21

Development
and Aging

Staying fit may improve quality of life as we age.

Who Should Make Life and Death Decisions for You?

Current Issue

I f you suffered severe brain damage and

could survive only on artificial life sup-
port, would you want to continue living?
provide fluids and nutrition.
Over the next four years
Terri was transferred to vari-
Who do you think will take care of you ous medical centers, where
and make decisions related to your life— she received ongoing
and death? If you think you’re too young testing, speech and occupa-
to worry about such issues, consider the tional therapy, and other at-
story of Terri Schiavo. tempts at rehabilitation.
In February 1990, 26-year-old Terri
suffered cardiac arrest, possibly caused
A Legal Battle and
by potassium imbalance due to bulimia.
She was rushed to a hospital. However,
National Debate Over
lack of oxygen had damaged her brain the Right to Die
permanently, and she lapsed into a In June 1990 Michael Schi-
coma. A feeding tube was inserted avo, Terri’s husband, was Family photo of Terri Schiavo before she had
a heart attack.
through the skin and into her stomach to appointed her legal guardian.
At the time Terri’s parents, Robert and One way around the
Mary Schindler, did not object. But eight uncertainty of a living will is to
years later, a decision made by Michael combine it with a durable power
Schiavo regarding Terri’s life became the of attorney for health care, also
focus of a bitter court battle that escalated called a “health care proxy
into a national debate about the right to appointment.” This is a legal doc-
live versus the right to die. ument in which you give a named
In 1998, when Terri had been in a person the right to make all med-
coma for eight years, Michael petitioned ical decisions on your behalf
the court to authorize the removal of Terri’s should you be unable to make
feeding tube and allow her to die. Immedi- A supporter lights a prayer candle for Terri. them for yourself. Combined with a
ately thereafter, Robert and Mary Schindler living will, a durable power of attor-
counterpetitioned to stop Michael from ney offers reasonable assurance
By the time Terri’s feeding tube was re-
ending their daughter’s life. that your wishes will be carried out. But
moved again in 2003, the case had caught
If Terri had made a living will (a legal you’ve got to trust that person, and you
the public’s (and politicians’) attention.
document that expresses one’s preferences need to be sure that they understand
Eventually the argument went all the way
regarding life support and resuscitation), your wishes well.
to the U.S. Supreme Court, which declined
the decision to terminate her life support Currently more advanced types of
to hear the case.
might have been relatively straightforward. directives are being studied as well.
Terri Schiavo’s feeding tube was
In this case, however, the decision became One type, called a Lifecare Advance
removed for the last time on March 18,
an immensely difficult undertaking. What Directive, focuses more on general health
2005. On March 31, at age 41, she died.
would Terri have wanted? When a person’s outcomes than on lists of specific
An autopsy showed that her brain had
wishes are not clear, who should make the medical conditions and treatments. A
shrunk to 50% of its normal weight and
decision? Do we have an ethical obligation Lifecare Advance Directive is less likely
confirmed the medical diagnosis of persis-
to keep someone alive as long as possible? to come in conflict with good medical
tent vegetative state. She could not have
And when, exactly, is someone no longer practice, but on the other hand it may
recovered.
“alive”? be too vague to describe a specific situa-
In Terri’s court case, Michael and two tion. Some legal experts are recommend-
relatives testified that prior to her heart Protecting Yourself from ing that you use all three types (living
attack, Terri had commented about a seri- a Similar Fate will, durable power of attorney, and life-
ously ill person, “I wouldn’t want to live Given the choice, many of us would care advance directive) if you want
like that. I would want to just die.” In re- choose not to live 15 years in a persistent virtual certainty that your wishes will be
sponse, Terri’s parents told the court that vegetative state. How can we protect our- carried out. Skeptics say that all of
Terri was a devout Catholic who would selves from such a fate? As mentioned these advance directives are more about
want to live. They insisted their daughter previously, one way is to sign a living will, protecting the medical and legal profes-
had the right to live; her husband insisted also known as an “advance health care di- sions than about doing the right thing
she had the right to die. A medical report rective.” A living will describes what med- for the patient.
submitted to the court indicated that Terri ical procedures you would want performed And what if you continue to do nothing
was in a “persistent vegetative state” with on you, and under what circumstances. It and then become unable to express your
no chance of improvement. is intended to be used only in the event wishes, as happened to Terri Schiavo? In
In 2000 a circuit court judge ruled that that you become unable to make decisions order of priority, decision-making authority
the feeding tube should be removed, on the for yourself. However, a comprehensive liv- generally rests first with your spouse, then
grounds that Terri would have chosen to do ing will is difficult to write because it is with your parents. Heaven help you if they
so. Her parents initiated a series of legal hard to envision virtually every possibility disagree, or if you and your spouse have
appeals, countered by appeals from Michael that might befall you. Furthermore, in a split up but have not yet gotten a divorce!
Schiavo. In 2001 the feeding tube was re- situation in which your wishes come in The more you make your wishes clear, the
moved, but it was reinserted two days later conflict with good medical practice, doc- better off you’ll be.
when the Schindlers filed a lawsuit. tors can refuse to honor it.

