You are on page 1of 70


Continuity of Life
Chapter 28, The Reproductive System, discusses how the male and female reproductive
organs (gonads) produce and store specialized reproductive cells (gametes) that comine to
form new individuals, and how the gonads also secrete hormones that play ma!or roles in the
maintenance of normal se"ual function#
Chapter 29, $evelopment and %nheritance, descries how genetic programming,
environmental factors, and various physiological processes affect the events following the
union of male and female gametes, from prenatal development, through childhood and
adolescence, and into maturity and senescence (aging)#
The End of Chapter &uestions within this unit include critical thin'ing &uestions aout oth
normal and anormal functions# (or comprehensive e"ercises covering material in the unit as
a whole, see the Clinical )rolems at the end of the corresponding unit in the Applications
Manual *+,-#
The Reproductive System
Introduction to the Reproductive System 0121
The Reproductive System of the Male 0121
The Testes 0121
Spermatogenesis 0126
Key 012/
The +natomy of a Spermatozoon 012/
Key 0123
The ,ale Reproductive Tract 0141
The +ccessory 5lands 0140
Semen 0142
The 6"ternal 5enitalia 0144
7ormones and ,ale Reproductive (unction 0148
The Reproductive System of the emale 014/
232459324.doc 1
The 9varies 0143
Key 018.
The :terine Tues 018.
The :terus 0182
The ;agina 018<
The 6"ternal 5enitalia 018/
The ,ammary 5lands 0183
7ormones and the (emale Reproductive Cycle 0160
Summary= 7ormonal Regulation of the (emale Reproductive Cycle 016.
Key 0168
The !hysiolo"y of Se#ual Intercourse 0168
,ale Se"ual (unction 0168
(emale Se"ual (unction 0166
$"in" and the Reproductive System 0166
,enopause 0166
The ,ale Climacteric 016<
Key 016<
Inte"ration %ith &ther Systems 016<
Clinical )atterns 016<
' S(MM$R) T$*+E 28,- ' .&RM&/ES & T.E RE!R&0(CTI1E S)STEM
The Reproductive System in )erspective 0163
Chapter Revie% 016/
Clinical /otes
$76+ 0146
)rostatic 7ypertrophy and )rostate Cancer 014<
>reast Cancer 0161
Introduction to the Reproductive System
? Specify the principal components of the human reproductive system and summarize their
232459324.doc 2
%n this chapter we e"amine the anatomy and physiology of the human reproductive system,
which is the only system that is not essential to the life of the individual# This system does,
however, ensure the continued e"istence of the human species@y proA
ducing, storing, nourishing, and transporting functional male and female reproductive cells,
or "ametes (5+,Ats)# The reproductive system includes the following components=
# ? 2onads (59Anadz), or reproductive organs that produce gametes and
# ? $ucts that receive and transport the gametes#
# ? +ccessory glands and organs that secrete fluids into the ducts of the
reproductive system or into other e"cretory ducts#
)erineal structures that are collectively 'nown as the e#ternal "enitalia (!enAiAT+BAl
%n oth males and females, the ducts are connected to chamers and passageways that open
to the e"terior of the ody# The structures involved constitute the reproductive tract# The
male and female reproductive systems are functionally &uite different,
however# %n adult males, the testes (T6SAt zC singular, testis), or male gonads, secrete se"
hormones called androgens (principally testosterone, which, together with other se"
hormones, was introduced in Chapter 0/)# lp3 422 The testes also produce the male
gametes, called spermato5oa (sperAmaAt
232459324.doc 3
AuhC singular, spermatozoon), or sperm@oneAhalf illion each day# $uring emission,
mature spermatozoa travel along a lengthy duct system, where they are mi"ed with the
secretions of accessory glands# The mi"
ture created is 'nown as semen (S
Amen)# $uring ejaculation, semen is e"pelled from the ody#
%n adult females, the ovaries, or female gonads, typically release only one immature gamete,
an oocyte, per month# This im
mature gamete travels along one of two short uterine tubes, which end in the muscular
organ called the uterus (
AterAus)# %f a sperm
reaches the oocyte and initiates the process of fertilization, the oocyte matures into an ovum
(plural, ova)# + short passageway, the vagina (vaAE Anuh), connects the uterus with the
e"terior# 6!aculation introduces semen into the vagina during sexual intercourse,
%Band the spermatozoa then ascend the female reproductive tract# %f fertilization occurs, the
uterus will enclose and support a developing embryo as it grows into a fetus and prepares
for irth#
Fe"t we will e"amine the anatomy of the male and female reproductive systems further, and
will consider the physiological and hormonal mechanisms responsile for the regulation of
reproductive function# 6arlier chapters introduced the anatomical reference points used in
the discussions that followC you may find it helpful to review the figures on the pelvic girdle
((igures /G< and /G/?, pp# .46, .4<), perineal musculature ((igure 00G0.?, p# 24/), pelvic
innervation ((igure 02G0.?, p# 426), and regional lood supply ((igures .0G.8 and .0G.3?,
pp# <44, <43)#
The Reproductive System of the Male
# ? $escrie the components of the male reproductive system#
# ? 9utline the processes of meiosis and spermatogenesis
232459324.doc 4
in the testes#
# ? 6"plain the roles played y the male reproductive tract and accessory glands
in the functional maturation, nourishment, storage, and transport of spermatozoa#
# ? Specify the normal composition of semen#
# ? Summarize the hormonal mechanisms that regulate male reproductive functions#
The principal structures of the male reproductive system are shown in (igure ./G0?#
)roceeding from a testis, the spermatozoa travel within the epididymis (epAiA$%$AiAmus)C the
ductus deferens ($:HAtus $6(AeArenz), or vas deferensC the ejaculatory ductC and
the urethra efore leaving the ody# +ccessory organs@the seminal (S6,AiAnal) vesicles,
the prostate ()R9SAt t) gland, and the 6B
bulbourethral (ulA
Athral) glands@secrete various fluids into the e!aculatory ducts and urethra# The e"ternal
Atum), which encloses the testes, and the penis ()
Anis), an erectile organ through which the distal
consist of the scrotum (SHR
portion of the urethra passes#
The Testes
6ach testis has the shape of a flattened egg that is roughly 8 cm (. in#) long, 2 cm (0#. in#)
wide, and .#8 cm (0 in#) thic'# 6ach has a weight of 01G08 g (1#28G1#82 oz)# The testes
232459324.doc 5
hang within the scrotum, a fleshy pouch suspended inferior to the perineum, anterior to the
anus and posterior to the ase of the penis (see (igure ./G0?)# +TL+S= 6mryology
Summary .0= The $evelopment of the Reproductive System
$escent of the Testes
$uring development of the fetus, the testes form inside the ody cavity ad!acent to the
'idneys# + undle of connectiveAtissue
fiers@called the "u6ernaculum testis (gooAurAF+HA
Alum T6SAtis)@e"tends from each testis to the posterior wall of a small
anterior and inferior poc'et of the peritoneum ((igure ./G.a?)# +s the fetus grows, the
guernacula do not get any longer, so they loc' the testes in position# +s a result, the
relative position of each testis changes as the ody enlarges= The testis gradually moves
inferiorly and anteriorly toward the anterior adominal wall# $uring the seventh
developmental month, fetal growth continues at a rapid pace, and circulating hormones
stimulate a contraction of the guernaculum testis# 9ver this period, each testis moves
through the adominal musculature, accompanied y small poc'ets of the peritoneal cavity#
This process is called the descent of the testes#
%n cryptorchidism ('ripAT9RA'iAdizmC crypto, hidden I orchis, testis), one or oth of the
testes have not descended into the scrotum y the time of irth# Typically, the cryptorchid
(adominal) testes are lodged in the adominal cavity or within the inguinal canal#
Cryptorchidism occurs in aout 2 percent of fullAterm deliveries and in roughly 21 percent of
premature irths# %n most instances, normal descent occurs a few wee's later, ut the
condition can e surgically corrected if it persists# Corrective measures should e ta'en
efore puberty (se"ual maturation), ecause a cryptorchid testis will not produce
spermatozoa# %f oth testes are cryptorchid, the individual will e sterile (infertile) and
unale to father children# %f the testes cannot e moved into the scrotum, in most cases they
will e removed, ecause aout 01 percent of males with uncorrected cryptorchid testes
develop testicular cancer# This surgical procedure is called an orchiectomy (orA'
232459324.doc 6
+s each testis moves through the ody wall, it is accompanied y the ductus deferens and
the testicular lood vessels, nerves, and lymphatic vessels# Together, these structures form
the ody of the spermatic cord, which we will discuss ne"t#
The Spermatic Cords
The spermatic cords are paired structures e"tending etween the adominopelvic cavity
and the testes ((igure ./G2?)# 6ach spermatic cord consists of layers of fascia and muscle
enclosing the ductus deferens and the lood vessels, nerves, and lymphatic vessels that
supply the testes# The lood vessels include the deferential artery, a testicular artery, and
the pampiniform (pamA)%FAiAformC pampinus, tendril I forma, form) ple#us of a testicular
vein# >ranches of the genitofemoral nerve from the lumar ple"us provide innervation# 6ach
spermatic cord egins at the entrance to the inguinal canal (a passageway through the
adominal musculature)# +fter passing through the inguinal canal, the spermatic cord
descends into the scrotum#
The inguinal canals form during development as the testes descend into the scrotumC at that
time, these canals lin' the scrotal cavities with the peritoneal cavity# %n normal adult males,
the inguinal canals are closed, ut the presence of the spermatic cords creates wea' points in
the adominal wall that remain throughout life# +s a result, inguinal hernias@protrusions of
visceral tissues or organs into the inguinal canal@are relatively common in males# Fote that
the inguinal canals in females are very small, containing only the ilioinguinal nerves and the
round ligaments of the uterusC the adominal wall is nearly intact, so inguinal hernias in
women are very rare#
The Scrotum and the )osition of the Testes
The scrotum is divided internally into two chamers# The partition etween the two is
mar'ed y a raised thic'ening in the scro
tal surface 'nown as the raphe (R
232459324.doc 7
) (see (igure ./G2?)# 6ach testis lies in a separate chamer, or scrotal cavity# >ecause the
scrotal cavities are separated y a partition, infection or inflammation of one testis does not
ordinarily spread to the other# + narrow space separates the inner surface of the scrotum
from the outer surface of the testis# The tunica va"inalis (T99AniA'a va!AiAF+LAis), a
serous memrane, lines the scrotal cavity and reduces friction etween the opposing parietal
(scrotal) and visceral (testicular) surfaces# The tunica vaginalis is an isolated portion of the
peritoneum that lost its connection with the peritoneal cavity after the testes descended,
when the inguinal canal closed#
The scrotum consists of a thin layer of s'in and the underlying superficial fascia# The dermis
contains a layer of smooth mus
cle, the dartos ($+RAt
s) muscle# Resting muscle tone in the dartos muscle elevates the testes and causes the
characteristic wrinA
'ling of the scrotal surface# + layer of s'eletal muscle, the cremaster ('r A,+SAter) muscle,
lies deep to the dermis# Contraction of the cremaster muscle during se"ual arousal or in
response to decreased testicular temperature tenses the scrotum and pulls the testes closer to
the ody# Formal development of spermatozoa in the testes re&uires temperatures aout
0#0JC (.J() lower than those elsewhere in the ody# The cremaster and dartos muscles rela"
or contract to move the testes away from or toward the ody as needed to maintain
acceptale testicular temperatures# Khen air or ody temperature rises, these muscles rela"
and the testes move away from the ody# Sudden cooling of the scrotum, as occurs during
entry into a cold swimming pool, results in contractions that pull the testes closer to the
ody and 'eep testicular temperatures from falling#
Structure of the Testes
$eep to the tunica vaginalis covering the testis is the tunica al6u"inea (alA
232459324.doc 8
Auh), a dense layer of connective tissue rich
in collagen fiers ((igure ./G4a?)# These fiers are continuous with those surrounding the
ad!acent epididymis and e"tend into the sustance of the testis# There they form firous
partitions, or septa, that converge toward the region nearest the entrance to the epididymis#
The connective tissues in this region support the lood vessels and lymphatic vessels that
supply and drain the testis, and the efferent ductules, which transport spermatozoa to the
7istology of the Testes
The septa sudivide the testis into a series of lo6ules (see (igure ./G4a?)# $istriuted
among the loules are roughly /11 slender, tightly coiled seminiferous (semAiAF%(AerAus)
tu6ules ((igures ./G4 and ./G8?)# 6ach tuule averages aout /1 cm (2. in#) in
length, and a typical testis contains nearly oneAhalf mile of seminiferous tuules# Sperm
production occurs within these tuules#
6ach seminiferous tuule forms a loop that is connected to a maze of passageways 'nown as
the rete (R
C rete, a net) testis
(see (igure ./G4?)# (ifteen to .1 large efferent ductules connect the rete testis to the
>ecause the seminiferous tuules are tightly coiled, most histological preparations show
them in transverse section ((igure ./G8a?)# 6ach tuule is surrounded y a delicate
connective tissue capsule ((igure ./G8?), and areolar tissue fills the spaces etween the
tuules# Kithin those spaces are numerous lood vessels and large interstitial cells (cells of
Leydig)# %nterstitial cells are responsile for the production of androgens, the dominant se"
hormones in males# Testosterone is the most important androgen#
Spermatozoa are produced y the process of spermato"enesis (sperAmaAt
232459324.doc 9
AE6FAeAsis)# Spermatogenesis egins at the outer
most layer of cells in the seminiferous tuules and proceeds toward the lumen ((igure ./G
8c?)# +t each step in this process, the
daughter cells move closer to the lumen# (irst, stem cells called spermato"onia (sperAmaAt
Auh) divide y mitosis to
produce two daughter cells, one of which remains at that location as a spermatogonium
while the other differentiates into a pri
mary spermatocyte# !rimary spermatocytes (sperA,+At
L involved only in the production of gametes (spermatozoa in males, ova in females)# )rimary
spermatocytes give rise to secondary spermatocytes that differentiate into spermatids
(S)6RAmaAtidz)@immature gametes that suse&uently differentiate into spermatozoa# The
spermatozoa lose contact with the wall of the seminiferous tuule and enter the fluid in the
6ach seminiferous tuule contains spermatogonia, spermatocytes at various stages of
meiosis, spermatids, spermatozoa, and
ts) are the cells that egin meiosis, a specialized form of cell divisionAs
large sustentacular (susAtenAT+HA
Alar) cells (or Sertoli cells)# Sustentacular cells are attached to the tuular capsule and
to the lumen etween the other types of cells (see (igure ./G8,c?)#
232459324.doc 10
Spermatogenesis involves three integrated processes=
-3 Mitosis. Spermatogonia undergo cell divisions throughout adult life# (Mou can review the
description of mitosis and cell division in Chapter 2# lpp3 97,98) 9ne daughter cell from
each division remains in place while the other is pushed toward the
lumen of the seminiferous tuule# The displaced cells differentiate into primary
spermatocytes, which prepare to egin meiosis#
23 Meiosis. ,eiosis (m
L chromosomes, half the normal set# +s a result, the fusion of the nuclei of a male
gamete and a female gamete produces a cell that has the normal numer of chromosomes
(46), rather than twice that numer# %n the seminiferous tuules, meiotic divisions that egin
with primary spermatocytes produce spermatids, the undifferentiated male gametes#
23 83 Spermiogenesis. Spermatids are small, relatively unspecialized cells# %n
spermiogenesis, spermatids differentiate into physically mature spermatozoa, which are
among the most highly specialized cells in the ody# Spermiogenesis involves ma!or changes
in a spermatidNs internal and e"ternal structures#
,itosis and ,eiosis
%n oth males and females, mitosis and meiosis differ significantly in terms of the events
occurring in the nucleus# +s you may recall from Chapter 2, somatic cells contain .2 pairs of
chromosomes# 6ach pair consists of one chromosome provided y the father, and another
provided y the mother, at the time of fertilization# ,itosis is part of the process of somatic
cell division, producing two daughter cells each containing identical pairs of chromosomesC
the pattern is illustrated in (igure ./G6a?# >ecause daughter cells contain oth memers of
each chromosome pair (for a total of 46 chromosomes), they are called diploid ($%)AloydC
diplo, doule) cells# ,eiosis ((igure ./G6?) involves two cycles of cell division (meiosis I
and meiosis II) and produces four cells, each of which contains .2 individual chromosomes#
>ecause these cells contain only one memer of each pair of chromosomes, they are called
haploid (7+)AloydC haplo, single) cells# The events in the nucleus shown in (igure ./G6?
