as possible. Her reproductive tract must provide a hospitable environment

Table 28–1 Hormones of the Reproductive System

Hormone Source Regulation of Secretion Primary Effects
Gonadotropin-releasing Hypothalamus Males: inhibited by testosterone and possibly Males: stimulates FSH secretion and LH
hormone (GnRH) by inhibin synthesis
Females: GnRH pulse frequency increased Females: stimulates FSH secretion and LH
by estrogens, decreased by progesterone synthesis
Follicle-stimulating Anterior lobe of the pituitary Males: stimulated by GnRH, inhibited by Males: stimulates spermatogenesis and
hormone (FSH) gland inhibin spermiogenesis through effects on nurse cells
Females: stimulated by GnRH, inhibited by Females: stimulates follicle development,
inhibin estrogen production, and oocyte maturation
Luteinizing Anterior lobe of the pituitary Males: stimulated by GnRH Males: stimulates interstitial cells to secrete
hormone (LH) gland Females: production stimulated by GnRH, testosterone
secretion by the combination of high GnRH Females: stimulates ovulation, formation of
pulse frequencies and high estrogen levels corpus luteum, and progesterone secretion
Androgens (primarily Interstitial endocrine cells Stimulated by LH Establish and maintain male secondary sex
testosterone and of testes characteristics and sexual behavior; promote
dihydrotestosterone) maturation of sperm; inhibit GnRH secretion
Estrogens (primarily Granulosa and thecal Stimulated by FSH Stimulate LH secretion (at high levels);
estradiol) endocrine cells of developing establish and maintain female secondary sex
28 follicles; corpus luteum characteristics and sexual behavior; stimulate
repair and growth of endometrium; increase
frequency of GnRH pulses
Progesterone Granulosa cells from Stimulated by LH Stimulate endometrial growth and glandular
midcycle through functional secretion; reduce frequency of GnRH pulses
life of corpus luteum
Inhibin Nurse cells of testes and Stimulated by factors released by developing Inhibits secretion of FSH (and possibly of
granulosa cells of ovaries sperm (male) and developing follicles (female) GnRH)

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 Chapter 28 The Reproductive System   1143

Build Your Knowledge

Figure 28–27 Integration of the REPRODUCTIVE system with the other body systems presented so far.

Integumentary System Nervous System

• The Integumentary System covers • The Nervous System controls sexual

external genitalia; provides sensations behaviors and sexual function
that stimulate sexual behaviors; mammary
• The reproductive system hormones affect
gland secretions nourish the newborn
CNS development and sexual behaviors
• The reproductive system hormones affect
the distribution of body hair and Endocrine System
subcutaneous fat
• The Endocrine System produces
hypothalamic regulatory hormones and
Respiratory System
pituitary hormones that regulate sexual
development and function; oxytocin
• The Respiratory System provides oxygen
stimulates smooth muscle contractions
and removes carbon dioxide generated
in uterus and mammary glands
by tissues of reproductive system and (in
pregnant women) by embryonic and fetal • The reproductive system produces
tissues steroid sex hormones and inhibin that
inhibit secretory activities of hypothala-
• The reproductive system changes
mus and pituitary gland
respiratory rate and depth during sexual
arousal, under control of the nervous
system Lymphatic System

• The Lymphatic System provides IgA for

Cardiovascular System secretions by epithelial glands; assists in
repairs and defense against infection
• The Cardiovascular System distributes
reproductive hormones; provides nutrients, • The reproductive system secretes
oxygen, and waste removal for fetus; local lysozymes and bactericidal chemicals that
blood pressure changes responsible for provide innate (nonspecific) defense
physical changes during sexual arousal against reproductive tract infections

• The reproductive system produces Urinary System

estrogens that may help maintain
healthy vessels and slow • The Urinary System in males carries
development of atherosclerosis semen to exterior in urethra; kidneys
excrete wastes generated by
Digestive System reproductive tissues and (in pregnant
women) by a growing embryo and fetus
• The Digestive System provides additional • The reproductive system secretions may
nutrients required to support gamete have antibacterial activity that helps
production and (in pregnant women) prevent urethral infections in males
embryonic and fetal development