Questions to consider
1 If you were in Terri Schiavo’s situation,
The facts...

■ In 1990, 26-year-old Terri Schiavo suffered a cardiac arrest and lapsed into
would you want to be allowed to die?
a coma.
2 Do you have a living will or a durable
■ Terri’s husband and her parents disagreed about whether Terri would have
power of attorney? If not, who do you
wanted to be kept on life support indefinitely.
think will make decisions for you if you
■ Terry died 15 years later, after a prolonged legal battle. An autopsy revealed are unable to speak for yourself?
that she could not have recovered.
■ Signing a living will or giving someone you trust a durable power of attorney
for health care can reduce the chances that this will happen.
485
 The journeys of egg and sperm
» Humans
Key concepts

develop from a single fertilized egg to

a complex organism of over 10 trillion cells in It’s remarkable that sperm and egg ever find each other at
just nine months. A complex pattern of cell all, given the journey that each must take. An egg is released
differentiation and cell division results in the from one of the two ovaries at ovulation, and the sperm are
formation of the various tissues, organs, and deposited in the vagina near the cervix during intercourse. The
organ systems. egg moves slowly and passively down the oviduct, propelled by
cilia that line the tube and sweep it gently downward.
» Birth represents the first step toward indepen-
dence. At birth the circulatory, respiratory,
Sperm are not much more than a head containing the
all-important DNA, a midpiece containing mitochondria for
and digestive systems function independently energy production during the sperm’s long journey, and a
for the first time. flagellum for movement (Figure 21.1a). The several hundred
» Adolescence is the transition from childhood to
adulthood. Adolescence is marked by matura-
million sperm encounter many hazards on their journey to-
ward the egg. First they must pass through the mucus that
tion of the reproductive systems and sexual blocks the cervical opening and cross the vast (to them) ex-
responsiveness. panse of the uterus. Then they must locate and enter the cor-
rect oviduct (half don’t). Sperm swim randomly at a rate of
» Aging is a process that begins in early adulthood
and continues until death. Aging is charac-
1/4 inch per minute. Along the way they must tolerate the
strongly acidic pH of the vagina, and avoid bacteria and the
terized by the slow decline in function of most occasional white blood cells roaming the uterine lining.
tissues, organs, and organ systems. The aging A sperm’s journey can take hours, and only a few thou-
process can be slowed, but it cannot yet be sand or a few hundred make it successfully to the upper
prevented. oviduct; the rest are lost along the way and die. Fertilization
» Death, defined as the cessation of life, is the
final transition. The death of a person is not
typically takes place in the upper third of the oviduct approx-
imately 6–24 hours after intercourse, provided that an egg is
necessarily accompanied by the immediate present (Figure 21.1b). If no egg is present, the sperm may
death of all organs, tissues, or cells. live for a couple of days and then die. If no sperm arrive, the
egg dies in about 24 hours.

One sperm fertilizes the egg

The odds of success for a single sperm are still not very
good, for only one of the several hundred million sperm

L ife is a journey that begins at fertilization and ends

around the time of our last breath. It’s an interesting
and complicated process in which one developmental
will fertilize the egg. This is important, for otherwise
the zygote would end up with an abnormal number of
chromosomes. The process of fertilization has evolved to
stage follows the next in a carefully choreographed dance. ensure that only one sperm can succeed. Recall that before
The changes that occur during development are nothing fertilization, the egg is really a secondary oocyte that has
short of amazing: in just nine months a single undifferentiated started but not finished the second stage of meiosis
cell develops into a human baby with complex organ systems (Figure 21.1c). The second stage of meiosis is not completed
comprising more than 10 trillion cells. In less than 20 years, until a sperm makes contact with the egg and fertilization
this helpless baby becomes a fully functioning adult capable of begins. At this point the secondary oocyte is surrounded
producing still more human babies. Then, starting at about age by a protective covering called the zona pellucida and by a
40, there is a slow decline of function for reasons we don’t yet layer of granulosa cells derived from the follicle called the
fully understand, ending inevitably in death. corona radiata. The egg is relatively large (nearly 2,000
The amazing journey of life is what this chapter is all times the mass of a sperm) because it contains a great deal
about. We begin at the beginning when sperm meets egg, of cytoplasm. The cytoplasm of the egg must support
and end at the ending when life finally ceases. nearly two weeks’ worth of cell divisions until the “pre-
embryo,” as it is called, makes contact with the uterine
lining and begins to receive nutrients from the mother.
21.1 Fertilization begins when sperm When a sperm encounters the egg, the tip of the sperm
head, called the acrosome, releases powerful enzymes
and egg unite (Figure 21.2). These enzymes digest a path for the sperm
The possibility that fertilization will occur begins with inter- between the granulosa cells of the corona radiata and
course. Fertilization is a process that begins when a sperm through the zona pellucida to the oocyte plasma membrane.
and egg unite and ends when the zygote (the new diploid Several sperm may be making this journey at the same time.
cell) is formed. When the first sperm makes contact with the oocyte plasma

