r

754 Chapter Twenty-Seven The Reproductive System and Early Development

HISTOLOGY
Female Reproductive System fertilized by a sperm. Components of the female reproductive systen
include the ovaries, uterine tubes, uterns, vagina, and mammar:
The female reproductive system has two important functions: pro- glands. Exercises 27. l to 27.4 explore the anatomic details of th
duce the female gametes (ova) and support, protect, and nourish ovary, uterine tube, uterus, and vagina. Mammary glands are covere1
the developing embryo/fetus that is fanned when ao ovum becomes in Exercise 27.12.

EXERCISE 27 .1

HISTOLOGY OF THE OVARY (gennen, a sprout) is a simple cuboidal epithelium that

forms the outermost covering of the ovary. The name
The ovary is the primary reproductive organ in the female and germinal refers to the fact that scientists once thought
functions in the production of the ova (eggs) and the female sex that the germ cells (oocytes) were formed from this
steroid hormones estrogen and progesterone. The ovaries con- epithelial layer. Most ovarian cancers arise in the germinal
tain several ovarian follicles, all at various stages of develop- epithelium. Deep to the germinal epithelium is the tunica
ment. Each follicle contains a developing oocyte (table 27.1). albuginea (tunica, a coat, + albugineus, white spot), which
The structure and function of the ovary is best viewed at the is composed of dense irregular connective tissue.
histological level because characteristics of ovarian follicles are
evident at each stage of development. 4. Focus on the cortex and identify follicles in each stage of
follicular development listed in table 27.2.
1. Obtain a histology slide of an ovary and place it on the
microscope stage. 5. Identify the following structures on the slide of the ovary,
using tables 27.1 and 27.2 and figures 27.1 and 27.2 as
2. Bring the tissue sample into focus on low power and guides:
identify the outer cortex and inner medulla of the ovary
(figure 27.1). Next, move the microscope stage so the □ corpus albicans □ secondary follicle
ovarian cortex is at the center of the field of view and
□ corpus luteum □ secondary oocyte
then change to high power.
□ germinal epithelium □ tunica albuginea
3. Observe the outermost region of the ovarian cortex.
Locate the two layers of tissue that compose the outer
□ primary follicle □ vesicular (Graafian)
follicle
coverings of the ovary: the gemlinal epithelium and the □ primary oocyte
tunica albuginea (figure 27.2). The germinal epithelium □ primordial follicle

Table 27.1 Developmental Stages of an Oocyte

Cell Name/ Description and Function Stage of Mitosis/ Ploidy Word Origin
Oocyte Stage Meiosis
Oogonia Primitive germ cells that undergo mitosis in the second to fifth Formed by mitosis; Diploid (2n) oon, egg,+ gonia,
months of embryonic life form oogonia that will subsequently undergo mitosis generation
develop into primary oocytes. Approximately 70% of the to form primary
primary oocytes will degenerate in a process called atresia. oocytes
Primary Oocyte Oogonia has replicated DNA and has begun the process of Arrested in Diploid (2n) primary, first,
meiosis and arrest in prophase of meiosis I. Primary oocytes Prophase I of + oon, egg,+
will not undergo subsequent meiotic divisions unless the me1os1s kytos, cell
follicle is stimulated to mature by follicle-stimulating hormone
(FSH) and luteinizing hormone (LH).
Secondary Oocyte Under the influence of FSH and LH (after puberty), primary Arrested in Haploid (ln) secondary,
oocytes complete meiosis I producing two cells of unequal Metaphase II of second,+ oon,
sizes. The larger cell becomes a secondary oocyte and the meiosis egg,+ kytos, cell
smaller one becomes a polar body, which degenerates.
The secondary oocyte will not undergo subsequent meiotic
divisions unless fertilization occurs
Definitive Oocyte Upon fertilization, the secondary oocyte undergoes the Formed after Haploid (ln) oon, egg,+ kytos,
second meiotic division to become an ovum secondary oocyte cell
is fertilized and
completes meiosis
~ - - - - - Secondary follicle
Granulosa c e l l s - - - - - - - - - -- - - - - - - - - - Zona pellucida
Antrum
Primary oocyte - - - - - - - - - - - - - - - - - ~
Primary follicles - - - - - - - - - - ~

Medulla

Primordial follicles

Tunica albuginea 1
Germinal epithelium _ __ /,

Ovarian ligament
~ - - - -- - Corona radiata
- - - - - - Z ona pellucida
----;-;r.;r-------- Ovulated secondary
Corpus albicans Corpus Developing oocyte
luteum corpus luteum
(a)

Ovarian ligament Primordial follicles Germinal epithelium and

tunica albuginea

_.,.;..:::....,-------1- Developing
follicles

"'
:::;
. .J ~-- ---------------------------------'
(b)

Figure 27.1 The Ovary. (a) The ovary is the primary reproductive organ of the female and produces the female gametes, the ova.
(bl Histological structure of the ovary demonstrating the outer cortex containing follicles in various stages of development, and the inner medulla,
which consists mainly of blood vessels and nerves.
(b) ©McGraw-Hill Education/Al Telser, photographer

6. Next move the microscope stage so a vesicular follicle is 0 antrum D secondary oocyte
in the center of the field of view. A vesicular (Graafian) 0 corona radiata D theca externa
follicle (figure 27.3) is a mature follicle that is ready to be
released during ovulation, after stimulation by a surge in 0 cumulus oophorus D theca interna
luteinizing hormone (LH), which is secreted by the anterior 0 granulosa cells D zona pellucida
pituitary gland. Table 27.3 lists the histological featu res 7. identify a corpus luteum and corpus albicans on the slide,
of a vesicular follicle. Identify the following parts of the
using table 27 .2 as a guide.
vesicular follicle, using table 27.3 and figure 27.3 as guides: ( continued on next paj
756 Chapter Twenty-Seven The Reproductive System and Early Development

(continued from previous page)

Table 27.2 Developmental Stages of Ovarian Follicles

Follicle Stage Primordial Follicle Primary Follicle Secondary (Maturing) Follicle

Photograph

©McGraw-Hill Education/Al Tclser, photographer ©McGraw-Hill Education/Al Telser, photographer ©Ed Reschke/Getty Images

Description Contains a primary oocyte surrounded Contains a primary oocyte surrounded Contains a full-size primary oocyte
and Function by a single layer of flattened follicular by one or more layers of cuboid al surrounded by an extracellular
cells follicular cells glycoprotein coat (the zona pell ucida),
which separates it from the coro na
radiata. Contains one or more fluid-
filled antra. Thecal ce lls deve lop into ,
internal and external layers.

Oocyte Stage Primary oocyte Primary oocyte Primary oocyte

Word Origins primus, first,+ ordior, to begin, primus, first, + ordior, to begin,+ secunda, second,+ ordior, to begin,
+ folliculus, a small sac fol/icu/us, a small sac + follicu/us, a small sac

Germinal epithelium

Tunica albuginea 'i~~~~ Theca externa

Theca interna
Granulosa cells
Antrum

Corona radiata
Ovarian cortex
Zona pellucida
Secondary oocyte

Cumulus oophorus

Figure 27.2 Germinal Epithelium and Tunica Albuginea

of the Ovary. The germinal epithelium is the most common origin
site for ovarian cancers. The tunic a albuginea is a "white coat" of
dense irregular connective tissue that surrounds the entire ovary Figure 27 .3 Vesicular (Graafian) Follicle. A vesic ul ar
internal to the germinal epithelium. follicle contains a secondary oocyte and a single antrum .
©McGraw-Hill Education/Al Tclscr, photographer ©McGraw-Hil l Education/Christine Eckel, photographer

INTEGRATE

LEARNING STRATEGY -$> contain a single layer of flattened cells, whe reas primary follicles con-
tain one or more layers of cuboida l cells. Next, look for the presence
There are several distinguishing features used to identify follicles in of fluid-filled antra. Secondary follicles contain one or more fluid-filled
various developmental stages. First, look for the number of cell layers antra, whereas vesicular follicles are characterized by the presence of
and the shape of the cells that surround the oocyte. Primord ial follicles a single large antrum.
Chapter Twenty-Seven The Reproductive System and Early Development

Vesicular (Graafian) Follicle Corpus Luteum Corpus Albicans

.,.," 8
C\I
::; "'
::;
-' - " " - ~ - " ' - ~ ' - - " " - """"-"'-=--~-'--"' -'
©Ed Reschke ©McGraw-Hill Education/Al Telser, photographer ©McGraw-Hill Education/Al Telser, photographer

Has nearly the same structure and fun ction A "yellow body" that gets its color from the If fertilization does not occ ur, t he corpus
as a secondary follicle, but contains a steroid hormones it secretes, which are luteum stops secretin g hormones after two
secondary oocyte and only a single large lipids. After ovulation, the theca interna cells weeks and becomes a smalle r, inactive
antrum enlarge and conti_nue to secrete the steroid "white body" consisting mainly of scar tissue
hormones estrogen and progesterone. In the
cente r of the corpus luteum is a large blood
clot.