• The reproductive system in pregnant Muscular System

women with a developing fetus crowds
digestive organs, causes constipation, • Muscular System contractions eject semen
and increases appetite from male reproductive tract
• The reproductive system hormone
testosterone accelerates skeletal muscle
Skeletal System
growth
• The Skeletal System (pelvic girdle)
protects reproductive organs of females, Reproductive System
portion of ductus deferens and accessory
glands in males The reproductive system secretes hormones
with effects on growth and metabolism. It:
• The reproductive system hormones • produces, stores, nourishes, and
stimulate bone growth and maintenance, transports male and female gametes
and at puberty accelerate growth and • supports the developing embryo and fetus
closure of epiphyseal cartilages in the uterus

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 1144 UNIT 6 Continuity of Life

for the survival and movement of sperm, and for the subse- and psychological factors. Many of these factors require intersys-
quent fertilization of the oocyte. For these steps to occur, the tem cooperation. In addition, the hormones that control and
reproductive, digestive, endocrine, nervous, cardiovascular, and coordinate sexual function have direct effects on the organs and
urinary systems must all be functioning normally. tissues of other systems. For example, testosterone and estradiol
Even when all else is normal and fertilization occurs at the affect both muscular development and bone density. Build Your
proper time and place, a healthy infant will not be produced Knowledge Figure 28–27 summarizes the functional relation-
unless the zygote—the union of an ovum with a sperm form- ships between the reproductive system and other body systems.
ing a single cell the size of a pinhead—manages to develop into
a full-term fetus that typically weighs about 3 kg (6.6 lb). In
Checkpoint
Chapter 29 we will consider the process of development, focus-
ing on the mechanisms that determine both the structure of 24. Describe the interaction between the reproductive system
the body and the distinctive characteristics of each individual. and the cardiovascular system.
Even though the reproductive system's primary function— 25. Describe the interaction between the reproductive system
producing offspring—doesn't play a role in maintaining homeo- and the skeletal system.
stasis, reproduction depends on a variety of physical, physiological, See the blue Answers tab at the back of the book.

28 Chapter Review
Study Outline
An Introduction to the Reproductive System p. 1102 > Pearson Mastering Access more chapter
study tools online in the Pearson Mastering A&P Study Area:
1. The reproductive system is the only system that is not essential
■ Chapter Quizzes, Chapter Practice Test, MP3 Tutor Sessions,
to the life of an individual, but it ensures continuation of our
and Clinical Case Studies
species.
■ Practice Anatomy Lab 3.0 ■ A&P Flix
™

28-1 Male and female reproductive system structures ■ Interactive Physiology ■ PhysioEx
produce gametes that combine to form a new individual
p. 1102
2. The human reproductive system produces, stores, nourishes,
and transports functional gametes (reproductive cells). Fertil- 7. The dartos muscle tightens the scrotum, giving it a wrinkled
ization is the fusion of male and female gametes. appearance as it elevates the testes. The cremaster has skeletal
3. The reproductive system includes gonads (testes or ova- muscle fibers that pull the testes close to the body.
ries), ducts, accessory glands and organs, and the external 8. The tunica albuginea surrounds each testis. Septa extend
genitalia. from the tunica albuginea to the region of the testis closest to
4. In males, the testes produce sperm, which are expelled from the entrance to the epididymis, creating a series of lobules.
the body in semen during ejaculation. The ovaries of a sexually (Figure 28–3)
mature female produce oocytes (immature ova) that travel 9. Seminiferous tubules within each lobule are the sites of
along uterine tubes toward the uterus. The vagina connects the sperm production. From there, sperm pass through the rete
uterus with the exterior of the body. testis. Seminiferous tubules connect to a straight tubule.
28-2 The structures of the male reproductive system Efferent ductules connect the rete testis to the epididymis.
consist of the testes, duct system, accessory glands, and Between the seminiferous tubules are interstitial cells, which
penis p. 1103 secrete sex hormones. (Figures 28–3, 28–4)
10. From the testis, the sperm enter the epididymis, an elongated
5. Sperm travel along the epididymis, the ductus deferens, the ejacu- tubule with head, body, and tail regions. The epididymis
28 latory duct, and the urethra before leaving the body. Accessory monitors and adjusts the composition of the fluid in the
organs (notably the seminal glands, prostate, and bulbo-urethral seminiferous tubules, serves as a recycling center for damaged
gland) secrete fluids into the ejaculatory ducts and the urethra. sperm, stores and protects sperm, and facilitates their func-
The scrotum encloses the testes, and the penis is an erectile tional maturation. (Figure 28–5)
organ. (Figure 28–1) 11. The ductus deferens, or vas deferens, begins at the epididymis
6. The raphe marks the boundary between the two chambers in and passes through the inguinal canal as part of the spermatic
the scrotum. (Figure 28–3) cord. Near the prostate, the ductus deferens enlarges to form