486
 Chapter 21 Development and Aging 487

Optimal site
of fertilization
Oviduct
Uterus

Secondary
Acrosome oocyte

Head
Corona
Sperm Ovary radiata
Midpiece
Cytoplasm
Tail
Nucleus

Plasma
membrane
Cervix
First polar
body
Vagina

Zona
pellucida

a) The male gamete, or sperm. The b) Fertilization. Fertilization generally c) The female gamete. The female gamete is
size of a sperm relative to the secondary takes place in the upper third of a secondary oocyte that is in an arrested
oocyte has been greatly exaggerated. the oviduct. state of stage II of meiosis.

Figure 21.1 Female and male gametes and fertilization.

Sperm
Corona radiata

Secondary oocyte
First polar body

Sperm nucleus

Acrosome
All other
sperm
denied
entrance
Acrosome
releases enzymes

Granulosa cells
of corona radiata

Zona pellucida
Oocyte and sperm
plasma membranes fuse
Oocyte plasma
membrane
Granules release
enzymes that make the
Granule zona pellucida impenetrable
Oocyte Sperm nucleus engulfed
cytoplasm by oocyte cytoplasm

Figure 21.2 Fertilization. Fertilization begins when a sperm makes contact with the secondary
oocyte and releases enzymes that digest a path through the zona pellucida. Once a sperm makes
contact with the oocyte plasma membrane and enters the oocyte, granules are released from
the oocyte that make the zona pellucida impenetrable to other sperm.
488 Chapter 21 Development and Aging

Egg nucleus

Sperm nucleus Sperm nucleus Fusion of egg and

sperm nuclei

Second
meiotic
division

First polar body Polar bodies

may also divide

a) A sperm nucleus has just entered the b) Completion of meiosis yields an c) The nuclei of the ovum
secondary oocyte, triggering ovum (egg) with a haploid nucleus and sperm fuse, and the
completion of meiosis II. and another polar body. If it has not polar bodies die. A single
already regressed, the original polar diploid cell called the zygote
body may divide as well. is formed, and fertilization
is complete.

Figure 21.3 Completion of fertilization.

membrane, special protein “keys” of the sperm recognize re- Twins may be fraternal or identical
ceptor protein “locks” in the oocyte plasma membrane, en- Twins occur once in about every 90 births. Twins are the most
suring that only human sperm can penetrate a human egg. common form of multiple births. Occasionally we hear of
The combination of lock and key causes the plasma mem- sextuplets or septuplets (six or seven newborns at once), but
branes of egg and sperm to fuse so the nucleus of the sperm these are rare and almost always result from the use of fertil-
can enter the egg. ity drugs. Even natural triplets are rare (1 in 8,000 births).
Fertilization begins when the sperm’s nucleus enters the Two different processes lead to twins (Figure 21.4).
egg. The entry of one sperm triggers the release of enzymes Fraternal twins arise from the ovulation of more than one
from granules located just inside the egg. These enzymes pro- oocyte in a particular cycle. The oocytes are as different geneti-
duce changes in the zona pellucida that make it impenetrable cally as any two oocytes ovulated at different cycles, and each is
to all other sperm. One sperm nucleus, and only one, will fertilized by a different sperm. Fraternal twins are as different
enter the egg. as any children by the same parents. Indeed, because they are
The process of fertilization is not yet complete, derived from different sperm they can be of different genders.
however, because the “egg” is still a secondary oocyte. Identical twins, sometimes called maternal twins, are al-
Entry of the sperm nucleus triggers the completion of ways genetically identical. Identical twins arise from a single
meiosis II and formation of the haploid ovum, or mature zygote. Recall that up until about the 8-cell stage, all cells of
egg, and another polar body. Fertilization is considered
complete when the haploid nuclei of sperm and mature
ovum join, forming a single diploid cell (the zygote) with
46 chromosomes (Figure 21.3). Some people refer to the
end of fertilization as conception and to the product of
conception as the conceptus. The follicular cells that
surrounded the secondary oocyte are shed and the polar
bodies die, leaving just the fertilized cell, or zygote.