Secondary oocyte NA NA
vesicular, a blister, + fol/icu/us, a sma ll sac corpus, body,+ Juteus, yellow corpus, body, + a/bus, white

Table 27.3 Components of a Vesicular Follicle

Structure Description and Function Word Origin

Antrum The fluid-filled space in the center of the folli cle antron, a cave

Corona Radiata A single layer of columnar cells derived from the cumulus oophorus that atta ch t o the corona, a crown,+ radiatus,
zona pellucida of the oocyte to shine

Cumulus A " mound" of granulosa cells that supports and surrounds the secondary oocyte within cumulus, a heap,+ oophoron,
Oophorus the follicle ovary

Granulosa Cells Epithelial cells lining the folli cle that will bec ome the luteal ce lls of th e corpus luteum granu/um, a small grain
after ovulation. They secrete the liquor folliculi, which is the fluid that fills the antrum.

Theca Externa The external fibrous layer of a well-developed vesicu lar follicle. The cells and fibers theca, a box, + externus,
are arranged in concentric layers. external

Theca lnterna Th e inner cellular layer of the vesicular follicle; these cells sec rete androgen th at is theca, a box,+ internus,
converted to estrogen by granulosa ce lls internal

Zona Pellucida A thick coat of glycoproteins that surrounds the oocyte zona, zone,+ pellucidus, clear
760 Chapter Twenty-Seven The Reproductive System and Early Development

EXERCISE 27 .3
'

HISTOLOGY OF THE UTERINE WALL three phases of the menstrual cycle and demonstrates the appearance
of the endometrium during each phase.
The uterus is a hollow, pear-shaped, muscular organ whose p1i-
mary function is to support, protect, and nourish the developing 1. Obtain a histology slide that includes a portion of the uterine
embryo/fetus. wall. Place it on the microscope stage and bring the tissue
The uterine wall is composed of three layers: endometrium, sample into focus on low power.
myometrium, and perimetrium. The inner lining of the uterus,
the endometrium, goes through its own cycle of growth during 2. Identify the following structures on the slide of the uterus,
the ovarian cycle. Each time a woman's ovary undergoes a single using tables 27 .5 and 27 .6 as guides:
ovarian cycle, the endornetrium becomes prepared for the possibil- D endometrium (basal layer) D myometrium
ity that a fertilized egg will become implanted. If no implantation
D endometrium (functional layer) D perimetrium
occurs, the innermost layer of the endometrium, the functional
layer, sloughs off in the process of menstruation (menstruus , 3. Move the microscope stage so the functional layer of the
monthly). Table 27.5 describes the layers of the uterine wall and endometii um is in the center of the field of view. Switch
the components that make up each layer. Table 27.6 summarizes to high power and identify uterine glands.

Table 27.5 Phases of the Menstrual Cycle

la) Menstrual Phase (b) Proliferative Phase le) Secretory Phase

Functional Functional Functional

layer layer layer
Endometrium
Endometrium
Endometrium Basal
Basal
layer layer
Basal
layer

n - - - -- Myometrium

~ - - - Myometrium

©Educational Images LTD/CustomMedical/Newscom ©Biophoto Associates/Science Source ©Biophoto Associates/Science Source

Days 1-5; Degeneration of the corpus luteum Days 5-14; The basal layer of the endometrium Days ·14-28; Begins at ovulation.
causes prog esterone and estrogen levels to begins to regenerate the functional layer Progesterone secreted by the corpus
drop. The functional layer of the endometrium luteum within the ovary stimulates the
becomes necrotic due to constriction of the uterine glands to begin secretion.
spiral arteries and is shed.

Table 27.6 Wall Layers of the Uterus

Wall Layer Description and Function Word Origin

Endometrium The mucous membrane composing the inner layer of the uterine wa ll. Consists of endon, within, + metra, uterus
simple columnar epithelium and a lamina propria with simple tubular uterine glands.
The structure, thickness, and state of the endometrium undergoes marked changes
during the menstrual cycle.

Functional Layer The apical layer of th e endometrium. Most of this layer is shed during menstruation. stratum, a layer, + functus,
(Stratum Functionalis) to perform

Basa/Layer The basal layer of the endometrium. It undergoes minimal changes during the stratum, a layer, + basalis, basal
(Stratum Basa/is) menstrual cycle and serves as the basis for regrowth of the more apical stratum
functionalis.

Myometrium The muscular wall of the uterus composed of three layers of smooth muscle mys, muscle,+ metra, uterus
Perimetrium The outermost covering of the uterus; a serous membrane formed from peritoneum peri-, around, + metra, uterus
Chapter Twenty-Seven The Reproductive System and Early Development
1
EXERCISE 27 .4

HISTOLOGY OF THE VAGINAL WALL

The vagina (vagina, sheath) is a muscular tube that functions
as the organ of copulation, serves as the passageway for men-
ses, and serves as the birth canal during parturition (parturio,

- to be in labor).
1. Obtain a histology slide demonstrating a portion of
the vaginal wall. Place it on the microscope stage and
bring the tissue sample into focus on low power. The
walls of the vagina are composed of a mucosa and
muscularis; the mucosa is lined with a nonkeratinized
stratified squamous epithelium (figure 27.S).

- 2. Identify the following structures on the slide of the

vagina, using figure 27.5 as a guide:
D lamina propria D nonkeratinized
D mucosa stratified squamous
epithelium
D muscularis

3. Optional Activity: IR 14: Reproductive System-

Watch the "Female Reproductive System Overview"
animation to review the female reproductive organs and
their histological features. Figure 27.5 Histology of the Vaginal Epithelium.
The epithelium is nonkeratinized stratified squamous.
©McGraw-Hill Education/Al Telser, photographer

Male Reproductive System

are to produce the male gamete (sperm), produce testosterone, and
The male reproductive system consists of the testes, epididymis, duc- provide nourishment for the sperm and a mechanism for the sperm
tus deferens, seminal vesicles, prostate gland, bulbourethral glands, to be delivered to the female reproductive tract. Testes also produce
and penis. The primary functions of the male reproductive system testosterone hormone.

EXERCISE 27 .5

.. HISTOLOGY OF THE SEMINIFEROUS developmental stages of the sperm, and table 27.8 describes the
structw·e and function of the accessory cell types located within
TUBULES the testes.
The testes are the primary reproductive organ in the male, and 1. Obtain a slide of the testes and place it on the microscope
function in the production of sperm and the male sex: steroid stage.
hormone testosterone. Within each testis are several hundred,
coiled seminiferous tubules (figure 27.6), which are the site of 2. Observe the slide on low power and identify several
sperm production, or spermatogenesis (sperma, seed, + genesis, cross sections of the seminiferous tubules. Move the
origin). Within each tubule, spermatogonia undergo succes- microscope stage so that one tubule is at the center of the
sive meiotic divisions as they move from the basal to the api- field of view, and change to high power.
cal surface of the tubule epithelium. Table 27.7 describes the (continued on next page)
762 Chapter Twenty-Seven The Reproductive System and Early Development

(continued from previous page)

Seminiferous tubule

Spermatogonium

Seminiferous tubules .,.,.m..rfil'""- Interstitial cell

l::'tf;j,~~r- Sustentacular cell
Tunica albuginea "ltf~~- Sperm cell nucleus

Lumen

(a) Testis, sagittal section

(b) Seminiferous tubules l

Primary - - -t--------..,.---- .,___

spermatocytes

Lumen - --+-

--...'-=---.,;,,...,e,;....-+--- Sustentacular cells

- ....-.r't-- Spermatid
- ; =~ - Spermatozoa

(c)

Figure 27.6 Testes and Seminiferous Tubules. (a) Testis and epididymis showing seminiferous tu bules within the testes and
tunica al bu ginea surrounding the testis. (b) Medium-power histological appearan ce of seminiferous tubules. (c) High-power histological view of
seminiferous tubules demonstrating stages of spermatogenesis and sustentacular cells.
(a) From Anatomy & Physiology Revealed, ©McGraw-Hill Education/The University of Toledo, photography and dissec tion; (b) ©McGraw-Hill Education/Al Telser,
photographer; (c) ©McGraw-Hill Education/Christine Ecke l, photographer