232459324.doc 11
are the same for the formation of spermatozoa or ova#
+s a cell prepares to egin meiosis, $F+ replication occurs within the nucleus !ust as it does
in a cell preparing to undergo mitosis# This similarity continues as prophase % arrivesC the
chromosomes condense and ecome visile with a light microscope# +s in mitosis, each
chromosome consists of two duplicate chromatids#
+t this point, the close similarities etween meiosis and mitosis end# %n meiosis, the
corresponding maternal and paternal chromosomes now come together, an event 'nown as
synapsis (siAF+)Asis)# Synapsis involves .2 pairs of chromosomesC each memer of each
pair consists of two chromatids# + matched set of four chromatids is called a tetrad (T6TA
radC tetras, four) (see (igure ./G6?)# Some e"change of genetic material can occur etween
the chromatids of a chromosome pair at this stage of meiosis# Such an e"change, called
crossing over, increases genetic variation among offspringC we will discuss it in Chapter .3#
,eiosis includes two division cycles, referred to as meiosis I and meiosis II# The stages
within each phase are identified as prophase %, metaphase %%, and so on# The nuclear
envelope disappears at the end of prophase %# +s metaphase % egins, the tetrads line up
along the metaphase plate# +s anaphase % egins, the tetrads rea' up@the maternal and
paternal chromosomes separate# This is a ma!or difference etween mitosis and meiosis= %n
mitosis, each daughter cell receives one of the two copies of every chromosome, maternal
and paternalC in meiosis %, each daughter cell receives oth copies of either the maternal
chromosome or the paternal chromosome from each tetrad# (Compare the two parts of
(igure ./G6?#)
+s anaphase proceeds, the maternal and paternal components are randomly and
independently distriuted# That is, as each tetrad splits, one cannot predict which daughter
cell will receive copies of the maternal chromosome, and which will receive copies
Asis) is a special form of cell division involved in gamete production# %n humans, gametes
contain .2
of the paternal chromosome# +s a result, telophase % ends with the formation of two
daughter cells containing uni&ue cominations of maternal and paternal chromosomes# >oth
cells contain .2 chromosomes# >ecause the first meiotic division reduces the numer of
232459324.doc 12
chromosomes from 46 to .2, it is called a reductional division# 6ach of these chromosomes
still consists of two duplicate chromatids# The duplicates will separate during meiosis %%#
The interphase separating meiosis % and meiosis %% is very rief, and no $F+ is replicated
during that period# 6ach cell proceeds through prophase %%, metaphase %%, and anaphase %%#
$uring anaphase %%, the duplicate chromatids separate# Telophase %% thus yields four cells,
each containing .2 chromosomes# >ecause the numer of chromosomes has not changed,
meiosis %% is an e9uational division# +lthough chromosomes are evenly distriuted among
these four cells, the cytoplasm may not e# %n males, meiosis produces four immature
gametes that are identical in sizeC each will develop into a functional sperm# %n females,
meiosis produces one huge ovum and three tiny, nonfunctional polar odies# Ke will
e"amine the details of spermatogenesis here and will consider oogensis in a later section#
%n spermatogenesis ((igure ./G<?), the mitotic division of each diploid spermatogonium
produces two daughter cells# 9ne is a spermatogonium that remains in contact with the asal
lamina, and the other is a primary spermatocyte that is displaced toward the lumen# +s
meiosis egins, each primary spermatocyte contains 46 individual chromosomes# +t the end
of meiosis %, the daughter cells are called secondary spermatocytes# 6very secondary
spermatocyte contains .2 chromosomes, each of which consists of a pair of duplicate
chromatids# The secondary spermatocytes soon enter prophase %%# The completion of
metaphase %%, anaphase %%, and telophase %% yields four haploid spermatids, each containing
.2 chromosomes#
(or each primary spermatocyte that enters meiosis, four spermatids are produced# >ecause
cyto'inesis (cytoplasmic division) is not completed in meiosis % or meiosis %%, the four
spermatids initially remain interconnected y ridges of cytoplasm# These connections assist
in the transfer of nutrients and hormones etween the cells, helping ensure that the cells
develop in synchrony# The ridges are not ro'en until the last stages of physical maturation#
-:: Keys ' ,eiosis produces gametes that contain half the numer of chromosomes found
in somatic cells# (or each cell
entering meiosis, the testes produce four spermatozoa, whereas the ovaries produce a single
%n spermio"enesis, the last step of spermatogenesis, each spermatid matures into a single
232459324.doc 13
spermatozoon, or sperm (see (igure ./G<?)# $eveloping spermatocytes undergoing meiosis,
and spermatids undergoing spermiogenesis, are not free in the seminiferous tuules# %nstead,
they are surrounded y the cytoplasm of the sustentacular cells# +s spermiogenesis
proceeds, the spermatids gradually develop the appearance of mature spermatozoa# +t
spermiation, a spermatozoon loses its attachment to the sustentacular cell and enters the
lumen of the seminiferous tuule# The entire process, from spermatogonial division to
spermiation, ta'es appro"imately nine wee's#
Sustentacular Cells Sustentacular cells, also called Sertoli cells, play a 'ey role in
spermatogenesis# These cells have si" important functions that directly or indirectly affect
mitosis, meiosis, and spermiogenesis within the seminiferous tuules=
-3 Maintenance of the lood!Testis arrier. The seminiferous tuules are isolated
from the general circulation y a 6lood,testis 6arrier, comparale in function to the loodG
rain arrier# lp3 ;<8 Sustentacular cells are !oined y tight !unctions, forming a layer that
divides the seminiferous tuule into an outer basal compartment, which contains the
spermatogonia, and an inner luminal compartment (or adluminal compartment), where
meiosis and spermiogenesis occur (see (igure ./G8c?)# Transport across the sustentacular
cells is tightly regulated, so conditions in the luminal compartment remain very stale# The
fluid in the lumen of a seminiferous tuule is produced y the sustentacular cells, which also
regulate the fluidNs composition# This fluid is very different from the surrounding interstitial
fluidC it is high in androgens, estrogens, potassium, and amino acids# The loodGtestis arrier
is essential to preserving the differences etween the tuular fluid and the interstitial fluid# %n
addition, this
arrier prevents immune system cells from detecting and attac'ing the developing
spermatozoa, which have in their cell memranes spermAspecific antigens not found in
somatic cell memranes and thus might e identified as Oforeign#P
23 23 Support of Mitosis and Meiosis. Spermatogenesis depends on the stimulation of
sustentacular cells y circulating follicleAstimulating hormone ((S7) and testosterone#
Stimulated sustentacular cells then promote the division of spermatogonia and the meiotic
divisions of spermatocytes#
83 83 Support of Spermiogenesis. Spermiogenesis re&uires the presence of sustentacular
cells# These cells surround and enfold the spermatids, providing nutrients and chemical
232459324.doc 14
stimuli that promote their development# Sustentacular cells also phagocytize cytoplasm that
is shed y spermatids as they develop into spermatozoa#
;3 ;3 Secretion of Inhibin. Sustentacular cells secrete the peptide hormone inhibin (inA
7%>Ain) in response to factors released y developing spermatozoa# %nhiin depresses the
pituitary production of (S7, and perhaps the hypothalamic secretion of gonadotropinA
releasing hormone (5nR7)# The faster the rate of sperm production, the more inhiin is
secreted# >y regulating (S7 and 5nR7 secretion, sustentacular cells provide feedac'
control of spermatogenesis#
<3 <3 Secretion of Androgen"inding #rotein. Androgen"binding protein $A#% inds
androgens (primarily testosterone) in the fluid contents of the seminiferous tuules# This
protein is thought to e important in oth elevating the concentration of androgens within
the seminiferous tuules and stimulating spermiogenesis# The production of +>) is
stimulated y (S7#
43 43 Secretion of M&llerian"Inhibiting 'actor. M&llerian"inhibiting factor $MI'% is
secreted y sustentacular cells in the developing testes# This hormone causes regression of
the fetal M&llerian ducts, passageways that participate in the formation of the uterine
tues and the uterus in females# %n males, inade&uate ,%( production during fetal
development leads to the retention of these ducts and the failure of the testes to descend into
the scrotum# +,= Testicular Cancer
The $natomy of a Spermato5oon
6ach spermatozoon has three distinct regions= the head, the middle piece, and the tail
((igure ./G/a?)# %n humans, the head is a
flattened ellipse containing a nucleus with densely pac'ed chromosomes# +t the tip of the
head is the acrosomal (a'AroBAS9Amal) cap, a memranous compartment containing
enzymes essential to fertilization# $uring spermiogenesis, saccules of the spermatidNs 5olgi
apparatus fuse and flatten into an acrosomal vesicle, which ultimately forms the acrosomal
cap of the spermatozoon#
+ short nec= attaches the head to the middle piece# The nec' contains oth centrioles of
the original spermatid# The microtuules of the distal centriole are continuous with those of
232459324.doc 15
the middle piece and tail# ,itochondria in the middle piece are arranged in a spiral around
the microtuules# ,itochondrial activity provides the +T) re&uired to move the tail#
The tail is the only flagellum in the human ody# + flagellum, a whipli'e organelle, moves a
cell from one place to another# Khereas cilia eat in a predictale, waveli'e fashion, the
flagellum of a spermatozoon has a comple", cor'screw motion#
:nli'e other, less specialized cells, a mature spermatozoon lac's an endoplasmic reticulum,
a 5olgi apparatus, lysosomes, pero"isomes, inclusions, and many other intracellular
structures# The loss of these organelles reduces the cellNs size and massC it is essentially a
moile carrier for the enclosed chromosomes, and e"tra weight would slow it down#
>ecause the cell lac's glycogen or other energy reserves, it must asor nutrients (primarily
fructose) from the surrounding fluid#
-:: Keys ' Spermatogenesis egins at puerty and continues until relatively late in life (past
age <1)# %t is a continuous
process, and all stages of meiosis can e oserved within the seminiferous tuules#
The Male Reproductive Tract
The testes produce physically mature spermatozoa that are incapale of successfully
fertilizing an oocyte# The other portions of the male reproductive system are responsile for
the functional maturation, nourishment, storage, and transport of spermatozoa#
The 6pididymis
Late in their development, spermatozoa detach from the sustentacular cells and lie within the
lumen of the seminiferous tuule# They have most of the physical characteristics of mature
spermatozoa, ut are functionally immature and incapale of coordinated locomotion or
fertilization# (luid currents, created y cilia lining the efferent ductules, transport the
immoile gametes into the epididymis (see (igure ./G4a?)# The epididymis, the start of the
male reproductive tract, is a coiled tue ound to the posterior order of the testis#
The epididymis can e felt through the s'in of the scrotum# + tuule almost < m (.2 ft) long,
the epididymis is coiled and twisted so as to ta'e up very little space# %t has a head, a ody,
and a tail ((igure ./G3a?)# The superior head is the portion of the epididymis pro"imal to
the testis# The head receives spermatozoa from the efferent ductules#
The 6ody egins distal to the last efferent ductule and e"tends inferiorly along the posterior
margin of the testis# Fear the inferior order of the testis, the numer of coils decreases,
232459324.doc 16
mar'ing the start of the tail# The tail recurves and ascends to its connection with the ductus
deferens# Spermatozoa are stored primarily within the tail of the epididymis#
The epididymis has three functions=
-3 -3 It Monitors and Adjusts the (omposition of the 'luid #roduced by the
Seminiferous Tubules. The pseudostratified columnar epithelial lining of the epididymis
ears distinctive stereocilia ((igure ./G3?), which increase the surface area availale for
asorption from, and secretion into, the fluid in the tuule#
23 23 It Acts as a )ecycling (enter for *amaged Spermatozoa. Cellular deris and
damaged spermatozoa are asored in the epididymis, and the products of enzymatic
rea'down are released into the surrounding interstitial fluids for pic'up y the epididymal
lood vessels#
83 83 It Stores and #rotects Spermatozoa and 'acilitates Their 'unctional Maturation.