Quick Check An egg can occasionally break out of its zona

pellucida before the sperm reach it. If no zona pellucida is
present when the sperm reach the egg, what might go wrong
during fertilization?
a) Fraternal twins arise when two b) Identiclal twins arise when a
eggs are ovulated and fertilized zygote divides in two during
in the same monthly cycle. development.

Answers to questions can be found in Appendix A. Figure 21.4 Twins.

 Health &
Wellness

Prenatal Diagnostic Techniques

F ifty years ago, when a woman became

pregnant there was little that the par-
ents or physician could do except hope
chromosomes and perform certain bio-
chemical tests. Amniocentesis is relatively
safe and may be recommended if there is
advantage of chorionic villi sampling is that
it can be done as early as five weeks of
pregnancy. Because there is a slight but
that the baby would be born healthy. Late a family history of a genetic disease or the real risk of injury to the embryo, the tech-
in pregnancy the physician could hear the risk of Down syndrome. Nearly 40 differ- nique is recommended only when a family
fetal heartbeat, but that was about it. ent fetal defects can be diagnosed using history indicates a risk of genetic defects.
Given that nearly 7% of all newborns had amniocentesis. A drawback is that amnio- Percutaneous umbilical blood sampling
a birth defect of some kind, pregnancy was centesis cannot be performed until about (PUBS) is a technique for collecting fetal
a stressful period of wait-and-see. Today, the fifteenth week of gestation, and test blood directly from the umbilical cord.
fetal cells can be collected for genetic results may not be available until the mid- Under ultrasound guidance, a needle is in-
testing fairly early in fetal development, dle of the second trimester. This is close serted through the abdominal wall and into
even early enough to permit a couple to to the time that termination of pregnancy a vein in the umbilical cord. Cord blood
terminate a pregnancy if that is their wish will no longer be an option. obtained by PUBS can be tested for sickle-
after reviewing the test results. In chorionic villi sampling, a thin, flexi- cell anemia, hemophilia, anemia, and Rh
In amniocentesis, a needle is inserted ble tube attached to a syringe is inserted problems, as well as genetic abnormalities.
through the abdominal wall and a sample through the vagina and into the uterus, and The risk of miscarriage is slightly higher
of amniotic fluid containing fetal cells is a small sample of chorionic villi tissue is than with amniocentesis or chorionic villi
collected. The harvested fetal cells are collected. Chorionic villi tissue originates sampling, so again, this technique may be
then grown in the laboratory until there from the fetus (review Figure 21.8), so advisable only when there is a known risk
are enough of them to examine the most of the cells are fetal cells. The great of certain diseases.

the developing pre-embryo are identical. If the ball of cells cells that is about the same size as the original zygote.
breaks into two groups before differentiation has begun, two Growth does not occur because the ball of cells is travel-
complete and similar individuals may be formed. Identical ing down the oviduct at this time. With no attachment to
twins are always of the same gender and are usually closely the mother, the only energy available is that stored in the
alike in phenotypic appearance. cytoplasm of the cells plus a little glycogen found within
the oviduct itself.
Recap Sperm deposited in the vagina must swim through the ■ Morphogenesis. Throughout development the organism
uterus and up the correct oviduct to meet the egg. Fertilization undergoes dramatic changes in shape and form. Starting
begins when one sperm’s nucleus enters the oocyte and ends as a ball of identical cells at day four, the organism
when the haploid nuclei of sperm and egg fuse, creating a new becomes (in succession) several layers of different types of
diploid cell. Fraternal twins result from the fertilization of two cells, a pre-embryo with a tail and head, an embryo with
separate eggs. Identical twins occur when a single fertilized recognizable human features, and finally a fetus with a
egg divides in two before differentiation has begun. nearly complete human form. This process of physical
change is called morphogenesis (morpho meaning “form;
21.2 Development: Cleavage, shape” and genesis meaning “origin; production”).
■ Differentiation. Individual cells, too, are beginning to
morphogenesis, differentiation, take on specialized forms and functions, a process we
and growth know as differentiation (review Chapter 17). Differentia-
tion of cells, and the development of organs and organ
During development, rapid and dramatic changes in size and
systems as a result of cell differentiation, is the primary
form take place. This involves four processes.
cause of morphogenesis.
■ Cleavage. Cleavage is a series of cell divisions without ■ Growth. Starting about the time that the developing or-
cell growth or differentiation during the first four days fol- ganism becomes embedded in the endometrial lining
lowing fertilization. Cleavage produces a ball of identical of the uterus and begins to receive nutrients from the
489
490 Chapter 21 Development and Aging

