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Raven Johnson McGraw-Hill Biology-11

Biology book part 11
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0% found this document useful (0 votes)
209 views42 pages

Raven Johnson McGraw-Hill Biology-11

Biology book part 11
Copyright
© © All Rights Reserved
We take content rights seriously. If you suspect this is your content, claim it here.
Available Formats
Download as PDF, TXT or read online on Scribd
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Sperm Zygote

Y Testes
Ovum XY
Develop in early
embryo

X Seminiferous
SRY tubules

Leydig
Indifferent cells
gonads

X No SRY
Ovaries

Ovum XX
X (Follicles do not
develop until
third trimester)
Sperm Zygote

FIGURE 59.3
Sex determination in mammals is made by a region of the Y chromosome designated SRY. Testes are formed when the Y
chromosome and SRY are present; ovaries are formed when they are absent.

There are some deep-sea fish that are hermaphro- onic gonads are said to be “indifferent.” If the embryo is a
dites—both male and female at the same time. Numerous male, it will have a Y chromosome with a gene whose prod-
fish genera include species in which individuals can uct converts the indifferent gonads into testes. In females,
change their sex, a process called sequential hermaphro- which lack a Y chromosome, this gene and the protein it
ditism. Among coral reef fish, for example, both protog- encodes are absent, and the gonads become ovaries. Recent
yny (“first female,” a change from female to male) and evidence suggests that the sex-determining gene may be
protandry (“first male,” a change from male to female) one known as SRY (for “sex-determining region of the Y
occur. In fish that practice protogyny (figure 59.2b), the chromosome”) (figure 59.3). The SRY gene appears to have
sex change appears to be under social control. These fish been highly conserved during the evolution of different
commonly live in large groups, or schools, where success- vertebrate groups.
ful reproduction is typically limited to one or a few large, Once testes form in the embryo, the testes secrete
dominant males. If those males are removed, the largest testosterone and other hormones that promote the devel-
female rapidly changes sex and becomes a dominant opment of the male external genitalia and accessory repro-
male. ductive organs. If the embryo lacks testes (the ovaries are
nonfunctional at this stage), the embryo develops female
external genitalia and sex accessory organs. In other words,
Sex Determination
all mammalian embryos will develop female sex accessory
Among the fish just described, and in some species of rep- organs and external genitalia unless they are masculinized
tiles, environmental changes can cause changes in the sex of by the secretions of the testes.
the animal. In mammals, the sex is determined early in em-
bryonic development. The reproductive systems of human
males and females appear similar for the first 40 days after Sexual reproduction is most common among animals,
conception. During this time, the cells that will give rise to but many reproduce asexually by fission, budding, or
ova or sperm migrate from the yolk sac to the embryonic parthenogenesis. Sexual reproduction generally involves
the fusion of gametes derived from different individuals
gonads, which have the potential to become either ovaries
of a species, but some species are hermaphroditic.
in females or testes in males. For this reason, the embry-

Chapter 59 Sex and Reproduction 1197


59.2 The evolution of reproduction among the vertebrates has led to
internalization of fertilization and development.
Fertilization and Development ductive tract. By this means, fertilization still occurs in a
nondesiccating environment, even when the adult ani-
Vertebrate sexual reproduction evolved in the ocean before mals are fully terrestrial. The vertebrates that practice in-
vertebrates colonized the land. The females of most species ternal fertilization have three strategies for embryonic
of marine bony fish produce eggs or ova in batches and re- and fetal development:
lease them into the water. The males generally release their
1. Oviparity. This is found in some bony fish, most
sperm into the water containing the eggs, where the union
reptiles, some cartilaginous fish, some amphibians, a
of the free gametes occurs. This process is known as exter-
few mammals, and all birds. The eggs, after being fer-
nal fertilization.
tilized internally, are deposited outside the mother’s
Although seawater is not a hostile environment for ga-
body to complete their development.
metes, it does cause the gametes to disperse rapidly, so
2. Ovoviviparity. This is found in some bony fish (in-
their release by females and males must be almost simul-
cluding mollies, guppies, and mosquito fish), some
taneous. Thus, most marine fish restrict the release of
cartilaginous fish, and many reptiles. The fertilized
their eggs and sperm to a few brief and well-defined peri-
eggs are retained within the mother to complete their
ods. Some reproduce just once a year, while others do so
development, but the embryos still obtain all of their
more frequently. There are few seasonal cues in the
nourishment from the egg yolk. The young are fully
ocean that organisms can use as signals for synchronizing
developed when they are hatched and released from
reproduction, but one all-pervasive signal is the cycle of
the mother.
the moon. Once each month, the moon approaches
3. Viviparity. This is found in most cartilaginous
closer to the earth than usual, and when it does, its in-
fish, some amphibians, a few reptiles, and almost all
creased gravitational attraction causes somewhat higher
mammals. The young develop within the mother
tides. Many marine organisms sense the tidal changes and
and obtain nourishment directly from their moth-
entrain the production and release of their gametes to the
er’s blood, rather than from the egg yolk (fig-
lunar cycle.
ure 59.4).
The invasion of land posed the new danger of desicca-
tion, a problem that was especially severe for the small
and vulnerable gametes. On land, the gametes could not
simply be released near each other, as they would soon Fertilization is external in most fish but internal in most
dry up and perish. Consequently, there was intense selec- other vertebrates. Depending upon the relationship of
the developing embryo to the mother and egg, those
tive pressure for terrestrial vertebrates (as well as some
vertebrates with internal fertilization may be classified
groups of fish) to evolve internal fertilization, that is,
as oviparous, ovoviviparous, or viviparous.
the introduction of male gametes into the female repro-

FIGURE 59.4
Viviparous fish
carry live, mobile
young within their
bodies. The young
complete their
development within
the body of the
mother and are then
released as small but
competent adults.
Here a lemon shark
has just given birth
to a young shark,
which is still
attached by the
umbilical cord.

1198 Part XIV Regulating the Animal Body


Fish and Amphibians
Most fish and amphibians, unlike other vertebrates, repro-
duce by means of external fertilization.

Fish
Fertilization in most species of bony fish (teleosts) is exter-
nal, and the eggs contain only enough yolk to sustain the
developing embryo for a short time. After the initial supply
of yolk has been exhausted, the young fish must seek its
food from the waters around it. Development is speedy,
and the young that survive mature rapidly. Although thou-
sands of eggs are fertilized in a single mating, many of the
resulting individuals succumb to microbial infection or pre-
dation, and few grow to maturity.
In marked contrast to the bony fish, fertilization in most
cartilaginous fish is internal. The male introduces sperm
into the female through a modified pelvic fin. Development
of the young in these vertebrates is generally viviparous.

Amphibians
The amphibians invaded the land without fully adapting FIGURE 59.5
to the terrestrial environment, and their life cycle is still The eggs of frogs are fertilized externally. When frogs mate,
tied to the water. Fertilization is external in most amphib- as these two are doing, the clasp of the male induces the female to
ians, just as it is in most species of bony fish. Gametes release a large mass of mature eggs, over which the male
discharges his sperm.
from both males and females are released through the
cloaca. Among the frogs and toads, the male grasps the fe-
male and discharges fluid containing the sperm onto the
eggs as they are released into the water (figure 59.5). Al-
though the eggs of most amphibians develop in the water,
there are some interesting exceptions. In two species of
frogs, for example, the eggs develop in the vocal sacs and
stomach, and the young frogs leave through their moth-
er’s mouth (figure 59.6)!
The time required for development of amphibians is
much longer than that for fish, but amphibian eggs do not
(a)
include a significantly greater amount of yolk. Instead, the (c)
process of development in most amphibians is divided into
embryonic, larval, and adult stages, in a way reminiscent of
the life cycles found in some insects. The embryo develops
within the egg, obtaining nutrients from the yolk. After
hatching from the egg, the aquatic larva then functions as a
free-swimming, food-gathering machine, often for a con-
siderable period of time. The larvae may increase in size
rapidly; some tadpoles, which are the larvae of frogs and
(b) (d)
toads, grow in a matter of weeks from creatures no bigger
than the tip of a pencil into individuals as big as a goldfish.
When the larva has grown to a sufficient size, it undergoes FIGURE 59.6
Different ways young develop in frogs. (a) In the poison arrow
a developmental transition, or metamorphosis, into the ter-
frog, the male carries the tadpoles on his back. (b) In the female
restrial adult form.
Surinam frog, froglets develop from eggs in special brooding
pouches on the back. (c) In the South American pygmy marsupial
The eggs of most bony fish and amphibians are
frog, the female carries the developing larvae in a pouch on her
fertilized externally. In amphibians the eggs develop
back. (d) Tadpoles of the Darwin’s frog develop into froglets in
into a larval stage that undergoes metamorphosis.
the vocal pouch of the male and emerge from the mouth.

Chapter 59 Sex and Reproduction 1199


Reptiles and Birds
Most reptiles and all birds are
oviparous—after the eggs are fertilized
internally, they are deposited outside of
the mother’s body to complete their de-
velopment. Like most vertebrates that
fertilize internally, male reptiles utilize a
tubular organ, the penis, to inject sperm
into the female (figure 59.7). The penis,
containing erectile tissue, can become
quite rigid and penetrate far into the fe-
male reproductive tract. Most reptiles
are oviparous, laying eggs and then
abandoning them. These eggs are sur-
rounded by a leathery shell that is de-
posited as the egg passes through the
oviduct, the part of the female reproduc-
tive tract leading from the ovary. A few
species of reptiles are ovoviviparous or
viviparous, forming eggs that develop
into embryos within the body of the
mother. FIGURE 59.7
All birds practice internal fertilization, The introduction of sperm by the male into the female’s body is called copulation.
though most male birds lack a penis. In Reptiles such as these turtles were the first terrestrial vertebrates to develop this form of
some of the larger birds (including reproduction, which is particularly suited to a terrestrial environment.
swans, geese, and ostriches), however,
the male cloaca extends to form a false
penis. As the egg passes along the oviduct, glands secrete
albumin proteins (the egg white) and the hard, calcareous
shell that distinguishes bird eggs from reptilian eggs. While
modern reptiles are poikilotherms (animals whose body
temperature varies with the temperature of their environ-
ment), birds are homeotherms (animals that maintain a rel-
atively constant body temperature independent of environ-
mental temperatures). Hence, most birds incubate their
eggs after laying them to keep them warm (figure 59.8).
The young that hatch from the eggs of most bird species
are unable to survive unaided, as their development is still
incomplete. These young birds are fed and nurtured by
their parents, and they grow to maturity gradually.
The shelled eggs of reptiles and birds constitute one of
the most important adaptations of these vertebrates to life
on land, because shelled eggs can be laid in dry places.
Such eggs are known as amniotic eggs because the embryo
develops within a fluid-filled cavity surrounded by a mem- FIGURE 59.8
brane called the amnion. The amnion is an extraembry- Crested penguins incubating their egg. This nesting pair is
onic membrane—that is, a membrane formed from embry- changing the parental guard in a stylized ritual.
onic cells but located outside the body of the embryo.
Other extraembryonic membranes in amniotic eggs in-
clude the chorion, which lines the inside of the eggshell,
the yolk sac, and the allantois. In contrast, the eggs of fish
and amphibians contain only one extraembryonic mem- Most reptiles and all birds are oviparous, laying
amniotic eggs that are protected by watertight
brane, the yolk sac. The viviparous mammals, including
membranes from desiccation. Birds, being
humans, also have extraembryonic membranes that will be
homeotherms, must keep the eggs warm by incubation.
described in chapter 60.

1200 Part XIV Regulating the Animal Body


Mammals The most primitive mammals, the monotremes (con-
sisting solely of the duck-billed platypus and the
Some mammals are seasonal breeders, reproducing only echidna), are oviparous, like the reptiles from which they
once a year, while others have shorter reproductive cycles. evolved. They incubate their eggs in a nest (figure 59.9a)
Among the latter, the females generally undergo the repro- or specialized pouch, and the young hatchlings obtain
ductive cycles, while the males are more constant in their milk from their mother’s mammary glands by licking her
reproductive activity. Cycling in females involves the peri- skin, as monotremes lack nipples. All other mammals are
odic release of a mature ovum from the ovary in a process viviparous, and are divided into two subcategories based
known as ovulation. Most female mammals are “in heat,” on how they nourish their young. The marsupials, a
or sexually receptive to males, only around the time of ovu- group that includes opossums and kangaroos, give birth
lation. This period of sexual receptivity is called estrus, to fetuses that are incompletely developed. The fetuses
and the reproductive cycle is therefore called an estrous complete their development in a pouch of their mother’s
cycle. The females continue to cycle until they become skin, where they can obtain nourishment from nipples of
pregnant. the mammary glands (figure 59.9b). The placental mam-
In the estrous cycle of most mammals, changes in the se- mals (figure 59.9c) retain their young for a much longer
cretion of follicle-stimulating hormone (FSH) and luteiniz- period of development within the mother’s uterus. The
ing hormone (LH) by the anterior pituitary gland cause fetuses are nourished by a structure known as the pla-
changes in egg cell development and hormone secretion in centa, which is derived from both an extraembryonic
the ovaries. Humans and apes have menstrual cycles that membrane (the chorion) and the mother’s uterine lining.
are similar to the estrous cycles of other mammals in their Because the fetal and maternal blood vessels are in very
cyclic pattern of hormone secretion and ovulation. Unlike close proximity in the placenta, the fetus can obtain nu-
mammals with estrous cycles, however, human and ape fe- trients by diffusion from the mother’s blood. The func-
males bleed when they shed the inner lining of their uterus, tioning of the placenta is discussed in more detail in
a process called menstruation, and may engage in copula- chapter 60.
tion at any time during the cycle.
Rabbits and cats differ from most other mammals in that Among mammals that are not seasonal breeders, the
they are induced ovulators. Instead of ovulating in a cyclic females undergo shorter cyclic variations in ovarian
fashion regardless of sexual activity, the females ovulate function. These are estrous cycles in most mammals
and menstrual cycles in humans and apes. Some
only after copulation as a result of a reflex stimulation of
mammals are induced ovulators, ovulating in response
LH secretion (described later). This makes these animals
to copulation.
extremely fertile.