Table 27.7 Developmental Stages of Sperm

Cell Name Description and Function Stage of Mitosis/Meiosis Ploidy Word Origin
Spermatogonia Primitive male gametes that are formed by mitosis Formed by mitosis; undergo mitosis Diploid (2n) sperma, seed,
from male germline stem cells to form primary spermatocytes + gonia, generation
Primary Male gamete that has replicated its DNA in DNA is replicated in prophase of Diploid (2n) primary, first,
Spermatocyte preparation for meiosis. It will subsequently undergo meiosis I + sperma, seed,
meiosis I to form two secondary spermatocytes. + kytos, cell
Secondary Male gamete that has completed meiosis I and Formed after the first meiotic Haploid (1 n) secondary, second,
Spermatocyte will subsequently undergo meiosis II to form two division + sperma, seed,
spermatids + kytos, cell
Spermatid Male gamete that has completed meiosis, but has not Formed after the second meiotic Haploid (1 n) sperma, seed, + -id,
undergone spermiogenesis, the process of becoming division a young specimen
a mature spermatozoon
Spermatozoon A mature male gamete that has undergone Formed by maturation of Haploid (1n) sperma, seed,
(Sperm) spermiogenesis spermatids + zoon, animal
I Table 27.8 Accessory Cells of the Testis
I , Cell Name Description and Function Word Origin

I Interstitial (Leydig)
Cells
Produ ce the steroid hormone testostero ne, which is required for proper sperm
development and for the development of male second ary sex characteristics
inter-, between,+ sisto, to sta

I Sustentacular (Sertoli)
Cells
Surround multiple developing spermatocytes to provide them with support and
nourishment. Phagocytize excess cytoplasm from developing spermatocytes.
Form the blood-testis barrier, whic h protects the developing spermatocytes from
sustento, to hold upright

I antigens that circulate in the blood. Produce androgen-binding protein (ABP),

which concentrates testosterone around the developing spermatocytes.

I
I 3. Identify the following structures, using tables 27.7 and
27.8 and figure 27.6 as guides:

I 0 interstitial cell 0 spermatids

I • 0
0
primary spermatocyte
seminiferous tubule
D
D
spermatogonia
sustentacular cell

I 4. Sketch a seminiferous lubule as seen through the

microscope in the space provided. Be sure to label the

I structures listed in step 3 in the drawing.

5. Optional Activity: IR 14: Reproductive System-

I Watch the "Spermatogenesis" animation to visualize the

formation of sperm in the seminiferous tubules.

I
I EXERCISE 27 .6

I HISTOLOGY OF THE EPIDIDYMIS

After spermatozoa are formed within the seminiferous tubules,
2. Identify the following structures on the slide of the
epididymis, using table 27.9 and figure 27.7 as guides:

I they are transported through the straight tubules, rete testis, and
efferent ductules to enter the long, coiled tube of the epididymis
D pseudostratified
ciliated columnar
D
D
sperm cells
stereocilia

I (epi-, upon, + didymos, a twin [related to didymoi, testes]). Sper-

matozoa undergo the process of maturation and are stored in the
epithelial cells

3. Sketch the epididymis as seen through the microscope in

I
epididymis until ejaculation takes place. One of the most charac-
teristic features of the epididymis is the presence of thousands of the space provided. Be sure to label the structures listed
sperm in the lumen of the tube (figure 27.7). Table 27.9 describes in step 2 in the draw ing.

I the structure and function of the male accessory reproductive

structures, including the epididymis.

I 1. Obtain a slide of the epididymis and place it on the

microscope stage. Bring the tissue sample into focus

I
on low power and identify several cross sections of the
epididymis. Move the microscope stage so that one part
of the lumen of the epididymis is at the center of the

I field of view, and then change to high power. Notice

the sperm inside the lumen of the tubule, and how they
do not come right up against the apical surface of the
I columnar epithelial cells. This is because of the stereocilia
on the apical surface of the epithelial cells. Stereocilia
I (stereo, solid, + cilium, eyelid) are single, long microvilli
(ironically, they are not cilia at all!) that increase the

I surface area of the epithelial cells for the purpose of

secreting substances that nourish the sperm and absorbing

L substances from the sperm as they undergo maturation. ( continued on next pag
764 Chapter Twenty-Seven The Reproductive System and Early Development

(continued from previous page)

Cross section - - ---1aa'

of duct of
epididymis

Sperm in - - - - -
lumen of
duct of
epididymis
Stereocilia -- --
Pseudostratified
ciliated columnar
epithelial cells

(a) (b)

Figure 27. 7 Cross Section Through the Epididymis. Several cross sections through the seminiferous tubules are visible. (a) Note the
numerous sperm w ithin the lumen. (b) A higher magnification view demonstrates pseudostratified columnar epithelium with stereocilia.
(a ) ©Ed Reschke/Getty Images; (b) ©McGraw-Hill Education/Christine Eckel, photographer

Table 27.9 Male Accessory Reproductive Structures

Structure Epithelium Function Word Origin

Epididymis Pseudostratified columnar Site of storage and maturation of sperm. Walls contain epi-, upon, + didymos, a
epithelium with cilia some smooth muscle that w ill propel sperm into the twin (re lated to didymoi,
ductus deferens during ejaculation. testes)

Ouctus Pseudostratified column ar A thick, muscular tube whose walls undergo peristaltic ductus, to lead, + defero, to
Oeferens (Vas epithelium with stereoc ilia contractions during ejaculation to propel sperm into the carry away
Oeferens)
urethra

Prostate Gland Glandular epithelium Accessory reproductive gland that contributes prostates, one standing
resembling pseudostratified approximately 25% of the volume of semen. The fluid before
or simple columnar produced is rich in Vitamin C (citric acid) and enzymes
(such as PSA, prostate-specific antigen) that are
important for proper sperm function .

Seminal Pseudostratified columnar Accessory reproductive gland that contributes seminal, relating to sperm, +
Vesicles approximately 60% of the volume of semen. The fluid vesicula, a blister
produced is rich in fructose, w hich is a sou rce of energy
for the sperm. They also produce prostaglandins, which
are important in promoting sperm motility and may
stimulate uterine contractions.

Bulbourethral Glandular epithelium Accessory reproductive gland that produces a bu/bus, bulb, + urethral,
Glands consisting of simple and mucus-like lubricating substance during sexual arousal relating to the urethra
pseudostratified columnar (prior to ejacul ation) t hat lubricates the urethra and
neutralizes the acidity of the urine, thus preparing the
way for the spermatozoa to pass
tbtj bC ♦ CU i .CC h C]> •

EXERCISE 27. 7

HISTOLOGY OF THE DUCTUS DEFERENS 3. Sketch a cross section of the ductus deferens as seen
through the microscope in the space provided. Be sure t

I The ductus deferens (vas deferens) is a continuation of the epi-

didymis and is the route by which sperm travel from the epididy-
mis to the urethra during ejaculation (ejaculo, to shoot out). It
label the structures listed in step 2 in the drawing.

I is a highly muscular tube lined with pseudostratified columnar

epithelium with cilia (figure 27.8).

I I. Obtain a slide demonstrating the ductus deferens and

place it on the microscope stage. Bring the tissue sample

I into focus on low power and identify the cross section of

the ductus deferens (figure 27.8). Move the microscope

I stage so the lu men of the ductus deferens is at the center

of the field of view, and then change to high power.

I 2. Identify the following structures on the slide of the ductus

deferens, using table 27.9 and figure 27.8 as guides:

I D
D
cilia
lumen

I D
D
mucosa
pseudostratified ciliated colwnnar epithelial cells

I D smooth muscle

I
I -
- - Ductus deferens

~~

I
I
I Cilia - -- - -- - ......;,.,.....e,~.,...."""......

I
Pseudostratified - - - - P~ '-'ac?e,:",,:,ii
ciliated columnar
epithelial cells
Lumen - - -- - - --F.~
Mucosa--- - - -1¥:"'i&,B:c

Longitudinal - - -- --+-<'-"""P--,----'C(-,

I smooth muscle
layer

I Circular - - - --
smooth muscle
layer
- -CT,-- - =•

I (a) (b)

I Figure 27 .8 Ductus Deferens. (a) Cross sect ion t hrough t he entire du ctus deferens, demonstrat ing thick layers of smooth muscle
surrounding th e lumen. (b) Close-up view of the pseudostratified ciliated columnar epithelium.