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 Chapter 28 The Reproductive System   1145

the ampulla of ductus deferens. The junction of the base Normally, the uterus bends anteriorly near its base (anteflex-
of the seminal gland and the ampulla creates the ejaculatory ion). The broad ligament, uterosacral ligaments, round
duct, which empties into the urethra. (Figures 28–5, 28–6) ligaments, and lateral ligaments stabilize the uterus.
12. The urethra extends from the urinary bladder to the tip of the (Figure 28–16)
penis. The urethra can be divided into prostatic, membranous, 26. Major anatomical landmarks of the uterus include the body,
and spongy regions. isthmus, cervix, external os (external orifice), uterine cavity,
13. Each seminal gland (seminal vesicle) is an active secre- cervical canal, and internal os (internal orifice). The uterine
tory gland that contributes about 60 percent of the volume wall consists of an inner endometrium, a muscular myome-
of semen. Its secretions contain fructose (which is easily me- trium, and a superficial perimetrium (an incomplete serous
tabolized by sperm), bicarbonate ions, prostaglandins, and fi- layer). (Figures 28–16, 28–17)
brinogen. The prostate secretes slightly acidic prostatic fluid. 27. The vagina is a muscular tube extending between the uterus
Alkaline mucus secreted by the bulbo-urethral glands has and the external genitalia; it is lined by a nonkeratinized strati-
lubricating properties. (Figures 28–6, 28–7) fied squamous epithelium. A thin epithelial fold, the hymen,
14. A typical ejaculation releases 2–5 mL of semen (ejaculate), partially blocks the entrance to the vagina until physical dis-
which contains 20–100 million sperm per milliliter. The fluid tortion ruptures the membrane. (Figures 28–18, 28–19)
component of semen is seminal fluid. 28. The components of the vulva are the vestibule, labia minora,
15. The skin overlying the penis resembles that of the scrotum. urethral glands, clitoris, mons pubis, labia majora, and
Most of the body of the penis consists of three masses of erec- lesser and greater vestibular glands. (Figure 28–19)
tile tissue. Beneath the superficial fascia are two corpora cav- 29. A newborn infant is nourished from milk secreted by the ma-
ernosa and a single corpus spongiosum, which surrounds ternal mammary gland in each breast. (Figure 28–20)
the urethra. Dilation of the blood vessels within the erectile
tissue produces an erection. (Figure 28–7) 28-5 Oogenesis occurs in the ovaries, and hormones from
the hypothalamus, pituitary gland, and ovaries control
28-3 Spermatogenesis occurs in the testes, and female reproductive functions p. 1128
hormones from the hypothalamus, pituitary gland, and 30. Ovaries are the site of oogenesis (ovum production), which
testes control male reproductive functions p. 1111 occurs in ovarian follicles. Follicle development proceeds
16. Seminiferous tubules contain spermatogonia, stem cells in- from primordial ovarian follicles through primary, second-
volved in spermatogenesis (the production of sperm). ary, and tertiary ovarian follicles. A dominant tertiary ovar-
17. Nurse (Sertoli) cells sustain and promote the development of ian follicle forms during the ovarian cycle. The ovarian cycle
sperm. (Figures 28–8, 28–9, 28–10) is divided into a follicular (preovulatory) phase and a luteal
18. Each sperm has a head capped by an acrosome, a middle (postovulatory) phase. (Figures 28–21, 28–22)
piece, and a tail. (Figure 28–11) 31. At ovulation, a secondary oocyte and the attached follicular
19. Important regulatory hormones include FSH (follicle-stimu- cells of the corona radiata are released through the ruptured
lating hormone), LH (luteinizing hormone), and GnRH (gonad- ovarian wall. The follicular cells remaining within the ovary
otropin-releasing hormone). Testosterone is the most important form the corpus luteum, which later degenerates into scar tis-
androgen. (Spotlight Figure 28–12) sue called a corpus albicans. (Figure 28–22)
32. A typical 28-day uterine, or menstrual, cycle begins with the
28-4 The structures of the female reproductive system onset of menstruation, also called menses, and the destruc-
consist of the ovaries, uterine tubes, uterus, vagina, and tion of the functional layer of the endometrium. This process
external genitalia p. 1118 of menstruation continues from 1 to 7 days. (Figure 28–23)
20. Principal organs of the female reproductive system include 33. After the menstrual phase, the proliferative phase begins,
the ovaries, uterine tubes, uterus, vagina, and external genitalia. and the functional layer thickens and undergoes repair. The
(Figure 28–13) proliferative phase is followed by the secretory phase, during
21. The ovaries, uterine tubes, and uterus are enclosed within the which uterine glands enlarge. Menstruation begins at men-
broad ligament. The mesovarium supports and stabilizes arche and continues until menopause. (Figure 28–23)
each ovary. (Figure 28–14) 34. Hormonal regulation of the female reproductive cycle
22. The ovaries are held in position by the ovarian ligament and involves the coordination of the ovarian and uterine cycles.
the suspensory ligament. Major blood vessels enter the ovary (Spotlight Figure 28–24)
at the ovarian hilum. Each ovary is covered by a tunica albu- 35. Estradiol, the most important estrogen, is the dominant hor-
ginea. (Figure 28–14) mone of the follicular phase. Ovulation occurs in response to
23. Each uterine tube has an infundibulum with fimbriae a midcycle surge in LH. (Spotlight Figure 28–24; Figure 28–25)
(fingerlike projections), an ampulla, and an isthmus. Each 36. The hypothalamic secretion of GnRH occurs in pulses that
uterine tube opens into the uterine cavity. For fertilization to trigger the pituitary secretion of FSH and LH. FSH initiates
occur, a secondary oocyte must encounter sperm during the follicular development, and activated follicles and ovarian 28
first 12–24 hours of its passage from the infundibulum to the interstitial cells produce estrogens. High estrogen levels stimu-
uterus. (Figure 28–15) late LH secretion, increase pituitary sensitivity to GnRH, and
24. Peg cells lining the uterine tube secrete a fluid that completes increase the GnRH pulse frequency. Progesterone is the main
the capacitation of sperm. hormone of the luteal phase. Changes in estrogen and proges-
25. The uterus provides mechanical protection, nutritional sup- terone levels are responsible for maintaining the uterine cycle.
port, and waste removal for the developing embryo and fetus. (Spotlight Figure 28–24)