(a) (b) (c)

FIGURE 59.9
Reproduction in mammals. (a) Monotremes, like the duck-billed platypus shown here, lay eggs in a nest. (b) Marsupials, such as this
kangaroo, give birth to small fetuses which complete their development in a pouch. (c) In placental mammals, like this domestic cat, the
young remain inside the mother’s uterus for a longer period of time and are born relatively more developed.

Chapter 59 Sex and Reproduction 1201


59.3 Male and female reproductive systems are specialized for different functions.

Structure and Function of the Male Production of Sperm


Reproductive System The wall of the seminiferous tubule consists of germinal
cells, which become sperm by meiosis, and supporting
The structures of the human male reproductive system, Sertoli cells. The germinal cells near the outer surface of
typical of mammals, are illustrated in figure 59.10. If the seminiferous tubule are diploid (with 46 chromo-
testes form in the human embryo, they develop seminifer- somes in humans), while those located closer to the
ous tubules beginning at around 43 to 50 days after con- lumen of the tubule are haploid (with 23 chromosomes
ception. The seminiferous tubules are the sites of sperm each). Each parent cell duplicates by mitosis, and one of
production. At about 9 to 10 weeks, the Leydig cells, lo- the two daughter cells then undergoes meiosis to form
cated in the interstitial tissue between the seminiferous sperm; the other remains as a parent cell. In that way, the
tubules, begin to secrete testosterone (the major male sex male never runs out of parent cells to produce sperm.
hormone, or androgen). Testosterone secretion during Adult males produce an average of 100 to 200 million
embryonic development converts indifferent structures sperm each day and can continue to do so throughout
into the male external genitalia, the penis and the scrotum, most of the rest of their lives.
a sac that contains the testes. In the absence of testos- The diploid daughter cell that begins meiosis is called
terone, these structures develop into the female external a primary spermatocyte. It has 23 pairs of homologous
genitalia. chromosomes (in humans) and each chromosome is du-
In an adult, each testis is composed primarily of the plicated, with two chromatids. The first meiotic division
highly convoluted seminiferous tubules (figure 59.11). separates the homologous chromosomes, producing two
Although the testes are actually formed within the ab- haploid secondary spermatocytes. However, each chromo-
dominal cavity, shortly before birth they descend through some still consists of two duplicate chromatids. Each of
an opening called the inguinal canal into the scrotum, these cells then undergoes the second meiotic division to
which suspends them outside the abdominal cavity. The separate the chromatids and produce two haploid cells,
scrotum maintains the testes at around 34°C, slightly the spermatids. Therefore, a total of four haploid sper-
lower than the core body temperature (37°C). This lower matids are produced by each primary spermatocyte (fig-
temperature is required for normal sperm development ure 59.11). All of these cells constitute the germinal ep-
in humans. ithelium of the seminiferous tubules because they
“germinate” the gametes.
In addition to the germinal epithelium, the walls of
the seminiferous tubules contain nongerminal cells
Bladder Ureter
known as Sertoli cells. The Sertoli cells nurse the devel-
oping sperm and secrete products required for spermato-
genesis (sperm production). They also help convert the
spermatids into spermatozoa by engulfing their extra
cytoplasm.
Seminal Spermatozoa, or sperm, are relatively simple cells, con-
vesicle
sisting of a head, body, and tail (figure 59.12). The head
Urethra encloses a compact nucleus and is capped by a vesicle
Ejaculatory called an acrosome, which is derived from the Golgi com-
duct
plex. The acrosome contains enzymes that aid in the pen-
etration of the protective layers surrounding the egg. The
Prostate
gland
body and tail provide a propulsive mechanism: within the
Vas deferens tail is a flagellum, while inside the body are a centriole,
Epididymis Cowper's
(bulbourethral) which acts as a basal body for the flagellum, and mito-
Testis gland chondria, which generate the energy needed for flagellar
Penis Scrotum movement.

FIGURE 59.10
Organization of the human male reproductive system. The
penis and scrotum are the external genitalia, the testes are the
gonads, and the other organs are sex accessory organs, aiding the
production and ejaculation of semen.

1202 Part XIV Regulating the Animal Body


Epididymis Sertoli cell
Testis
Germinal cell
(diploid)
Coiled
seminiferous
tubules

Primary
spermatocyte
(diploid)
MEIOSIS I
Secondary
spermatocytes
(haploid)
MEIOSIS II
Vas deferens
Spermatids
(haploid)

Spermatozoa

Cross-section of
seminiferous tubule

FIGURE 59.11
The testis and spermatogenesis. Inside the testis, the seminiferous tubules are the sites of spermatogenesis. Germinal cells in the
seminiferous tubules give rise to spermatozoa by meiosis. Sertoli cells are nongerminal cells within the walls of the seminiferous tubules.
They assist spermatogenesis in several ways, such as helping to convert spermatids into spermatozoa. A primary spermatocyte is diploid. At
the end of the first meiotic division, homologous chromosomes have separated, and two haploid secondary spermatocytes form. The
second meiotic division separates the sister chromatids and results in the formation of four haploid spermatids.

Centriole Body
Acrosome

Mitochondrion

Nucleus
Head
Flagellum

Tail
(b)
(a) (b)

FIGURE 59.12
Human sperm. (a) A scanning electron micrograph. (b) A diagram of the main components of a sperm cell.

Chapter 59 Sex and Reproduction 1203


Male Accessory Sex Organs
After the sperm are produced within the seminiferous
tubules, they are delivered into a long, coiled tube called
the epididymis (figure 59.13). The sperm are not motile Epididymis
when they arrive in the epididymis, and they must remain
there for at least 18 hours before their motility develops.
From the epididymis, the sperm enter another long tube,
the vas deferens, which passes into the abdominal cavity via
the inguinal canal. Testis
The vas deferens from each testis joins with one of the
ducts from a pair of glands called the seminal vesicles (see
figure 59.10), which produce a fructose-rich fluid. From
this point, the vas deferens continues as the ejaculatory
duct and enters the prostate gland at the base of the urinary
bladder. In humans, the prostate gland is about the size of a Vas
golf ball and is spongy in texture. It contributes about 60% deferens
of the bulk of the semen, the fluid that contains the prod-
ucts of the testes, fluid from the seminal vesicles, and the
products of the prostate gland. Within the prostate gland,
the ejaculatory duct merges with the urethra from the uri-
nary bladder. The urethra carries the semen out of the
body through the tip of the penis. A pair of pea-sized bul-
bourethral glands secrete a fluid that lines the urethra and
lubricates the tip of the penis prior to coitus (sexual inter-
course).
In addition to the urethra, there are two columns of
FIGURE 59.13
erectile tissue, the corpora cavernosa, along the dorsal Photograph of the human testis. The dark, round object in the
side of the penis and one column, the corpus spongiosum, center of the photograph is a testis, within which sperm are
along the ventral side (figure 59.14). Penile erection is formed. Cupped around it is the epididymis, a highly coiled
produced by neurons in the parasympathetic division of passageway in which sperm complete their maturation. Mature
the autonomic nervous system. As a result of the release sperm are stored in the vas deferens, a long tube that extends from
of nitric oxide by these neurons, arterioles in the penis di- the epididymis.
late, causing the erectile tissue to become engorged with
blood and turgid. This increased pressure in the erectile
tissue compresses the veins, so blood flows into the penis
but cannot flow out. The drug sildenafil (Viagra) pro-
longs erection by stimulating release of nitric oxide in the Dorsal veins Corpus
spongiosum
penis. Some mammals, such as the walrus, have a bone in
Artery
the penis that contributes to its stiffness during erection, Urethra
but humans do not.
The result of erection and continued sexual stimulation
is ejaculation, the ejection from the penis of about 5 milli-
liters of semen containing an average of 300 million sperm.
Successful fertilization requires such a high sperm count
because the odds against any one sperm cell successfully
Deep
completing the journey to the egg and fertilizing it are ex- artery
traordinarily high, and the acrosomes of several sperm need
to interact with the egg before a single sperm can penetrate
the egg. Males with fewer than 20 million sperm per milli- Corpora
liter are generally considered sterile. Despite their large cavernosa
numbers, sperm constitute only about 1% of the volume of
the semen ejaculated.
FIGURE 59.14
A penis in cross-section (left) and longitudinal section (right).
Note that the urethra runs through the corpus spongiosum.

1204 Part XIV Regulating the Animal Body


Table 59.1 Mammalian Reproductive Hormones

MALE
Follicle-stimulating hormone (FSH) Stimulates spermatogenesis
Luteinizing hormone (LH) Stimulates secretion of testosterone by Leydig cells
Testosterone Stimulates development and maintenance of male secondary sexual characteristics and accessory
sex organs

FEMALE
Follicle-stimulating hormone (FSH) Stimulates growth of ovarian follicles and secretion of estradiol
Luteinizing hormone (LH) Stimulates ovulation, conversion of ovarian follicles into corpus luteum, and secretion of
estradiol and progesterone by corpus luteum
Estradiol Stimulates development and maintenance of female secondary sexual characteristics;
prompts monthly preparation of uterus for pregnancy
Progesterone Completes preparation of uterus for pregnancy; helps maintain female secondary sexual
characteristics
Oxytocin Stimulates contraction of uterus and milk-ejection reflex
Prolactin Stimulates milk production

Hormonal Control of Male Reproduction Hypothalamus


Inhibition –
As we saw in chapter 56, the anterior pituitary gland se-
cretes two gonadotropic hormones: FSH and LH. Al- GnRH
though these hormones are named for their actions in the
female, they are also involved in regulating male reproduc-
tive function (table 59.1). In males, FSH stimulates the Ser- Anterior
toli cells to facilitate sperm development, and LH stimu- pituitary
lates the Leydig cells to secrete testosterone. gland
The principle of negative feedback inhibition discussed in
chapter 56 applies to the control of FSH and LH secretion
(figure 59.15). The hypothalamic hormone, gonadotropin-
releasing hormone (GnRH), stimulates the anterior pituitary Inhibition – Inhibition –
gland to secrete both FSH and LH. FSH causes the Sertoli
cells to release a peptide hormone called inhibin that specifi- LH FSH
cally inhibits FSH secretion. Similarly, LH stimulates testos-
terone secretion, and testosterone feeds back to inhibit the
Testosterone Inhibin
release of LH, both directly at the anterior pituitary gland
and indirectly by reducing GnRH release. The importance
of negative feedback inhibition can be demonstrated by re- Leydig Sertoli
cells cells
moving the testes; in the absence of testosterone and inhibin,
the secretion of FSH and LH from the anterior pituitary is Maintains
greatly increased. secondary Spermatogenesis
Testes
sex characteristics
An adult male produces sperm continuously by meiotic
division of the germinal cells lining the seminiferous FIGURE 59.15
tubules. Semen consists of sperm from the testes and Hormonal interactions between the testes and anterior
fluid contributed by the seminal vesicles and prostate pituitary. LH stimulates the Leydig cells to secrete testosterone,
gland. Production of sperm and secretion of and FSH stimulates the Sertoli cells of the seminiferous tubules to
testosterone from the testes are controlled by FSH and secrete inhibin. Testosterone and inhibin, in turn, exert negative
LH from the anterior pituitary. feedback inhibition on the secretion of LH and FSH, respectively.

Chapter 59 Sex and Reproduction 1205


Structure and Function of the maintain the female accessory sex organs: the fallopian
tubes, uterus, and vagina.
Female Reproductive System
The structures of the reproductive system in a human fe- Female Accessory Sex Organs
male are shown in figure 59.16. In contrast to the testes,
the ovaries develop much more slowly. In the absence of The fallopian tubes (also called uterine tubes or oviducts)
testosterone, the female embryo develops a clitoris and transport ova from the ovaries to the uterus. In humans,
labia majora from the same embryonic structures that the uterus is a muscular, pear-shaped organ that narrows to
produce a penis and scrotum in males. Thus clitoris and form a neck, the cervix, which leads to the vagina (figure
penis, and the labia majora and scrotum, are said to be 59.17a). The uterus is lined with a simple columnar epithe-
homologous structures. The clitoris, like the penis, contains lial membrane called the endometrium. The surface of the
corpora cavernosa and is therefore erectile. The ovaries endometrium is shed during menstruation, while the un-
contain microscopic structures called ovarian follicles, derlying portion remains to generate a new surface during
which each contain an egg cell and smaller granulosa the next cycle.
cells. The ovarian follicles are the functional units of the Mammals other than primates have more complex fe-
ovary. male reproductive tracts, where part of the uterus divides to
At puberty, the granulosa cells begin to secrete the form uterine “horns,” each of which leads to an oviduct
major female sex hormone estradiol (also called estrogen), (figure 59.17b, c). In cats, dogs, and cows, for example,
triggering menarche, the onset of menstrual cycling. there is one cervix but two uterine horns separated by a
Estradiol also stimulates the formation of the female sec- septum, or wall. Marsupials, such as opossums, carry the
ondary sexual characteristics, including breast develop- split even further, with two unconnected uterine horns, two
ment and the production of pubic hair. In addition, estra- cervices, and two vaginas. A male marsupial has a forked
diol and another steroid hormone, progesterone, help to penis that can enter both vaginas simultaneously.

Fallopian tube

Ovary

Uterus

Bladder Cervix

Rectum
Clitoris

Urethra

Vagina

FIGURE 59.16
Organization of the human female reproductive system. The ovaries are the gonads, the fallopian tubes receive the ovulated ova, and
the uterus is the womb, the site of development of an embryo if the egg cell becomes fertilized.

1206 Part XIV Regulating the Animal Body


Oviducts
Ovary Ovary

Uterine horns Uterine horns


Oviduct

Uterus
Ovary
Cervix

Vagina Cervices
Cervix

Vagina Vagina

FIGURE 59.17
A comparison of mammalian uteruses. (a) Humans and other primates; (b) cats, dogs, and cows; and (c) rats, mice, and rabbits.