+ spermatozoon passes through the epididymis in aout two wee's and completes its
functional maturation at that time# 9ver this period, spermatozoa e"ist in a sheltered
environment that is precisely regulated y the surrounding epithelial cells# +lthough
spermatozoa leaving the epididymis are mature, they remain immoile# To ecome motile
(actively swimming) and fully functional, spermatozoa must undergo a process called
capacitation# Capacitation normally occurs in two steps= (0) Spermatozoa ecome motile
when they are mi"ed with secretions of the seminal vesicles, and (.) they ecome capale of
successful fertilization when e"posed to conditions in the female reproductive tract# The
epididymis secretes a sustance (as yet unidentified) that prevents premature capacitation#
Transport along the epididymis involves a comination of fluid movement and peristaltic
contractions of smooth muscle in the walls of the epididymis# +fter passing along the tail of
the epididymis, the spermatozoa enter the ductus deferens#
The $uctus $eferens
6ach ductus deferens, or vas deferens, is 41G48 cm (06G0/ in#) long# %t egins at the tail of
the epididymis (see (igure ./G3a?) and, as part of the spermatic cord, ascends through the
inguinal canal (see (igure ./G2?, p# 0122)# %nside the adominal cavity, the ductus deferens
passes posteriorly, curving inferiorly along the lateral surface of the urinary ladder toward
the superior and posterior margin of the prostate gland (see (igure ./G0?)# Eust efore the
232459324.doc 17
ductus deferens reaches the prostate gland and seminal
vesicles, its lumen enlarges# This e"panded portion is 'nown as the ampulla (amA):LAuh)
of the ductus deferens
((igure ./G01a?)#
The wall of the ductus deferens contains a thic' layer of smooth muscle ((igure ./G01?)#
)eristaltic contractions in this layer propel spermatozoa and fluid along the duct, which is
lined y a pseudostratified ciliated columnar epithelium# %n addition to transporting
spermatozoa, the ductus deferens can store spermatozoa for several months# $uring this
time, the spermatozoa remain in a state of suspended animation and have low metaolic
The !unction of the ampulla with the duct of the seminal vesicle mar's the start of the
e>aculatory duct# This short passageway (. cm, or less than 0 in#) penetrates the muscular
wall of the prostate gland and empties into the urethra near the opening of the e!aculatory
duct from the opposite side (see (igures ./G0 and ./G01a?)#
The :rethra
%n males, the urethra e"tends 0/G.1 cm (<G/ in#) from the urinary ladder to the tip of the
penis (see (igure ./G0?)# %t is divided into prostatic, membranous, and spongy regions# The
male urethra is a passageway used y oth the urinary and reproductive systems#
The $ccessory 2lands
The fluids contriuted y the seminiferous tuules and the epididymis account for only aout
8 percent of the volume of semen# The fluid component of semen is a mi"ture of secretions
@each with distinctive iochemical characteristics@ from many glands# %mportant glands
include the seminal vesicles, the prostate gland, and the bulbourethral glands, all of which
occur only in males# +mong the ma!or functions of these glands are (0) activating
spermatozoaC (.) providing the nutrients spermatozoa need for motilityC (2) propelling
spermatozoa and fluids along the reproductive tract, mainly y peristaltic contractionsC and
(4) producing uffers that counteract the acidity of the urethral and vaginal environments#
The Seminal ;esicles
The ductus deferens on each side ends at the !unction etween the ampulla and the duct that
drains the seminal vesicle (see (igure ./G01a?)# The seminal vesicles are glands emedded
232459324.doc 18
in connective tissue on either side of the midline, sandwiched etween the posterior wall of
the urinary ladder and the rectum# 6ach seminal vesicle is a tuular gland with a total length
of aout 08 cm (6 in#)# The ody of the gland has many short side ranches# The entire
assemlage is coiled and folded into a compact, tapered mass roughly 8 cm Q .#8 cm (. in# Q
0 in#)#
Seminal vesicles are e"tremely active secretory glands with an epithelial lining that contains
e"tensive folds ((igure ./G01c?)# The seminal vesicles contriute aout 61 percent of the
volume of semen# +lthough the vesicular fluid generally has the same osmotic concentration
as that of lood plasma, the compositions of the two fluids are &uite different# %n particular,
the secretion of the seminal vesicles contains (0) higher concentrations of fructose, which is
easily metaolized y spermatozoaC (.) prostaglandins, which can stimulate smooth muscle
contractions along the male and female reproductive tractsC and (2) firinogen, which after
e!aculation forms a temporary clot within the vagina# The secretions of the seminal vesicles
are slightly al'aline, helping to neutralize acids in the secretions of the prostate gland and
within the vagina# Khen mi"ed with the secretions of the seminal vesicles, previously
inactive ut functional spermatozoa undergo the first step in capacitation and egin eating
their flagella, ecoming highly motile#
The secretions of the seminal vesicles are discharged into the e!aculatory duct at emission,
when peristaltic contractions are under way in the ductus deferens, seminal vesicles, and
prostate gland# These contractions are under the control of the sympathetic nervous system#
The )rostate 5land
The prostate "land is a small, muscular, rounded organ aout 4 cm (0#6 in#) in diameter#
The prostate gland encircles the pro"i
mal portion of the urethra as it leaves the urinary ladder (see (igure ./G01?)# The glandular
tissue of the prostate ((igure ./G01d?) consists of a cluster of 21G81 compound
tuuloalveolar glands# lp3 --7 These glands are surrounded y and wrapped in a thic'
lan'et of smooth muscle fiers#
The prostate gland produces prostatic fluid, a slightly acidic solution that contriutes .1G
21 percent of the volume of semen# %n addition to several other compounds of uncertain
significance, prostatic secretions contain seminalplasmin (semAiAnalA)L+DAmin), an
antiiotic that may help prevent urinary tract infections in males# These secretions are
232459324.doc 19
e!ected into the prostatic urethra y peristaltic contractions of the muscular prostate wall#
)rostatic inflammation, or prostatitis (prosAtaAT Atis), can occur in males at any age, ut it
most commonly afflicts older men#
%B)rostatitis can result from acterial infections ut also occurs in the apparent asence of
pathogens# Symptoms can resemle those of prostate cancer# %ndividuals with prostatitis
may complain of pain in the lower ac', perineum, or rectum, in some cases accompanied y
painful urination and the discharge of mucous secretions from the e"ternal urethral orifice#
+ntiiotic therapy is effective in treating most cases that result from acterial infection#
The >ulourethral 5lands
The paired 6ul6ourethral "lands, or (o+per,s glands, are situated at the ase of the penis,
covered y the fascia of the urogenital diaphragm ((igure ./G01a and ./G00a?)# The
ulourethral glands are round, with diameters approaching 01 mm (less than
1#8 in#)# The duct of each gland travels alongside the penile urethra for 2G4 cm (0#.G0#6 in#)
efore emptying into the urethral lumen# The ulourethral glands are compound,
tuuloalveolar mucous glands ((igure ./G01e?) that secrete a thic', al'aline mucus# The
secretion helps neutralize any urinary acids that may remain in the urethra, and it luricates
the glans, or tip of the penis#
+ typical e!aculation releases .G8 ml of semenC an anormally low volume may indicate
prolems with the prostate gland or seminal vesicles# Khen sampled for analysis, semen is
collected after a 26Ahour period of se"ual astinence# The volume of fluid produced y an
e!aculation, called the e>aculate, typically contains the following=
# ? Spermato5oa3 The normal sperm count ranges from .1 million to 011
million spermatozoa per milliliter of semen# ,ost individuals with lower sperm counts are
infertile, ecause too few spermatozoa survive the ascent of the female reproductive tract to
perform fertilization# + low sperm count may reflect inflammation of the epididymis, ductus
deferens, or prostate gland# %n a fertile male, at least 61 percent of the spermatozoa in the
sample are normal in appearanceC common anormalities are malformed heads and OtwinP
spermatozoa that did not separate at the time of spermiation# The normal sperm will e
swimming actively#
# ? Seminal luid3 Seminal fluid, the fluid component of semen, is a mi"ture of
232459324.doc 20
glandular secretions with a distinct ionic and nutrient composition# + typical sample of
seminal fluid contains the comined secretions of the seminal vesicles (61 percent), the
prostate gland (21 percent), the sustentacular cells and epididymis (8 percent), and the
ulourethral glands (less than 8 percent)#
# ? En5ymes3 Several important enzymes are present in seminal fluid, including
(0) a protease that may help dissolve mucous secretions in the vaginaC (.) seminalplasmin, an
antiiotic prostatic enzyme that 'ills a variety of acteria, including -scherichia coliC (2) a
prostatic enzyme that coagulates the semen within a few minutes after e!aculation y
converting firinogen to firinC and (4) fibrinolysin, which li&uefies the clotted semen after
08G21 minutes#
+ complete chemical analysis of semen appears in +ppendi" %;#
The E#ternal 2enitalia
The male e"ternal genitalia consist of the scrotum and penis# The structure of the scrotum
has already een descried (p# 0122)# The penis is a tuular organ through which the distal
portion of the urethra passes (see (igure ./G00a?)# %t conducts urine to the e"terior and
introduces semen into the femaleNs vagina during se"ual intercourse# The penis is divided
into three regions= the root, the ody, and the glans ((igure ./G00c?)# The root of the penis
is the fi"ed portion that attaches the penis to the ody wall# This connection occurs within
the urogenital triangle immediately inferior to the puic symphysis# The 6ody ?shaft@ of the
penis is the tuular, movale portion of the organ# The "lans of the penis is the e"panded
distal end that surrounds the e"ternal urethral orifice# The nec. is the narrow portion of the
penis etween the shaft and the glans#
The s'in overlying the penis resemles that of the scrotum# The dermis contains a layer of
smooth muscle that is a continuation of the dartos muscle of the scrotum, and the underlying
areolar tissue allows the thin s'in to move without distorting underlying structures# The
sucutaneous layer also contains superficial arteries, veins, and lymphatic vessels#
+ fold of s'in called the prepuce ()R6B
s), or, surrounds the tip of the penis# The prepuce attaches to the relatively
232459324.doc 21
narrow nec' of the penis and continues over the glans# !reputial (pr
Aal) "lands in the s'in of the nec' and the inner
surface of the prepuce secrete a wa"y material 'nown as sme"ma (S,65Ama)#
:nfortunately, smegma can e an e"cellent nutrient source for acteria# ,ild inflammation
and infections in this locale are common, especially if the area is not washed thoroughly and
fre&uently# 9ne way to avoid troule is circumcision (serA'umAS%D7Aun), the surgical
removal of the prepuce# %n Kestern societies (especially the :nited States), this procedure is
generally performed shortly after irth# +lthough the practice of circumcision remains
controversial, strong religious and cultural iases and epidemiological evidence suggest that
it will continue#
$eep to the areolar tissue, a dense networ' of elastic fiers encircles the internal structures
of the penis# ,ost of the ody of the penis consists of three cylindrical columns of erectile
tissue ((igure ./G00?)# 6rectile tissue consists of a threeAdimensional maze of vascular
channels incompletely separated y partitions of elastic connective tissue and smooth muscle
fiers# %n the resting state, the arterial ranches are constricted and the muscular partitions
are tense# This comination restricts lood flow into the erectile tissue# The parasympathetic
innervation of the penile arteries involves neurons that release nitric o"ide (F9) at their
synaptic 'nos# The smooth muscles in the arterial walls rela" when F9 is released, at which
time the vessels dilate, lood flow increases, the vascular channels ecome engorged with
lood, and erection of the penis occurs# The flaccid (nonerect) penis hangs inferior to the
puic symphysis and anterior to the scrotum, ut during erection the penis stiffens and
assumes a more upright position#
The anterior surface of the flaccid penis covers two cylindrical masses of erectile tissue= the
corpora cavernosa (H9RAporAa
232459324.doc 22
AsuhC singular, corpus cavernosum)# The two are separated y a thin septum and encircled y
a dense collagenous sheath
(see (igure ./G00?)# The corpora cavernosa diverge at their ases, forming the crura
(crura, legsC singular, crus) of the penis (see
(igure ./G00a?)# 6ach crus is ound to the ramus of the ischium and puis y tough
connectiveAtissue ligaments# The corpora cavernosa e"tend along the length of the penis as
far as its nec'# The erectile tissue within each corpus cavernosum surrounds a central artery
(see (igure ./G00?)#
The relatively slender corpus spon"iosum (sponA!
Asum) surrounds the penile urethra (see (igure ./G00a,?)# This erec
tile ody e"tends from the superficial fascia of the urogenital diaphragm to the tip of the
penis, where it e"pands to form the glans# The sheath surrounding the corpus spongiosum
contains more elastic fiers than does that of the corpora cavernosa, and the erectile tissue
contains a pair of small arteries#
$natomy 84: ' Review the anatomy of the male reproductive system on the $natomy 84:
C0AR&MB Reproductive SysAtemCMale Reproductive System3
.ormones and Male Reproductive unction
The hormonal interactions that regulate male reproductive function are diagrammed in
(igure ./G0.?# The ma!or reproductive hormones were introduced in Chapter 0/# lp3 422
The anterior loe of the pituitary gland releases follicle"stimulating hormone ?S.@ and
luteinizing hormone ?+.@# The pituitary release of these hormones occurs in response to
gonadotropin"releasing hormone ?2nR.@, a peptide synthesized in the hypothalamus and
carried to the anterior loe y the hypophyseal portal system#
232459324.doc 23
The hormone 5nR7 is secreted in pulses rather than continuously# %n adult males, small
pulses occur at 61G31Aminute intervals# +s levels of 5nR7 change, so do the rates of
secretion of (S7 and L7 (and testosterone, which is released in response to L7)# :nli'e the
situation in women, which we will consider later in the chapter, the 5nR7 pulse fre&uency
in adult males remains relatively steady from hour to hour, day to day, and year to year# +s a
result, plasma levels of (S7, L7, and testosterone remain within a relatively narrow range
until relatively late in life (see p# 016<)#
(S7 and Spermatogenesis
%n males, (S7 targets primarily the sustentacular cells of the seminiferous tuules# :nder
(S7 stimulation, and in the presence of testosterone from the interstitial cells, sustentacular
cells (0) promote spermatogenesis and spermiogenesis and (.) secrete androgenAinding
protein (+>))#
The rate of spermatogenesis is regulated y a negative feedac' mechanism involving
5nR7, (S7, and inhiin# :nder 5nR7 stimulation, (S7 promotes spermatogenesis along
the seminiferous tuules# +s spermatogenesis accelerates, however, so does the rate of
inhiin secretion y the sustentacular cells of the testes (see (igure ./G0.?)# %nhiin inhiits
(S7 production in the anterior loe of the pituitary gland and may also suppress the
secretion of 5nR7 at the hypothalamus#
The net effect is that when (S7 levels ecome elevated, inhiin production increases until
(S7 levels return to normal# %f (S7 levels decline, inhiin production falls, so the rate of
(S7 production accelerates#
L7 and +ndrogen )roduction
%n males, L7 induces the secretion of testosterone and other androgens y the interstitial
cells of the testes# Testosterone, the most important androgen, has numerous functions= (0)
stimulating spermatogenesis and promoting the functional maturation of spermatozoa,
through its effects on sustentacular cellsC (.) affecting central nervous system (CFS)
function, including the liido (se"ual drive) and related ehaviorsC (2) stimulating
metaolism throughout the ody, especially pathways concerned with protein synthesis,
lood cell formation, and muscle growthC (4) estalishing and maintaining male secondary
se" characteristics, such as the distriution of facial hair, increased muscle mass and ody
size, and the &uantity and location of characteristic adipose tissue depositsC and (8)
232459324.doc 24
maintaining the accessory glands and organs of the male reproductive tract#
Testosterone functions li'e other steroid hormones, circulating in the loodstream while
ound to one of two types of transport proteins= (0) gonadal steroid"binding globulin
(5>5), which carries roughly twoAthirds of the circulating testosterone, and
(.) the alumins, which ind the remaining oneAthird# Testosterone diffuses across the cell
memrane of target cells and inds to an intracellular receptor# The hormoneGreceptor
comple" then inds to the $F+ in the nucleus# %n many target tissues, some of the arriving
testosterone is converted to dihydrotestosterone ?0.T@# + small amount of $7T diffuses
ac' out of the cell and into the loodstream, and $7T levels are usually aout 01 percent
of circulating testosterone levels# $ihydrotestosterone can also enter peripheral cells and
ind to the same hormone receptors targeted y testosterone# %n addition, some tissues
(notaly those of the e"ternal genitalia) respond to $7T rather than to testosterone, and
other tissues (including the prostate gland) are more sensitive to $7T than to testosterone#
Testosterone production egins around the seventh wee' of fetal development and reaches a
prenatal pea' after roughly si" months# 9ver this period, the secretion of ,RllerianA
inhiiting factor y developing sustentacular cells leads to the regression of the ,Rllerian
ducts# The early surge in testosterone levels stimulates the differentiation of the male duct
system and accessory organs and affects CFS development# The estA'nown CFS effects
occur in the developing hypothalamus# There, testosterone apparently programs the
hypothalamic centers that are involved with (0) 5nR7 production and the regulation of
pituitary (S7 and L7 secretion, (.) se"ual ehaviors, and (2) se"ual drive# +s a result of
this prenatal e"posure to testosterone, the hypothalamic centers will respond appropriately
when the individual ecomes se"ually mature# The factors responsile for regulating the fetal
production of testosterone are not 'nown#
Testosterone levels are low at irth# :ntil puerty, ac'ground testosterone levels, although