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 1146 UNIT 6 Continuity of Life

28-6 The autonomic nervous system influences male and internal and external reproductive structures are similar in
female sexual function p. 1137 both sexes. The presence of testosterone stimulates the dif-
ferentiation and development of male reproductive structures.
37. During sexual arousal in males, erotic thoughts, sensory
Without testosterone, the sexually indifferent embryo will de-
stimulation, or both lead to parasympathetic activity that
velop female reproductive structures. (Figure 28–26)
produces erection. Stimuli accompanying sexual intercourse
41. The descent of the testes through the inguinal canals begins
lead to emission and ejaculation. Contractions of the bulbo-
during the seventh month of fetal development.
spongiosus muscles are associated with orgasm. During reso-
42. During the male climacteric, at ages 50–60, circulating testos-
lution, the heart rate and blood pressure decrease and blood
terone levels decrease, and FSH and LH levels increase.
begins to leave the erectile tissue and the erection begins to
43. Menopause (the time that ovulation and menstruation stop)
subside.
typically occurs at ages 45–55. The production of GnRH, FSH,
38. The events of female sexual function resemble those of male
and LH rises, whereas circulating concentrations of estrogen
sexual function, with parasympathetic arousal and skeletal
and progesterone decrease.
muscle contractions associated with orgasm.
39. Infertility is the inability to conceive or carry a pregnancy to 28-8 The reproductive system secretes hormones
term after 1 year of unprotected intercourse. affecting growth and metabolism of all body systems
p. 1142
28-7 Changes in levels of reproductive hormones cause
functional changes throughout the life span p. 1139 44. Normal human reproduction depends on a variety of physi-
cal, physiological, and psychological factors, many of which
40. At weeks 5–6 of embryonic development, an embryo is said
require intersystem cooperation. (Figure 28–27; Table 28–1)
to be sexually indifferent, because the gonads and other