mother, the organism begins to grow in size. The Zygote

growth of a human infant from fertilization to
birth is truly spectacular—from a single cell too Diploid
small to be seen to over 10 trillion cells with a com- nucleus
bined weight of 6 or 7 pounds. Every time the cells
divide, the two daughter cells double in mass to Zona
pellucida
prepare for the next cell division. The only time cell
Uterine
division is not accompanied by cell growth is the tube
initial period of cleavage.
Pregnancy is considered to comprise three periods
called trimesters. Each trimester is approximately three
months long, and characteristic events in embryonic and Morula
fetal development take place during each.
Ovary
Zona
Recap The four processes associated with development pellucida
are (1) cleavage, a series of cell divisions producing a ball regresses
of identical cells; (2) morphogenesis, a sequence of physi- Blastocyst
cal changes; (3) differentiation, as cells assume Inner cell
specialized forms and functions; and (4) growth in size. mass
Pregnancy is divided into three trimesters. Hollow cavity
Trophoblast
21.3 Pre-embryonic development:
The first two weeks Figure 21.5 Pre-embryonic development leading up to implantation. After
a series of cleavages the morula enters the uterus and develops into a blasto-
The period of prenatal (“before birth”) human develop-
cyst comprising an outer layer of cells called the trophoblast, a hollow cavity,
ment can be divided into three stages called pre-embry-
and an inner mass of cells. The blastocyst attaches to the uterine wall.
onic, embryonic, and fetal development. Characteristic
processes and changes take place in each stage. How is cleavage different from the normal cycles of cell division seen
During pre-embryonic development (the first two in adult tissues? (Hint: Compare the final size of the blastocyst with
weeks), the conceptus makes its way down the oviduct, the original zygote.)
embeds itself in the endometrial lining of the uterus,
begins to receive nutrients from the mother, and starts to
grow. Cells differentiate into several tissue layers, and the first During the second week the inner cell mass begins to sepa-
stages of morphogenesis begin. rate from the surface of the blastocyst, creating a second hollow
Throughout the pre-embryonic period, the conceptus cavity. The second cavity will become the amniotic cavity filled
is known informally as a pre-embryo because many (if with amniotic fluid. At this point the cell mass, now called the
not most) of the cells are destined to become part of the embryonic disk, differentiates into two cell types called ecto-
placenta, not the embryo. While the conceptus is still in derm and endoderm. The appearance of an amniotic cavity
the oviduct, a series of successive cleavages yields a ball of and of ectoderm and endoderm in the embryonic disk marks
about 32 identical cells called the morula (Figure 21.5). the end of the pre-embryonic period.
The term morula means “little mulberry,” aptly describing Pre-embryonic development is a hazardous time in
its spherical, clustered appearance. which a lot can go wrong. Usually a woman is not even
On about the fourth day the morula is swept into the aware that she is pregnant at this time, and so she may con-
uterus, where it undergoes the first stages of differentiation tinue risky behaviors such as smoking and drinking alcohol.
and morphogenesis. Over the next several days it becomes a On rare occasions the blastocyst implants outside the
blastocyst, a hollow ball comprising (1) an outer sphere of uterine cavity, resulting in an ectopic pregnancy. Most
cells called a trophoblast, (2) a hollow central cavity, and (3) ectopic pregnancies occur in the oviduct, but on rare
a group of cells called the inner cell mass. Only the inner cell occasions they may occur in the abdominal cavity, in an
mass is destined to become the embryo. ovary, or near the cervix. Ectopic pregnancies usually are not
About day six or seven, the trophoblast cells make con- successful because sites other than the uterus either are not
tact with the endometrial lining and secrete enzymes that large enough to accommodate a full-term baby or result in
digest endometrial cells. The creation of a path for the poor placental development. Occasionally an ectopic
blastocyst causes it to burrow inward (Figure 21.6). The pregnancy in an oviduct is successful, but only if the infant is
process by which the blastocyst becomes buried within the delivered surgically as soon as it is able to survive with inten-
endometrium is called implantation. sive care in a hospital.
 Chapter 21 Development and Aging 491

Amniotic cavity
■ Mesoderm. The middle layer, or
mesoderm, becomes muscle,
connective tissue and bone, kidneys
Ectoderm and ureters, bone marrow, testes or
Endometrium Cells derived ovaries, the lining of the blood ves-
from trophoblast Endoderm sels, and other organs and tissues.
■ Endoderm. The innermost layer is
the endoderm. It gives rise to the
liver and pancreas, the alveoli of the
lungs, the linings of the urinary
bladder, urethra and vagina, and sev-
Inner Amniotic Yolk sac Embryonic
cell
eral glands.
cavity disk
mass
Trophoblast Quick Check A certain type of birth
Blastocyst
defect involves an absence of impor-
Figure 21.6 Implantation and the end of pre-embryonic development. Implantation takes tant nerves in various organs. In
place during days 7–14, at the same time that the inner cell mass of the blastocyst is developing many of these patients, the skin and
into the embryonic disk. The inner cell mass separates from the outer ball to become the embry- hair is unusually pigmented as well.
onic disk, comprising two cell types. The embryonic disk is destined to become the embryo. Which germ layer is the likely cause
What does the trophoblast secrete that enables implantation to occur? of this birth defect? Explain your rea-
soning.