I
(a ) ©McGraw-Hill Education/Al Telser, photographer; (b) © McGraw-H ill Education/Christine Eckel, photographer

I
I
766 Chapter Tuenty-Seven The Reproductive System and Early Development

EXERCISE 27 .8
3. Sketch the seminal vesicles as seen through the
HISTOLOGY OF THE SEMINAL VESICLES microscope in the space provided. Be sure to label the
Near the posterior wall of the bladder, each ductus defer- structures listed in step 2 in the drawing.
ens comes together with the duct from a seminal vesicle
(figure 27.9). These accessory reproductive glands are lined
with pseudostratified columnar epithelium and produce sub-
stances (including fructose, which is a source of energy for the
sperm) that are an important component of semen. Semen is the
tluid expelled from the penis during ejaculation (table 27.9).
1. Obtain a slide of the seminal vesicles and place it on
the microscope stage. Bring the tissue sample into focus
on low power. Identify cross sections of the seminal
vesicles. Move the microscope stage so the lumen of a
portion of the seminal vesicle is at the center of the field
of view, and then change to high power.
2. Identify the following structures on the slide of the seminal
vesicles, using table 27.9 and figure 27.9 as guides:
_____ x
D lumen
D rnucosalfolds
D muscular wall

(a)

Figure 27.9 The Seminal Vesicles. (a) Low magnification. (b) Medium magnification.
(a) & (b) ©McGraw-Hill Education/Al Telser, photographer
EXERCISE 27.9

I HISTOLOGY OF THE PROSTATE GLAND

I The prostate gland is an accessory reproductive gland located

immediately inferior to the urinary bladder. Like the seminal

I '
vesicles, the prostate produces substances that will become part
of semen and that are important for proper sperm function.

I
). Obtain a slide demonstrating the prostate gland and place
it on the microscope stage. Bring the tissue sample into
focus on low power and identify the prostatic urethra

I and the prostate gland itself (figure 27.10). Move the

microscope stage so a part of the gland relatively far

I away from the urethra is at the center of the field of view,

and then change to a higher power. Prostatic calculi - -41:--- 41=~
(corpora aranacea)
~ ~~~~

I 2. Identify the following structure on the slide of the

prostate, using table 27.9 and figure 27. 10 as guides:

I D tubuloalveolar glands
3. As men age, calcifications often form in the prostate. Tubuloalveolar - ----.. ~ :..:ca~

I Such calcifications are called prostatic calculi (corpora

aranacea) (figure 27. 10). Scan the slide and see if any of
glands

I
these are present in the specimen.

I Figure 27.10 The Prostate Gland.

calculi (corpora aranac ea ).
©McGraw-Hill Education/Al Telser, photographer
Not e the ma ny prost

I
I
I
I
I

I
I \

I -'
I
I
I
I
768 Chapter Tuenty-Seven The Reproductive System and Early Development

EXERCISE 27 .1 0

HISTOLOGY OF THE PENIS J. Obtain a slide demonstrating a cross section of the penis
and place it on the microscope stage. Observe at low
The penis is the male copulatory organ and is homologous to the power.
female clitoris. It is composed of three erectile bodies, the paired
corpora cavernosa and the single corpus spongiosum, which 2. Identify the following structures on the slide of the penis,
contains the male urethra (figure 27.11). In the center of each using figure 27.11 as a guide:
corpus cavemosum is a central artery that supplies blood to the
penis. During an erection, the veins draining the penis become □ central artery □ dorsal vein

constricted and venous blood fills up the erectile tissues. □ corpus cavernosum □ superficial fascia
(connective tissue)
□ corpus spongiosum

□ dorsal surface □ urethra

□ ventral surface
3. Sketch a cross section of the penis as seen through the
microscope in the space provided. Be sure to label the
structures listed in step 2 in the drawing.

Dorsal

Dorsal vein
Superficial
fascia
Tunica albuginea
of corpus
cavernosum
Corpora
cavernosa

Central
artery
Corpus spongiosum
Urethra
Tunica albuginea _ __ __ x
of corpus 0
X
st
spongiosum ::e _J

4. Optional Activity: IR 14: Reproductive System-

Ventral
Watch the "Male Reproductive System Overview"
animation to review the male reproductive structures and
Figure 27 .11 Cross Section Through the Shaft of the
Penis. The penis contains two paired corpora cavernosa dorsally, their relationships.
and a single corpus spongiosum ventrally. Note the male urethra in the
center of the corpus spongiosum.
©McGraw-Hill Education/Christine Eckel, photographer

CLINICAL VIEW
stimulated by the sympathetic nervous system. Erectile dysfunction involves
Erectile Dysfunction the interruption of the arousal response, and is most commonly caused by
Erectile dysfunction (ED) occurs when a male is unable to achieve or main- neurological, cardiovascular, or hormonal disturbances. Interestingly, some
tain an erection. Stimulation of the parasympathetic nervous system causes treatments for those suffering from erectile dysfunction, including sildenafil
central arteries within the corpora cavernosa to dilate. This dilation increases (i.e., Viagra), were first introduced as antihypertensive drugs. Sildenafil
local blood flow to the erectile tissues. Concomitant constriction of veins at reduces systemic blood pressure by dilating blood vessels and decreasing
the base of the penis prevents the drainage of blood from the erectile tissues. total peripheral resistance. As concerns an erection, sildenafil dilates the
The resulting engorging of the erectile tissues within the penis with blood central arteries within the penis, which facilitates the process of achieving
causes an erection. The penis remains erect until ejaculation, a process and maintaining an erection.
Chapter Tuenty-Seven The Reproductive System and Early Development 769

GRO~S ANATOMY
of these organs to gain an appreciation of their location within the
Female Reproductive System
female pelvic cavity, and their relationships with each other. Note
The internal detail, wall structures, and f~nctions _of most of ~he that there are numerous supporting ligaments that anchor the ovaries,
l.:rnale reproductive organs were covered m the histology section uterine tubes, and uterus in place within the pelvic cavity. Primary
,ii this chapter. This section involves observing the gross structure anatomic features of the female breast will then be observed.

-,;:
EXERCISE
_, 27 .11

GRO SS A NATOMY OF THE OVARY, UTERINE TUBES, UTERUS,

AND SUPPORTING LIGAMENTS
The ovary, uterine tubes, and uterus are all contained within the 2. Identify the structures listed in figures 27.12 and 27.13
pelvic cavity of the female and are held in place by a number on the cadaver or on classroom models, using table 27.10
of supporting ligaments, many of which form from folds of the and the textbook as guides. Then label them in
peiitoneum as it drapes over these structures. Table 27.10 sum- figures 27.12 and 27.13.
marizes the structure, function, and location of the supporting
3. Optional Activity: ~ 14: Reproductive System-
ligaments. In the Quiz section, select the "Structures: Female"
1. Observe a female human cadaver or classroom models option to test yourself on gross anatomy of female
of the female pelvis and female reproductive organs. structures.
Also observe a classroom model demonstrating a sagittal
section through a female pelvis.

\"
TableJ 7.10 Ligaments Supporting the Ovary, Uterine Tubes, and Uterus

Supporting Ligament Description and Function Word Origin

Broad Ligament Fold of peritoneum draped over the superior surface of the uterus. Portions of the broad, wide, + ligamentum,
broad ligament form the mesosalpinx and the mesovarium. a bandage

Mesosalpinx The mesentery of the uterine tube, which is formed as a fold of the most superior mesa-, a mesentery•like
part of the broad ligament structure,+salpinx, trumpet
(tube)

Mesovarium The mesentery of the ovary, which is formed as a posterior extension of the broad mesa·, a mesentery-like
ligament structure,+ovarium, ova ry

Ovarian Ligament Ligament contained within folds of the broad ligament. It extends from the medial ovarium, ovary,+ ligamentum,
part of the ovary to the superolateral surface of the body of the uterus. a bandage

Round Ligament of the Ligament attached to the superolateral surface of the body of the uterus. It extends /igamentum, a bandage,+
Uterus laterally to the deep inguinal ring, passes through the inguinal canal, and attaches uterus, uterus
to the skin of the labia majora.