still relatively low, are higher in males than in females# Testosterone secretion accelerates
mar'edly at puerty, initiating se"ual maturation and the appearance of secondary se"
characteristics# %n adult males, negative feedac' controls the level of testosterone
production# +oveAnormal testosterone levels inhiit the release of 5nR7 y the
hypothalamus, causing a reduction in L7 secretion and lowering testosterone levels (see
(igure ./G0.?)#
232459324.doc 25
The plasma of adult males also contains relatively small amounts of estradiol (. ng S dl,
versus 8.8 ng S dl of testosterone)# Seventy percent of the estradiol is formed from
circulating testosteroneC the rest is secreted, primarily y the interstitial and sustentacular
cells of the testes# The conversion of testosterone to estradiol is performed y an enzyme
called aromatase# (or un'nown reasons, estradiol production increases in older men#
Clinical /ote
*ehydroepiandrosterone, or 0.E$, is the primary androgen secreted y the zona
reticularis of the adrenal corte"# lp3 4-< +s noted in
Chapter 0/, these androgens, which are secreted in small amounts, are converted to
testosterone (or estrogens) y other tissues#
The significance of this small adrenal androgen contriution in oth se"es remains unclear,
ut $76+ is eing promoted as a wonder
drug for increasing vitality, strength, and muscle mass, and food supplements prepared from
wild ,e"ican yams are now eing ad
vertised as containing O$76+ precursors#P These claims are falseC the compounds contained
in these supplements have no effect on
circulating $76+ levels# The current recommendations are that $76+ use e restricted to
controlled, supervised clinical trials, and
that no one under age 41 use the drug# The effects of longAterm high doses of $76+ remain
largely un'nownC however, recall from
Chapter 0/ that the longAterm effects of androgen ause can e &uite serious# lp3 429 7igh
levels of $76+ in women have een
lin'ed to an increased ris' of ovarian cancer as well as to masculinization, due to the
conversion of $76+ to testosterone# The %9C (%nternational 9lympic Committee), FC++,
and F(L have anned the use of $76+ for muscle enhancement#
Concept Chec=
9n a warm day, would the cremaster muscle e contracted or rela"edT KhyT
Khat will happen if the arteries within the penis dilateT
Khat effect would low levels of (S7 have on sperm productionT
$ns%ers 6e"in on p3 $,-
The Reproductive System
232459324.doc 26
of the emale
# ? $escrie the components of the female reproductive system#
# ? 9utline the processes of meiosis and oogenesis in the ovaries#
# ? %dentify the phases and events of the ovarian and uterine cycles#
# ? $escrie the structure, histology, and functions of the vagina#
# ? Summarize the anatomical, physiological, and hormonal aspects of the female
reproductive cycle#
+ womanNs reproductive system produces se" hormones and functional gametes, and it must
also e ale to protect and support a developing emryo and nourish a neworn infant# The
principal organs of the female reproductive system are the ovaries, the uterine tubes, the
uterus, the vagina, and the components of the e"ternal genitalia ((igure ./G02?)# +s in
males, a variety of accessory glands release secretions into the female reproductive tract#
The ovaries, uterine tues, and uterus are enclosed within an e"tensive mesentery 'nown as
the 6road li"ament# The uterine tues run along the superior order of the road ligament
and open into the pelvic cavity lateral to the ovaries# The mesov
arium (mesA
Aum), a thic'ened fold of mesentery, supports and stailizes the position of each ovary (see
(igure ./G04a?)#
The road ligament attaches to the sides and floor of the pelvic cavity, where it ecomes
continuous with the parietal peritoneum# The road ligament thus sudivides this part of the
peritoneal cavity# The poc'et formed etween the posterior wall of the uterus
and the anterior surface of the colon is the rectouterine (re'At
232459324.doc 27
AterAin) pouch (see (igure ./G02?)C the poc'et formed etween
the uterus and the posterior wall of the ladder is the vesicouterine (vesAiA'
AterAin) pouch# These sudivisions are most ap
parent in sagittal section#
Several other ligaments assist the road ligament in supporting and stailizing the position of
the uterus and associated reproductive organs# These ligaments lie within the mesentery
sheet of the road ligament and are connected to the ovaries or uterus# The road ligament
limits sideAtoAside movement and rotation, and the other ligaments (descried in our
discussion of the ovaries and uterus) prevent superiorGinferior movement#
The &varies
The paired ovaries are small, lumpy, almondAshaped organs near the lateral walls of the
pelvic cavity ((igure ./G04?)# The ovaries perform three main functions= (0) production of
immature female gametes, or oocytes, (.) secretion of female se" hormones, including
estrogens and progestins, and (2) secretion of inhiin, involved in the feedac' control of
pituitary (S7 production#
The position of each ovary is stailized y the mesovarium and y a pair of supporting
ligaments= the ovarian ligament and the suspensory ligament (see (igure ./G04a?)# The
ovarian li"ament e"tends from the uterus, near the attachment of the uterine tue, to the
medial surface of the ovary# The suspensory li"ament e"tends from the lateral surface of
the ovary past the open end of the uterine tue to the pelvic wall# The suspensory ligament
contains the ma!or lood vessels of the ovary= the ovarian artery and ovarian vein# These
vessels are connected to the ovary at the ovarian hilum, where the ovary attaches to the
mesovarium ((igure ./G04?)#
+ typical ovary is aout 8 cm long, .#8 cm wide, and / mm thic' (. in# Q 0 in# Q 1#22 in#)
232459324.doc 28
and weighs 6G/ g (roughly
1#.8 oz)# +n ovary is pin' or yellowish and has a nodular consistency# The visceral
peritoneum, or germinal epithelium, covering the surface of each ovary consists of a layer of
columnar epithelial cells that overlies a dense connectiveAtissue layer called the tunica
al6u"inea (see (igure ./G04?)# Ke can divide the interior tissues, or stroma, of the ovary
into a superficial cortex and a deeper medulla# 5ametes are produced in the corte"#
9vum production, or oo"enesis (
AE6FAeAsis), egins efore a womanNs irth, accelerates at puerty, and ends at menopause#
>etween puerty and menopause, oogenesis occurs on a monthly asis as part of the
ovarian cycle#
9ogenesis is summarized in (igure ./G08?# :nli'e spermatogonia, the oo"onia (
Auh), or stem cells of females,
complete their mitotic divisions efore irth# >etween the third and seventh months of fetal
development, the daughter cells, or
primary oocytes (9
232459324.doc 29
L comes to a halt# The primary oocytes remain in a state of suspended development until the
individual reaches puerty, when rising levels of (S7 trigger the start of the ovarian cycle#
6ach month thereafter, some of the primary oocytes are stimulated to undergo further
development# Fot all primary oocytes produced during development survive until puerty#
The ovaries have roughly . million primordial follicles at irth, each containing a primary
oocyte# >y the time of puerty, the numer has dropped to aout
ts), prepare to undergo meiosis# They proceed as far as the prophase of meiosis %, ut then
the process A
411,111# The rest of the primordial follicles degenerate in a process called atresia (aATR
+lthough the nuclear events in the ovaries during meiosis are the same as those in the testes,
the process differs in two important details=
-3 -3 The cytoplasm of the primary oocyte is unevenly distriuted during the two
meiotic divisions# 9ogenesis produces one functional ovum, which contains most of the
original cytoplasm, and two or three polar 6odies, nonfunctional cells that later disintegrate
(see (igure ./G08?)#
23 23 The ovary releases a secondary oocyte rather than a mature ovum# The secondary
oocyte is suspended in metaphase of meiosis %%C meiosis will not e completed unless and
until fertilization occurs#
The 9varian Cycle
&varian follicles are specialized structures in the corte" of the ovaries where oth oocyte
growth and meiosis % occur# )rimary oocytes are located in the outer portion of the ovarian
corte", near the tunica aluginea, in clusters called egg nests ((igure ./G06?)# 6ach primary
232459324.doc 30
oocyte within an egg nest is surrounded y a single s&uamous layer of follicle cells# The
primary oocyte and its follicle cells form a primordial follicle# +fter se"ual maturation, a
different group of primordial follicles is activated each month# This monthly process is
'nown as the ovarian cycle#
The ovarian cycle can e divided into a follicular phase, or preovulatory phase, and a
luteal phase, or postovulatory phase# %mportant steps in the ovarian cycle can e
summarized as follows (see (igure ./G06?)=
Step / The 'ormation of #rimary 'ollicles. The ovarian cycle egins as activated primordial
follicles develop into primary follicles# %n a primary follicle, the follicle cells enlarge and
undergo repeated divisions that create several layers of follicle cells around the oocyte#
These follicle cells, which ecome rounded in appearance, are now called "ranulosa cells#
+s layers of granulosa cells develop around the primary oocyte, microvilli from the
surrounding granulosa cells intermingle with those of the primary oocyte# The microvilli are
surrounded y a layer of glycoproteinsC the entire region is called the 5ona pellucida (D9Ana
peAL99AsidAuh)# The microvilli increase the surface area availale for the transfer of
materials from the granu
losa cells to the rapidly enlarging primary oocyte#
The conversion from primordial to primary follicles and suse&uent follicular development
occurs under stimulation of (S7 from the anterior loe of the pituitary gland# +s the
granulosa cells enlarge and multiply, ad!acent cells in the ovarian stroma form a layer of
thecal cells around the follicle# Thecal cells and granulosa cells wor' together to produce
se" hormones called estrogens#
Step 0 The 'ormation of Secondary 'ollicles. +lthough many primordial follicles develop
into primary follicles, only a few will proceed to this step# The transformation egins as the
wall of the follicle thic'ens and the granulosa cells egin secreting small amounts of fluid#
This follicular fluid, or li1uor folliculi, accumulates in small poc'ets that gradually e"pand
and separate the inner and outer layers of the follicle# +t this stage, the comple" is 'nown as
a secondary follicle# +lthough the primary oocyte continues to grow at a steady pace, the
follicle as a whole now enlarges rapidly ecause follicular fluid accumulates#
Step 2 The 'ormation of a Tertiary 'ollicle. 6ight to 01 days after the start of the ovarian
232459324.doc 31
cycle, the ovaries generally contain only a single secondary follicle destined for further
development# >y the 01th to the 04th day of the cycle, that follicle has ecome a
tertiary follicle, or mature 3raafian (5R+(AAan) follicle, roughly 08 mm in diameter# This
comple" spans the entire width of the ovarian corte" and distorts the ovarian capsule,
creating a prominent ulge in the surface of the ovary# The oocyte pro!ects into the antrum
(+FAtrum), or e"panded central chamer of the follicle# The antrum is surrounded y a mass
of granulosa cells#
:ntil this time, the primary oocyte has een suspended in prophase of meiosis %# +s the
development of the tertiary follicle ends, L7 levels egin rising, prompting the primary
oocyte to complete meiosis %# %nstead of producing two secondary oocytes, the first meiotic
division yields a secondary oocyte and a small, nonfunctional polar ody# The secondary
oocyte then enters meiosis %%, ut stops once again upon reaching metaphase# ,eiosis %% will
not e completed unless fertilization occurs#
5enerally, on day 04 of a ./Aday cycle, the secondary oocyte and the attached granulosa
cells lose their connections with the follicular wall and drift free within the antrum# The
granulosa cells still associated with the secondary oocyte form a protective
layer 'nown as the corona radiata ('oAR
Anuh r
Step 4 5vulation. +t ovulation, the tertiary follicle releases the secondary oocyte# The
distended follicular wall suddenly ruptures, e!ecting the follicular contents, including the
secondary oocyte and corona radiata, into the pelvic cavity# The stic'y follicular fluid 'eeps
the corona radiata attached to the surface of the ovary, where direct contact with pro!ections
surrounding the entrance to the uterine tue or with fluid currents estalished y the ciliated
epithelium lining the uterus can transfer the secondary oocyte to the uterine tue# 9vulation
232459324.doc 32
mar's the end of the follicular phase of the ovarian cycle and the start of the luteal phase#
Step 6 The 'ormation and *egeneration of the (orpus Luteum. The empty tertiary follicle
initially collapses, and ruptured vessels leed into the antrum# The remaining granulosa cells
then invade the area, proliferating to create an endocrine structure 'nown
as the corpus luteum (L99At Aum), named for its yellow color (lutea, yellow)# This process
occurs under L7 stimulation# The cholesterol contained in the corpus luteum is used to
manufacture steroid hormones 'nown as pro"estins (pr
principally the steroid pro"esterone (pr
n)# +lthough the corpus luteum also secretes moderate amounts of estrogens,
levels are not as high as they were at ovulation, and progesterone is the principal hormone in
the luteal phase# )rogesteroneNs primary function is to prepare the uterus for pregnancy y
stimulating the maturation of the uterine lining and the secretions of uterine glands#
Step 7 8nless 'ertilization 5ccurs9 the (orpus Luteum egins to *egenerate )oughly /0
*ays after 5vulation. )rogesterone and estrogen levels then fall mar'edly# (irolasts
invade the nonfunctional corpus luteum, producing a 'not of pale scar tissue called a corpus
al6icans (+LAiA'anz)# The disintegration, or involution, of the corpus luteum mar's the
end of the ovarian cycle# + new ovarian cycle then egins with the activation of another
group of primordial follicles#
+ge and 9ogenesis
+lthough many primordial follicles may have developed into primary follicles, and several
primary follicles may have een converted to secondary follicles, generally only a single
oocyte is released into the pelvic cavity at ovulation# The rest undergo atresia# +t puerty,
each ovary contains aout .11,111 primordial follicles# (orty years later, few if any follicles
remain, although only aout 811 will have een ovulated in the interim# +,= 9varian Cancer
-:: Keys ' 9ogenesis egins during emryonic development, and primary oocyte
232459324.doc 33
production is completed efore irth# 6ach month after puerty, the ovarian cycle produces
one or more secondary oocytes from the preAe"isting population of primary oocytes# The
numer of viale and responsive primary oocytes declines mar'edly over time, until ovarian
cycles end at age 48G88#
The (terine Tu6es
6ach uterine tu6e ('allopian tube or oviduct) is a hollow, muscular tue measuring
roughly 02 cm (8#. in#) in length (see (igures ./G02 and (igure ./G04?)# 6ach uterine tue
is divided into three segments ((igure ./G0<a?)=
-3 The Infundibulum. The end closest to the ovary forms an e"panded funnel, or
infundi6ulum, with numerous fingerli'e pro
!ections that e"tend into the pelvic cavity# The pro!ections are called fim6riae ((%,Ar
)# The inner surfaces of the in
fundiulum are lined with cilia that eat toward the middle segment of the uterine tue,
called the ampulla#
-3 23 The Ampulla. The thic'ness of the smooth muscle layers in the wall of the
ampulla, the middle segment of the uterine tue, gradually increases as the tue approaches
the uterus#
23 83 The Isthmus. The ampulla leads to the isthmus (%SAmus), a short segment
connected to the uterine wall#
7istology of the :terine Tue
The epithelium lining the uterine tue is composed of ciliated columnar epithelial cells with
scattered mucinAsecreting cells ((igure ./G0<c?)# The mucosa is surrounded y concentric
layers of smooth muscle ((igure ./G0<?)# 9ocyte transport involves a comination of
ciliary movement and peristaltic contractions in the walls of the uterine tue# + few hours
efore ovulation, sympathetic and parasympathetic nerves from the hypogastric ple"us Oturn
onP this eating pattern and initiate peristalsis# %t normally ta'es three to four days for an
232459324.doc 34
oocyte to travel from the infundiulum to the uterine cavity# %f fertilization is to occur, the
secondary oocyte must encounter spermatozoa during the first 0.G.4 hours of its passage#
(ertilization typically occurs near the oundary etween the ampulla and isthmus of the
uterine tue#
%n addition to its transport function, the uterine tue provides a nutrientArich environment
that contains lipids and glycogen# This mi"ture supplies nutrients to oth spermatozoa and a
developing pre"embryo (the cell cluster produced y the initial divisions following
fertilization)# :nfertilized oocytes degenerate in the terminal portions of the uterine tues or
within the uterus# +,= )elvic %nflammatory $isease
The (terus
The uterus provides mechanical protection, nutritional support, and waste removal for the
developing embryo (wee's 0G/) and fetus (wee' 3 through delivery)# %n addition,
contractions in the muscular wall of the uterus are important in e!ecting the fetus at irth#
The uterus is a small, pearAshaped organ ((igure ./G0/a?) aout <#8 cm (2 in#) long and
with a ma"imum diameter of 8 cm (. in#)# %t weighs 21G41 g (0G0#4 oz)# %n its normal
position, the uterus ends anteriorly near its ase (see (igure ./G02?, p# 014/),
a condition 'nown as anteflexion (anAt A(L6HAshun)# %n this position, the uterus covers the
superior and posterior surfaces of the urinary ladder# %f the uterus ends ac'ward toward
the sacrum, the condition is termed retroflexion (reAtr
Retrofle"ion, which occurs in aout .1 percent of adult women, has no clinical significance#
(+ retrofle"ed uterus generally ecomes antefle"ed spontaneously during the third month of
Suspensory Ligaments of the :terus
%n addition to the road ligament, three pairs of suspensory ligaments stailize the position
of the uterus and limit its range of movement ((igure ./G0/?)# The uterosacral
232459324.doc 35
A'ral) li"aments e"tend from the lateral surfaces of the uterus to the anterior face of the
sacrum, 'eeping the ody
of the uterus from moving inferiorly and anteriorly# The round li"aments arise on the lateral
margins of the uterus !ust posterior and inferior to the attachments of the uterine tues#
These ligaments e"tend through the inguinal canal and end in the connective tissues of the
e"ternal genitalia# The round ligaments restrict primarily posterior movement of the uterus#
The lateral (cardinal) li"aments e"tend from the ase of the uterus and vagina to the lateral