Menstrual and Estrous Cycles


At birth, a female’s ovaries contain some 2 million follicles,
each with an ovum that has begun meiosis but which is ar-
rested in prophase of the first meiotic division. At this
stage, the ova are called primary oocytes. Some of these
primary-oocyte-containing follicles are stimulated to de-
velop during each cycle. The human menstrual (Latin mens,
“month”) cycle lasts approximately one month (28 days on
the average) and can be divided in terms of ovarian activity
into a follicular phase and luteal phase, with the two phases
separated by the event of ovulation.

Follicular Phase
Secondary
During the follicular phase, a few follicles are stimulated to oocyte
grow under FSH stimulation, but only one achieves full
maturity as a tertiary, or Graafian, follicle (figure 59.18).
This follicle forms a thin-walled blister on the surface of
the ovary. The primary oocyte within the Graafian follicle
completes the first meiotic division during the follicular Granulosa
cells
phase. Instead of forming two equally large daughter cells,
however, it produces one large daughter cell, the secondary
oocyte, and one tiny daughter cell, called a polar body.
Thus, the secondary oocyte acquires almost all of the cyto-
plasm from the primary oocyte, increasing its chances of
sustaining the early embryo should the oocyte be fertilized. FIGURE 59.18
The polar body, on the other hand, often disintegrates. A mature Graafian follicle in a cat ovary (50). Note the ring
The secondary oocyte then begins the second meiotic divi- of granulosa cells that surrounds the secondary oocyte. This ring
sion, but its progress is arrested at metaphase II. It is in this will remain around the egg cell when it is ovulated, and sperm
form that the egg cell is discharged from the ovary at ovu- must tunnel through the ring in order to reach the plasma
lation, and it does not complete the second meiotic division membrane of the egg cell.
unless it becomes fertilized in the fallopian tube.

Chapter 59 Sex and Reproduction 1207


FIGURE 59.19 Primary follicles
The meiotic events
Fallopian tube
of oogenesis in Germinal cell
humans. A primary (diploid)
oocyte is diploid. At Fertilization Developing
the completion of the follicle
first meiotic division,
one division product Primary
is eliminated as a oocyte
polar body, while the (diploid) Mature follicle
with secondary
other, the secondary MEIOSIS I oocyte
oocyte, is released
First polar body Secondary Ruptured
during ovulation. The follicle
secondary oocyte does oocyte
(haploid) Corpus luteum
not complete the MEIOSIS II
second meiotic
division until after
fertilization; that Ovum
(haploid)
division yields a Second
second polar body and polar
body
a single haploid egg,
or ovum. Fusion of
the haploid egg with a
haploid sperm during
fertilization produces
a diploid zygote.

Ovulation Fallopian tube


The increasing level of estradiol in the Uterus
blood during the follicular phase stimu- First mitosis
lates the anterior pituitary gland to se- Cleavage
crete LH about midcycle. This sudden
secretion of LH causes the fully devel- Morula
oped Graafian follicle to burst in the
process of ovulation, releasing its sec- Blastocyst
ondary oocyte. The released oocyte en- Fertilization
ters the abdominal cavity near the fim-
briae, the feathery projections Developing follicles
surrounding the opening to the fallopian
tube. The ciliated epithelial cells lining Implantation
the fallopian tube propel the oocyte
through the fallopian tube toward the Corpus
luteum
uterus. If it is not fertilized, the oocyte
Ovary
will disintegrate within a day following
ovulation. If it is fertilized, the stimulus Fimbria Ovulation
of fertilization allows it to complete the
second meiotic division, forming a fully
mature ovum and a second polar body. FIGURE 59.20
Fusion of the two nuclei from the ovum The journey of an egg. Produced within a follicle and released at ovulation, an egg is
and the sperm produces a diploid zygote swept into a fallopian tube and carried along by waves of ciliary motion in the tube walls.
(figure 59.19). Fertilization normally oc- Sperm journeying upward from the vagina fertilize the egg within the fallopian tube. The
curs in the upper one-third of the fallop- resulting zygote undergoes several mitotic divisions while still in the tube, so that by the
time it enters the uterus, it is a hollow sphere of cells called a blastocyst. The blastocyst
ian tube, and in a human the zygote
implants within the wall of the uterus, where it continues its development. (The egg and its
takes approximately three days to reach
subsequent stages have been enlarged for clarification.)
the uterus, then another two to three
days to implant in the endometrium (fig-
ure 59.20).
1208 Part XIV Regulating the Animal Body
Luteal Phase
After ovulation, LH stimulates the empty Graafian follicle
to develop into a structure called the corpus luteum (Latin, Levels of
“yellow body”). For this reason, the second half of the gonadotropic LH
hormones in blood Pituitary
menstrual cycle is referred to as the luteal phase of the gland
cycle. The corpus luteum secretes both estradiol and an-
FSH
other steroid hormone, progesterone. The high blood lev-
els of estradiol and progesterone during the luteal phase FSH
now exert negative feedback inhibition of FSH and LH se-
cretion by the anterior pituitary gland. This inhibition dur-
ing the luteal phase is in contrast to the stimulation exerted
by estradiol on LH secretion at midcycle, which caused 0 7 14 21 28 days
ovulation. The inhibitory effect of estradiol and proges- Ovarian cycle
terone on FSH and LH secretion after ovulation acts as a
natural contraceptive mechanism, preventing both the de-
velopment of additional follicles and continued ovulation.
Luteal
During the follicular phase the granulosa cells secrete in- Developing follicles Ovulation Corpus luteum regression
creasing amounts of estradiol, which stimulates the growth
Follicular phase Luteal phase
of the endometrium. Hence, this portion of the cycle is also
referred to as the proliferative phase of the endometrium.
During the luteal phase of the cycle, the combination of Hormone blood levels
estradiol and progesterone cause the endometrium to be-
come more vascular, glandular, and enriched with glycogen
deposits. Because of the endometrium’s glandular appear-
ance, this portion of the cycle is known as the secretory
phase of the endometrium (figure 59.21). Estradiol
In the absence of fertilization, the corpus luteum triggers
Progesterone
its own atrophy, or regression, toward the end of the luteal
phase. It does this by secreting hormones (estradiol and prog- 0 7 14 21 28 days
esterone) that inhibit the secretion of LH, the hormone Endometrial changes
needed for its survival. In many mammals, atrophy of the cor- during menstrual cycle
pus luteum is assisted by luteolysin, a paracrine regulator be-
lieved to be a prostaglandin. The disappearance of the corpus
luteum results in an abrupt decline in the blood concentra-
tion of estradiol and progesterone at the end of the luteal
phase, causing the built-up endometrium to be sloughed off
with accompanying bleeding. This process is called menstru- Menstrual Proliferative Ovulation Secretory Menstrual
ation, and the portion of the cycle in which it occurs is known phase phase phase phase
as the menstrual phase of the endometrium. 0 7 14 21 28 days
If the ovulated oocyte is fertilized, however, regression of
the corpus luteum and subsequent menstruation is averted
FIGURE 59.21
by the tiny embryo! It does this by secreting human chori-
The human menstrual cycle. The growth and thickening of the
onic gonadotropin (hCG), an LH-like hormone produced endometrial (uterine) lining is stimulated by estradiol and
by the chorionic membrane of the embryo. By maintaining progesterone. The decline in the levels of these two hormones
the corpus luteum, hCG keeps the levels of estradiol and triggers menstruation, the sloughing off of built-up endometrial
progesterone high and thereby prevents menstruation, tissue.
which would terminate the pregnancy. Because hCG comes
from the embryonic chorion and not the mother, it is the
hormone that is tested for in all pregnancy tests. The ovarian follicles develop under FSH stimulation,
Menstruation is absent in mammals with an estrous and one follicle ovulates under LH stimulation. During
cycle. Although such mammals do cyclically shed cells from the follicular and luteal phases, the hormones secreted
the endometrium, they don’t bleed in the process. The es- by the ovaries stimulate the development of the
trous cycle is divided into four phases: proestrus, estrus, endometrium, so an embryo can implant there if
metestrus, and diestrus, which correspond to the prolifera- fertilization has occurred. A secondary oocyte is
released from an ovary at ovulation, and it only
tive, mid-cycle, secretory, and menstrual phases of the en-
completes meiosis if it is fertilized.
dometrium in the menstrual cycle.

Chapter 59 Sex and Reproduction 1209


59.4 The physiology of human sexual intercourse is becoming better known.

Physiology of Human Sexual becomes very sensitive and withdraws up into a sheath or
“hood.” Once it has withdrawn, the clitoris is stimulated
Intercourse indirectly when the thrusting movements of the penis rub
Few physical activities are more pleasurable to humans the clitoral hood against the clitoris. The nervous stimula-
than sexual intercourse. The sex drive is one of the tion produced by the repeated movements of the penis
strongest drives directing human behavior, and as such, it is within the vagina elicits a continuous response in the auto-
circumscribed by many rules and customs. Sexual inter- nomic nervous system, greatly intensifying the physiologi-
course acts as a channel for the strongest of human emo- cal changes initiated during the excitement phase. In the fe-
tions such as love, tenderness, and personal commitment. male, pelvic thrusts may begin, while in the male the penis
Few subjects are at the same time more private and of more reaches its greatest length and rigidity.
general interest. Here we will limit ourselves to a very nar-
row aspect of sexual behavior, its immediate physiological Orgasm
effects. The emotional consequences are no less real, but
they are beyond the scope of this book. The climax of intercourse is reached when the stimulation
Until relatively recently, the physiology of human sexual is sufficient to initiate a series of reflexive muscular con-
activity was largely unknown. Perhaps because of the tractions. The nerve impulses producing these contractions
prevalence of strong social taboos against the open discus- are associated with other activity within the central nervous
sion of sexual matters, no research was carried out on the system, activity that we experience as intense pleasure. In
subject, and detailed information was lacking. Over the past females, the contractions are initiated by impulses in the
40 years, however, investigations by William Masters and hypothalamus, which causes the posterior pituitary gland to
Virginia Johnson, as well as an army of researchers who release large amounts of oxytocin. This hormone, in turn,
followed them, have revealed much about the biological causes the muscles in the uterus and around the vaginal
nature of human sexual activity. opening to contract and the cervix to be pulled upward.
The sexual act is referred to by a variety of names, in- Contractions occur at intervals of about one per second.
cluding sexual intercourse, copulation, and coitus, as well as There may be one to several intense peaks of contractions
a host of informal terms. It is common to partition the (orgasms), or the peaks may be more numerous but less in-
physiological events that accompany intercourse into four tense.
phases—excitement, plateau, orgasm, and resolution— Analogous contractions take place in the male. The first
although there are no clear divisions between these phases. contractions, which occur in the vas deferens and prostate
gland, cause emission, the peristaltic movement of sperm
and seminal fluid into a collecting zone of the urethra lo-
Excitement cated at the base of the penis. Shortly thereafter, violent
The sexual response is initiated by the nervous system. In contractions of the muscles at the base of the penis result in
both males and females, commands from the brain increase ejaculation of the collected semen through the penis. As in
the respiratory rate, heart rate, and blood pressure. The the female, the contractions are spaced about one second
nipples commonly harden and become more sensitive. apart, although in the male they continue for only a few
Other changes increase the diameter of blood vessels, lead- seconds and are almost invariably restricted to a single in-
ing to increased circulation. In some people, these changes tense wave.
may produce a reddening of the skin around the face,
breasts, and genitals (the sex flush). Increased circulation Resolution
also leads to vasocongestion, producing erection of the
male’s penis and similar swelling of the female’s clitoris. After ejaculation, males rapidly lose their erection and
The female experiences changes that prepare the vagina for enter a refractory period lasting 20 minutes or longer, in
sexual intercourse: the labia majora and labia minora, lips which sexual arousal is difficult to achieve and ejaculation is
of tissue that cover the opening to the vagina, swell and almost impossible. By contrast, many women can be
separate due to the increased circulation; the vaginal walls aroused again almost immediately. After intercourse, the
become moist; and the muscles encasing the vagina relax. bodies of both men and women return over a period of sev-
eral minutes to their normal physiological state.