Suspensory Ligament of A fold of peritoneum draping over the ovarian artery and vein superolateral to the suspensio-, to hang up, +
the Ovary ovary. Anchors the ovary to the lateral body wall. ligamentum, a bandage

Transverse Cervical Ligament extending laterally from the cervix and vagina, connecting them to the transverse-, across,+
(Cardinal) Ligament pelvic wall cervix, neck, + ligamentum, a
bandage

Uterosacra/ Ligaments Ligament connecting the inferior part of the uterus to the sacrum posteriorly utero·, the uterus,+ sacral,
the sac rum

(continued on next page)

770 Chapter Tuenty-Seven The Reproductive System and Early Development

( continued from previous page)

, - - - - - - - - - - - - - - - - - 12
3 Fundus of
uterus
Lumen of
uterus
r--::~~~ - - - -13
l '9rc'c::- - - - -- 14
7J!-'wH.:Fii-- - - - - - - 15

4- - - - - - l , - ~~ ~-~
[[11--_ _ ____L_J~_c:_------J:::;..c==~
Ovary- - - -- ,-,-- ----,;_-===:.-
6 _ _ _ _ _ _ _ _ _ _ _ _-"'"---""----"' +
16

Isthmus of the uterus - - - - - -- - - - - - ----

8 _ _ _ __ _ _ _ _____.. A'--"1
• ~ ~ '-+,- - - - - - - - -20
- - -4-- - - - - - - - - 2 1
"5-'~ -i.-H-- - - - - - - - - 22
[]]>---_ _ _ _ _ _ _ _ _ _ _ _ ____,.
f
10 l
11 _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ __ !+---+!
(a) Posterior view

Ovary

I
\.
--~,-;~
(b) Lateral sectional view
n~--- - - - - - - - - - ~
~ - - - - - - - - - - - H i l u m of ovary

Figure 27.12 Supporting Ligaments of the Ovary, Uterine Tubes, and Uterus as Seen from a Posterior View. (a) Posterior
view. (b) Lateral view. Use the terms listed to fill in the numbered labels in the figure. Numbers indicating ligaments are highlighted in green.
Some answers may be used more than once.

□ ampulla of uterine tube □ fimbria □ myometrium □ transverse cervical ligament

□ body of uterus □ infundibulum of uterine tube □ ovarian ligament □ uterine blood vessels

□ broad ligament □ internal os □ perimetrium □ uterine part of uterine tube

□ cervical canal □ isthmus of uterine tube □ round ligament of the uterus □ uterine tu be

□ endometrium □ mesosalpinx □ suspensory ligament of the

ovary
□ uterosacral ligament

□ external os □ mesovarium □ vagina

Chapter Twenty-Seven The Reproductive System and Early Development 771

2 - - -- - -- -- - - --.l!j
3 ____________ ~

4 _ __ __ _ _ _ _ _ __ _ _ __

_ _ _ _ _ _ _ _ _ _ _ __ _ _ _ __ 8
lschiopubic ramus-- - -- - -- - - -

___________________ g
5 _ _ _ _ _ _ _ _ _ _ _ _ _ __ __

6 _ _ __ _ _ _ _ _ _ _ _ _ _ __

(a) Sagittal view

_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ 17

_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ 18

~-- -- - - - - - - - - - - - -- 19
_ _ __ __ _ _ _ _ _ _ __ _ _ _ _ 20

(b) Midsagittal view

Figure 27.13 Classroom Model of the Female Pelvic Cavity. (a) Sagitta l view with pelvic struct ures intact. (bl Midsagittal view.
Use the terms listed to fill in the numbered labels in the figure. Some answers may be used more t han once.
[J anus □ f imbria of uterine tube □ rectouterine pouc h □ uterine tube

□ bulb of t he vestibule □ labia majora □ re ctum □ uterus

□ cervix of uterus □ labia minora □ round ligament of the uterus □ vagina

□ clitoris □ ovary □ ureter □ vagi nal orifice

□ external urethral orifice □ pubic symphysis □urinary bladder □ vesicouterine pouch

Model# Hl0 (1000281 ] ©3B Scientific GmbH, Germany, 2015 www.3bscientific.com/Photo by Christine Eckel, Ph.D.
772 Chapter Tuenty-Seven The Reproductive System and Early Development

EXERCISE 27 .12

GROSS ANATOMY OF THE FEMALE 1. Observe the breast of a prosected female human cadaver
or a classroom model of the female breast.
BREAST
2. Identify the structures listed in figure 27.14 on the
The female breast consists largely of fatty tissue and suspen-
cadaver or the classroom model of the female breast,
sory ligaments. Imbedded within are numerous modified sweat
using table 27 .11 and the textbook as guides. Then label
glands, the mammary glands (mamma, breast), which are com-
them in figure 27. 14.
pound tubuloal veolar exocrine glands. These glands enlarge
greatly during pregnancy, enabling them to produce milk to
nourish the new baby. Table 27 .11 describes the structures that
compose the female breast.

Table 27. 11 The Female Breast

Structure Description and Function Word Origin

Alveoli Secretory units of the mammary glands, which produce milk alveolus, a concave vessel,
a bowl

Areola Th e pigmented area of skin surrounding the nipple areola, area

Areolar Glands Sebaceous glands deep to the skin of the areola; produce sebum, which keeps the skin of areolar, relating to the
the areola moist, particularly during lactation areola of the breast

Lactiferous Ducts Ducts that form from the confluence of small ducts draining milk from the alveoli and facto-, milk+ ductus, to lead
lobules

Lactiferous 10-20 large channels that form from the confluence of several lactiferous ducts; the spaces facto-, milk+ sinus, cavity
Sinuses where milk is stored prior to release from the nipple

Lobes Large subdivisions of the mammary glands /obos, lobe

Lobules Smaller subdivisions of the mammary glands, which contain the alveoli /obu/us, a small lobe

Nipple A cylindrical projection in the center of the breast that contains the openings of the neb, beak or nose
lactiferous ducts

Suspensory Bands of connective tissue that anchor the breast skin and tissue to the deep fascia suspensio, to hang up,+
Ligaments overlying the pectoralis major muscle ligamentum, a band

INTEGRATE

CONCEPT CONNECTION gland, which further increases milk production. Oxytocin also stimulates
Lactation, the process of milk production and release from the mam- myoepithelial cells within mammary glands to contract, thereby releasing
mary glands, involves both positive and negative feedback mechanisms. milk from the breast Milk moves through the lactiferous ducts and sinuses,
Lactation is regulated by hormones that are present during pregnancy and exits the breast through openings in the nipple. Positive feedback
and after childbirth. During pregnancy, estrogen and progesterone levels mechanisms stimulate the production and ejection of milk. Negative
are elevated, which stimulates proliferation of the acini and branching of feedback mechanisms may inhibit GnRH release by the hypothalamus,
the lactiferous ducts within mammary glands. Following birth, estrogen thereby inhibiting FSH and LH release from the anterior pituitary gland
and progesterone levels drop, however prolactin levels are elevated, and preventing ovulation. Even so, women should use alternative methods
which stimulates milk production. The baby suckling on the mother's of birth control while breast feeding because some women continue to
breast stimulates the release of oxytocin from the posterior pituitary ovulate during this time.
_ __ _ __ _ _ _ _ _ _ _ _ _ _ _ _ ?

_ _ _ _ _ __ __ _ _ _ _ _ _ _ _ _ 8

____________________ 9

./
(a) Anteromedial view

,------------------16

11

12 17

13

(b) Sagittal view

Figure 27.14 The Female Breast. (a) Anterior view. (b) Sagittal view . Use the terms listed to fill in the numbered labels in the figure.
Some answers may be used more than once.