walls of the pelvis# These ligaments tend to prevent inferior movement of the uterus#
+dditional mechanical support is provided y the muscles and fascia of the pelvic floor#
%nternal +natomy of the :terus
Ke can divide the uterus into two anatomical regions (see (igure ./G0/a?)= the ody and
the cervi"# The uterine 6ody, or corpus, is the largest portion of the uterus# The fundus is
the rounded portion of the ody superior to the attachment of the uterine tues# The ody
ends at a constriction 'nown as the isthmus of the uterus# The cervi# (S6RAvi's) is the
inferior portion of the uterus that e"tends from the isthmus to the vagina#
The tuular cervi" pro!ects aout 0#.8 cm (1#8 in#) into the vagina# Kithin the vagina, the
distal end of the cervi" forms a curving surface that surrounds the cervical os (os, an
opening or mouth), or external orifice of the uterus# The cervical os leads into the cervical
canal, a constricted passageway that opens into the uterine cavity of the ody at the
internal os, or internal orifice#
The uterus receives lood from ranches of the uterine arteries, which arise from ranches
of the internal iliac arteries, and from the ovarian arteries, which arise from the adominal
aorta inferior to the renal arteries# The arteries to the uterus are e"tensively interconnected,
ensuring a reliale flow of lood to the organ despite changes in its position and shape
during pregnancy# Fumerous veins and lymphatic vessels also drain each portion of the
uterus# The organ is innervated y autonomic fiers from the hypogastric ple"us
232459324.doc 36
(sympathetic) and from sacral segments S2 and S4 (parasympathetic)# Sensory information
reaches the central nervous system within the dorsal roots of spinal nerves T00 and T0.#
The most delicate anesthetic procedures used dur
ing laor and delivery, 'nown as segmental bloc.s, target only spinal nerves T01
The :terine Kall
The dimensions of the uterus are highly variale# %n women of reproductive age who have
not given irth, the uterine wall is aout
0#8 cm (1#6 in#) thic'# The wall has a thic', outer, muscular myometrium (m
L Aum), or mucosa ((igure ./G03?)# The fundus and the posterior surA
AumC myoA, muscle metra, uterus)
and a thin, inner, glandular endometrium (enAd
face of the uterine ody and isthmus are covered y a serous memrane that is continuous
232459324.doc 37
with the peritoneal lining# This incomplete serosa is called the perimetrium#
The endometrium contriutes aout 01 percent to the mass of the uterus# The glandular and
vascular tissues of the endometrium support the physiological demands of the growing fetus#
;ast numers of uterine glands open onto the endometrial surface and e"tend deep into the
lamina propria, almost to the myometrium# :nder the influence of estrogen, the uterine
glands, lood vessels, and epithelium change with the phases of the monthly uterine cycle#
The myometrium, the thic'est portion of the uterine wall, constitutes almost 31 percent of
the mass of the uterus# Smooth muscle in the myometrium is arranged into longitudinal,
circular, and oli&ue layers# The smooth muscle tissue of the myometrium provides much of
the force needed to move a fetus out of the uterus and into the vagina#
7istology of the :terus Ke can divide the endometrium into a functional 5one@the layer
closest to the uterine cavity@and a 6asilar 5one, ad!acent to the myometrium (see (igure
./G03?)# The functional zone contains most of the uterine glands and contriutes most of
the endometrial thic'ness# %t is this zone that undergoes the dramatic changes in thic'ness
and structure during the menstrual cycle# The asilar zone attaches the endometrium to the
myometrium and contains the terminal ranches of the tuular endometrial glands#
Kithin the myometrium, ranches of the uterine arteries form arcuate arteries, which
encircle the endometrium (see (igure ./G03a?)# Radial arteries supply strai"ht arteries,
which deliver lood to the asilar zone of the endometrium, and spiral arteries, which
supply the functional zone#
The structure of the asilar zone remains relatively constant over time, ut that of the
functional zone undergoes cyclical changes in response to se" hormone levels# These cyclical
changes produce the characteristic histological features of the uterine cycle#
Cervical cancer is the most common cancer of the reproductive system in women ages 08G
24# 6ach year roughly 02,111 :#S# women are diagnosed with invasive cervical cancer, and
appro"imately oneAthird of them eventually die from the condition# +nother 28,111 women
are diagnosed with a less aggressive form of cervical cancer# Cervical and other uterine
tumors and cancers are discussed in the Applications Manual# +,= :terine Tumors and
The :terine Cycle
The uterine cycle, or menstrual (,6FAstrooAal) cycle, is a repeating series of changes in
232459324.doc 38
the structure of the endometrium ((igure ./G.1?)# The uterine cycle averages ./ days in
length, ut it can range from .0 to 28 days in healthy women of reproductive age# Ke can
divide the uterine cycle into three phases= (0) menses, (.) the proliferative phase, and (2) the
secretory phase# The phases occur in response to hormones associated with the regulation of
the ovarian cycle# ,enses and the proliferative phase occur during the follicular phase of the
ovarian cycleC the secretory phase corresponds to the luteal phase of the cycle# Ke will
consider the regulatory mechanism involved in a later section#
The uterine cycle egins with the onset of menses (,6FAs z), an interval mar'ed y the
degeneration of the functional zone of the endometrium ((igure ./G.1a?)# This degeneration
occurs in patches and is caused y constriction of the spiral arteries, which reduces lood
flow to areas of endometrium# $eprived of o"ygen and nutrients, the secretory glands and
other tissues in the functional zone egin to deteriorate# 6ventually, the wea'ened arterial
walls rupture, and lood pours into the connective tissues of the functional zone# >lood cells
and degenerating tissues then rea' away and enter the uterine lumen, to e lost y passage
through the cervical os and into the vagina# 9nly the functional zone is affected, ecause the
deeper, asilar zone is provided with lood from the straight arteries, which remain
The sloughing off of tissue is gradual, and at each site repairs egin almost at once#
Fevertheless, efore menses has ended,
the entire functional zone has een lost# The process of endometrial sloughing, called
menstruation (menAstrooA
Ashun), gen
erally lasts from one to seven days# 9ver this period roughly 28 to 81 ml of lood are lost#
The process can e relatively painless# )ainful menstruation, or dysmenorrhea, can result
from uterine inflammation, myometrial contractions (OcrampsP), or from conditions
involving ad!acent pelvic structures#
232459324.doc 39
The )roliferative )hase The asilar zone, including the asal parts of the uterine glands,
survives menses intact# %n the days after menses, the epithelial cells of the uterine glands
multiply and spread across the endometrial surface, restoring the integrity of the uterine
epithelium ((igure ./G.1?)# (urther growth and vascularization result in the complete
restoration of the functional zone# +s this reorganization proceeds, the endometrium is in
the proliferative phase# Restoration occurs at the same time as the enlargement of primary
and secondary follicles in the ovary# The proliferative phase is stimulated and sustained y
estrogens secreted y the developing ovarian follicles#
>y the time ovulation occurs, the functional zone is several millimeters thic', and prominent
mucous glands e"tend to the order with the asilar zone# +t this time, the endometrial
glands are manufacturing a mucus rich in glycogen# This specialized mucus appears to e
essential for the survival of the fertilized egg through its earliest developmental stages#
(These stages will e considered in Chapter .3#) The entire functional zone is highly
vascularized, with small arteries spiraling toward the inner surface from larger arteries in the
The Secretory )hase $uring the secretory phase of the uterine cycle, the endometrial
glands enlarge, accelerating their rates of secretion, and the arteries that supply the uterine
wall elongate and spiral through the tissues of the functional zone ((igure ./G.1c?)# This
activity occurs under the comined stimulatory effects of progestins and estrogens from the
corpus luteum# The secretory phase egins at the time of ovulation and persists as long as
the corpus luteum remains intact#
Secretory activities pea' aout 0. days after ovulation# 9ver the ne"t day or two, glandular
activity declines, and the uterine cycle ends as the corpus luteum stops producing
stimulatory hormones# + new cycle then egins with the onset of menses and the
disintegration of the functional zone# The secretory phase generally lasts 04 days# +s a
result, you can identify the date of ovulation y counting ac'ward 04 days from the first
day of menses# +,= 6ndometriosis
,enarche and ,enopause The uterine cycle egins at puerty# The first cycle, 'nown as
menarche (meAF+RA'
C men, month
232459324.doc 40
arche eginning), typically occurs at age 00G0.# The cycles continue until menopause
Apawz), the termination of the
uterine cycle, at age 48G88# 9ver the interim, the regular appearance of uterine cycles is
interrupted only y circumstances such as illness, stress, starvation, or pregnancy#
%f menarche does not appear y age 06, or if the normal uterine cycle of an adult woman
ecomes interrupted for si" months
or more, the condition of amenorrhea (
Auh) e"ists# #rimary amenorrhea is the failure to initiate menses# This con
dition may indicate developmental anormalities, such as nonfunctional ovaries, the asence
of a uterus, or an endocrine or ge
netic disorder# %t can also result from malnutrition= )uerty is delayed if leptin levels are too
low# lp3 42; Transient secondary amenorrhea can e caused y severe physical or emotional
stresses# %n effect, the reproductive system gets Oswitched off#P (actors that cause either
type of amenorrhea include drastic weight loss, anore"ia nervosa, and severe depression or
grief# +menorrhea has also een oserved in marathon runners and other women engaged in
training programs that re&uire sustained high levels of e"ertion and severely reduce ody
lipid reserves#
The 1a"ina
The va"ina is an elastic, muscular tue e"tending etween the cervi" and the vestibule, a
space ounded y the female e"ternal genitalia (see (igure ./G02?)# The vagina is typically
<#8G3 cm (2G2#6 in#) long, ut its diameter varies ecause it is highly distensile#
232459324.doc 41
+t the pro"imal end of the vagina, the cervi" pro!ects into the va"inal canal# The shallow
recess surrounding the cervical protrusion is 'nown as the forni# ((9RAni's)# The vagina
lies parallel to the rectum, and the two are in close contact posteriorly# +nteriorly, the
urethra e"tends along the superior wall of the vagina from the urinary ladder to the e"ternal
urethral orifice, which opens into the vestiule# The primary lood supply of the vagina is via
the va"inal 6ranches of the internal iliac (or uterine) ar
teries and veins# %nnervation is from the hypogastric ple"us, sacral nerves S. GS4, and
ranches of the pudendal nerve# lpp3 ;8<, 7;<, 7<:
The vagina has three ma!or functions= %t (0) serves as a passageway for the elimination of
menstrual fluids, (.) receives the penis during se"ual intercourse, and holds spermatozoa
prior to their passage into the uterus, and (2) forms the inferior portion of the birth canal,
through which the fetus passes during delivery#
+natomy and 7istology of the ;agina
%n sectional view, the lumen of the vagina appears constricted, forming a rough 7# The
vaginal walls contain a networ' of lood vessels and layers of smooth muscle ((igure ./G
.0?)# The lining is moistened y secretions of the cervical glands and y the movement of
water across the permeale epithelium# Throughout childhood the vagina and vestiule are
usually separated y the hymen
(7 Amen), an elastic epithelial fold that partially or completely loc's the entrance to the
vaginaC an intact hymen is typically rup
tured during se"ual intercourse or tampon usage# The two bulbospongiosus muscles e"tend
along either side of the vaginal entrance, which is constricted y their contractions# lp3 8;9
These muscles cover the vestibular bulbs, masses of erectile tissue on either side of the
vaginal entrance# The vestiular uls have the same emryological origins as the corpus
spongiosum of the penis in males#
The vaginal lumen is lined y a non'eratinized stratified s&uamous epithelium (see (igure
./G.0?)# %n the rela"ed state, this epithelium forms folds called rugae# The underlying lamina
propria is thic' and elastic, and it contains small lood vessels, nerves, and lymph nodes#
The vaginal mucosa is surrounded y an elastic muscularis layer consisting of layers of
smooth muscle fiers arranged in circular and longitudinal undles continuous with the
uterine myometrium# The portion of the vagina ad!acent to the uterus has a serosal covering
232459324.doc 42
that is continuous with the pelvic peritoneum# +long the rest of the vagina, the muscularis
layer is surrounded y an adventitia of firous connective tissue#
The vagina contains a population of resident acteria, usually harmless, supported y
nutrients in the cervical mucus# The metaolic activity of these acteria creates an acidic
environment, which restricts the growth of many pathogens# 1a"initis (va!A
B% The hormonal changes associated with the ovarian cycle also affect the vaginal epithelium# >y e"amining a vaginal smear
iAF Atis), an inflammation of the vaginal canal, is caused y fungi, acteria, or parasites# %n
addition to any discomfort that may result, the condition may affect the survival of
spermatozoa and therey reduce fertility# +n acidic environment also inhiits the motility of
spermC for this reason, the uffers in semen are important to successful fertilization# +,=
sample of epithelial cells shed at the surface of the vagina@a clinician can estimate the
corresponding stage in the ovarian and uterine cycles# This diagnostic procedure is an
e"ample of exfoliative cytology# lp3 --;
The E#ternal 2enitalia
The area containing the female e"ternal genitalia is the vulva (;:LAvuh), or pudendum (puA
$6FAdumC (igure ./G..?)# The vagina
opens into the vesti6ule, a central space ounded y small folds 'nown as the la6ia minora
Auh miAF9RAuhC singular,
labium minus)# The laia minora are covered with a smooth, hairless s'in# The urethra opens
into the vestiule !ust anterior to the vaginal entrance# The paraurethral "lands, or S.ene,s
glands, discharge into the urethra near the e"ternal urethral opening# +nterior to this
opening, the clitoris (HL%TAoAris) pro!ects into the vestiule# + small, rounded tissue
pro!ection, the clitoris is derived from the same emryonic structures as the penis in males#
%nternally, it contains erectile tissue comparale to the corpora cavernosa of the penis# The
clitoris engorges with lood during se"ual arousal# + small erectile glans sits atop itC
232459324.doc 43
e"tensions of the laia minora encircle the ody of the clitoris, forming its prepuce, or hood#
+TL+S= 6mryology Summary .0= The $evelopment of the Reproductive System
+ variale numer of small lesser vesti6ular "lands discharge their secretions onto the
e"posed surface of the vestiule, 'eeping it moist# $uring se"ual arousal, a pair of ducts
discharges the secretions of the "reater vesti6ular "lands (artholin,s glands) into the
vestiule near the posterolateral margins of the vaginal entrance# These mucous glands have
the same emryological origins as the ulourethral glands of males#
The outer limits of the vulva are estalished y the mons puis and the laia ma!ora# The
ulge of the mons pu6is is created y adipose tissue deep to the s'in superficial to the puic
symphysis# +dipose tissue also accumulates within the la6ia ma>ora (singular, labium
majus), prominent folds of s'in that encircle and partially conceal the laia minora and
ad!acent structures# The outer margins of the laia ma!ora and the mons puis are covered
with coarse hair, ut the inner surfaces of the laia ma!ora are relatively hairless# Seaceous
glands and scattered apocrine sweat glands release their secretions onto the inner surface of
the laia ma!ora, moistening and luricating them#
The Mammary 2lands
+ neworn infant cannot fend for itself, and several of its 'ey systems have yet to complete
development# 9ver the initial period
of ad!ustment to an independent e"istence, the infant can gain nourishment from the mil'
secreted y the maternal mammary "lands# ,il' production, or lactation (la'AT+BAshun),
occurs in these glands# %n females, mammary glands are specialized organs of the
integumentary system that are controlled mainly y hormones of the reproductive system
and y the placenta, a temporary structure that provides the emryo and fetus with
9n each side, a mammary gland lies in the sucutaneous tissue of the pectoral fat pad deep
to the s'in of the chest
((igure ./G.2a?)# 6ach reast ears a nipple, a small conical pro!ection where the ducts of
the underlying mammary gland open onto the ody surface# The reddishArown s'in around
each nipple is the areola (aAR6BAoBAluh)# Large seaceous glands deep to the areolar
surface give it a grainy te"ture#
The glandular tissue of a mammary gland consists of separate loes, each containing several
232459324.doc 44
secretory loules# $ucts leaving the loules converge, giving rise to a single lactiferous
(la'AT%(AerAus) duct in each loe# Fear the nipple, that lactiferous duct enlarges, forming an
e"panded chamer called a lactiferous sinus# Typically, 08G.1 lactiferous sinuses open onto
the surface of each nipple# $ense connective tissue surrounds the duct system and forms
partitions that e"tend etween the loes and the loules# These ands of connective tissue,
the suspensory ligaments of the breast, originate in the dermis of the overlying s'in# + layer
of areolar tissue separates the mammary gland comple" from the underlying pectoralis
muscles# >ranches of the internal thoracic artery (see (igure .0G.0?, p# <23) supply lood
to each mammary gland#
(igure ./G.2,c? compares the histological organizations of inactive and active mammary
glands# +n inactive, or resting, mammary gland is dominated y a duct system rather than y
active glandular cells# The size of the mammary glands in a nonpregnant woman reflects
primarily the amount of adipose tissue present, not the amount of glandular tissue# The
secretory apparatus normally does not complete its development unless pregnancy occurs#
+n active mammary gland is a tuuloalveolar gland, consisting of multiple glandular tues
that end in secretory alveoli# Ke will discuss the hormonal mechanisms involved in lactation