Plateau Sexual intercourse is a physiological series of events


The penetration of the vagina by the thrusting penis con- leading to the ultimate deposition of sperm within the
female reproductive tract. The phases are similar in
tinuously stimulates nerve endings both in the tip of the
males and females.
penis and in the clitoris. The clitoris, which is now swollen,

1210 Part XIV Regulating the Animal Body


Birth Control
In most vertebrates, copulation is associated
solely with reproduction. Reflexive behavior
that is deeply ingrained in the female limits
sexual receptivity to those periods of the sex-
ual cycle when she is fertile. In humans and a
few species of apes, the female can be sexually
receptive throughout her reproductive cycle,
and this extended receptivity to sexual inter- (a) (b)
course serves a second important function—it
reinforces pair-bonding, the emotional rela-
tionship between two individuals living to-
gether.
Not all human couples want to initiate a
pregnancy every time they have sexual inter-
course, yet sexual intercourse may be a nec-
essary and important part of their emotional
lives together. The solution to this dilemma
is to find a way to avoid reproduction with-
out avoiding sexual intercourse; this ap- (c) (d)
proach is commonly called birth control or FIGURE 59.22
contraception. A variety of approaches dif- Four common methods of birth control. (a) Condom; (b) diaphragm and
fering in effectiveness and in their accept- spermicidal jelly; (c) oral contraceptives; (d) Depo-Provera.
ability to different couples are commonly
taken to achieve birth control (figure 59.22
and table 59.2).
the dimensions of individual cervices vary, a cervical cap
or diaphragm must be fitted by a physician. Failure rates
Abstinence average 4 to 25% for diaphragms, perhaps because of the
The simplest and most reliable way to avoid pregnancy is propensity to insert them carelessly when in a hurry. Fail-
not to have sexual intercourse at all. Of all methods of birth ure rates for cervical caps are somewhat lower.
control, this is the most certain. It is also the most limiting,
because it denies a couple the emotional support of a sexual
Sperm Destruction
relationship.
A third general approach to birth control is to eliminate the
sperm after ejaculation. This can be achieved in principle
Sperm Blockage by washing out the vagina immediately after intercourse,
If sperm cannot reach the uterus, fertilization cannot before the sperm have a chance to enter the uterus. Such a
occur. One way to prevent the delivery of sperm is to en- procedure is called a douche (French, “wash”). The douche
case the penis within a thin sheath, or condom. Many method is difficult to apply well, because it involves a rapid
males do not favor the use of condoms, which tend to de- dash to the bathroom immediately after ejaculation and a
crease their sensory pleasure during intercourse. In prin- very thorough washing. Its failure rate is as high as 40%.
ciple, this method is easy to apply and foolproof, but in Alternatively, sperm delivered to the vagina can be de-
practice it has a failure rate of 3 to 15% because of incor- stroyed there with spermicidal jellies or foams. These treat-
rect use or inconsistent use. Nevertheless, it is the most ments generally require application immediately before in-
commonly employed form of birth control in the United tercourse. Their failure rates vary from 10 to 25%. The use
States. Condoms are also widely used to prevent the of a spermicide with a condom increases the effectiveness
transmission of AIDS and other sexually transmitted dis- over each method used independently.
eases (STDs). Over a billion condoms were sold in the
United States last year.
Prevention of Ovulation
A second way to prevent the entry of sperm into the
uterus is to place a cover over the cervix. The cover may Since about 1960, a widespread form of birth control in the
be a relatively tight-fitting cervical cap, which is worn for United States has been the daily ingestion of birth control
days at a time, or a rubber dome called a diaphragm, pills, or oral contraceptives, by women. These pills contain
which is inserted immediately before intercourse. Because analogues of progesterone, sometimes in combination with

Chapter 59 Sex and Reproduction 1211


Table 59.2 Methods of Birth Control
Failure
Device Action Rate* Advantages Disadvantages

Oral Hormones (progesterone 1–5, Convenient; highly effective; Must be taken regularly;
contraceptive analogue alone or in depending provides significant possible minor side effects which
combination with other on type noncontraceptive health new formulations have
hormones) primarily prevent benefits, such as protection reduced; not for women with
ovulation against ovarian and endometrial cardiovascular risks (mostly
cancers smokers over age 35)
Condom Thin sheath for penis that 3–15 Easy to use; effective; Requires male cooperation; may
collects semen; “female inexpensive; protects against diminish spontaneity; may
condoms” sheath vaginal walls some sexually transmitted deteriorate on the shelf
diseases
Diaphragm Soft rubber cup covers 4–25 No dangerous side effects; Requires careful fitting; some
entrance to uterus, prevents reliable if used properly; inconvenience associated with
sperm from reaching egg, provides some protection insertion and removal; may be
holds spermicide against sexually transmitted dislodged during intercourse
diseases and cervical cancer
Intrauterine Small plastic or metal device 1–5 Convenient; highly effective; Can cause excess menstrual
device (IUD) placed in the uterus; infrequent replacement bleeding and pain; risk of
prevents implantation; perforation, infection, expulsion,
some contain copper, pelvic inflammatory disease, and
others release hormones infertility; not recommended for
those who eventually intend to
conceive or are not monogamous;
dangerous in pregnancy
Cervical cap Miniature diaphragm covers Probably No dangerous side effects; fairly Problems with fitting and
cervix closely, prevents sperm similar to effective; can remain in place insertion; comes in limited
from reaching egg, holds that of longer than diaphragm number of sizes
spermicide diaphragm
Foams, creams, Chemical spermicides 10–25 Can be used by anyone who Relatively unreliable; sometimes
jellies, vaginal inserted in vagina before is not allergic; protect against messy; must be used 5–10 minutes
suppositories intercourse that prevent some sexually transmitted before each act of intercourse
sperm from entering uterus diseases; no known side effects
Implant Capsules surgically implanted .03 Very safe, convenient, and Irregular or absent periods;
(levonorgestrel; under skin slowly release effective; very long-lasting minor surgical procedure needed
Norplant) hormone that blocks (5 years); may have for insertion and removal; some
ovulation nonreproductive health benefits scarring may occur
like those of oral contraceptives

Injectable Injection every 3 months of 1 Convenient and highly Animal studies suggest it may
contraceptive a hormone that is slowly effective; no serious side effects cause cancer, though new studies
(medroxy- released and prevents other than occasional heavy in humans are mostly encouraging;
progesterone; ovulation menstrual bleeding occasional heavy menstrual
Depo-Provera) bleeding

*Failure rate is expressed as pregnancies per 100 actual users per year.
Source: Data from American College of Obstetricians and Gynecologists: Contraception, Patient Education Pamphlet No. AP005.ACOG, Washington,
D.C., 1990.

estrogens. As described earlier, progesterone and estradiol pills for three weeks; during the fourth week, she takes pills
act by negative feedback to inhibit the secretion of FSH without hormones (placebos), allowing the levels of those
and LH during the luteal phase of the menstrual cycle, hormones in her blood to fall, which causes menstruation.
thereby preventing follicle development and ovulation. Oral contraceptives provide a very effective means of birth
They also cause a buildup of the endometrium. The hor- control, with a failure rate of only 1 to 5%. In a variation of
mones in birth control pills have the same effects. Because the oral contraceptive, hormone-containing capsules are
the pills block ovulation, no ovum is available to be fertil- implanted beneath the skin. These implanted capsules have
ized. A woman generally takes the hormone-containing failure rates below 1%.

1212 Part XIV Regulating the Animal Body


Vas deferens within
spermatic cord

Vas deferens Ovary


cut and tied

(a)

Fallopian tube Uterus


cut and tied

FIGURE 59.23
Birth control through sterilization. (a)
(b)
Vasectomy; (b) tubal ligation.

A small number of women using birth control pills or Another method of preventing embryo implantation is
implants experience undesirable side effects, such as blood the “morning after pill,” which contains 50 times the dose
clotting and nausea. These side effects have been reduced of estrogen present in birth control pills. The pill works by
in newer generations of birth control pills, which contain temporarily stopping ovum development, by preventing
less estrogen and different analogues of progesterone. fertilization, or by stopping the implantation of a fertilized
Moreover, these new oral contraceptives provide a number ovum. Its failure rate is 1 to 10%, but many women are un-
of benefits, including reduced risks of endometrial and easy about taking such high hormone doses, as side effects
ovarian cancer, cardiovascular disease, and osteoporosis (for can be severe. This is not recommended as a regular
older women). However, they may increase the risk of con- method of birth control but rather as a method of emer-
tracting breast cancer and cervical cancer. The risks in- gency contraception.
volved with birth control pills increase in women who
smoke and increase greatly in women over 35 who smoke.
Sterilization
The current consensus is that, for many women, the health
benefits of oral contraceptives outweigh their risks, al- A completely effective means of birth control is steriliza-
though a physician must help each woman determine the tion, the surgical removal of portions of the tubes that
relative risks and benefits. transport the gametes from the gonads (figure 59.23). Ster-
ilization may be performed on either males or females, pre-
venting sperm from entering the semen in males and pre-
Prevention of Embryo Implantation venting an ovulated oocyte from reaching the uterus in
females. In males, sterilization involves a vasectomy, the re-
The insertion of a coil or other irregularly shaped object
moval of a portion of the vas deferens from each testis. In
into the uterus is an effective means of birth control, be-
females, the comparable operation involves the removal of
cause the irritation it produces in the uterus prevents the
a section of each fallopian tube.
implantation of an embryo within the uterine wall. Such in-
trauterine devices (IUDs) have a failure rate of only 1 to
5%. Their high degree of effectiveness probably reflects Fertilization can be prevented by a variety of birth
their convenience; once they are inserted, they can be for- control methods, including barrier contraceptives,
gotten. The great disadvantage of this method is that al- hormonal inhibition, surgery, and abstinence. Efficacy
most a third of the women who attempt to use IUDs expe- rates vary from method to method.
rience cramps, pain, and sometimes bleeding and therefore
must discontinue using them.

Chapter 59 Sex and Reproduction 1213


Chapter 59 www.mhhe.com/raven6e www.biocourse.com

Summary Questions Media Resources


59.1 Animals employ both sexual and asexual reproductive strategies.

• Parthenogenesis is a form of asexual reproduction 1. How are oviparity, • Introduction to


ovoviviparity, and viviparity reproduction
that is practiced by many insects and some lizards.
different?
• Among mammals, the sex is determined by the
presence of a Y chromosome in males and its absence
in females.

59.2 The evolution of reproduction among the vertebrates has led to


internalization of fertilization and development.

• Most bony fish practice external fertilization, 2. How does fetal development • On Science articles:
releasing eggs and sperm into the water where differ in the monotremes, Interactions
marsupials, and placental • Student Research:
fertilization occurs. Amphibians have external Reproductive biology
fertilization and the young go through a larval stage mammals?
of house mice
before metamorphosis. Evolution of uterine
function
• Reptiles and birds are oviparous, the young
developing in eggs that are deposited externally. Most
mammals are viviparous, the young developing within
the mother.

59.3 Male and female reproductive systems are specialized for different functions.

• Sperm leave the testes and pass through the 3. Briefly describe the function • Art Activities:
epididymis and vas deferens; the ejaculatory duct of seminal vesicles, prostate Sperm and egg
gland, and bulbourethral glands. anatomy
merges with the urethra, which empties at the tip of Male reproductive
the penis. 4. When do the ova in a female system
• An egg cell released from the ovary in ovulation is mammal begin meiosis? When Penis anatomy
do they complete the first Female reproductive
drawn by fimbria into the fallopian tube, which system
meiotic division?
conducts the egg cell to the lining of the uterus, or Breast anatomy
endometrium, where it implants if fertilized. 5. What hormone is secreted by
the granulosa cells in a Graafian • Spermatogenesis
• If fertilization does not occur, the corpus luteum follicle? What effect does this • Menstruation
regresses at the end of the cycle and the resulting fall hormone have on the • Female reproductive
in estradiol and progesterone secretion cause endometrium? cycle
• Oogenesis
menstruation to occur in humans and apes.

59.4 The physiology of human sexual intercourse is becoming better known.

• The physiological events that occur in the human 6. What are the four phases in • Penile erection
sexual response are grouped into four phases: the physiological events of sexual • Vasectomy
intercourse in humans? During • Tubal ligation
excitement, plateau, orgasm, and resolution.
the first phase, what events occur
• Males and females have similar phases, but males specifically in males, and what
enter a refractory period following orgasm that is events occur specifically in
absent in many women. females?
• There are a variety of methods of birth control 7. How do birth control pills
available that range in ease of use, effectiveness, and prevent pregnancy?
permanence.

1214 Part XIV Regulating the Animal Body


60
Vertebrate
Development

Concept Outline
60.1 Fertilization is the initial event in development.
Stages of Development. Fertilization of an egg cell by a
sperm occurs in three stages: penetration, activation of the
egg cell, and fusion of the two haploid nuclei.

60.2 Cell cleavage and the formation of a blastula set


the stage for later development.
Cell Cleavage Patterns. The cytoplasm of the zygote is
divided into smaller cells by a mitotic cell division in a
process called cleavage.

60.3 Gastrulation forms the three germ layers of the


embryo.
The Process of Gastrulation. Cells of the blastula
invaginate and involute to produce an outer ectoderm, an
inner endoderm layer, and a third layer, the mesoderm.

60.4 Body architecture is determined during the next


stages of embryonic development. FIGURE 60.1
Developmental Processes during Neurulation. The Development is the process that determines an organism’s
mesoderm of chordates forms a notochord, and the form and function. A human fetus at 18 weeks is not yet
overlying ectoderm rolls to produce a neural tube. halfway through the 38 weeks—about 9 months—it will spend
How Cells Communicate during Development. Cell- within its mother, but it has already developed many distinct
to-cell contact plays a major role in selecting the paths behaviors, such as the sucking reflex that is so important to
along which cells develop. survival after birth.
Embryonic Development and Vertebrate Evolution.
The embryonic development of a mammal includes stages
that are characteristic of more primitive vertebrates.
Extraembryonic Membranes. Embryonic cells form
several membranes outside of the embryo that provide
R eproduction in all but a few vertebrates unites two
haploid gametes to form a single diploid cell called a
zygote. The zygote develops by a process of cell division
protection, nourishment, and gas exchange for the embryo. and differentiation into a complex multicellular organism,
composed of many different tissues and organs (figure
60.5 Human development is divided into trimesters. 60.1). Although the process of development is a continuous
First Trimester. A blastocyst implants into the mother’s series of events with some of the details varying among dif-
endometrium, and the formation of body organs begins ferent vertebrate groups, we will examine vertebrate devel-
during the third and fourth week. opment in six stages. In this chapter, we will consider the
Second and Third Trimesters. All of the major organs stages of development and conclude with a description of
of the body form during the first trimester and so further the events that occur during human development.
growth and development take place during this time.
Birth and Postnatal Development. Birth occurs as a
result of uterine contractions; the human brain continues to
grow significantly after birth.
1215
60.1 Fertilization is the initial event in development.