□ adipose tissue □ deep fascia □ lobule

□ alveoli □ lactiferous ducts □ nipple

□ areola □ lactiferous sinus □ suspensory ligaments

□ areolar gland □ lobe

(continued on next poge)

774 Chapter Twenty-Seven The Reproductive System and Early Development

(continued from previous page)

CLINICAL VIEW feel "lumpy." Specifically, fibrocystic changes in the breast consist
of fluid-filled cysts and are often surrounded by dense fibrous tissue.
Breast Cancer Screening and Breast This tissue, though often large and dense, is typically mobile (moves
Self-Examination (BSE) around easily), and often mirrors itself on the opposite breast. Th at is,
if you find such changes in the lower-right quadrant of the right breast,
Clinical Breast Exam (CBE) you might also find it on the lower-right quadrant of the left breast. On
All women of reproductive age are encouraged to make annual visits to the contrary, bre ast cancers tend to feel much firmer, like a kernel of
their gynecologist for a clinical breast exam (CBE) and a pap smear to unpopped popcorn, and they are immobile (they do not move wh en you
test for early cervical changes that are risk factors for cervica l cancer. touch them). They are generally painless and sometimes cause dimpling
The physician performing the CBE is palpating for abnormal lumps in the of the skin overlying the tumor.
breast. However, the physician can also help teach a woman how to The following are three procedures for performing a breast
perform breast self-exam (BSE) so she can monitor the condition of her self-exam:
breasts monthly instead of just during yearly visits. The vast majority of
1. In the showe r: Use soapy hands to palpate the breast while
breast cancers are discovered by the patient herself.
raising the arm on the same side as the breast you are
palpating. Starting at the periphery of the breast, use two or
Mammography three fingers to make small circles that progressively move
A mammogram is an x-ray of the breast that is used to look for areas of towa rd the nipple. Squeeze the nipple to look for discharge. Be
extra density or calcifications, which can be early signs of breast cancer. sure to palpate all the way up into the axilla, as the majority of
Most women are advised to start having mammograms at age 40, but tumors arise in the outer/upper quadrant towa rd the axillary
region of the breast.
those with a family history of the disease are often advised to have their
first mammogram at age 35. Mammography is good at locating very small 2. Lying down: Use the same procedure as in the shower. It is good
tumors (especially DCIS), which may not be palpable on self-examinations to do an exam lying down because it makes the breasts lie flat,
which may make it easier to feel certain lumps.
or clinical examinations.
3. Standing in front of a mirror: Place your hands on your hips, tilt
Breast Self-Exam (BSE) your elbows forward (anterior), and look for any indentations in
the skin of the breast or "orange peel"-looking skin, which can
Women should perform a breast self-exam at least once a month to
be indicative of an underlying tumor.
look for any unusual bumps, lumps, or thickening of breast tissue. Upon
palpation, a typical breast feels a bit lumpy, with some firmer areas If you discover anything that concerns you, make an appointment with
and some softer areas. Th e firmer/harder areas can be thickenings of your gynecologist as soon as possible to discuss your findings so the
the suspensory ligaments, or fibrocystic changes to the breast. The physician can help determine if what you felt was something benign or
softer areas are typically just adipose tissue. There are a number of something that nee(.ls further exploration. Always remember that breast
benign (noncancerous) changes to the breast that can make breasts cancer is highly curable when caught early.

Male Reproductive System majority of semen. The bulbouretbral glands are small glands located
within the urogenital diaphragm that produce a mucus-like substance
The testes, the primary male reproductive organs, function in the during sexual arousal that neutralizes the acidity of the male ure-
production of sperm and testosterone. Sperm produced in the testes thra and provides lubrication to ease the passage of semen during
are stored in the epididymis and transported via the ductus deferens ejaculation.
to the male urethra during ejaculation. The seminal vesicles and Exercises 27.5 to 27.10 involved examining the histological
prostate gland are accessory reproductive glands that produce sub- appearance of many of these structures. The following exercise
stances that nourish and protect the sperm and also compose the vast involves observing the male reproductive structures at the gross level.
l EXERCISE 27 .1 3
Chapter Twenty-Seven The Reproductive System and Early Development 775

GROSS ANATOMY OF THE SCROTUM, 2. Identify the structures listed in figures 27.15 and 27.16 on
the cadaver or on classroom models, using the textbook as
TESTIS, SPERMATIC CORD, AND PENIS a guide. Then label them in figures 27 .15 and 27 .16.
1. Observe a prosected male human cadaver or classroom 3. Optional Activity: ~ 14: Reproductive System- In
models of the male reproductive organs. the Quiz section, select the "Structures: Male" option to
test your knowledge of male reproductive gross anatomy.

....

....
•
1k
• 5- - -- - - -- - -- - - -++----- ---=~

..
'

.,

f.

'
•
'

1--- - - - - -- -- -- - -- - - - - - 2 0

- □
Figure 27.15 M ale Reproductive Tract Structures.

ampulla of du ctus deferens □ crus of penis

Poste rior view. Use t he t erms list ed to fill in the numbered labels in the fig ure.

□ mem branous urethra □ spongy ur ethra

□ bulb of pe nis □ du ctus def erens □ pro stat e gl and □ t esti s

□ bulbourethral gland □ ejaculato ry duct □ prost atic urethra □ ureter

~
□ corpus cavernos um □ epididymis □ seminal vesicle □ urinary bladder

□ co rpus spongio sum □ gl ans penis □ seminiferous tubules □ urogenital diaphragm

(continued on next page)

776 Chapter Twenty-Seven The Reproductive System and Early Development

( continued from previous poge)

h ---i- - - - -- - - - - -- - - - - -- 7
4 ,---'!lli--- -- -- -- - - - - - -- - - - 8

1071-- - - - - - - 1 2
_ __ __ __ _ __ 11J

- -- =---- - -- - - -- - -- - -- - - -- - - - 1 3
5 - - - - - - - - - - - - - - - - - - - - ' i~ - - -- -- -- -- -- - -- - -- - - --14
~ ~ r ~ ~ :::::::=-_ _ ___ ___ ___ _ _ _ _ ___ 15
6--- - - - -- - - - - - -- -- ~
(a) Sagittal view

16

17 26

27
18
28
19 29
30
20
31
21
32
22
33
23
24 34

25 35
(b) Mldsagittal view

Figure 27.16 Classroom Model of the Male Pelvic Cavity. (a) Sagittal view with pelvic structures intact. (b) Midsagittal view. Use
the terms listed to fill in the numbered labels in the figure. Some answers may be used more than once.
0 anus D glans penis O pubic symphysis 0 testicular artery and vein
D corpus cavernosum D internal urethral sphincter 0 rectum □ testis

□ corpus spongiosum □ membranous urethra 0 scrotum □ tunic a albuginea of testis

□ ductus deferens (ampullal □ penis 0 seminal vesicle D tunica vaginalis of testis

0 ductus deferens 0 prepuce 0 seminiferous tubules □ ureter
D ejaculatory duct □ prostate gland 0 spermatic cord □ urinary bladder
0 epididymis prostatic urethra n snnnnv llrP.thr;, n 11rnnnn:+ol rli»nhr ... nm
Chapter twency-se,en 1 fie iteproaue,e► t CJD

PHYSIOLOGY
.. Reproductive Physiology Table 27.12 depicts and describes each phase of meiosis and the
defining features of each phase.
.. Gametogenesis, the production of the gametes (sperm and second- Human somatic cells contain 23 pairs of chromosomes, or
ary oocytes) is accomplished by meiosis. Meiosis is similar in many homologous pairs. These include 22 pairs of autosomal chromo-
ways to mitosis, but there are two nuclear divisions (I and II), which somes, and one pair of sex chromosomes. Human gametes (egg and
result in a reduction of chromosome number from a diploid number sperm) contain 22 autosomal chromosomes and one sex chromo-
(2n, or 46) to a haploid number (n, or 23). Another difference in some, an X chromosome for oocytes and an X or Y chromosome for
meiosis is the process of synapsis (crossing over) that occurs when sperm. Meiosis yields four genetically different, haploid daughter
homologous chromosomes are adjacent to each other. Synapsis celJs. The process of fertilization brings haploid sperm together
is a process that introduces variability in the genetic code, which with a haploid secondary oocyte. When a sperm cell has fertilized
ensures that daughter cells are genetically different from parent cells. an oocyte, the resulting cell (called a zygote) is once again diploid.