in Chapter .3#
$natomy 84: ' Review the anatomy of the female reproductive system on the $natomy
84: C0AR&MB Reproductive SysAtemCemale Reproductive System3
Concept Chec=
+s the result of infections such as gonorrhea, scar tissue can loc' the lumen of each
uterine tue# 7ow would this loc'age affect a womanNs aility to conceiveT Khat is the
advantage of the acidic p7 of the vaginaT
Khich layer of the uterus is sloughed off during menstruationT
Kould the loc'age of a single lactiferous sinus interfere with the delivery of mil' to the
nippleT 6"plain#
$ns%ers 6e"in on p3 $,-
.ormones and the emale Reproductive Cycle
The activity of the female reproductive tract is under hormonal control that involves an
interplay etween secretions of oth the pituitary gland and the gonads# >ut the regulatory
pattern in females is much more complicated than in males, ecause it must coordinate the
232459324.doc 45
ovarian and uterine cycles# Circulating hormones control the female reproductive cycle,
coordinating the ovarian and uterine cycles to ensure proper reproductive function# %f the
two cycles are not properly coordinated, infertility results# + woman who fails to ovulate
cannot conceive, even if her uterus is perfectly normal# + woman who ovulates normally,
ut whose uterus is not ready to support an emryo, will also e infertile# >ecause the
processes are comple" and difficult to study, many of the iochemical details of the female
reproductive cycle still elude us, ut the general patterns are reasonaly clear#
+s in males, 5nR7 from the hypothalamus regulates reproductive function in females#
7owever, in females, the 5nR7 pulse fre&uency and amplitude (amount secreted per pulse)
change throughout the course of the ovarian cycle# %f the hypothalamus were a radio station,
the pulse fre&uency would correspond to the radio fre&uency itNs transmitting on, and the
amplitude would e the volume# Ke will consider changes in pulse fre&uency, as their effects
are oth dramatic and reasonaly well understood# Changes in 5nR7 pulse fre&uency are
controlled primarily y circulating levels of estrogens and progestins# 6strogens increase the
5nR7 pulse fre&uency, and progestins decrease it#
The endocrine cells of the anterior loe of the pituitary gland respond as if each group of
endocrine cells is monitoring different fre&uencies# +s a result, each group of cells is
sensitive to some 5nR7 pulse fre&uencies and insensitive to others# (or e"ample, consider
the gonadotropes9 the cells responsile for (S7 and L7 production# +t one pulse fre&uency,
the gonadotropes respond preferentially and secrete (S7, whereas at another fre&uency, L7
is the primary hormone released# (S7 and L7 production also occurs in pulses that follow
the rhythm of 5nR7 pulses# %f 5nR7 is asent or is supplied at a constant rate (without
pulses), (S7 and L7 secretion will stop in a matter of hours#
7ormones and the (ollicular )hase
(ollicular development egins under (S7 stimulationC each month some of the primordial
follicles egin to develop into primary follicles# +s the follicles enlarge, thecal cells start
producing androstenedione, a steroid hormone that is a 'ey intermediate in the synthesis of
estrogens and androgens# +ndrostenedione is asored y the granulosa cells and converted
to estrogens# %n addition, small &uantities of estrogens are secreted y interstitial cells
scattered throughout the ovarian stroma# Circulating estrogens are ound primarily to
alumins, with lesser amounts carried y gonadal steroidAinding gloulin (5>5)#
232459324.doc 46
9f the three estrogens circulating in the loodstream@estradiol, estrone, and estriol@the
one that is most aundant and has the most pronounced effects on target tissues is estradiol
(esAtraA$ Aol)# %t is the dominant hormone prior to ovulation# %n estraA
%Bdiol synthesis ((igure ./G.4?), androstenedione is first converted to testosterone, which
the enzyme aromatase converts to estradiol# The synthesis of oth estrone and estriol
proceeds directly from androstenedione#
6strogens have multiple functions that affect the activities of many tissues and organs
throughout the ody# +mong the important general functions of estrogens are (0)
stimulating one and muscle growth, (.) maintaining female secondary se" characteristics,
such as ody hair distriution and the location of adipose tissue deposits, (2) affecting
central nervous system (CFS) activity (especially in the hypothalamus, where estrogens
increase the se"ual drive), (4) maintaining functional accessory reproductive glands and
organs, and (8) initiating the repair and growth of the endometrium# (igure ./G.8?, which
diagrams the hormonal regulation of ovarian activity, includes an overview of the effects of
estrogens on various aspects of reproductive function#
SummaryB .ormonal Re"ulation of the emale Reproductive Cycle
(igure ./G.6? shows the changes in circulating hormone levels that accompany the ovarian
cycle# 6arly in the follicular phase, estrogen levels are low and the 5nR7 pulse fre&uency is
06G.4 per day (one pulse every 61G31 minutes) ((igure ./G.6a?)# +t this fre&uency, (S7 is
the dominant hormone released y the anterior pituitary glandC the estrogens released y
developing follicles inhiit L7 secretion ((igure ./G.6?)# +s secondary follicles develop,
(S7 levels decline due to the negative feedac' effects of inhiin# (ollicular development
and maturation continue, however, supported y the comination of estrogens, (S7, and
+s one or more tertiary follicles egin forming, the concentration of circulating estrogens
rises steeply# +s a result, the 5nR7 pulse fre&uency increases to aout 26 per day (one
pulse every 21G61 minutes)# The increased pulse fre&uency stimulates L7 secretion# %n
addition, at roughly day 01 of the cycle, the effect of estrogen on L7 secretion changes from
inhiition to stimulation# The switchover occurs only after rising estrogen levels have
e"ceeded a specific threshold value for aout 26 hours# (The threshold value and the time
re&uired vary among individuals#) 7igh estrogen levels also increase gonadotrope sensitivity
232459324.doc 47
to 5nR7# +t aout day 04, the estrogen level has pea'ed, the gonadotropes are at ma"imum
sensitivity, and the 5nR7 pulses are arriving aout every 21 minutes# The result is a massive
release of L7 from the anterior pituitary gland# This sudden surge in L7 concentration
triggers (0) the completion of meiosis % y the primary oocyte, (.) the rupture of the
follicular wall, and (2) ovulation# Typically, ovulation occurs 24G2/ hours after the L7
surge egins, roughly 3 hours after the L7 pea'#
7ormones and the Luteal )hase
The high L7 levels that trigger ovulation also promote progesterone secretion and the
formation of the corpus luteum# +s progesterone levels rise and estrogen levels fall, the
5nR7 pulse fre&uency declines sharply, soon reaching 0G4 pulses per day# This fre&uency of
5nR7 pulses stimulates L7 secretion more than it does (S7 secretion, and the L7
maintains the structure and secretory function of the corpus luteum#
+lthough moderate amounts of estrogens are secreted y the corpus luteum, progesterone is
the main hormone of the luteal phase# %ts primary function is to continue the preparation of
the uterus for pregnancy y enhancing the lood supply to the functional zone and
stimulating the secretion of endometrial glands# )rogesterone levels remain high for the ne"t
wee', ut unless pregnancy occurs, the corpus luteum egins to degenerate# Roughly 0.
days after ovulation, the corpus luteum ecomes nonfunctional, and progesterone and
estrogen levels fall mar'edly# The lood supply to the functional zone is restricted, and the
endometrial tissues egin to deteriorate# +s progesterone and estrogen levels drop, the
5nR7 pulse fre&uency increases, stimulating (S7 secretion y the anterior loe of the
pituitary gland, and the ovarian cycle egins again#
The hormonal changes involved with the ovarian cycle in turn affect the activities of other
reproductive tissues and organs# +t the uterus, the hormonal changes maintain the uterine
7ormones and the :terine Cycle
(igure ./G.6e? depicts the changes in the endometrium during a single uterine cycle# The
declines in progesterone and estrogen levels that accompany the degeneration of the corpus
luteum (see (igure ./G.6c,d?) result in menses# The shedding of endometrial tissue
continues for several days, until rising estrogen levels stimulate the repair and regeneration
of the functional zone of the endometrium# The proliferative phase continues until rising
232459324.doc 48
progesterone levels mar' the arrival of the secretory phase# The comination of estrogen and
progesterone then causes the enlargement of the endometrial glands and an increase in their
secretory activities#
7ormones and >ody Temperature
The monthly hormonal fluctuations cause physiological changes that affect core ody
temperature# $uring the follicular phase, when estrogen is the dominant hormone, the basal
body temperature, or the resting ody temperature measured upon awa'ening in the
morning, is aout 1#2JC lower than it is during the luteal phase, when progesterone
dominates ((igure ./G.6d?)# +t the time of ovulation, the asal ody temperature declines
noticealy, ma'ing the rise in temperature over the ne"t day even more noticeale ((igure
./G.6f?)# +s a result, y 'eeping records of ody temperature over a few uterine cycles, a
woman can often determine the precise day of ovulation# This information can e important
for individuals who wish to avoid or promote a pregnancy, ecause fertilization typically
occurs within a day of ovulation# Thereafter, oocyte viaility and the li'elihood of successful
fertilization decrease mar'edly# +,= ),S )remenstrual Syndrome
-:: Keys ' Cyclic changes in (S7 and L7 levels are responsile for the maintenance of the
ovarian cycleC the hormones produced y the ovaries in turn regulate the uterine cycle#
%nade&uate hormone levels, inappropriate or inade&uate responses to circulating hormones,
or poor coordination and timing of hormone production or secondary oocyte release will
reduce or eliminate the chances of pregnancy#
Concept Chec=
Khat changes would you e"pect to oserve in the ovarian cycle if the L7 surge did not
Khat effect would a loc'age of progesterone receptors in the uterus have on the
Khat event in the uterine cycle occurs when the levels of estrogens and progesterone
$ns%ers 6e"in on p3 $,-
The !hysiolo"y of Se#ual Intercourse
? $iscuss the physiology of se"ual intercourse as it affects the reproductive systems of males
232459324.doc 49
and females#
Se#ual intercourse, also 'nown as coitus (H9AiAtus) or copulation, introduces semen into
the female reproductive tract# Ke will
now consider the process as it affects the reproductive systems of males and females# +,=
>irth Control Strategies
Male Se#ual unction
Se"ual function in males is coordinated y comple" neural refle"es that we do not yet
understand completely# The refle" pathways utilize the sympathetic and parasympathetic
divisions of the autonomic nervous system# $uring se"ual arousal, erotic thoughts, the
stimulation of sensory nerves in the genital region, or oth lead to an increase in
parasympathetic outflow over the pelvic nerves# This outflow in turn leads to erection of the
penis (discussed on p# 0144)# The integument covering the glans of the penis contains
numerous sensory receptors, and erection tenses the s'in and increases their sensitivity#
Suse&uent stimulation can initiate the secretion of the ulourethral glands, luricating the
penile urethra and the surface of the glans#
$uring intercourse, the sensory receptors of the penis are rhythmically stimulated# This
stimulation eventually results in the coordinated processes of emission and e!aculation#
Emission occurs under sympathetic stimulation# The process egins when the peristaltic
contractions of the ampulla push fluid and spermatozoa into the prostatic urethra# The
seminal vesicles then egin contracting, and the contractions increase in force and duration
over the ne"t few seconds# )eristaltic contractions also appear in the walls of the prostate
gland# The comination moves the seminal mi"ture into the memranous and penile portions
of the urethra# Khile the contractions are proceeding, sympathetic commands also cause the
contraction of the urinary ladder and the internal urethral sphincter# The comination of
elevated pressure inside the ladder and the contraction of the sphincter effectively prevents
the passage of semen into the ladder#
E>aculation occurs as powerful, rhythmic contractions appear in the ischiocavernosus and
bulbospongiosus muscles, two superficial s'eletal muscles of the pelvic floor# The
ischiocavernosus muscles insert along the sides of the penisC their contractions serve
primarily to stiffen that organ# The ulospongiosus muscle wraps around the ase of the
232459324.doc 50
penisC the contraction of this muscle pushes semen toward the e"ternal urethral opening# The
contractions of oth muscles are controlled y somatic motor neurons in the inferior lumar
and superior sacral segments of the spinal cord# (The positions of these muscles are shown
in (igure 00G0.?, p# 24/#)
6!aculation is associated with intensely pleasurale sensations, an e"perience 'nown as male
or"asm (9RAgazm)# Several other noteworthy physiological changes occur at this time,
including pronounced ut temporary increases in heart rate and lood pressure# +fter
e!aculation, lood egins to leave the erectile tissue, and the erection egins to suside# This
susidence, called detumescence (deAtuBA,6SAens), is mediated y the sympathetic
nervous system#
%n sum, arousal, erection, emission, and e!aculation are controlled y a comple" interplay
etween the sympathetic and parasympathetic divisions of the autonomic nervous system#
7igher centers, including the cereral corte", can facilitate or inhiit many of the important
refle"es, therey modifying the patterns of se"ual function# +ny physical or psychological
factor that affects a single component of the system can result in male se"ual dysfunction,
also called impotence#
%mpotence is defined as an inaility to achieve or maintain an erection# ;arious physical
causes may e responsile for impotence, ecause erection involves vascular changes as well
as neural commands# (or e"ample, low lood pressure in the arteries supplying the penis,
due to a circulatory loc'age such as a pla&ue, will reduce the aility to attain an erection#
$rugs, alcohol, trauma, or illnesses that affect the autonomic nervous system or the central
nervous system can have the same effect# >ut male se"ual performance can also e strongly
affected y the psychological state of the individual# Temporary periods of impotence are
relatively common in healthy individuals who are e"periencing severe stresses or emotional
prolems# $epression, an"iety, and fear of impotence are e"amples of emotional factors that
can result in se"ual dysfunction# The prescription drug ;iagra, which enhances and prolongs
the effects of nitric o"ide on the erectile tissue of the penis, has proven useful in treating
many cases of impotence#
emale Se#ual unction
The phases of female se"ual function are comparale to those of male se"ual function#
$uring se"ual arousal, parasympathetic activation leads to engorgement of the erectile
232459324.doc 51
tissues of the clitoris and increased secretion of cervical mucous glands and the greater
vestiular glands# Clitoral erection increases the receptorsN sensitivity to stimulation, and the
cervical and vestiular glands luricate the vaginal walls# + networ' of lood vessels in the
vaginal walls ecomes filled with lood at this time, and the vaginal surfaces are also
moistened y fluid that moves across the epithelium from underlying connective tissues#
(This process accelerates during intercourse as the result of mechanical stimulation#)
)arasympathetic stimulation also causes contraction of sucutaneous smooth muscle of the
nipples, ma'ing them more sensitive to touch and pressure#
$uring se"ual intercourse, rhythmic contact of the penis with the clitoris and vaginal walls@
reinforced y touch sensations from the reasts and other stimuli (visual, olfactory, and
auditory)@provides stimulation that leads to orgasm# (emale orgasm is accompanied y
peristaltic contractions of the uterine and vaginal walls and, y means of impulses traveling
over the pudendal nerves, rhythmic contractions of the ulospongiosus and
ischiocavernosus muscles# The latter contractions give rise to the intensely pleasurale
sensations of orgasm#
Se"ual activity carries with it the ris' of infection with a variety of microorganisms# The
conse&uences of such an infection may range from merely inconvenient to potentially lethal#
Se#ually transmitted diseases ?ST0s@ are transferred from individual to individual,
primarily or e"clusively y se"ual intercourse# +t least two dozen acterial, viral, and fungal
infections are currently recognized as ST$s# The acterium (hlamydia can cause pelvic
inflammatory disease ?!I0@ and infertilityC +%$S, caused y a virus, is deadly# The
incidence of ST$s has een increasing in the :nited States since 03/4C an estimated 08
million new cases are diagnosed each year# )overty, intravenous drug use, prostitution, and
the appearance of drugAresistant pathogens all contriute to the prolem# The Applications
Manual contains a detailed discussion of the most common forms of ST$, including
gonorrhea, syphilis, herpes, genital +arts, and chancroid# +,= Se"ually Transmitted
$"in" and the Reproductive System
? $escrie the changes in the reproductive system that occur with aging#
The aging process affects all ody systems, including the reproductive systems of men and
232459324.doc 52
women ali'e# +s noted earlier in the chapter, these systems ecome fully functional at
puerty# Thereafter, the most stri'ing ageArelated changes in the female reproductive system
occur at menopause# Comparale ageArelated changes in the male reproductive system occur
more gradually and over a longer period of time#
,enopause is usually defined as the time that ovulation and menstruation cease# ,enopause
typically occurs at age 48G88, ut in the years immediately preceding it, the ovarian and
uterine cycles ecome irregular# This interval is called perimenopause# + shortage of
primordial follicles is the underlying cause of the irregular cycles# %t has een estimated that
almost < million potential oocytes are in fetal ovaries after five months of development, ut
the numer drops to aout . million at irth, and to a few hundred thousand at puerty#
Kith the arrival of perimenopause, the numer of follicles responding each month egins to
drop mar'edly# +s the numer of availale follicles decreases, estrogen levels decline and
may not rise enough to trigger ovulation# >y age 81, there are often no primordial follicles