Stages of Development protein layer called the zona pellucida. The head of each
sperm is capped by an organelle called the acrosome, which
In vertebrates, as in all sexual animals, the first step in de- contains glycoprotein-digesting enzymes. These enzymes
velopment is the union of male and female gametes, a become exposed as the sperm begin to work their way into
process called fertilization. Fertilization is typically ex- the layer of granulosa cells, and the activity of the enzymes
ternal in fish and amphibians, which reproduce in water, enables the sperm to tunnel their way through the zona
and internal in all other vertebrates. In internal fertiliza- pellucida to the egg’s plasma membrane. In sea urchins,
tion, small, motile sperm are introduced into the female egg cytoplasm bulges out at this point, engulfing the head
reproductive tract during mating. The sperm swim up the of the sperm and permitting the sperm nucleus to enter the
reproductive tract until they encounter a mature egg or cytoplasm of the egg (figure 60.3).
oocyte in an oviduct, where fertilization occurs. Fertiliza-
tion consists of three stages: penetration, activation, and
Activation
fusion.
The series of events initiated by sperm penetration are col-
lectively called egg activation. In some frogs, reptiles, and
Penetration
birds, more than one sperm may penetrate the egg, but
As described in chapter 59, the secondary oocyte is released only one is successful in fertilizing it. In mammals, by con-
from a fully developed Graafian follicle at ovulation. It is trast, the penetration of the first sperm initiates changes in
surrounded by the same layer of small granulosa cells that the egg membrane that prevent the entry of other sperm.
surrounded it within the follicle (figure 60.2). Between the As the sperm makes contact with the oocyte membrane,
granulosa cells and the egg’s plasma membrane is a glyco- there is a change in the membrane potential (see chapter 54

First polar
Cytoplasm body
of oocyte Granulosa cell

Second meiotic
spindle

Zona pellucida

(b)

Oocyte Granulosa cells

Plasma membrane
of oocyte

(a)

FIGURE 60.2
Mammalian reproductive cells. (a) A sperm must penetrate a layer of
granulosa cells and then a layer of glycoprotein called the zona pellucida,
before it reaches the oocyte membrane. This penetration is aided by
digestive enzymes in the acrosome of the sperm. These scanning electron
micrographs show (b) a human oocyte (90×) surrounded by numerous
granulosa cells, and (c) a human sperm on an egg (3000×). (c)

1216 Part XIV Regulating the Animal Body


Sperm
nucleus

Jelly
coat
Acrosome Cortical granule
secreting contents
into perivitelline
space

Vitelline
membrane
Acrosomal
process Sperm nucleus
Egg plasma
membrane

Altered vitelline
membrane prevents
further sperm
penetration

(a) (b)

FIGURE 60.3
Sperm penetration of a sea urchin egg. (a) The stages of penetration. (b) An electron micrograph (50,000×) of penetration. Penetration
in both invertebrate and vertebrate eggs is similar.

for discussion of membrane potential) that prevents other


sperm from fusing with the oocyte membrane. In addition
to these changes, sperm penetration can have three other Sperm
effects on the egg. First, in mammals it stimulates the chro- Gray
mosomes in the egg nucleus to complete the second mei- crescent
otic division, producing two egg nuclei. One of these nuclei
is extruded from the egg as a second polar body (see chap-
ter 59), leaving a single haploid egg nucleus within the egg. Movement of pigment
Second, sperm penetration in some animals triggers opposite sperm entry
movements of the egg cytoplasm around the point of sperm
FIGURE 60.4
entry. These movements ultimately establish the bilateral
Gray crescent formation in frog eggs. The gray crescent
symmetry of the developing animal. In frogs, for example,
appears opposite the point of penetration by the sperm.
sperm penetration causes an outer pigmented cap of egg
cytoplasm to rotate toward the point of entry, uncovering a
gray crescent of interior cytoplasm opposite the point of
penetration (figure 60.4). The position of the gray crescent
Nuclei Fusion
determines the orientation of the first cell division. A line
drawn between the point of sperm entry and the gray cres- The third stage of fertilization is the fusion of the entering
cent would bisect the right and left halves of the future sperm nucleus with the haploid egg nucleus to form the
adult. Third, activation is characterized by a sharp increase diploid nucleus of the zygote. This fusion is triggered by
in protein synthesis and an increase in metabolic activity in the activation of the egg. If a sperm nucleus is microin-
general. Experiments demonstrate that the protein synthe- jected into an egg without activating the egg, the two nu-
sis in the activated oocyte is coded by mRNA that was pre- clei will not fuse. The nature of the signals that are ex-
viously produced and already present in the cytoplasm of changed between the two nuclei, or sent from one to the
the unfertilized egg cell. other, is not known.
In some vertebrates, it is possible to activate an egg
without the entry of a sperm, simply by pricking the egg The three stages of fertilization are penetration,
membrane. An egg that is activated in this way may go on activation, and nuclei fusion. Penetration initiates a
to develop parthenogenetically. A few kinds of amphibians, complex series of developmental events, including
major movements of cytoplasm, which eventually lead
fish, and reptiles rely entirely on parthenogenetic repro-
to the fusion of the egg and sperm nuclei.
duction in nature, as we mentioned in chapter 59.
Chapter 60 Vertebrate Development 1217
60.2 Cell cleavage and the formation of a blastula set the stage for later
development.
Cell Cleavage Patterns Nucleus

Following fertilization, the second major event in verte- Yolk


(a) Lancelet
brate reproduction is the rapid division of the zygote into a
larger and larger number of smaller and smaller cells (table
60.1). This period of division, called cleavage, is not ac- Nucleus
companied by an increase in the overall size of the embryo.
The resulting tightly packed mass of about 32 cells is called Yolk
(b) Frog
a morula, and each individual cell in the morula is referred
to as a blastomere. As the blastomeres continue to divide,
they secrete a fluid into the center of the morula. Eventu- Nucleus
ally, a hollow ball of 500 to 2000 cells, the blastula, is
Yolk
formed. The fluid-filled cavity within the blastula is known Air
as the blastocoel. bubble
Shell
The pattern of cleavage division is influenced by the
presence and location of yolk, which is abundant in the
Albumen
eggs of many vertebrates (figure 60.5). As we discussed in
the previous chapter, vertebrates have embraced a variety
of reproductive strategies involving different patterns of (c) Chicken
yolk utilization.
FIGURE 60.5
Yolk distribution in three kinds of eggs. (a) In the lancelet, a
Primitive Chordates primitive chordate, the egg consists of a central nucleus surrounded
by a small amount of yolk. (b) In a frog egg there is much more
When eggs contain little or no yolk, cleavage occurs yolk, and the nucleus is displaced toward one pole. (c) Bird eggs are
throughout the whole egg (figure 60.6). This pattern of complexly organized, with the nucleus just under the surface of a
cleavage, called holoblastic cleavage, was characteristic of large, central yolk.
the ancestors of the vertebrates and is
still seen in groups such as the
lancelets and agnathans. In these ani-
mals, holoblastic cleavage results in the
formation of a symmetrical blastula
composed of cells of approximately
equal size.

Amphibians and Advanced Fish


The eggs of bony fish and frogs con-
tain much more cytoplasmic yolk in
one hemisphere than the other. Be-
cause yolk-rich cells divide much more
slowly than those that have little yolk,
holoblastic cleavage in these eggs re-
sults in a very asymmetrical blastula
(figure 60.7), with large cells contain-
ing a lot of yolk at one pole and a con-
centrated mass of small cells contain-
ing very little yolk at the other. In
these blastulas, the pole that is rich in
yolk is called the vegetal pole, while FIGURE 60.6 FIGURE 60.7
the pole that is relatively poor in yolk Holoblastic cleavage (3000×). In this Dividing frog eggs. The closest cells in
is called the animal pole. type of cleavage, cell division occurs this photo (those near the animal pole)
throughout the entire egg. divide faster and are smaller than those
near the vegetal pole.

1218 Part XIV Regulating the Animal Body


Table 60.1 Stages of Vertebrate Development (Mammal)

Fertilization The haploid male and female gametes fuse to form a diploid
zygote.

Cleavage The zygote rapidly divides into many cells, with no overall
increase in size. These divisions affect future development, since
different cells receive different portions of the egg cytoplasm and,
hence, different regulatory signals.

Ectoderm Gastrulation The cells of the embryo move, forming three primary cell layers:
ectoderm, mesoderm, and endoderm.
Mesoderm

Endoderm

Neural groove

Neurulation In all chordates, the first organ to form is the notochord; second
Notochord
is the dorsal nerve cord.

Neural crest
Neural crest During neurulation, the neural crest is produced as the neural
cell formation tube is formed. The neural crest gives rise to several uniquely
Neural tube
vertebrate structures.
Notochord

Organogenesis Cells from the three primary layers combine in various ways to
produce the organs of the body.

Chapter 60 Vertebrate Development 1219


Reptiles and Birds
The eggs produced by reptiles, birds, and some fish are
composed almost entirely of yolk, with a small amount of
cytoplasm concentrated at one pole. Cleavage in these eggs Yolk
occurs only in the tiny disc of polar cytoplasm, called the
blastodisc, that lies astride the large ball of yolk material.
This type of cleavage pattern is called meroblastic cleavage Cleaving
(figure 60.8). The resulting embryo is not spherical, but embryonic
rather has the form of a thin cap perched on the yolk. cells

Mammals
Mammalian eggs are in many ways similar to the reptilian
eggs from which they evolved, except that they contain
very little yolk. Because cleavage is not impeded by yolk in FIGURE 60.8
mammalian eggs, it is holoblastic, forming a ball of cells Meroblastic cleavage (400×). In this type of cleavage, only a
surrounding a blastocoel. However, an inner cell mass is portion of the egg actively divides to form a mass of cells.
concentrated at one pole (figure 60.9). This inner cell mass
is analogous to the blastodisc of reptiles and birds, and it
goes on to form the developing embryo. The outer sphere
Inner cell Blastocoel
of cells, called a trophoblast, is analogous to the cells that
mass Blastodisc
form the membranes underlying the tough outer shell of
the reptilian egg. These cells have changed during the Yolk
course of mammalian evolution to carry out a very different
function: part of the trophoblast enters the endometrium
(the epithelial lining of the uterus) and contributes to the
placenta, the organ that permits exchanges between the Trophoblast
fetal and maternal bloods. While part of the placenta is
FIGURE 60.9
composed of fetal tissue (the trophoblast), part is composed
The embryos of mammals and birds are more similar than
of the modified endometrial tissue (called the decidua they seem. (a) A mammalian blastula is composed of a sphere of
basalis) of the mother’s uterus. The placenta will be dis- cells, the trophoblast, surrounding a cavity, the blastocoel, and an
cussed in more detail in a later section. inner cell mass. (b) An avian (bird) blastula consists of a cap of
cells, the blastodisc, resting atop a large yolk mass. The blastodisc
will form an upper and a lower layer with a compressed blastocoel
The Blastula in between.
Viewed from the outside, the blastula looks like a simple
ball of cells all resembling each other. In many animals, this
appearance is misleading, as unequal distribution of devel- cells in the body—and probably all of them. Biologists have
opmental signals from the egg produces some degree of long sought to grow these cells, called embryonic stem cells,
mosaic development, so that the cells are already commit- in tissue culture, as such stem cells might in principle be
ted to different developmental paths. In mammals, how- used to produce tissues for human transplant operations.
ever, it appears that all of the blastomeres receive equiva- Injected into a patient, for example, they might be able to
lent sets of signals, and body form is determined by respond to local signals and produce new tissue (see chap-
cell-cell interactions. In a mammalian blastula, called a blas- ter 19). The first success in growing stem cells in culture
tocyst, each cell is in contact with a different set of neigh- was reported in 1998, when researchers isolated cells from
boring cells, and these interactions with neighboring cells the inner cell mass of human blastocysts and successfully
are a major factor influencing the developmental fate of grew them in tissue culture. These stem cells continue to
each cell. This positional information is of particular im- grow and divide in culture indefinitely, unlike ordinary
portance in the orientation of mammalian embryos, setting body cells, which divide only 50 or so times and then die.
up different patterns of development along three embry-
onic axes: anterior-posterior, dorsal-ventral, and proximal-
distal. A series of rapid cell divisions called cleavage
transforms the zygote into a hollow ball of cells, the
For a short period of time, just before they implant in
blastula. The cleavage pattern is influenced by the
the uterus, the cells of the mammalian blastocyst have the
amount of yolk and its distribution in the egg.
power to develop into many of the 210 different types of

1220 Part XIV Regulating the Animal Body


60.3 Gastrulation forms the three germ layers of the embryo.

The Process of Gastrulation Table 60.2 Developmental Fates of the


Primary Cell Layers
The first visible results of cytoplasmic distribution and cell
position within the blastula can be seen immediately after Ectoderm Epidermis, central nervous system, sense
organs, neural crest
the completion of cleavage. Certain groups of cells invagi-
nate (dent inward) and involute (roll inward) from the sur- Mesoderm Skeleton, muscles, blood vessels, heart, gonads
face of the blastula in a carefully orchestrated activity called Endoderm Lining of digestive and respiratory tracts; liver,
gastrulation. The events of gastrulation determine the pancreas
basic developmental pattern of the vertebrate embryo. By
the end of gastrulation, the cells of the embryo have re-
arranged into three primary germ layers: ectoderm,
mesoderm, and endoderm. The cells in each layer have
Gastrulation in Primitive Chordates
very different developmental fates. In general, the ecto-
derm is destined to form the epidermis and neural tissue; In primitive chordates such as lancelets, which develop
the mesoderm gives rise to connective tissue, skeleton, from symmetrical blastulas, gastrulation begins as the sur-
muscle, and vascular elements; and the endoderm forms the face of the blastula invaginates into the blastocoel. About
lining of the gut and its derivatives (table 60.2). half of the blastula’s cells move into the interior of the blas-
How is cell movement during gastrulation brought tula, forming a structure that looks something like an in-
about? Apparently, migrating cells creep over stationary dented tennis ball. Eventually, the inward-moving wall of
cells by means of actin filament contractions that change cells pushes up against the opposite side of the blastula and
the shapes of the migrating cells affecting an invagination then stops moving. The resulting two-layered, cup-shaped
of blastula tissue. Each cell that moves possesses particular embryo is the gastrula (figure 60.10). The hollow structure
cell surface polysaccharides, which adhere to similar poly- resulting from the invagination is called the archenteron,
saccharides on the surfaces of the other moving cells. This and it is the progenitor of the gut. The opening of the
interaction between cell surface molecules enables the mi- archenteron, the future anus, is known as the blastopore.
grating cells to adhere to one another and move as a single This process produces an embryo with two cell layers:
mass (see chapter 7). an outer ectoderm and an inner endoderm. Soon afterward,
Just as the pattern of cleavage divisions in different a third cell layer, the mesoderm, forms between the ecto-
groups of vertebrates depends heavily on the amount and derm and endoderm. In lancelets, the mesoderm forms
distribution of yolk in the egg, so the pattern of gastrula- from pouches that pinch off the endoderm. The appear-
tion among vertebrate groups depends on the shape of the ance of these three primary cell layers sets the stage for all
blastulas produced during cleavage. subsequent tissue and organ differentiation.