Table 27.12 Phases of Meiosis

Description
Phase Word Origin
Prophase Homologous Replicated Nu cle ar pro -, before

\J .:.r\
maternal chromosome envelope
and paternal Sister breaks down;
chromosomes <h,omatids chromosomes
pair; crossover
(genetic
va ria bility}
1 gather

-t
occurs

Tetrad

Metaphase Homologous Spindle fiber Chromosomes meta-, mid dle

chromosomes attached to centromere attach to spindle
attach to sp indle fibers and line
___ .._ I Centromere
fibers and line - '-
the cell equator
the equator of
the cell
~ v '

Equator

(continued on next
778 Chapter Twenty-Seven The Reproductive System and Early Development

Table 27.12 Phases of Meiosis (continued)

Description
Phase Meiosis I Meiosis II
Anaphase Separation of Homologous replicated Sister chromatids ana-, upward
chromosome chromosomes separate pulled apart
pairs; reduction (single- stranded
division chromosomes)

Telophase Completion of Completion of telo-, end

nuclear division; nuclear division;
nuclear envelope nuclear envelope
reforms reforms
Cytokinesis Division of the Division of the cyto-, ce ll
cytoplasm; each cytoplasm;
cell contains 23 each of 4 cells
chromosomes contains 23
single- stranded
chromosomes
Cells separate into four
haploid daughter cells
...
,I
,:I
/,
l

Oogenesis continuation of meiosis. These "arrested" oocytes are housed within

The process of producing a mature egg (ovum) in females requires both ovarian follicles. Primordial follicles contain primary oocytes, which
mitotic and meiotic division. Primordial germ cells, or oogonia (singu- are available to be selected each month once a female reaches puberty,
lar, oogonium), divide by mitosis. Primary oocytes start the process of the time at which monthly hormone fluctuations begin. Only a few
meiosis. Oogenesis involves forming secondary oocytes from primary primordial follicles will mature into primary follicles, and fewer still
oocytes. However, rather than proceeding rapidly through the two will become secondary follicles. At ovulation a large vesicular follicle
phases of meiosis, primary and secondary oocytes experience periods bursts and releases a secondary oocyte from the follicle's fluid-filled
of "arrest" where meiosis is stopped until hormones or events trigger sac. The secondary oocyte will then be drawn into the uterine tubes to
cnaptct I ♦♦ CUCj' OCiCI I IP IOS:l&piO 0

potentially be fertilized by sperm. The final phases of meiosis occur (uterine cycle). The overall purpose is to produce viable eggs (ova)
.. after fertilization (see table 27. 12). for fertilization and to provide an attractive place for implantation
'
....
The result of fertilization is formation of a zygote, which will
subsequently undergo mitosis to form a mass of cells that may
of the ferti lized egg. The release of gonadotropin-releasing hormone
(GnRH) from the hypothalamus stimulates the release of luteinizing
implant in the uterine wall. Development in utero continues, as hormone (LH) and follicle-stimulating hormone (FSH) from the
described in exercise 27.16. Figure 27.17 depicts the overall pro- anterior pituitary gland. It is LH and FSH that stimulate the devel-
cess of oogenesis, incl uding identification of the various phases and opment of the follicles during the follicular phase of the ovarian
arrests of meiosis, corresponding stages of the ovarian follicles, and cycle. As the follicles develop, the granulosa cells surrounding the
... histological images of each stage. follicle secrete estrogen. Estrogen stimulates the cells in the uterine
Hormones are responsible for regulating the monthly changes endometrium to proliferate during the proliferative phase of the
that occur both in the ovaries (ovarian cycle) and in the uterus uterine cycle.

Before birth 0ogenesis Follicle development Micrograph

(embryonic and fetal period) {development of oocytes) examples

- 46 Oogonium

i Follicle cells

-
- - - - Primary oocyte - - - - - -
Primordial follicle
(arrested in prophase I)

Childhood
Meiosis I
arrested

Monthly, from puberty

to menopause
Primary follicle

tf
46 , - - - - Primary oocyte --------1
(arrested in
prophase I)
I\Secondary
Meiosis II - ~ . , ,.~ tollicle
arrested

2 3'
• '
) - (arrested
Secondary oocyte
in metaphase II)
. Vesicular
follicle

First
polar body
(degenerates) Ovulated
L ~ulaUoo

! /_
23 ~ tf
, - Sperm
- e-io-s-is_ l_l _ _ ~
~M
secondary
oocyte

~ completed (if
fertilization occurs)

Corpus luteum

23 c,
✓
Second
- -Ovum
polar body
(degenerates) c:JP Corpus albicans

Figure 27 .17 Oogenesis.

(]), (2), (6) & (7) ©McGraw-Hill Education/Al Telser, photographe r; (3) ©Ed Reschke/Getty Images; (4) ©Ed Reschke; (5) ©Dr. Francisco Gaytan
780 Chapter Tuenty-Seven The Reproductive System and Early Development

Once ovulation occurs, the ruptured vesicular follicle becomes responsible for maintalllillg a viable pregnancy. Human chori-
the corpus luteum. This structure begins producing progesterone and onic gonadotropin (hCG), the hormone secreted by the developing
estrogen throughout the luteal phase of the ovarian cycle. Increasing embryo, prevents the degradation of the corpus luteum in the event
levels of progesterone result in thickening and increased vasculariza- that fertilization and implantation occur. If fertilization and implan-
tion of the endometrium. In addition, the corpus luteum maintains tation do not occur, the corpus luteum degenerates into the corpus
elevated progesterone and estrogen levels in the event fertilization albicans. This degeneration also decreases levels of progesterone
occurs, but prior to the formation of the placenta, the structure and estrogen. As the progesterone levels drop, blood vessels (spiral

Primary Secondary Vesicular Ovulation Corpus luteum Regression Corpus

follicle follicle follicle forms albicans

Days
~ - - - 3 - - - 5 - - ~ 7_ _ _9_ _ _1_1_ _ _1~3 t ~1-5_ __1_1_ _ _1_9_ _ _2_1--r-__2_3_ _ _2_5_ __2_1_ __,

Follicular phase Ovulation Luteal phase

Gonadotropln levels

FSH
LH

Days 3 5 7 9 11 13 t 15 17 19 21 23 25 27

Ovulation

Ovarian hormone levels

Estrogen

Progesterone

Days 3 5 7 9 11 13 15 17 19 21 23 25 27

Uterine cycle

Days 13

Menstrual phase Proliferative phase Secretory phase

arterioles) in the uterine wall constrict, and the endometriaJ layer is Spennatogenesis
shed for menses. Typically, these changes occur in 28-day cycles,
The production of sperm, spermatogenesis, in males is similar
as depicted in figure 27.18. However, cycles can vary in length.
oogenesis in females. It is interesting that the overall process
Research suggests that the duration of the luteal phase remains more
regulated by the same hormones (GnRH, LH, and FSH) that regula
consistent at roughly 14 days, whereas the duration of the foliicular
oogenesis in females, with the exception of males secreting test
phase may vary widely among women.
terone in place of estrogen. GnRH from the hypothalamus stimula
the release of LH and FSH from the anterior pituitary gland.
INTEGRATE stimulates interstitial cells in the seminiferous tubules of the tes
to release testosterone, and FSH stimulates sustentacular cells
release androgen binding protein (ABP) and inhibin. The elevat
CONCEPT CONNECTION levels of testosterone are stimulatory for sperm production, yet
Estrogen released by follicular cells acts directly on the hypothalamus inhibitory for GnRH production by the hypothalamus. The proc
and the anterior pituitary gland to regulate the amount of gonadotropin- begins as sperrnatogonia, or primordial germ cells, actively under
releasing hormone (GnRH) released from the hypothalamus, and of mitosis and remain at the basement membrane of the seminifera
luteinizing hormone (LH) and follicle-stimulating hormone (FSH) released tubules. These spermatogonia divide to form primary sperma
from the anterior pituitary gland. Prior to ovulation, the increasing estro- cytes, which are diploid cells. Primary spermatocytes then unde
gen is stimulatory (positive feedback) for the release of GnRH, LH, and meiosis. Unlike females, males have the ability to continuously fo
FSH. However, once ovulation occurs, estrogen is inhibitory (negative new primary spermatocytes for gamete formation. After meosis
feedback) to the release of these hormones. It is a peak in LH that allows the spermatocyte is co nsidered secondary and haploid, with o
ovulation to occur; therefore, this switch from positive to negative feed- 23 chromosomes. Once the secondary spermatocyte undergoes me
back limits the number of eggs released each month. Incidentally, fertility sis II, it is considered a spermatid and is located closer to the lull]
drugs will often override these feedback mechanisms, thereby releasing of the seminferous tubule . Each of the four spermatids produced~
multiple secondary oocytes with the chance for multiples (twins, triplets, undergoes the maturation process, or spermiogenesis. This proc
or more). The release of these hormones was first introduced in the involves the development of a head, or chromosome-containing ac
chapter on the endocrine system, chapter 19. The concepts of negative some, a midpiece containing abundant mitochondria, and a tail (
and positive feedback appear throughout the book, and are critical to gellum) of the functional spermatozoon (plural, spermatozoa).
understanding homeostasis. spermatozoon continues the maturation process in the epididy
before becoming a viable sperm cell.