left to respond to (S7# %n premature menopause, this depletion occurs efore age
,enopause is accompanied y a decline in circulating concentrations of estrogens and
progesterone, and a sharp and sustained rise in the production of 5nR7, (S7, and L7# The
decline in estrogen levels leads to reductions in the size of the uterus and reasts,
accompanied y a thinning of the urethral and vaginal epithelia# The reduced estrogen
concentrations have also een lin'ed to the development of osteoporosis, presumaly
ecause one deposition proceeds at a slower rate# + variety of neural effects are reported
as well, including Ohot flashes,P an"iety, and depression# 7ot flashes typically egin while
estrogen levels are declining, and cease when estrogen levels reach minimal values# These
intervals of elevated ody temperature are associated with surges in L7 production# The
hormonal mechanisms involved in other CFS effects of menopause are poorly understood#
%n addition, the ris's of atherosclerosis and other forms of cardiovascular disease increase
after menopause#
The ma!ority of women e"perience only mild symptoms, ut some individuals e"perience
acutely unpleasant symptoms in perimenopause or during or after menopause# (or most of
those women, hormone replacement therapy (7RT) involving a comination of estrogens
232459324.doc 53
and progestins can control the unpleasant neural and vascular changes associated with
menopause# The hormones are administered as pills, y in!ection, or y transdermal
Oestrogen patches#P 7owever, recent studies suggest that ta'ing estrogenAreplacement
therapy for more than 8 years increases the ris' of heart disease, reast cancer, +lzheimers
disease, lood clots, and stro'eC 7RT should e prescried with caution, only after a full
discussion and assessment of the potential ris's and enefits, and ta'en for as short a time as
The Male Climacteric
Changes in the male reproductive system occur more gradually than do those in the female
reproductive system# The period of declining reproductive function, which corresponds to
perimenopause in women, is 'nown as the male climacteric or andropause# Levels of
circulating testosterone egin to decline etween the ages of 81 and 61, and levels of
circulating (S7 and L7 increase# +lthough sperm production continues (men well into their
eighties can father children), older men e"perience a gradual reduction in se"ual activity#
This decrease may e lin'ed to declining testosterone levels# Some clinicians suggest the use
of testosterone replacement therapy to enhance the liido (se"ual drive) of elderly men, ut
this may increase the ris' of prostate disease#
-:: Keys ' Se" hormones have widespread effects on the ody# They affect rain
development and ehavioral drives, mus
cle mass, one mass and density, ody proportions, and the patterns of hair and ody fat
distriution# +s aging occurs, re
ductions in se" hormone levels affect appearance, strength, and a variety of physiological
Concept Chec=
+n inaility to contract the ischiocavernosus and ulospongiosus muscles would
interfere with which part(s) of the male se" actT
Khat changes occur in females during se"ual arousal as the result of increased
parasympathetic stimulationT
Khy does the level of (S7 rise and remain high during menopauseT
$ns%ers 6e"in on p3 $,-
Inte"ration %ith &ther Systems
232459324.doc 54
Formal human reproduction is a comple" process that re&uires the participation of multiple
systems# The hormones discussed in this chapter play a ma!or role in coordinating
reproductive events (Tale ./G0)# )hysical factors also play a role# The manNs sperm count
must e ade&uate, the semen must have the correct p7 and nutrients, and erection and
e!aculation must occur in the proper se&uenceC the womanNs ovarian and uterine cycles must
e properly coordinated, ovulation and oocyte transport must occur normally, and her
reproductive tract must provide a hospitale environment for the survival and movement of
sperm, and for the suse&uent fertilization of the oocyte# (or these steps to occur, the
reproductive, digestive, endocrine, nervous, cardiovascular, and urinary systems must all e
functioning normally#
6ven when all else is normal and fertilization occurs at the proper time and place, a healthy
infant will not e produced unless the zygote@a single cell the size of a pinhead@manages
to develop into a fullAterm fetus that typically weighs aout 2 'g (6#6 l)# %n Chapter .3 we
will consider the process of development, focusing on the mechanisms that determine oth
the structure of the ody and the distinctive characteristics of each individual#
6ven though the reproductive systemNs primary function@producing children@doesnNt play
a role in maintaining homeostasis, reproduction depends on a variety of physical,
physiological, and psychological factors, many of which re&uire intersystem cooperation# %n
addition, the hormones that control and coordinate se"ual function have direct effects on the
organs and tissues of other systems# (or e"ample, testosterone and estradiol affect oth
muscular development and one density# (igure ./G.<? summarizes the relationships
etween the reproductive system and other physiological systems#
Clinical !atterns
The male and female reproductive systems are comple", and reproductive disorders are
many and varied# ,a!or categories of reproductive disorders include the following=
# ? Tumors, such as testicular, prostate, ovarian, or uterine cancers#
# ? %nflammation and infection, such as prostatitis, pelvic inflammatory disease,
to"ic shoc' syndrome, and the various se"ually transmitted diseases#
# ? :terine disorders such as endometriosis, and hormonally related prolems
such as amenorrhea#
# ? Trauma, such as testicular torsion and inguinal hernias#
232459324.doc 55
# ? Congenital disorders, such as cryptorchidism#
,ost reproductive disorders are primary disorders that reflect prolems originating within
the reproductive system# 7owever, amenorrhea and premenstrual syndrome are e"amples of
secondary disorders that can result from prolems with the endocrine system, and impotence
can result from neural, hormonal, or vascular prolems# The Applications Manual discusses
the diagnosis and treatment of the ma!or classes of reproductive system disorders#
Chapter Revie%
Selected Clinical Terminolo"y
amenorrheaB The failure of menarche to appear efore age 06, or a cessation of
menstruation for si" months or more in an adult female of reproductive age# (p# 018<)
6reast cancerB + malignant, metastasizing tumor of the mammary gland that is the primary
cause of death for women ages 28G48#
# (p# 0161) cervical cancerB + malignant, metastasizing tumor of the cervi", and the
most common reproductive system cancer in women# (p# 0186) cryptorchidismB The failure
of one or oth testes to descend into the scrotum y the time of irth# (p# 0120)
dysmenorrheaB )ainful menstruation# (p# 018<) endometriosisB The growth of endometrial
tissue outside the uterus# *+,- fi6rocystic diseaseB Clusters of loular cysts within the
tissues of the mammary gland# (p# 0161) "onorrheaB + se"ually transmitted acterial
disease# *+,- impotenceB The inaility to achieve or maintain an erection# (p# 0168)
mammo"raphyB The use of "Arays to e"amine reast tissue# (p# 0161) mastectomyB The
surgical removal of part or all of a reast containing cancerous glandular tissue# (p# 0121)
orchiectomyB The surgical removal of a testis# (p# 0120) ovarian cancerB + malignant,
metastasizing tumor of the ovaries, and the most dangerous reproductive system cancer in
women# *+,- pelvic inflammatory disease ?!I0@B +n infection of the uterine tues# (p#
0166) prostate cancerB + malignant, metastasizing tumor of the prostate gland, and the
second most common cause of cancer deaths in males#
# (p# 014<) prostatectomyB The surgical removal of the prostate gland# (p# 014<)
prostateAspecific anti"en ?!S$@B +n antigen whose level in lood increases in men with
prostate cancer# (p# 014<) se#ually transmitted diseases ?ST0s@B $iseases transferred from
one individual to another primarily or e"clusively through se"ual
232459324.doc 56
contact# 6"amples include gonorrhea, syphilis, herpes genitalis, and +%$S# (p# 0166 and
*+,-) va"initisB +n infection of the vaginal canal y fungal or acterial pathogens# (p# 018/
and *+,-) vasectomyB The surgical removal of a segment of the ductus deferens, ma'ing it
impossile for spermatozoa to reach the distal portions
of the male reproductive tract# *+,-
Study &utline
Introduction to the Reproductive System p# 0121
-3 -3 The human reproductive system produces, stores, nourishes, and transports
functional "ametes (reproductive cells)# ertili5ation is the fusion of male and female
23 23 The reproductive system includes "onads ?testes or ovaries@, ducts,
accessory glands and organs, and the e#ternal "enitalia#
83 83 %n males, the testes produce spermato5oa, which are e"pelled from the ody
in semen during ejaculation# The ovaries of a se"ually mature female produce oocytes
(immature ova) that travel along uterine tubes toward the uterus# The vagina connects the
uterus with the e"terior of the ody#
The Reproductive System of the Male p# 0121
-3 The spermatozoa travel along the epididymis, the ductus deferens, the ejaculatory
duct, and the urethra efore leaving the ody# +ccessory organs (notaly the seminal
vesicles, prostate gland, and bulbourethral glands) secrete fluids into the e!aculatory ducts
and the urethra# The scrotum encloses the testes, and the penis is an erectile organ# $'igure
The Testes p# 0121
-3 23 The descent of the testes through the inguinal canals occurs during fetal
development# The testes remain connected to internal structures via the spermatic cords#
The raphe mar's the oundary etween the two chamers in the scrotum# $'igures 0:!09
23 83 The dartos muscle tightens the scrotum, giving it a wrin'led appearance as it
elevates the testesC the cremaster muscles are more sustantial muscle that pull the testes
232459324.doc 57
close to the ody#
83 ;3 The tunica al6u"inea surrounds each testis# Septa e"tend from the tunica
aluginea to the region of the testis closest to the entrance to the epididymis, creating a
series of lo6ules# $'igure 0:!4%
;3 <3 Seminiferous tu6ules within each loule are the sites of sperm production#
(rom there, spermatozoa pass through the rete testis# Efferent ductules connect the rete
testis to the epididymis# >etween the seminiferous tuules are interstitial cells, which
secrete se" hormones# $'igures 0:!49 0:!6%
Spermatogenesis p# 0126
43 Seminiferous tuules contain spermato"onia, stem cells involved in
spermato"enesis (the production of spermatozoa), and sustentacular cells, which sustain
and promote the development of spermatozoa# $'igures 0:!79 0:!;%
-:: Keys ' p# 012/
The +natomy of a Spermatozoon p# 012/
73 6ach spermato5oon has a head tipped y an acrosomal cap, a middle piece, and a
tail# $'igure 0:!:%
-:: Keys ' p# 0123
The ,ale Reproductive Tract p# 0141
-3 83 (rom the testis, the spermatozoa enter the epididymis, an elongate tuule
with head, 6ody, and tail regions# The epididymis monitors and ad!usts the composition of
the fluid in the seminiferous tuules, serves as a recycling center for damaged spermatozoa,
stores and protects spermatozoa, and facilitates their functional maturation# $'igure 0:!<%
23 93 The ductus deferens, or vas deferens, egins at the epididymis and passes
through the inguinal canal as part of the spermatic cord# Fear the prostate gland, the ductus
deferens enlarges to form the ampulla# The !unction of the ase of the seminal vesicle and
the ampulla creates the e>aculatory duct, which empties into the urethra# $'igures 0:!<9
83 -:3 The urethra e"tends from the urinary ladder to the tip of the penis# The urethra
can e divided into prostatic9 membranous, and spongy regions#
232459324.doc 58
The +ccessory 5lands p# 0140
--3 6ach seminal vesicle is an active secretory gland that contriutes aout 61 percent of
the volume of semenC its secretions contain fructose (which is easily metaolized y
spermatozoa), icaronate ions, prostaglandins, and firinogen# The prostate "land secretes
slightly acidic prostatic fluid# +l'aline mucus secreted y the 6ul6ourethral "lands has
luricating properties# $'igures 0:!/=9 0:!//%
Semen p# 0142
-23 + typical e!aculation releases .G8 ml of semen (e>aculate), which contains .1G011
million spermatozoa per milliliter# The fluid component of semen is seminal fluid#
The 6"ternal 5enitalia p# 0144
-83 The s'in overlying the penis resemles that of the scrotum# ,ost of the 6ody of the
penis consists of three masses of erectile tissue# >eneath the superficial fascia are two
corpora cavernosa and a single corpus spon"iosum, which surrounds the urethra# $ilation
of the lood vessels within the erectile tissue produces an erection# $'igure 0:!//%
+natomy 261 ' Reproductive SystemU,ale Reproductive System
7ormones and ,ale Reproductive (unction p# 0148
-;3 %mportant regulatory hormones include S. (follicle"stimulating hormone), +.
(luteinizing hormone), and 2nR. (gonadotropinreleasing hormone)# Testosterone is the
most important androgen# $'igure 0:!/0%
The Reproductive System of the emale p# 014/
-3 -3 )rincipal organs of the female reproductive system include the ovaries,
uterine tubes, uterus, vagina, and e"ternal genitalia# $'igure 0:!/2%
23 23 The ovaries, uterine tues, and uterus are enclosed within the 6road
li"ament# The mesovarium supports and stailizes each ovary#
$'igure 0:!/4%
The 9varies p# 0143
-3 83 The ovaries are held in position y the ovarian li"ament and the suspensory
li"ament# ,a!or lood vessels enter the ovary at the ovarian hilum# 6ach ovary is covered
y a tunica al6u"inea# $'igure 0:!/4%
23 ;3 &o"enesis (ovum production) occurs monthly in ovarian follicles as part of
232459324.doc 59
the ovarian cycle, which is divided into a follicular (preovulatory) phase and a luteal
(postovulatory) phase# $'igures 0:!/69 0:!/7%
83 <3 +s development proceeds, primordial, primary, secondary, and tertiary
follicles are produced in turn# +t ovulation, a secondary oocyte and the attached follicular
cells of the corona radiata are released through the ruptured ovarian wall# The follicular
cells remaining within the ovary form the corpus luteum, which later degenerates into scar
tissue called a corpus al6icans# $'igure 0:!/7%
-:: Keys ' p# 018.
The :terine Tues p# 018.
43 6ach uterine tu6e has an infundi6ulum with fim6riae (fingerli'e pro!ections), an
ampulla, and an isthmus# 6ach uterine tue opens into the uterine cavity# (or fertilization
to occur, a secondary oocyte must encounter spermatozoa during the first 0.G.4 hours of its
passage from the infundiulum to the uterus# $'igure 0:!/;%
The :terus p# 0182
-3 73 The uterus provides mechanical protection, nutritional support, and waste
removal for the developing emryo# Formally, the uterus ends anteriorly near its ase
(anteflexion)# %t is stailized y the road ligament, uterosacral li"aments, round
li"aments, and lateral li"aments# $'igure 0:!/:%
23 83 ,a!or anatomical landmar's of the uterus include the 6ody, isthmus, cervi#,
cervical os (external orifice), uterine cavity, cervical canal, and internal os (internal
orifice)# The uterine wall consists of an inner endometrium, a muscular myometrium, and
a superficial perimetrium (an incomplete serous layer)# $'igures 0:!/:9 0:!/<%
83 93 + typical ./Aday uterine, or menstrual, cycle egins with the onset of
menses and the destruction of the functional 5one of the endometrium# This process of
menstruation continues from one to seven days# $'igure 0:!0=%
;3 -:3 +fter menses, the proliferative phase egins, and the functional zone undergoes
repair and thic'ens# The proliferative phase is followed y the secretory phase, during
which endometrial glands enlarge# ,enstrual activity egins at menarche and continues
until menopause# $'igure 0:!0=%
232459324.doc 60
The ;agina p# 018<
--3 The va"ina is a muscular tue e"tending etween the uterus and the e"ternal genitalia# +
thin epithelial fold, the hymen, partially loc's the entrance to the vagina until physical
distortion (often associated with se"ual intercourse) ruptures the memrane# $'igure 0:!0/%
The 6"ternal 5enitalia p# 018/
-23 The components of the vulva are the vesti6ule, la6ia minora, paraurethral "lands,
clitoris, la6ia ma>ora, and lesser and "reater vesti6ular "lands# $'igure 0:!00%
The ,ammary 5lands p# 0183
-83 + neworn infant can gain nourishment from mil' secreted y maternal mammary
"lands# $'igure 0:!02%
+natomy 261 ' Reproductive SystemU(emale Reproductive System
7ormones and the (emale Reproductive Cycle p# 0160
-3 -;3 7ormonal regulation of the female reproductive cycle involves the coordination
of the ovarian and uterine cycles#
23 -<3 Estradiol, the most important estrogen, is the dominant hormone of the
follicular phase# 9vulation occurs in response to a midcycle surge in L7# $'igures 0:!049
83 -43 The hypothalamic secretion of 5nR7 occurs in pulses that trigger the pituitary
secretion of (S7 and L7# (S7 initiates follicular development, and activated follicles and
ovarian interstitial cells produce estrogens# 7igh estrogen levels stimulate L7 secretion,
increase pituitary sensitivity to 5nR7, and increase the 5nR7 pulse fre&uency#
!ro"esterone, one of the pro"estins, is the principal hormone of the luteal phase# Changes
in estrogen and progesterone levels are responsile for the maintenance of the uterine cycle#
$'igures 0:!069 0:!07%
Summary= 7ormonal Regulation of the (emale Reproductive Cycle p# 016.
-:: Keys ' p# 0168
The !hysiolo"y of Se#ual Intercourse p# 0168 ,ale Se"ual (unction p# 0168
-3 $uring se"ual arousal in males, erotic thoughts, sensory stimulation, or oth lead to
parasympathetic activity that produces erection# Stimuli accompanying se#ual intercourse
lead to emission and e>aculation# Contractions of the ulospongiosus muscles are
232459324.doc 61
associated with or"asm#
(emale Se"ual (unction p# 0166
23 The phases of female se"ual function resemle those of male se"ual function, with
parasympathetic arousal and s'eletal muscle contractions associated with orgasm#
$"in" and the Reproductive System p# 0166 ,enopause p# 0166
-3 -3 ,enopause (the time that ovulation and menstruation cease) typically occurs
at ages 48G88# The production of 5nR7, (S7, and L7 rise, whereas circulating
concentrations of estrogen and progesterone decline#
23 23 $uring the male climacteric, at ages 81G61, circulating testosterone levels
fall, and levels of (S7 and L7 rise#
The ,ale Climacteric p# 016<
-:: Keys ' p# 016<
Inte"ration %ith &ther Systems
$'igure 0:!0;% p# 016<
Revie% Duestions
My$E! ' +ccess more review material online at My$E!3 There youNll find learning
activities, case studies, &uizzes, %nteractive )hysiology e"ercises, and more to help you
succeed# To access the site, go to www#myaandp#com#
Ans+ers to the )evie+ >uestions begin on page A!/.