Ectoderm
Ectoderm Archenteron
Ectoderm

Endoderm

Endoderm Endoderm Blastopore


(a) (b) (c)

FIGURE 60.10
Gastrulation in a lancelet. In these chordates, the endoderm is formed by invagination of surface cells (a, b). This produces the primitive
gut, or archenteron (c). Mesoderm will later be formed from pouches off the endoderm.

Chapter 60 Vertebrate Development 1221


Gastrulation in Most Aquatic Vertebrates lancelets, the involuting cell layer eventually presses against
the inner surface of the opposite side of the embryo, elimi-
In the blastulas of amphibians and those aquatic vertebrates
nating the blastocoel and producing an archenteron with a
with asymmetrical yolk distribution, the yolk-laden cells of
blastopore. In this case, however, the blastopore is filled
the vegetal pole are fewer and much larger than the yolk-
with yolk-rich cells, forming the yolk plug. The outer layer
free cells of the animal pole. Consequently, gastrulation is
of cells resulting from these movements is the ectoderm,
more complex than it is in the lancelets. First, a layer of
and the inner layer is the endoderm. Other cells that invo-
surface cells invaginates to form a small, crescent-shaped
lute over the dorsal lip and ventral lip (the two lips of the
slit where the blastopore will soon be located. Next, cells
blastopore that are separated by the yolk plug) migrate be-
from the animal pole involute over the dorsal lip of the
tween the ectoderm and endoderm to form the third germ
blastopore (figure 60.11), at the same location as the gray
layer, the mesoderm (figure 60.11).
crescent of the fertilized egg (see figure 60.4). As in the

Animal pole Ectoderm


Ectoderm
Ectoderm Mesoderm
Archenteron Endoderm
Blastocoel
Blastocoel

Dorsal lip
Blastocoel
Yolk plug
Mesoderm Dorsal lip of
blastopore Ventral lip
Vegetal pole

(a) (b) (c)


Neural fold
Neural plate Neural plate

(d) (e)

FIGURE 60.11
Frog gastrulation. (a) A layer of cells from the animal pole moves toward the yolk cells ultimately involuting through the dorsal lip of
the blastopore. (b) Cells in the dorsal lip zone then involute into the hollow interior, or blastocoel, eventually pressing against the far
wall. The three primary tissues (ectoderm, mesoderm, and endoderm) become distinguished. Ectoderm is shown in blue, mesoderm in
red, and endoderm in yellow. (c) The movement of cells in the dorsal lip creates a new internal cavity, the archenteron, which opens to
the outside through the plug of yolk remaining at the point of invagination. (d) The neural plate later forms from ectoderm. (e) This will
next form a neural groove and then a neural tube as the embryo begins the process of neurulation. The cells of the neural ectoderm are
shown in green.

1222 Part XIV Regulating the Animal Body


Gastrulation in Reptiles, Birds, and
Mammals
Blastodisc
In the blastodisc of a bird or reptile and the
inner cell mass of a mammal, the develop-
ing embryo is a small cap of cells rather
Yolk
than a sphere. No yolk separates the two
sides of the embryo, and, as a result, the
lower cell layer is able to differentiate into
endoderm and the upper layer into ecto- (a)
derm without cell movement. Just after
these two primary cell layers form, the Ectoderm
mesoderm arises by invagination and invo-
Blastocoel
lution of cells from the upper layer. The
surface cells begin moving to the midline
where they involute and migrate laterally to
Endoderm
form a mesodermal layer between the ecto-
derm and endoderm. A furrow along the
longitudinal midline marks the site of this
(b)
involution (figures 60.12 and 60.13). This
furrow, analogous to an elongated blasto-
pore, is called the primitive streak. Ectoderm

Gastrulation in lancelets involves the


formation of ectoderm and endoderm Endoderm Primitive streak
by the invagination of the blastula, and
the mesoderm layer forms from Mesoderm
pouches pinched from the endoderm.
In those vertebrates with extensive
amounts of yolk, gastrulation requires (c)
the involution of surface cells into a
FIGURE 60.12
blastopore or primitive streak, and the
Gastrulation in birds. The upper layer of the blastodisc (a) differentiates into
mesoderm is derived from some of
ectoderm, the lower layer into endoderm (b). Among the cells that migrate into the
these involuted cells.
interior through the dorsal primitive streak are future mesodermal cells (c).

Inner cell Formation of Primitive streak


mass Amniotic cavity
extraembryonic
membranes
Ectoderm

Trophoblast Endoderm
Mesoderm

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

FIGURE 60.13
Mammalian gastrulation. (a) The amniotic cavity forms within the inner cell mass and its base. Layers of ectoderm and endoderm
differentiate (b and c) as in the avian blastodisc. (d) A primitive streak develops, through which cells destined to become mesoderm migrate
into the interior, again reminiscent of gastrulation in birds. The trophoblast has now moved further away from the embryo and begins to
play a role in forming the placenta.

Chapter 60 Vertebrate Development 1223


60.4 Body architecture is determined during the next stages of embryonic
development.
Developmental Processes during of the developing notochord, segmented blocks of meso-
derm tissue called somites form; more somites are added
Neurulation as development continues. Ultimately, the somites give rise
During the next step in vertebrate development, the three to the muscles, vertebrae, and connective tissues. The
primary cell layers begin their transformation into the mesoderm in the head region does not separate into dis-
body’s tissues and organs. The process of tissue differentia- crete somites but remains connected as somitomeres and
tion begins with the formation of two morphological fea- form the striated muscles of the face, jaws, and throat.
tures found only in chordates, the notochord and the hol- Some body organs, including the kidneys, adrenal glands,
low dorsal nerve cord. This development of the dorsal and gonads, develop within another strip of mesoderm that
nerve cord is called neurulation. runs alongside the somites. The remainder of the meso-
The notochord is first visible soon after gastrulation is derm moves out and around the endoderm and eventually
complete, forming from mesoderm along the dorsal mid- surrounds it completely. As a result of this movement, the
line of the embryo. It is a flexible rod located along the mesoderm becomes separated into two layers. The outer
dorsal midline in the embryos of all chordates, although its layer is associated with the body wall and the inner layer is
function is replaced by the vertebral column when it devel- associated with the gut. Between these two layers of meso-
ops from mesoderm in the vertebrates. After the notochord derm is the coelom (see chapter 45), which becomes the
has been laid down, a layer of ectodermal cells situated body cavity of the adult.
above the notochord invaginates, forming a long crease, the
neural groove, down the long axis of the embryo. The
The Neural Crest
edges of the neural groove then move toward each other
and fuse, creating a long hollow cylinder, the neural tube Neurulation occurs in all chordates, and the process in a
(figure 60.14), which runs beneath the surface of the em- lancelet is much the same as it is in a human. However, in
bryo’s back. The neural tube later differentiates into the vertebrates, just before the neural groove closes to form the
spinal cord and brain. neural tube, its edges pinch off, forming a small strip of
The dorsal lip of the blastopore induces the formation cells, the neural crest, which becomes incorporated into
of a notochord, and the presence of the notochord induces the roof of the neural tube (figure 60.14). The cells of the
the overlying ectoderm to differentiate into the neural neural crest later move to the sides of the developing em-
tube. The process of induction, when one embryonic re- bryo. The appearance of the neural crest was a key event in
gion of cells influences the development of an adjacent re- the evolution of the vertebrates because neural crest cells,
gion by changing its developmental pathway, was discussed after migrating to different parts of the embryo, ultimately
in chapter 17 and is further examined in the next section. develop into the structures characteristic of (though not
While the neural tube is forming from ectoderm, the necessarily unique to) the vertebrate body.
rest of the basic architecture of the body is being deter- The differentiation of neural crest cells depends on their
mined rapidly by changes in the mesoderm. On either side location. At the anterior end of the embryo, they merge

Neural plate
Neural fold
Neural fold Neural groove
Ectoderm Neural plate

Notochord
Mesoderm
Endoderm
Archenteron
(b) (c)
(a)

FIGURE 60.14
Mammalian neural tube formation. (a) The neural tube forms above the notochord when (b) cells of the neural plate fold together to
form the (c) neural groove.

1224 Part XIV Regulating the Animal Body


with the anterior portion of the brain, the forebrain. bars, with their internal blood supply, become highly
Nearby clusters of ectodermal cells associated with the branched and form the gills of the adult.
neural crest cells thicken into placodes, which are distinct Because the stiff bars of the gill chamber can be bent in-
from neural crest cells although they arise from similar cel- ward by powerful muscles controlled by nerves, the whole
lular interactions. Placodes subsequently develop into parts structure is a very efficient pump that drives water past the
of the sense organs in the head. The neural crest and asso- gills. The gills themselves act as highly efficient oxygen ex-
ciated placodes exist in two lateral strips, which is why the changers, greatly increasing the respiratory capacity of the
vertebrate sense organs that develop from them are paired. animals that possess them.
Neural crest cells located in more posterior positions
have very different developmental fates. These cells mi-
Elaboration of the Nervous System
grate away from the neural tube to other locations in the
head and trunk, where they form connections between the Some neural crest cells migrate ventrally toward the noto-
neural tube and the surrounding tissues. At these new loca- chord and form sensory neurons in the dorsal root ganglia
tions, they contribute to the development of a variety of (see chapter 54). Others become specialized as Schwann
structures that are particularly characteristic of the verte- cells, which insulate nerve fibers and permit the rapid con-
brates, several of which are discussed below. The migration duction of nerve impulses. Still others form the autonomic
of neural crest cells is unique in that it is not simply a ganglia and the adrenal medulla. Cells in the adrenal
change in the relative positions of cells, such as that seen in medulla secrete epinephrine when stimulated by the sym-
gastrulation. Instead, neural crest cells actually pass pathetic division of the autonomic nervous system during
through other tissues. the fight-or-flight reaction. The similarity in the chemical
nature of the hormone epinephrine and the neurotransmit-
ter norepinephrine, released by sympathetic neurons, is un-
The Gill Chamber derstandable—both adrenal medullary cells and sympa-
Primitive chordates such as lancelets are filter-feeders, thetic neurons derive from the neural crest.
using the rapid beating of cilia to draw water into their
bodies through slits in their pharynx. These pharyngeal Sensory Organs and Skull
slits evolved into the vertebrate gill chamber, a structure
that provides a greatly improved means of respiration. The A variety of sense organs develop from the placodes. In-
evolution of the gill chamber was certainly a key event in cluded among them are the olfactory (smell) and lateral line
the transition from filter-feeding to active predation. (primitive hearing) organs discussed in chapter 55. Neural
In the development of the gill chamber, some of the crest cells contribute to tooth development and to some of
neural crest cells form cartilaginous bars between the em- the facial and cranial bones of the skull.
bryonic pharyngeal slits. Other neural crest cells induce
portions of the mesoderm to form muscles along the carti- The appearance of the neural crest in the developing
lage, while still others form neurons that carry impulses be- embryo marked the beginning of the first truly
tween the nerve cord and these muscles. A major blood vertebrate phase of development, as many of the
structures characteristic of vertebrates derive directly
vessel called the aortic arch passes through each of the em-
or indirectly from neural crest cells.
bryonic bars. Lined by still more neural crest cells, these

Neural crest Neural tube


Neural crest
Neural crest
Somite

Coelom
Notochord
Neural tube

(d) (e) Archenteron


(digestive cavity)

FIGURE 60.14 (continued)


(d) The neural groove eventually closes to form a hollow tube. (e) As the tube closes, some of the cells from the dorsal margin of the neural
tube differentiate into the neural crest, which is characteristic of vertebrates.

Chapter 60 Vertebrate Development 1225


How Cells Communicate during las develops from the grey crescent zone and is the site of
origin of those mesoderm cells that later produce the no-
Development tochord.) The new location corresponded to that of the
In the process of vertebrate development, the relative posi- animal’s belly. What happened? The embryo developed
tion of particular cell layers determines, to a large extent, two notochords, a normal dorsal one and a second one
the organs that develop from them. By now, you may have along its belly!
wondered how these cell layers know where they are. For By using genetically different donor and host blastulas,
example, when cells of the ectoderm situated above the de- Spemann and Mangold were able to show that the noto-
veloping notochord give rise to the neural groove, how do chord produced by transplanting dorsal lip cells contained
these cells know they are above the notochord? host cells as well as transplanted ones. The transplanted
The solution to this puzzle is one of the outstanding dorsal lip cells had acted as organizers (see also chapter
accomplishments of experimental embryology, the study 17) of notochord development. As such, these cells stimu-
of how embryos form. The great German biologist Hans lated a developmental program in the belly cells of the
Spemann and his student Hilde Mangold solved it early in embryos in which they were transplanted: the develop-
this century. In their investigation they removed cells ment of the notochord. The belly cells clearly contained
from the dorsal lip of an amphibian blastula and trans- this developmental program but would not have expressed
planted them to a different location on another blastula it in the normal course of their development. The trans-
(figure 60.15). (The dorsal lip region of amphibian blastu- plantation of the dorsal lip cells caused them to do so.

Discard mesoderm
opposite dorsal lip

Dorsal lip

Donor mesoderm
from dorsal lip

Primary
neural fold Primary notochord Secondary notochord
and neural development and neural development

Secondary
neural development

FIGURE 60.15
Spemann and Mangold’s dorsal lip transplant experiment.

1226 Part XIV Regulating the Animal Body


Ectoderm
Wall of forebrain
Optic cup
Optic stalk Lens
invagination Lens vesicle Lens
Lens

Neural
cavity Optic nerve
Sensory
layer
Retina
Pigment
layer

FIGURE 60.16
Development of the eye by induction. An extension of the optic stalk grows until it contacts ectoderm, which induces a section of the
ectoderm to pinch off and form the lens. Other structures of the eye develop from the optic stalk.

