Raven Johnson McGraw-Hill Biology-11
Raven Johnson McGraw-Hill Biology-11
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-
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.
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.
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.
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.
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.
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
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.
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.
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.
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.
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%.
(a)
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.
• 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.
• 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.
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.
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)
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)
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.
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.
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
Ectoderm
Ectoderm Archenteron
Ectoderm
Endoderm
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.
Dorsal lip
Blastocoel
Yolk plug
Mesoderm Dorsal lip of
blastopore Ventral lip
Vegetal pole
(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.
Trophoblast Endoderm
Mesoderm
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.
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.
Coelom
Notochord
Neural tube
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.
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.
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.
Salamander
Tortoise
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
Union of
Embryo Amniotic folds Allantois Chorion amniotic folds Amnion Allantois Chorion Amnion Allantois
Yolk
Extra
embryonic
coelom Yolk sac
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.
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
Chorion
Amnion
Yolk sac
Villus of chorion
frondosum
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.
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.
FIGURE 60.23
The developing human. (a) Four weeks, (b) seven weeks, (c) three months, and (d) four months.
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.
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.
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.
• 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?
• 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.
• 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