Recap During pre-embryonic development, successive cleav- Extra-embryonic membranes

ages yield a morula. Early stages of differentiation and Early in embryonic development, four different extra-embryonic
morphogenesis cause the morula to become a blastocyst, membranes form that extend out from or surround the
which implants in the lining of the uterus. The embryonic disk embryo: the amnion, allantois, yolk sac, and chorion. Most
is destined to become the embryo. of the components of these membranes are either resorbed
during later development or discarded at birth.
The innermost layer is the amnion, also known as the
21.4 Embryonic development: Weeks “bag of waters” (Figure 21.8a). The amnion lines the amniotic
cavity, which is filled with amniotic fluid. The amniotic fluid is
three to eight derived from the mother’s interstitial fluid and is in continuous
From about the beginning of week three until the end of exchange with it. Later in development when the kidneys of
week eight the developing human is called an embryo. the fetus form, the fetus will urinate into the amniotic fluid
During this time growth, differentiation, and morphogene- and the urine will be removed via exchange with the mother’s
sis are especially rapid. All the organs and organ systems are
established, though most of them are not fully functional.
By the end of the embryonic period the embryo has taken Trophoblast Amniotic
on distinctly human features but is still only an inch long. cell cavity

Tissues and organs derive from three germ layers The embryonic
period begins when a third cell layer, called the mesoderm, Ectoderm
appears between the other two layers in the embryonic disk
(Figure 21.7). These three primary tissues, called the germ
layers, represent the precursor (or germ) cells for the four basic Mesoderm Embryonic
tissue types introduced in Chapter 4 (epithelial, muscle, disk
connective, and nervous) and all organs and organ systems in
Endoderm
the body. Differentiation and morphogenesis follow a
predictable pattern, so we know which tissues and organs are
derived from each germ layer. The three germ layers are
Endometrial
■ Ectoderm. The ectoderm is the outermost layer, the one cells
exposed to the amniotic cavity. Tissues derived from the Yolk sac
ectoderm become the epidermis of the skin, the nervous
Figure 21.7 The three primary germ layers. The appearance of a
system, hair, nails, enamel of teeth, parts of the eye, and third type of cell layer (mesoderm) between the ectoderm and endo-
several other organs and tissues (Table 21.1). derm in the embryonic disk marks the start of the embryonic period.
492 Chapter 21 Development and Aging

Table 21.1 Tissues, organs, and organ systems derived from the three primary germ layers
Ectoderm Mesoderm Endoderm

Epidermis of skin, including hair, nails, Dermis of skin Epithelial tissues lining the digestive tract
and glands (except mouth and anus), vagina, bladder,
All connective tissue, cartilage, and bone
and urethra
Mammary glands
All muscle tissue (heart, skeletal, smooth)
Alveoli of lungs
The nervous system, including the brain,
Bone marrow (blood cells)
spinal cord, and all nerves Liver and pancreas
Kidneys and ureters
Cornea, retina, and lens of eye Thyroid, parathyroid, and thymus glands
Testes, ovaries, and reproductive ducts
Enamel of teeth Anterior pituitary gland
Lining of blood vessels
Posterior pituitary gland Tonsils
Lymphatic vessels
Adrenal medulla Portions of the inner ear
Adrenal cortex
Epithelial lining of nose, mouth, and anus

Fetal portion Maternal portion

of placenta of placenta
(chorion) (endometrium)
Placenta

Yolk sac

Amniotic cavity Fetal arteriole

containing
amniotic fliud
Fetal venule
Umbilical cord

Amnion
Uterus
Umbilical cord
Amnion
Connection
to yolk sac
Cavity of uterus

Umbilical Umbilical Pool of Chorionic Maternal Maternal

arteries vein maternal villus vein artery
blood containing
fetal capillaries
a) The fetus is bathed in amniotic fluid within b) A closer view of portions of the placenta and umbilical cord, showing how
the amnion. Its only connection to the mother nutrients and gases are exchanged between maternal and fetal blood
is the umbilical cord. without mixing.