Review Questions See the blue Answers tab at the back of the book.

LEVEL 1 Reviewing Facts and Terms

1. All of the following structures are part of the spermatic cord except the
(a) ductus deferens, (b) pampiniform plexus, (c) genitofemoral nerve,
(d) testicular artery, (e) epididymis.
2. The uterus (a) undergoes the uterine cycle every month, (b) has a
thick, muscular myometrium, (c) has a tubular cervix that projects
into the vagina, (d) does/has all of these. i
3. In meiosis, sister chromatids separate during (a) prophase I,
(b) metaphase I, (c) anaphase I, (d) anaphase II.
a j
4. Gonadotropin-releasing hormone (a) stimulates FSH secretion and
LH synthesis in only males, (b) stimulates FSH secretion and LH
synthesis in only females, (c) stimulates FSH secretion and LH
b k
synthesis in both males and females, (d) is produced in the ovaries.
c l
5. The main hormone of the postovulatory phase is (a) progesterone,
d
(b) estradiol, (c) estrogen, (d) luteinizing hormone. e
6. Which accessory structures contribute to the composition of semen?
What are the functions of each structure? f
7. What types of cells in the testes are responsible for functions related to g
reproductive activity? What are the functions of each cell type? h
8. Which part of the nervous system controls sexual arousal?
9. List the functions of testosterone in males.
10. List and summarize the important steps in the ovarian cycle.
28 11. Describe the histology of the uterine wall.
12. What is the role of the clitoris in the female reproductive system? (a) (b) (c)
13. Trace the path of milk flow from its site of production to outside the (d) (e) (f)
female.
14. Identify the main structures of the male reproductive system in the (g) (h) (i)
diagram on the right. (j) (k) (l)

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 Chapter 28 The Reproductive System   1147

LEVEL 2 Reviewing Concepts LEVEL 3 Critical Thinking and Clinical Applications

15. The immune system does not identify the developing sperm as a 24. Andrew is 75 and is in good health. But he has recently started to
foreign matter because of the (a) blood testis barrier, (b) capacitation experience lower back pain and painful urination. A rectal examina-
of sperm, (c) secretion of dehydroepiandrosterone, (d) secretion of tion reveals an enlargement of the prostate. Explain why it is not
inhibin, (e) formation of acrosome. unusual for elderly men to develop this condition.
16. Summarize the roles of the hormones in the ovarian and uterine 25. In a condition known as endometriosis, endometrial cells are believed
cycles. to migrate from the body of the uterus into the uterine tubes or by
17. Compare the fates of parent cells undergoing meiosis in males and way of the uterine tubes into the peritoneal cavity, where they become
females. established. A major symptom of endometriosis is periodic pain. Why
18. Describe the erectile tissues of the penis. How does erection occur? does such pain occur?
19. Describe each of the three phases of a typical 28-day uterine cycle. 26. Birth control pills contain estradiol and progesterone, or progesterone
20. Describe the hormonal events associated with the ovarian cycle. alone, administered at programmed doses during the ovarian cycle
21. Describe the hormonal events associated with the uterine (menstrual) to prevent tertiary ovarian follicle maturation and ovulation. Explain
cycle. how such pills are effective.
22. What is meant by the descent of the testes? Is there a comparable 27. Female bodybuilders and women with eating disorders such as anorexia
process in females? nervosa commonly experience amenorrhea. What does this fact suggest
23. How does the aging process affect the reproductive systems of men about the relationship between body fat and menstruation? What effect
and women? would amenorrhea have on achieving a successful pregnancy?