These cells had indeed induced the ectoderm cells of the The Nature of Developmental Decisions
belly to form a notochord. This phenomenon as a whole
All of the cells of the body, with the exception of a few spe-
is known as induction.
cialized ones that have lost their nuclei, have an entire
The process of induction that Spemann discovered ap-
complement of genetic information. Despite the fact that
pears to be the basic mode of development in vertebrates.
all of its cells are genetically identical, an adult vertebrate
Inductions between the three primary tissue types—ecto-
contains hundreds of cell types, each expressing various as-
derm, mesoderm, and endoderm—are referred to as pri-
pects of the total genetic information for that individual.
mary inductions. Inductions between tissues that have al-
What factors determine which genes are to be expressed in
ready been differentiated are called secondary inductions.
a particular cell and which are not to be? In a liver cell,
The differentiation of the central nervous system during
what mechanism keeps the genetic information that speci-
neurulation by the interaction of dorsal ectoderm and dor-
fies nerve cell characteristics turned off? Does the differen-
sal mesoderm to form the neural tube is an example of pri-
tiation of that particular cell into a liver cell entail the phys-
mary induction. In contrast, the differentiation of the lens
ical loss of the information specifying other cell types? No,
of the vertebrate eye from ectoderm by interaction with tis-
it does not—but cells progressively lose the capacity to ex-
sue from the central nervous system is an example of sec-
press ever-larger portions of their genomes. Development is a
ondary induction.
process of progressive restriction of gene expression.
The eye develops as an extension of the forebrain, a stalk
Some cells become determined quite early in develop-
that grows outward until it comes into contact with the epi-
ment. For example, all of the egg cells of the human female
dermis (figure 60.16). At a point directly above the growing
are set aside very early in the life of the embryo, yet some
stalk, a layer of the epidermis pinches off, forming a trans-
of these cells will not achieve differentiation into functional
parent lens. When the optic stalks of the two eyes have just
oocytes for more than 40 years. To a large degree, a cell’s
started to project from the brain and the lenses have not yet
location in the developing embryo determines its fate. By
formed, one of the budding stalks can be removed and
changing a cell’s location, an experimenter can alter its de-
transplanted to a region underneath a different epidermis, velopmental destiny. However, this is only true up to a cer-
such as that of the belly. When Spemann performed this tain point in the cell’s development. At some stage, every
critical experiment, a lens still formed, this time from belly cell’s ultimate fate becomes fixed, a process referred to as
epidermis cells in the region above where the budding stalk commitment. Commitment is not irreversible (entire indi-
had been transplanted. viduals can be cloned from an individual specialized cell, as
What is the nature of the inducing signal that passes recounted in chapter 17), but rarely if ever reverses under
from one tissue to the other? If one imposes a nonporous ordinary circumstances.
barrier, such as a layer of cellophane, between the inducer
and the target tissue, no induction takes place. In contrast,
a porous filter, through which proteins can pass, does per- When a cell is “determined,” it is possible to predict its
mit induction to occur. The induction process was dis- developmental fate; when a cell is “committed,” that
cussed in detail in chapter 17. In brief, the inducer cells developmental fate cannot be altered. Determination
often occurs very early in development, commitment
produce a protein factor that binds to the cells of the target
somewhat later.
tissue, initiating changes in gene expression.

Chapter 60 Vertebrate Development 1227


Embryonic Development and Ontogeny Recapitulates Phylogeny
Vertebrate Evolution The patterns of development in the vertebrate groups that
evolved most recently reflect in many ways the simpler pat-
The primitive chordates that gave rise to vertebrates terns occurring among earlier forms. Thus, mammalian de-
were initially slow-moving, filter-feeding animals with velopment and bird development are elaborations of reptile
relatively low metabolic rates. Many of the unique verte- development, which is an elaboration of amphibian devel-
brate adaptations that contribute to their varied ecologi- opment, and so forth (figure 60.18). During the develop-
cal roles involve structures that arise from neural crest ment of a mammalian embryo, traces can be seen of ap-
cells. The vertebrates became fast-swimming predators pendages and organs that are apparently relicts of more
with much higher metabolic rates. This accelerated me- primitive chordates. For example, at certain stages a human
tabolism permitted a greater level of activity than was embryo possesses pharyngeal slits, which occur in all chor-
possible among the more primitive chordates. Other evo- dates and are homologous to the gill slits of fish. At later
lutionary changes associated with the derivatives of the stages, a human embryo also has a tail.
neural crest provided better detection of prey, a greatly In a sense, the patterns of development in chordate
improved ability to orient spatially during prey capture, groups has built up in incremental steps over the evolu-
and the means to respond quickly to sensory information. tionary history of those groups. The developmental in-
The evolution of the neural crest and of the structures structions for each new form seem to have been layered
derived from it were thus crucial steps in the evolution of on top of the previous instructions, contributing addi-
the vertebrates (figure 60.17).

Chordates Vertebrates
Zygote

Brain,
Pharynx Blastula spinal cord,
Lining of spinal nerves
respiratory
tract
Dorsal Neural
Endoderm Gastrula Ectoderm
nerve cord crest
Lining of
digestive
tract Epidermis, skin, Gill arches,
Major hair, epithelium, sensory ganglia,
glands inner ear, lens Schwann cells,
of eye adrenal medulla
Pancreas Liver

Mesoderm Notochord
Outer covering
of internal Circulatory
Integuments Heart
organs system

Blood Vessels
Lining of
thoracic and
Gonads Somites Skeleton
abdominal
cavities Segmented
Kidney muscles
Dermis

FIGURE 60.17
Derivation of the major tissue types. The three germ layers that form during gastrulation give rise to all organs and tissues in the body,
but the neural crest cells that form from ectodermal tissue give rise to structures that are prevalent in the vertebrate animal such as gill
arches and Schwann cells.

1228 Part XIV Regulating the Animal Body


Fish

Salamander

Tortoise

FIGURE 60.18 Chicken


Embryonic development
of vertebrates. Notice
that the early embryonic
stages of these vertebrates
bear a striking resem-
blance to each other, even
though the individuals are
from different classes (fish,
amphibians, reptiles, birds, Human
and mammals). All
vertebrates start out with
an enlarged head region,
gill slits, and a tail
regardless of whether
these characteristics are
retained in the adults.

tional steps in the developmental journey. This hypothe- pharyngeal slits of a mammalian embryo are not like the
sis, proposed in the nineteenth century by Ernst Haeckel, gill slits its ancestors had when they were adults. Rather,
is referred to as the “biogenic law.” It is usually stated as they are like the pharyngeal slits its ancestors had when
an aphorism: ontogeny recapitulates phylogeny; that is, em- they were embryos.
bryological development (ontogeny) involves the same
progression of changes that have occurred during evolu- Vertebrates seem to have evolved largely by the
tion (phylogeny). However, the biogenic law is not liter- addition of new instructions to the developmental
ally true when stated in this way because embryonic stages program. Development of a mammal thus proceeds
through a series of stages, and the earlier stages are
are not reflections of adult ancestors. Instead, the embry-
unchanged from those that occur in the development of
onic stages of a particular vertebrate often reflect the em- more primitive vertebrates.
bryonic stages of that vertebrate’s ancestors. Thus, the

Chapter 60 Vertebrate Development 1229


Extraembryonic Membranes development, the allantois of a bird embryo expands to
form a sac that eventually fuses with the overlying chorion,
As an adaptation to terrestrial life, the embryos of reptiles, just under the eggshell. The fusion of the allantois and
birds, and mammals develop within a fluid-filled amniotic chorion form a functioning unit in which embryonic blood
membrane (see chapter 48). The amniotic membrane and vessels, carried in the allantois, are brought close to the
several other membranes form from embryonic cells but porous eggshell for gas exchange. The allantois is thus the
are located outside of the body of the embryo. For this functioning “lung” of a bird embryo.
reason, they are known as extraembryonic membranes. In mammals, the embryonic cells form an inner cell
The extraembryonic membranes, later to become the mass that will become the body of the embryo and a layer
fetal membranes, include the amnion, chorion, yolk sac, of surrounding cells called the trophoblast (see figure 60.9).
and allantois. The trophoblast implants into the endometrial lining of its
In birds, the amnion and chorion arise from two folds mother’s uterus and becomes the chorionic membrane (fig-
that grow to completely surround the embryo (figure ure 60.20). The part of the chorion in contact with en-
60.19). The amnion is the inner membrane that surrounds dometrial tissue contributes to the placenta, as is described
the embryo and suspends it in amniotic fluid, thereby mim- in more detail in the next section. The allantois in mam-
icking the aquatic environments of fish and amphibian em- mals contributes blood vessels to the structure that will be-
bryos. The chorion is located next to the eggshell and is come the umbilical cord, so that fetal blood can be deliv-
separated from the other membranes by a cavity, the ex- ered to the placenta for gas exchange.
traembryonic coelom. The yolk sac plays a critical role in the
nutrition of bird and reptile embryos; it is also present in The extraembryonic membranes include the yolk sac,
mammals, though it does not nourish the embryo. The al- amnion, chorion, and allantois. These are derived from
lantois is derived as an outpouching of the gut and serves embryonic cells and function in a variety of ways to
to store the uric acid excreted in the urine of birds. During support embryonic development.

Union of
Embryo Amniotic folds Allantois Chorion amniotic folds Amnion Allantois Chorion Amnion Allantois

Yolk

Extra
embryonic
coelom Yolk sac

(a) (b) (c)

FIGURE 60.19
The extraembryonic membranes of a chick embryo. The membranes begin as amniotic folds from the embryo (a) that unite (b) to form
a separate amnion and chorion (c). The allantois continues to grow until it will eventually unite with the chorion just under the eggshell.

1230 Part XIV Regulating the Animal Body


Amnion

Syncitial
trophoblast Maternal
blood vessels

Chorion

Cellular
trophoblast Developing
chorionic villi
Ectoderm
Embryo Mesoderm

Endoderm

Body stalk
(umbilical cord)

Yolk sac
of embryo

Extraembryonic
coelom

Umbilical blood vessels

Chorion

Amnion

Yolk sac

Villus of chorion
frondosum

Maternal blood vessels

FIGURE 60.20
The extraembryonic membranes of a mammalian embryo. (a) After the embryo implants into the mother’s endometrium (6–7 days
after fertilization), the trophoblast becomes the chorion, and the yolk sac and amnion are produced. (b) The chorion develops extensions,
called villi, that interdigitate with surrounding endometrial tissue. The embryo is encased within an amniotic sac.

Chapter 60 Vertebrate Development 1231


60.5 Human development is divided into trimesters.

First Trimester forms and gives rise to the three primary germ layers.
Human development, from fertilization to birth, takes an
The development of the human embryo shows its evolu- average of 266 days. This time is commonly divided into
tionary origins. Without an evolutionary perspective, we three periods, called trimesters.
would be unable to account for the fact that human devel-
opment proceeds in much the same way as development in
The First Month
a bird. In both animals, the embryo develops from a flat-
tened collection of cells—the blastodisc in birds or the About 30 hours after fertilization, the zygote undergoes its
inner cell mass in humans. While the blastodisc of a bird is first cleavage; the second cleavage occurs about 30 hours
flattened because it is pressed against a mass of yolk, the after that. By the time the embryo reaches the uterus (6–7
inner cell mass of a human is flat despite the absence of a days after fertilization), it is a blastula, which in mammals is
yolk mass. In humans as well as in birds, a primitive streak referred to as a blastocyst. As we mentioned earlier, the

Chorion

Amnion

Umbilical
cord

Chorionic
frondosum
(fetal)
Placenta
Decidua
basalis
(maternal)

Maternal
Maternal Uterine
artery
vein wall

FIGURE 60.21
Structure of the placenta. The placenta contains a fetal component, the chorionic frondosum, and a maternal component, the decidua
basalis. Deoxygenated fetal blood from the umbilical arteries (shown in blue) enters the placenta, where it picks up oxygen and nutrients
from the mother’s blood. Oxygenated fetal blood returns in the umbilical vein (shown in red) to the fetus, where it picks up oxygen and
nutrients from the mother’s blood.

1232 Part XIV Regulating the Animal Body


blastocyst consists of an inner cell mass, which will become
the body of the embryo, and a surrounding layer of tro-
phoblast cells (see figure 60.9). The blastocyst begins to
grow rapidly and initiates the formation of the amnion and
the chorion. The blastocyst digests its way into the en-
dometrial lining of the uterus in the process known as im-
plantation.
During the second week after fertilization, the develop-
ing chorion forms branched extensions, the chorionic frondo-
sum (fetal placenta) that protrude into the endometrium
(figure 60.21). These extensions induce the surrounding
endometrial tissue to undergo changes and become the de-
cidua basalis (maternal placenta). Together, the chorionic
frondosum and decidua basalis form a single functioning
unit, the placenta (figure 60.22). Within the placenta, the
mother’s blood and the blood of the embryo come into
close proximity but do not mix (see figure 60.21). Oxygen
can thus diffuse from the mother to the embryo, and car-
bon dioxide can diffuse in the opposite direction. In addi-
tion to exchanging gases, the placenta provides nourish-
ment for the embryo, detoxifies certain molecules that may
pass into the embryonic circulation, and secretes hor-
mones. Certain substances such as alcohol, drugs, and an-
tibiotics are not stopped by the placenta and pass from the
FIGURE 60.22
mother’s bloodstream to the fetus. Placenta and fetus at seven weeks.
One of the hormones released by the placenta is human
chorionic gonadotropin (hCG), which was discussed in
chapter 59. This hormone is secreted by the trophoblast
cells even before they become the chorion, and is the hor-
mone assayed in pregnancy tests. Because its action is al-
most identical to that of luteinizing hormone (LH), hCG many women are unaware they are pregnant at this stage.
maintains the mother’s corpus luteum. The corpus luteum, Early pregnancy is a very critical time in development
in turn, continues to secrete estradiol and progesterone, because the proper course of events can be interrupted eas-
thereby preventing menstruation and further ovulations. ily. In the 1960s, for example, many pregnant women took
Gastrulation also takes place in the second week after the tranquilizer thalidomide to minimize the discomforts
fertilization. The primitive streak can be seen on the sur- associated with early pregnancy. Unfortunately, this drug
face of the embryo, and the three germ layers (ectoderm, had not been adequately tested. It interferes with limb bud
mesoderm, and endoderm) are differentiated. development, and its widespread use resulted in many de-
In the third week, neurulation occurs. This stage is formed babies. Organogenesis may also be disrupted dur-
marked by the formation of the neural tube along the dor- ing the first and second months of pregnancy if the mother
sal axis of the embryo, as well as by the appearance of the contracts rubella (German measles). Most spontaneous
first somites, which give rise to the muscles, vertebrae, and abortions occur during this period.
connective tissues. By the end of the week, over a dozen
somites are evident, and the blood vessels and gut have The Second Month
begun to develop. At this point, the embryo is about 2 mil-
limeters long. Morphogenesis (the formation of shape) takes place dur-
Organogenesis (the formation of body organs) begins ing the second month (figure 60.23b). The miniature limbs
during the fourth week (figure 60.23a). The eyes form. The of the embryo assume their adult shapes. The arms, legs,
tubular heart develops its four chambers and starts to pul- knees, elbows, fingers, and toes can all be seen—as well as a
sate rhythmically, as it will for the rest of the individual’s short bony tail! The bones of the embryonic tail, an evolu-
life. At 70 beats per minute, the heart is destined to beat tionary reminder of our past, later fuse to form the coccyx.
more than 2.5 billion times during a lifetime of 70 years. Within the abdominal cavity, the major organs, including
Over 30 pairs of somites are visible by the end of the fourth the liver, pancreas, and gallbladder, become evident. By the
week, and the arm and leg buds have begun to form. The end of the second month, the embryo has grown to about
embryo has increased in length to about 5 millimeters. Al- 25 millimeters in length, weighs about one gram, and be-
though the developmental scenario is now far advanced, gins to look distinctly human.