L6;6L 0 Reviewing (acts and Terms
3 -3 $eveloping spermatozoa are nourished y the
3 ?a@ interstitial cells ?6@ seminal vesicles
3 ?c@ sustentacular cells ?d@ Leydig cells
3 ?e@ epididymis
3 23 The ovaries are responsile for
3 ?a@ the production of female gametes
3 ?6@ the secretion of female se" hormones
3 ?c@ the secretion of inhiin
3 ?d@ a, , and c are correct
3 83 %n females, meiosis is not completed until
232459324.doc 62
3 ?a@ irth
3 ?6@ puerty
3 ?c@ fertilization occurs
3 ?d@ uterine implantation occurs
3 ;3 + sudden surge in L7 secretion causes the
3 ?a@ onset of menses
3 ?6@ rupture of the follicular wall and ovulation
3 ?c@ eginning of the proliferative phase
3 ?d@ end of the uterine cycle
3 <3 The principal hormone of the postovulatory phase is
3 ?a@ progesterone ?6@ estradiol
3 ?c@ estrogen ?d@ luteinizing hormone
23 43 Trace the duct system that the sperm traverses from the site of its production
to the e"terior of the ody#
83 73 Khich accessory organs and glands contriute to the composition of semenT
Khat are the functions of eachT
;3 83 Khat are the primary cell populations in the testes that are responsile for
functions related to reproductive activityT Khat are the functions of these cellsT
<3 93 %dentify the three regions of the male urethra#
43 -:3 List the functions of testosterone in males#
73 --3 List and summarize the important steps in the ovarian cycle#
83 -23 $escrie the histological composition of the uterine wall#
93 -83 Khat is the role of the clitoris in the female reproductive systemT
-:3 -;3 Trace the route that mil' ta'es from its site of production to the outside of the
L6;6L . Reviewing Concepts
3 -<3 +ll of the following are true of pelvic inflammatory disease, except that it
3 ?a@ is fre&uently caused y se"ually transmitted pathogens
3 ?6@ causes fever and adominal pain
3 ?c@ can lead to a ruptured urinary ladder
232459324.doc 63
3 ?d@ can possily lead to peritonitis
3 ?e@ can cause sterility
3 -43 %n the follicular phase of the ovarian cycle, the ovary is
3 ?a@ undergoing atresia
3 ?6@ forming a corpus luteum
3 ?c@ releasing a mature egg
3 ?d@ secreting progesterone
3 ?e@ maturing a follicle
23 -73 Khat are the main differences in gamete production etween males and femalesT
83 -83 $escrie the erectile tissues of the penis# 7ow does erection occurT
;3 -93 :sing an average cycle of ./ days, descrie each of the three phases of the
uterine cycle#
<3 2:3 $escrie the hormonal events associated with the ovarian cycle#
43 2-3 $escrie the hormonal events associated with the uterine cycle#
73 223 Summarize the steps that occur in se"ual arousal and orgasm# $o these processes
differ in males and femalesT
83 283 7ow does the aging process affect the reproductive systems of men and womenT
L6;6L 2 Critical Thin'ing and Clinical +pplications
-3 2;3 $iane has peritonitis (an inflammation of the peritoneum), which her physician
says resulted from a urinary tract infection# Khy might this condition occur more readily in
females than in malesT
23 2<3 %n a condition 'nown as endometriosis, endometrial cells are elieved to migrate
from the ody of the uterus into the uterine tues or y way of the uterine tues into the
peritoneal cavity where they ecome estalished# + ma!or symptom of endometriosis is
periodic pain# Khy do you thin' this occursT
83 243 Contraceptive pills contain estradiol or estradiol and progesterone that are given
at programmed doses during the ovarian cycle to prevent follicle maturation and ovulation#
7ow would this happenT
;3 273 Komen odyuilders and women with eating disorders such as anore"ia nervosa
commonly e"perience amenorrhea# Khat does this fact suggest aout the relation etween
232459324.doc 64
ody fat and menstruationT Khat might e the enefit of amenorrhea under such
Clinical /ote
)rostatic 7ypertrophy and )rostate Cancer
6nlargement of the prostate gland, or 6eni"n prostatic hypertrophy, typically occurs
spontaneously in men over age 81# The increase in size occurs as testosterone production y
the interstitial cells decreases# (or un'nown reasons, small masses called prostatic
concretions may form within the glands (see photo)# +t the same time, the interstitial cells
egin releasing small &uantities of estrogens into the loodstream# The comination of lower
testosterone levels and the presence of estrogens proaly stimulates prostatic growth# %n
severe cases, prostatic swelling constricts and loc's the urethra and constricts the rectum#
%f not corrected, the urinary ostruction can cause permanent 'idney damage#0 )artial
surgical removal is the most effective treatment# %n the procedure 'nown as a T(R!
(transurethral prostatectomy), an instrument pushed along the urethra restores normal
function y cutting away the swollen prostatic tissue# ,ost of the prostate gland remains in
place, and no e"ternal scars result#
)rostate cancer, a malignancy of the prostate gland, is the second most common cancer and
the second most common cause of cancer deaths in males# %n .114, appro"imately .21,001
new cases of prostate cancer were diagnosed in the :nited States, and aout .3,311 deaths
resulted from the ailment# ,ost patients are elderly# (The average age at diagnosis is <.#)
(or reasons that are poorly understood, prostate cancer rates for +sianA+merican males are
relatively low compared with those of either CaucasianA+mericans or +fricanA+mericans#
(or all age and ethnic groups, the rates of prostate cancer are rising sharply# The reason for
the increase is not 'nown# +ggressive diagnosis and treatment of localized prostate cancer in
elderly patients is controversial ecause many of these men have nonAmetastatic tumors, and
even if untreated are more li'ely to die of some other disease#
)rostate cancer normally originates in one of the secretory glands# +s the cancer progresses,
it produces a nodular lump or swelling on the surface of the prostate gland# )alpation of this
gland through the rectal wall@a procedure 'nown as a digital rectal exam ($R6)@is the
easiest diagnostic screening procedure# Transrectal prostatic ultrasound (TR:S) can e
232459324.doc 65
used to otain more detailed information aout the status of the prostate, ut at significantly
higher cost to the patient# >lood tests are also used for screening purposes# The most
sensitive is a lood test for prostateAspecific antigen ()S+)# 6levated levels of this antigen,
normally present in low concentrations, may indicate the presence of prostate cancer# The
serum enzyme assay, which chec's the level of the isozyme prostatic acid phosphatase,
detects prostate cancer in later stages of development# Screening with periodic )S+ tests is
now eing recommended for men over age 81#
%f cancer is detected efore it has spread to other organs, and the patient is elderly or has
other serious health prolems, Owatchful waitingP is an option# %n other cases, the usual
treatment is localized radiation or surgical removal of the prostate gland# This operation, a
prostatectomy (prosAtaAT6HAtoAm eB), can e effective in controlling the condition, ut
oth surgery and radiation can have undesirale side effects, including urinary incontinence
and loss of se"ual function# ,odified treatment procedures along with medications such as
;iagra can reduce the ris's and maintain normal se"ual function in perhaps three out of four
The prognosis is much worse for prostate cancer diagnosed after metastasis has occurred,
ecause metastasis rapidly involves the lymphatic system, lungs, one marrow, liver, or
adrenal glands# Survival rates at this stage are relatively low# Treatments for metastasized
prostate cancer include widespread irradiation, hormonal manipulation, lymph node removal,
and aggressive chemotherapy# >ecause the cancer cells are stimulated y testosterone,
treatment may involve castration or administering hormones that depress 5nR7 or L7
production# There are three treatment options= (0) an estrogen, typically diethylstilbestrol
$*-S%C (.) drugs that mimic 3n)?9 which when given in high doses produce a surge in L7
production followed y a sharp decline to very low levels (presumaly as the endocrine cells
adapt to the e"cessive stimulation)C and (2) drugs that bloc. the binding of androgens to the
receptors on target cells (including the new drugs flutamide and bicalutamide), which
prevent the stimulation of cancer cells y testosterone# The death rate for prostate cancer
may e falling in some countries, perhaps from earlier detection and more effective
' S(MM$R) T$*+E 28,- ' .&RM&/ES & T.E RE!R&0(CTI1E
232459324.doc 66
.ormone Source Re"ulation of Secretion !rimary Effects
2onadotropinAreleasin" 7ypothalamus Males= inhiited y testosterone and
Stimulates (S7 secretion and L7
hormone ?2nR.@ possily y inhiin synthesis in males and females
'emales= 5nR7 pulse fre&uency
increased y estrogens, decreased
y progestins
ollicleAstimulatin" +nterior loe of pituitary Males= stimulated y 5nR7, inhiited Males=
stimulates spermatogenesis
hormone ?S.@ gland y inhiin and spermiogenesis through
'emales= stimulated y 5nR7, effects on sustentacular cells
inhiited y inhiin 'emales= stimulates follicle development, estrogen production, and
oocyte maturation
+uteini5in" +nterior loe of pituitary Males= stimulated y 5nR7 Males= stimulates
interstitial cells to
hormone ?+.@ gland 'emales= production stimulated y secrete testosterone
5nR7, secretion y the comination 'emales= stimulates ovulation,
of high 5nR7 pulse fre&uencies and formation of corpus luteum,
high estrogen levels and progestin secretion
$ndro"ens (primarily %nterstitial cells of testes Stimulated y L7 6stalish and maintain
testosterone and se" characteristics and se"ual
dihydrotestosterone) ehaviorC promote maturation
of spermatozoaC inhiit 5nR7 secretion
Estro"ens (primarily 5ranulosa and thecal cells Stimulated y (S7 Stimulate L7 secretion
(at high levels)C
estradiol) of developing folliclesC estalish and maintain secondary se"
232459324.doc 67
corpus luteum characteristics and se"ual ehaviorC stimulate repair and growth of
endometriumC increase fre&uency of 5nR7 pulses
!ro"estins (primarily 5ranulosa cells from
progesterone) midcycle through
functional life of corpus
Stimulated y L7 Stimulate endometrial growth and glandular secretionC reduce fre&uency
of 5nR7 pulses
Inhi6in Sustentacular cells of Stimulated y factors released y %nhiits secretion of (S7
(and possily of testes and granulosa cells developing spermatozoa (male) 5nR7) of ovaries
and developing follicles (female)
# ? I2(RE 28,- The ,ale Reproductive System# + sagittal section of the male
reproductive organs# +TL+S= )late 64
# ? I2(RE 28,2 The $escent of the Testes# ?a@ Sagittal sectional views of the
positional changes involved in the descent of the right testis# >ecause the size of the
guernaculum testis remains constant (see the scale ar at the left) while the rest of the fetus
grows, the relative position of the testis shifts# ?6@ (rontal views showing the descent of the
testes and the formation of the spermatic cords#
# ? I2(RE 28,8 The ,ale Reproductive System in +nterior ;iew
# ? I2(RE 28,; The Structure of the Testes# ?a@ + frontal section# ?6@ + section
through a testis#
# ? I2(RE 28,< The Seminiferous Tuules# ?a@ + section through a coiled
seminiferous tuule# ?6@ + cross section through a single tuule#
?c@ Stages in spermatogenesis in the wall of a seminiferous tuule# Sustentacular cells
surround the stem cells of the tuule and support the developing spermatocytes and
? I2(RE 28,4 Chromosomes in ,itosis and ,eiosis# ?a@ The fates of three representative
chromosomes during mitosis# (See 'igure 2!06, pp. <:!<<#) ?6@ The fates of three
232459324.doc 68
representative chromosomes during the two stages of meiosis#
# ? I2(RE 28,7 Spermatogenesis# The events depicted occur in the seminiferous
tuules# The fates of three representative chromosomes are shownC compare with 'igure
# ? I2(RE 28,8 Spermiogenesis and Spermatozoon Structure# ?a@ The
differentiation of a spermatid into a spermatozoon# This differentiation process is completed
in appro"imately five wee's# ?6@ 7uman spermatozoa#
# ? I2(RE 28,9 The 6pididymis# ?a@ + diagrammatic view# ?6@ 6pithelial features,
especially the elongate stereocilia characteristic of the epididymis# +TL+S= )late 61a
# ? I2(RE 28,-: The $uctus $eferens and +ccessory 5lands# ?a@ + posterior view
of the prostate gland, showing sudivisions of the ductus deferens in relation to surrounding
structures# ?6@ The ductus deferens, showing the smooth muscle around the lumen# *VR# 5#
Hessel and R# 7# Hardon, Tissues and 5rgans@ A Text"Atlas of Scanning -lectron
Microscopy, K# 7# (reeman W Co#, 03<3# +ll Right Reserved#- Sections of ?c@ the seminal
vesicle, ?d@ the prostate gland, and ?e@ a ulourethral gland#
? I2(RE 28,-- The )enis# ?a@ + frontal section through the penis and associated
organs# ?6@ + sectional view through the penis# ?c@ +n anterior and lateral view of the penis,
showing positions of the erectile tissues# +TL+S= )late 61
ovaries# ?6@ + sectional view of the ovary, uterine tue, and associated mesenteries#
+TL+S= )late 6<
# ? I2(RE 28,-< 9ogenesis# %n oogenesis, a single primary oocyte produces an
ovum and two or three nonfunctional polar odies# Compare this schematic diagram with
'igure 0:!;, p# 012<#
# ? I2(RE 28,-4 The 9varian Cycle
# ? I2(RE 28,-7 The :terine Tues# ?a@ Regions of the uterine tues# ?6@ +
sectional view of the isthmus# ?c@ + colorized S6, of the ciliated lining of the uterine tue#
# ? (igure ./G0/ The :terus# ?$@ + )9ST6R%9R ;%6K K%T7 T76 L6(T
)9RT%9F 9( T76 :T6R:S, L6(T :T6R%F6 T:>6, +F$ L6(T 9;+RM S79KF
%F S6CT%9F# ?*@ T76 L%5+,6FTS T7+T ST+>%L%D6 T76 )9S%T%9F 9( T76
:T6R:S %F T76 )6L;%C C+;%TM# +TL+S= )late 66C 6<
# ? I2(RE 28,-9 The :terine Kall# ?a@ + diagrammatic sectional view of the
232459324.doc 69
uterine wall, showing the endometrial regions and the circulatory supply to the
endometrium# ?6@ The asic histological structure of the endometrium#
# ? I2(RE 28,2: The +ppearance of the 6ndometrium during the :terine Cycle#
The appearance of the endometrium ?a@ at menses, ?6@ during the proliferative phase, and
?c@ during the secretory phase of the uterine cycle#
# ? I2(RE 28,2- The 7istology of the ;agina
# ? I2(RE 28,22 The (emale 6"ternal 5enitalia
# ? I2(RE 28,28 The ,ammary 5lands# ?a@ The mammary gland of the left reast#
?6@ +n inactive mammary gland of a nonpregnant woman#
? I2(RE 28,-2 7ormonal (eedac' and the Regulation of ,ale Reproductive
? I2(RE 28,-8 The (emale Reproductive System# + sagittal section of the female
reproductive organs# +TL+S= )late 68
? I2(RE 28,-; The 9varies and Their Relationships to the :terine Tue and :terus#
?a@ + posterior view of the uterus, uterine tues, and
?c@ +n active mammary gland of a nursing woman# +TL+S= )late ./
# ? I2(RE 28,2; )athways of Steroid 7ormone Synthesis in ,ales and (emales#
+ll gonadal steroids are derived from cholesterol# %n men, the pathway ends with the
synthesis of testosterone, which may suse&uently e converted to dihydrotestosterone# %n
women, an additional step past testosterone leads to estradiol synthesis# The synthesis of
progesterone and the estrogens other than estradiol involve alternative pathways#
# ? I2(RE 28,2<
The 7ormonal Regulation of 9varian +ctivity
# ? I2(RE 28,24 The 7ormonal Regulation of the (emale Reproductive Cycle
# ? I2(RE 28,27 (unctional Relationships etween the Reproductive System and
9ther Systems
232459324.doc 70