EXERCISE 27 .14

A CLINICAL CASE IN REPRODUCTIVE hours, with little time to cook or clean at home. For that rea-
son, they were constantly eating on the go, often getting takeout
PHYSIOLOGY meals that were, admittedly, not the healthiest. Despite this hectic
The purpose of this exerdse is to apply the concept of reproduc- schedule, the couple was ready to have a child of their own. So,
tive physiology to a real scenario, or clinical case study. This they began trying to conceive. Both Elizabeth and Frank were
exercise requires information presented in this chapter, as well optimistic that they would be able to conceive despite having
as in previous chapters. Integrating multiple systems in a case many friends that struggled with infertility. No women in Eliza-
study reinforces a concept that is very important in physiology: beth's family had struggled before, so she had no real concerns.
homeostasis. Frank and Elizabeth's attempts to conceive were met with
Before beginning, state how the ovarian cycle may be influ- disappointment. They "tried" for 1 year with no luck, and ulti-
enced by increasing levels of testosterone. mately sought medical advice, because Elizabeth was con-
cerned that her age would soon make conception more difficult.
Both Frank and Elizabeth visited a fertility cli nic to investigate
their case further. Doctors first tested Frank's sperm count and
sperm viability, because this is. the simplest test to conduct.
There was no evidence of inadequate numbers of sperm or their
Consult with the laboratory instructor as to whether this inability to fertilize an egg. Frank and Elizabeth were relieved,
exercise will be completed individually, in pairs, or in groups. yet still frustrated that they were unsure as to the cause of their
Review the topics of the cardiovascular, lymphatic, respiratory, infertility.
digestive, urinary, and reproductive systems in the textbook and Doctors then made note of Elizabeth' s physical health. At
in previous chapters in this laboratory manual as needed for 5 feet, 0 inches, Elizabeth weighed 175 pounds. In addition to
assistance with the case study. being overweight, she also suffered from chronic hypertension
and an elevated heart rate. Doctors suggested a change in life-
Case Study style to reduce stress, which they hoped would help Elizabeth
Elizabeth, a 32-year-old newlywed, was eager to start a family. lose weight and improve her cardiovascular health. Doctors
She was career focused, as was her husband, Frank, and both also inquired about Elizabeth's reproductive health. Eliza-
thrived under high-stress conditions. Their jobs required Jong beth admitted that she had been prescribed birth control pills

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782 Chapter Twenty-Seven The Reproductive System and Early Development

( continued from previous page)

Answer the following questions regarding Elizabeth's
at 15 years old to help regulate her irregular (unpredictable) case:
periods. Since stopping the pill just 1 year ago, her periods a. Doctors often prescribe metformin, a drug that decreases
were again irregular, with cycle durations ranging from 31 to glucose transport in the GI tract, for patients suffering
51 days. Menses would last 7 days with no remarkable pain from PCOS. Describe how this would be beneficial,
or discomfort. She also noticed that she had an increased inci- given the symptoms of the disease.
dence of facial hair and acne, something that was quite unusual
for her. Based on these symptoms, doctors ordered blood
samples (table 27.13) to test for a condition called polycystic
ovarian syndrome (PCOS).
PCOS is an endocrine disorder, characterized by elevated
levels of testosterone, that is a major cause of infertility in b. Elizabeth's blood samples were taken on day 3 of the
females. Patients are often diabetic (type TT), and exhibit signs of menstrual cycle. Describe the relative levels of estrogen,
insulin resistance. Patients tend to be overweight and experience progesterone, LH, and FSH during this phase of the cycle.
irregular periods due to a disruption in the positive and nega-
tive feedback mechanisms that regulate the ovarian and uterine
cycles. Often, patients with PCOS do not ovulate, thereby pre-
venting fertilization and implantation. Instead of ovulating,
partially developed follicles may persist as "cysts" in the ovary.
Clinically, ultrasound is used to observe ovarian cysts, and a c. How might these hormone levels be different if samples
diagnosis of PCOS requires that one ovary has more than twelve were collected on day 14 (assume a 28-day cycle)?
follicles ranging 2-9 mm in diameter. It is proposed that regula-
tion of blood glucose levels and weight loss may allow ovula-
tion to resume, although the exact mechanisms are unclear. In
the event that dietary changes are unsuccessful, drugs can be
administered to regulate blood glucose levels as well as stimu-
late ovulation. d. Desc1ibe the potential consequences of elevated LH when
compared to FSH levels.
Table 27.13 Elizabeth's Blood Test Results
Blood Test Result Reference Range

Cholesterol (mg/di) 218 <200

e. Discuss why elevated testosterone may lead to anovulation.
FSH(mLU/L) 6 3-20 (Hint: Testosterone exhibits similar feedback mechanisms
in males and females).
Glucose (mg/di) 141 70-110

Insulin (pU/mL) 167 0-20

LH(mLU/L) 18 3-20
f. Do you think that Elizabeth is suffering from PCOS?
Testosterone (pg/ml) 15 1.1- 6.3

Triglycerides (mg/di) 291 40-150

Chapter twency-se,en 1 ne nep, oao O P
~ I

,' Fertilization and Development

The time period between fertilization and birth is considered the
the official start of gestation, a period that lasts 38 weeks in al
Weeks 3 through 8 constitute the embryonic period, a time in whic
the organism is called an embryo. During this time, major organs an
prenatal period. Fertilization yields a diploid cell, or zygote, that organ systems develop. In the ninth week of development, the organ
undergoes multiple mitotic divisions, known as cleavage events. ism is classified as a fetus. The fetal period lasts for the remainin
Two cells become four cells, and then eight cells, until there is a 30 weeks of gestation. During the fetal period, organs and system
mass of cells, called a blastocyst, which is ready for implantation in continue to grow and develop in preparation for birth and to sustai

- the uterine wall. This cleavage process takes approximately 2 weeks

and constitutes the pre-embryonic period. Implantation marks
life separate from the mother's womb. The following laboratory exe
cises explore the events of the pre-embryonic and embryonic period

EXERCISE 27 .15

EARLY DEVELOPMENT: FERTILIZATION

- AND ZVGOTE FORMATION

The following laboratory exercise focuses on the changes that depicting the chronology of events in pre-embryonic development,

- occur during the pre-embryonic period. Complete table 27.14,

Table 27.14
using the textbook as a guide.

Pre-Embryonic Period

- ---Sperm pronucleus

1-- 120 µm ----i

·'.>

---Nucleus

1-- 120 µm ----i

-
1--120 µm ----i 1--120 µm - i
4-cell stage 8-cell stage

1--120 µm-i
Morula

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-
784 Chapter Twenty-Seven The Reproductive System and Early Development

(continued from previous page)

Table 27.14 Pre-Embryonic Period (continued)

Appearance Developmental Stage
---- Location
----~--
Events

f--120 µm -----i

Cytotrophoblast - -~ n ~
Embryoblast - - -~ - ---,-"

EXERCISE 27 .16

EARLY DEVELOPMENT: EMBRYONIC 1. Label the structures of the 3-week-old embryo identified
in figure 27.19.
DEVELOPMENT
2. Complete table 27.15 depicting the chronology of events
This laboratory exercise focuses on the changes that occur in an in embryonic development.
organism during the embryonic period. Complete the following
using the textbook as a guide. 3. Label the structures of the 4-week-old embryo identified
in figure 27.20.

~ -~ -+- -- -- - - - - - - 5
',-,,--,' - - -- - - - -- - -- 6 1 - - - - - - - - - - 8
---,,._ _ _ _ __ _ __ _ _ _ 7

Figure 27.19 Structures of the 3-week-old Embryo. Use the terms listed to fill in the numbered la bels in the figure .

D amnion D chorion D embryo D placent a

D amniotic cavity D connecting stalk D functional layer of D yolk sac

uterus
Table 27.15 Stages of Embryonic Development

- Developmental Week Events

-
l f
Week3

- Ne"'"'
Neural 9""'"
fold

- 1.5 mm

j t;- Primitive streak

Superior/dorsal view

Week4

l
-Head

Heart
4.0mm ")--- Umbilical cord
Limb buds

j
Lateral view

Week5-8

l
30mm

j
Lateral view

f----------------6

;-------------7
___________ 8

_ _ _ _ _ __ _ _ _ _ g

- - - - - - - - - - - - 10

Figure 27 .20 Structures of the 4-week-old (Late) Embryo. Use the terms listed to fill in the numbered labels in the figure.

D abdominal wa ll 0 endoderm 0 mesentery 0 peritoneal cavity

D amnion 0 guttube 0 mesoderm 0 somite

D ectoderm 0 intermediate mesoderm