Chapter 60 Vertebrate Development 1233


(a) (b)

FIGURE 60.23
The developing human. (a) Four weeks, (b) seven weeks, (c) three months, and (d) four months.

The Third Month


Increasing hormone concentration

The nervous system and sense organs develop during the


third month, and the arms and legs start to move (figure hCG Estradiol
60.23c). The embryo begins to show facial expressions and
carries out primitive reflexes such as the startle reflex and
sucking. The eighth week marks the transition from em- Progesterone
bryo to fetus. At this time, all of the major organs of the
body have been established. What remains of development
is essentially growth.
At around 10 weeks, the secretion of human chorionic 0 1 2 3 4 5 6 7 8 9
gonadotropin (hCG) by the placenta declines, and the
Months of pregnancy
corpus luteum regresses as a result. However, menstrua-
tion does not occur because the placenta itself secretes
estradiol and progesterone (figure 60.24). In fact, the FIGURE 60.24
amounts of these two hormones secreted by the placenta Hormonal secretion by the placenta. The placenta secretes
far exceed the amounts that are ever secreted by the chorionic gonadotropin (hCG) for 10 weeks. Thereafter, it
ovaries. The high levels of estradiol and progesterone in secretes increasing amounts of estradiol and progesterone.
the blood during pregnancy continue to inhibit the re-
lease of FSH and LH, thereby preventing ovulation.
The embryo implants into the endometrium,
They also help maintain the uterus and eventually pre-
differentiates the germ layers, forms the
pare it for labor and delivery, and they stimulate the de-
extraembryonic membranes, and undergoes
velopment of the mammary glands in preparation for lac- organogenesis during the first month and
tation after delivery. morphogenesis during the second month.

1234 Part XIV Regulating the Animal Body


(c) (d)

Second and Third Trimesters Third Trimester

The second and third trimesters are characterized by the The third trimester is predominantly a period of growth
tremendous growth and development required for the via- rather than development. The weight of the fetus doubles
bility of the baby after its birth. several times, but this increase in bulk is not the only kind
of growth that occurs. Most of the major nerve tracts in the
brain, as well as many new neurons (nerve cells), are
Second Trimester formed during this period. The developing brain produces
Bones actively enlarge during the fourth month (figure neurons at an average rate estimated at more than 250,000
60.23d), and by the end of the month, the mother can feel per minute! Neurological growth is far from complete at
the baby kicking. During the fifth month, the head and the end of the third trimester, when birth takes place. If the
body grow a covering of fine hair. This hair, called lanugo, fetus remained in the uterus until its neurological develop-
is another evolutionary relict but is lost later in develop- ment was complete, it would grow too large for safe deliv-
ment. By the end of the fifth month, the rapid heartbeat of ery through the pelvis. Instead, the infant is born as soon as
the fetus can be heard with a stethoscope, although it can the probability of its survival is high, and its brain contin-
also be detected as early as 10 weeks with a fetal monitor. ues to develop and produce new neurons for months after
The fetus has grown to about 175 millimeters in length and birth.
attained a weight of about 225 grams. Growth begins in
earnest in the sixth month; by the end of that month, the The critical stages of human development take place
baby weighs 600 grams (1.3 lbs) and is over 300 millimeters quite early, and the following six months are essentially
a period of growth. The growth of the brain is not yet
(1 ft) long. However, most of its prebirth growth is still to
complete, however, by the end of the third trimester,
come. The baby cannot yet survive outside the uterus with- and must be completed postnatally.
out special medical intervention.

Chapter 60 Vertebrate Development 1235


Birth and Postnatal Intestine
Development
In some mammals, changing hormone levels in the Placenta
developing fetus initiate the process of birth. The
fetuses of these mammals have an extra layer of cells
in their adrenal cortex, called a fetal zone. Before
birth, the fetal pituitary gland secretes corticotropin,
which stimulates the fetal zone to secrete steroid Umbilical
cord
hormones. These corticosteroids then induce the
uterus of the mother to manufacture prostaglandins,
which trigger powerful contractions of the uterine
smooth muscles.
The adrenal glands of human fetuses lack a fetal
zone, and human birth does not seem to be initiated
by this mechanism. In a human, the uterus releases
prostaglandins, possibly as a result of the high levels
Wall of
of estradiol secreted by the placenta. Estradiol also uterus
stimulates the uterus to produce more oxytocin re-
ceptors, and as a result, the uterus becomes increas-
ingly sensitive to oxytocin. Prostaglandins begin the Vagina
uterine contractions, but then sensory feedback
from the uterus stimulates the release of oxytocin
from the mother’s posterior pituitary gland. Work- FIGURE 60.25
ing together, oxytocin and prostaglandins further Position of the fetus just before birth. A developing fetus is a major
stimulate uterine contractions, forcing the fetus addition to a woman’s anatomy. The stomach and intestines are pushed far
up, and there is often considerable discomfort from pressure on the lower
downward (figure 60.25). Initially, only a few con-
back. In a natural delivery, the fetus exits through the vagina, which must
tractions occur each hour, but the rate eventually in- dilate (expand) considerably to permit passage.
creases to one contraction every two to three min-
utes. Finally, strong contractions, aided by the mother’s
pushing, expel the fetus, which is now a newborn baby.
After birth, continuing uterine contractions expel the
placenta and associated membranes, collectively called the
afterbirth. The umbilical cord is still attached to the baby, Adipose
tissue
and to free the newborn, a doctor or midwife clamps and Rib
cuts the cord. Blood clotting and contraction of muscles in
the cord prevent excessive bleeding.

Nursing Intercostal
muscles
Milk production, or lactation, occurs in the alveoli of mam-
Mammary
mary glands when they are stimulated by the anterior pitu- (alveolar)
Pectoralis
itary hormone, prolactin. Milk from the alveoli is secreted minor duct
into a series of alveolar ducts, which are surrounded by
smooth muscle and lead to the nipple (figure 60.26). Dur- Lactiferous
ing pregnancy, high levels of progesterone stimulate the Pectoralis duct
major
development of the mammary alveoli, and high levels of
estradiol stimulate the development of the alveolar ducts.
However, estradiol blocks the actions of prolactin on the

FIGURE 60.26
A sagittal section of a mammary gland. The mammary alveoli
Lobule Containing
produce milk in response to stimulation by prolactin, and milk is
Lobe mammary
ejected through the lactiferous duct in response to stimulation by alveoli
oxytocin.

1236 Part XIV Regulating the Animal Body


mammary glands, and it inhibits prolactin secretion by pro- Chimpanzee Human
moting the release of prolactin-inhibiting hormone from
the hypothalamus. During pregnancy, therefore, the mam-
mary glands are prepared for lactation but prevented from
lactating.
When the placenta is discharged after birth, the concen- Fetus
trations of estradiol and progesterone in the mother’s
blood decline rapidly. This decline allows the anterior pitu-
itary gland to secrete prolactin, which stimulates the mam-
mary alveoli to produce milk. Sensory impulses associated
with the baby’s suckling trigger the posterior pituitary
gland to release oxytocin. Oxytocin stimulates contraction
of the smooth muscle surrounding the alveolar ducts, thus
causing milk to be ejected by the breast. This pathway is
known as the milk-ejection reflex. The secretion of oxy-
tocin during lactation also causes some uterine contrac- Infant
tions, as it did during labor. These contractions help to re-
store the tone of uterine muscles in mothers who are
breastfeeding.
The first milk produced after birth is a yellowish fluid
called colostrum, which is both nutritious and rich in ma-
ternal antibodies. Milk synthesis begins about three days
following the birth and is referred to as when milk “comes
in.” Many mothers nurse for a year or longer. During this
period, important pair-bonding occurs between the mother
and child. When nursing stops, the accumulation of milk in
the breasts signals the brain to stop prolactin secretion, and Child
milk production ceases.

Postnatal Development
Growth of the infant continues rapidly after birth. Babies
typically double their birth weight within two months. Be-
cause different organs grow at different rates and cease
growing at different times, the body proportions of infants
are different from those of adults. The head, for example, is
disproportionately large in newborns, but after birth it
Adult
grows more slowly than the rest of the body. Such a pattern
of growth, in which different components grow at different
rates, is referred to as allometric growth.
In most mammals, brain growth is mainly a fetal phe-
nomenon. In chimpanzees, for instance, the brain and the
cerebral portion of the skull grow very little after birth,
while the bones of the jaw continue to grow. As a result, FIGURE 60.27
the head of an adult chimpanzee looks very different from Allometric growth. In young chimpanzees, the jaw grows at a
that of a fetal chimpanzee (figure 60.27). In human infants, faster rate than the rest of the head. As a result, the adult chim-
on the other hand, the brain and cerebral skull grow at the panzee head shape differs greatly from its head shape as a
same rate as the jaw. Therefore, the jaw-skull proportions newborn. In humans, the difference in growth between the jaw
do not change after birth, and the head of a human adult and the rest of the head is much smaller, and the adult head shape
looks very similar to that of a human fetus. It is primarily is similar to that of the newborn.
for this reason that an early human fetus seems so remark-
ably adultlike. The fact that the human brain continues to
grow significantly for the first few years of postnatal life Birth occurs in response to uterine contractions
means that adequate nutrition and a rich, safe environment stimulated by oxytocin and prostaglandins. Lactation is
stimulated by prostaglandin, but the milk-ejection
are particularly crucial during this period for the full devel-
reflex requires the action of oxytocin.
opment of the person’s intellectual potential.

Chapter 60 Vertebrate Development 1237


Chapter 60 www.mhhe.com/raven6e www.biocourse.com

Summary Questions Media Resources


60.1 Fertilization is the initial event in development.

• Fertilization is the union of an egg and a sperm to 1. What are the three stages of • Fertilization
form a zygote. It is accomplished externally in most fertilization, and what happens
fish and amphibians, and internally in all other during each stage?
vertebrates.
• The three stages of fertilization are (1) penetration,
(2) activation, and (3) fusion.

60.2 Cell cleavage and the formation of a blastula set the stage for later development.

• Cleavage is the rapid division of the newly formed 2. What is the difference • Early development
zygote into a mass of cells, without any increase in between holoblastic cleavage and • Preembryonic
meroblastic cleavage? What is development
overall size.
responsible for an embryo
• The cleavages produce a hollow ball of cells, called undergoing one or the other
the blastula. type of cleavage?

60.3 Gastrulation forms the three germ layers of the embryo.

• During gastrulation, cells in the blastula change their 3. What is an archenteron, and • Cell differentation
relative positions, forming the three primary cell during what developmental stage
layers: ectoderm, mesoderm, and endoderm. does it form? What is the future
fate of this opening in
• In eggs with moderate or large amounts of yolk, cells vertebrates?
involute down and around the yolk, through a
4. How is gastrulation in
blastopore or primitive streak to form the three germ
amphibians different from that
layers. in lancelets?

60.4 Body architecture is determined during the next stages of embryonic development.

• Neurulation involves the formation of a structure 5. What structure unique to • Art Activity:
found only in chordates, the notochord and dorsal chordates forms during Human extra-
neurulation? embryonic membranes
hollow nerve cord.
• The formation of the neural crest is the first 6. What are the functions of
developmental event unique to vertebrates. Most of the amnion, chorion, and • Embryonic and fetal
allantois in birds and mammals? development
the distinctive structures associated with vertebrates
are derived from the neural crest.

60.5 Human development is divided into trimesters.

• Most of the critical events in human development 7. How does the placenta • Bioethics Case Study:
occur in the first month of pregnancy. Cleavage prevent menstruation during the Critically ill newborns
occurs during the first week, gastrulation during the first two months of pregnancy?
second week, neurulation during the third week, and 8. At what time during human
organogenesis during the fourth week. pregnancy does organogenesis • Human development
occur? • Pregnancy
• The second and third months of human development • Postnatal period
are devoted to morphogenesis and to the elaboration 9. Is neurological growth
of the nervous system and sensory organs. complete at birth?
10. What hormone stimulates
• During the last six months before birth, the human
lactation (milk production)?
fetus grows considerably, and the brain produces What hormone stimulates milk
large numbers of neurons and establishes major nerve ejection from the breast?
tracts.
1238 Part XIV Regulating the Animal Body

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