Figure 21.8 The placenta and umbilical cord.

blood. The amnion and amniotic fluid absorb physical shocks, cells until that job is taken over by other tissues, and it is
insulate the fetus, and keep it from drying out. the source of the germ cells that migrate into the gonads
The allantois is a temporary membrane that helps form (testes and ovaries) and give rise to the gametes (sperm
the blood vessels of the umbilical cord. It degenerates dur- and eggs).
ing the second month of development. The outermost layer, the chorion, is derived primarily
The yolk sac forms a small sac that hangs from the em- from the trophoblast of the early blastocyst. The chorion
bryo’s ventral surface. In many species the yolk sac serves a forms structures that will be part of the exchange mechan-
nutritive function, but in humans that function has been ism in the placenta. It is also the source of human chori-
taken over by the placenta, discussed shortly. Nevertheless, onic gonadotropin (hCG), a hormone that supports
the yolk sac is important in humans because it becomes pregnancy for the first three months until the placenta be-
part of the fetal digestive tract. It also produces fetal blood gins producing enough progesterone and estrogen. The
 Chapter 21 Development and Aging 493

presence (or absence) of hCG in the urine is the basis of menstruation does not occur, inhibit uterine contractions
most home pregnancy tests. during pregnancy, and help form a thick mucous plug over
the cervix to inhibit uterine infections.
The placenta and umbilical cord The two umbilical arteries and single umbilical vein are
As the embryo grows and develops, its relationship with the considered part of the embryonic circulation, meaning the
mother becomes more complex. The embryo cannot supply umbilical vein carries blood toward the embryonic heart and
itself with nutrients or get rid of its own wastes, so exchange the umbilical arteries carry blood away from the embryonic
vessels form between the embryo and the mother for the ex- heart and back to the placenta. Because the exchange of nutri-
change of nutrients and wastes without direct mixing of their ents by the placenta requires a functional embryonic heart, the
blood. This is the function of the placenta and the umbilical placenta and umbilical vessels do not become fully functional
cord. The placenta (Figure 21.8b) is the entire structure that until the embryonic heart develops at about five weeks. Until
forms from embryonic tissue (chorion and chorionic villi) then the developing chorion supplies the nutrient needs of
and maternal tissue (endometrium). The umbilical cord is the embryo by diffusion.
the two-way lifeline that connects the placenta to the
embryo’s circulation. Quick Check Name the four extra-embryonic membranes
The placenta develops because the cells of the chorion, and state the major function of each. Which one plays the
just like the cells of the trophoblast from which they derive, largest role in the formation of the placenta?
secrete enzymes that eat away at the endometrial tissues and
capillaries in the vicinity of the embryo. These enzymes rup-
ture the capillaries, causing the formation of small blood- The embryo develops rapidly
filled cavities. The developing chorion extends fingerlike At two weeks the embryo is fully embedded in the endo-
processes called chorionic villi into these pools of maternal metrium, which provides it with nutrients via the develop-
blood. Each chorionic villus contains small capillaries that ing chorion. At about this time the embryo begins to take
are connected to umbilical arteries and veins that are part of shape as cells of the three primary tissue layers migrate
the circulation of the fetus. In other words, the chorion dam- to other locations and begin to form the rudimentary or-
ages the endometrium to cause local bleeding, and then it gans and organ systems. First a small groove called the
taps that bleeding as a source of nutrients and oxygen for the primitive streak appears in the flat, round embryonic disk,
embryo and as a place to get rid of embryonic wastes, includ- and the embryonic disk begins to elongate along one axis
ing carbon dioxide. In effect, an infant behaves like a parasite (Figure 21.9).
within the mother. At three weeks certain precursors to embryonic struc-
The placenta is an effective filter. It permits the exchange tures emerge. A neural groove of ectoderm forms that will
of nutrients, gases, and antibodies between the maternal and later become the brain and spinal cord. Meanwhile, the
embryonic circulations but not the exchange of large proteins mesoderm begins to separate into several segments called
or blood cells. The placenta allows the embryo to take advan- somites that will become most of the bone, muscle, and
tage of the mother’s organ systems until its own organs skin. Prominent bumps called the pharyngeal arches
develop and become functional. appear at one end; they are destined to become part of the
However, the placenta may permit certain toxic face, neck, and mouth. By the end of week four the heart
substances and agents of disease to cross over to the fetus as is beginning to develop, the head begins to take shape, the
well. Examples include alcohol, cocaine, the HIV virus, and a position of the eyes becomes apparent, the neural groove
wide variety of prescription and nonprescription drugs. has closed into a neural tube, and four limb buds and a
Many of these substances can do a great deal of harm to the tail appear (Figure 21.10).
embryo when it is in the earliest stages of differentiation and The patterns of cell differentiation and organism mor-
morphogenesis, even though some of the prescription drugs phogenesis are so consistent among vertebrates that even
may be therapeutic for the mother (review Chapter 17). an expert would have a hard time telling whether a four-
Upon learning she is pregnant, a woman should review all week-old embryo of unknown origin was destined to
her prescription and nonprescription drugs with her become a fish, a mammal, or a bird. Because early differen-
physician. tiation and morphogenesis follow such a common path, it
The placenta is also an endocrine organ. Initially the is likely that these patterns developed early in the history of
placenta secretes hCG so that the corpus luteum will the vertebrates.
continue to secrete the progesterone and estrogen necessary Weeks five through eight mark the transition from a
to maintain the pregnancy (Chapter 16). Later the placenta general vertebrate form to one that is recognizably human.
secretes its own progesterone and estrogen, and the corpus The head grows in relation to the rest of the body, the eyes
luteum regresses. Progesterone and estrogen promote the and ears are visible, and the four limbs are formed with dis-
growth of the myometrium in preparation for intense tinct fingers and toes. A cartilaginous skeleton forms. The
contractions at birth, maintain the endometrial lining so heart and circulatory system complete their development
494 Chapter 21 Development and Aging