+ CLINICAL CASE Wrap-Up And Baby Makes Three?

A basic rule of thumb is that a couple that has been A man’s infertility is most often due to problems
trying to get pregnant for 1 year is a candidate for a with his sperm. The count may be too low or the
fertility work-up. Fertility problems can originate in the motility of the sperm may be poor. Checking the
female, in the male, in both, or for unknown reasons. quantity and quality of Tim’s sperm is simple. He
A woman’s ability to get pregnant decreases can provide a sample for analysis in the lab. Tim
with increasing age. Her fertility depends on how can change his running gear from briefs to boxers in
many oocytes she has left in her ovaries, how order to provide a cooler environment in the testes
healthy the oocytes are, and—most importantly—if for sperm production.
they are being released from the ovary. A healthy The Millers have youth and general good health
reproductive tract is needed for the oocyte to unite with a sperm, on their side. However, they do have risk factors for infertility that
and then for the fertilized ovum to be conveyed to a healthy uterus need to be addressed. Baby Miller may yet join the family!
to implant and complete gestation. 1. Trace the path of Susan’s oocyte to meet Tim’s sperm. Where
Is Susan ovulating? That is the first thing to check, as it is the will a fertilized ovum travel next?
most common reason for female infertility. At home she can take
2. When should Susan expect ovulation to occur during her cycle?
her morning temperature, which should spike with ovulation at day
How can we verify that she has ovulated?
14 and stay high through day 28 of her cycle under the influence
of progesterone. Is her reproductive tract blocked by scarring from See the blue Answers tab at the back of the book.
her earlier STI? If so, surgery can open a blocked uterine tube.

Related Clinical Terms

cervical dysplasia: Abnormal growth of epithelial cells in the uter- hydrocele: The accumulation of serous fluid in any body sac, but
ine cervix; may progress to cancer. especially in the tunica vaginalis of the testis or along the sper-
endometriosis: The growth of endometrial tissue outside the uterus. matic cord. 28
genital herpes: A sexually transmitted disease caused by a herpes hysterectomy: The surgical removal of the uterus.
virus and characterized by painful blisters in the genital area. menorrhagia: The condition of experiencing extremely heavy
gonorrhea: A sexually transmitted bacterial disease caused by Neis- bleeding at menstruation.
seria gonorrhoeae. Commonly called “the clap.” oophorectomy: The surgical removal of one or both ovaries.
gynecology: The branch of medicine that deals with the functions orchitis: Inflammation of one or both testicles.
and diseases specific to women and girls affecting the reproduc- ovarian cyst: A condition (usually harmless) in which fluid-filled
tive system. sacs develop in or on the ovary.

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 1148 UNIT 6 Continuity of Life

polycystic ovary syndrome (PCOS): A condition in women that salpingitis: Inflammation of a uterine tube.
is characterized by irregular or no menstrual periods, acne, obe- uterine fibroids (leiomyomas): Benign tumors of the uterus
sity, and excessive hair growth. composed of smooth muscle tissue that grows in the wall of the
premature ejaculation: A common complaint of ejaculating uterus of some women. Although not usually dangerous they
semen sooner than the man desires while achieving orgasm dur- can cause problems such as very heavy menstrual periods and
ing intercourse. An estimated 30 percent of men regularly experi- pain.
ence the problem. uterine prolapse: Condition that occurs when a woman's pelvic
premenstrual dysphoric disorder (PMDD): A collection of floor muscles and ligaments stretch and weaken and provide
physical and emotional symptoms that occurs 5 to 11 days be- inadequate support for the uterus, which then descends into the
fore a woman's period begins, and goes away once menstruation vaginal canal.
starts. Over 150 signs and symptoms have been associated with vasectomy: The surgical removal of a segment of each ductus (vas)
the condition. deferens, making it impossible for sperm to reach the distal por-
premenstrual syndrome: A condition occurring in the last half tions of the male reproductive tract.
of the menstrual cycle after ovulation that is a combination of vulvovaginal candidiasis: A common female vaginal infection
physical and mood disturbances that normally end with the caused by the yeast Candida, usually Candida albicans.
onset of the menstrual flow.

28

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