Primitive streak Neural groove Future brain Pharyngeal arches

Somites

Day 14 Day 15 Days 19–23 Day 25

Figure 21.9 Embryonic development during the third and early fourth week. Day 14 represents
the flat embryonic disk as it would look from above (from the amniotic cavity). On day 15 the
embryo begins to elongate and the primitive streak of ectoderm appears. Days 19–23 mark the ap-
pearance of a neural groove of ectoderm. Somites develop in the mesoderm that will become bone,
muscle, and skin. By day 25, pharyngeal arches that will contribute to structures of the head
become visible. The embryo on day 25 is viewed from the side.

and the umbilical cord and placenta become Week 4 Yolk sac
functional, so that now blood circulates Embryo Actual length
throughout the fetus. However, the blood Connecting stalk
cells for the fetal blood are produced by the
yolk sac because the blood-forming tissues
of the fetus are not yet mature. Nutrients Forebrain
and wastes are now exchanged more Future lens
efficiently and in greater quantities. The tail Pharyngeal arches
regresses. At the end of the embryonic Developing heart
period, the embryo is an inch long and Upper limb bud
weighs just 1 gram (Figure 21.11).
Somites
Although it is difficult to estimate how
many embryos fail to survive to the end of Neural tube forming
Lower limb bud
the embryonic period, some estimates place Tail
the number as high as 20%. Spontaneous ter-
mination of pregnancy followed by expulsion Figure 21.10 Week four of development. The future head is prominent, the
of the embryo is called a miscarriage, or position of the eyes becomes apparent, the heart and neural tube are forming, and
spontaneous abortion. Sometimes a woman limb buds appear.
never even realizes she was pregnant. Miscar-
riages are probably nature’s way of weeding out embryos with
genetic disorders that might prevent them from developing gases between embryo and mother, and secretes hormones.
normally. The umbilical cord joins the embryo to the placenta. By the
fifth week the embryo is becoming distinctly human in form,
and by eight weeks it is an inch long.
Web Animation Embryonic Development at
www.humanbiology.com
21.5 Gender development
Recap By the beginning of embryonic development the em- begins at six weeks
bryo comprises three primary germ layers, called ectoderm, Male and female external and internal genitalia do not
mesoderm, and endoderm, that ultimately give rise to fetal begin to develop until about six weeks. Until that time the
tissues and organs. Four extra-embryonic membranes embryo remains “sexually indifferent,” meaning that it still
(amnion, allantois, yolk sac, and chorion) serve varying has the capacity to develop into either a male or a female.
supportive functions. The placenta exchanges nutrients and However, the final outcome (male or female) is not in
 Chapter 21 Development and Aging 495

suppresses the development of the female internal and external

genitalia.) In males the undifferentiated urogenital groove closes
up to become the urethra within the penis. The penis elongates,
and the previously undifferentiated bud becomes the head of
the penis. The urogenital swellings develop into the scrotum
into which the testes later descend.
In the absence of a Y chromosome, female internal and ex-
ternal genitalia develop and the embryo becomes a female. The
urogenital groove expands to become the vagina, and the un-
differentiated bud becomes the clitoris. Meanwhile the ovaries,
fallopian tubes, and uterus all develop internally. The develop-
ment of female internal and external genitalia is the “default”
condition; it occurs in the absence of any hormones, whenever
a Y chromosome is not present. In other words, gender is deter-
mined by the presence or absence of a Y chromosome, not by
whether the embryo has one or two X chromosomes.

Recap Gender development begins at about 6 weeks. The

Figure 21.11 Week eight of development. The embryo is now presence of a Y chromosome signals the embryo to develop
distinctly human in appearance. The face, limbs, hands, and feet
into a male; the absence of a Y chromosome causes the
are well formed. The circulatory system is functional, and blood
now circulates throughout the embryo and to and from the placenta. embryo to develop into a female.
Actual length is 1 inch.

Bud
doubt; it was determined at the moment
Urogenital groove Labioscrotal swelling
of conception by the single sperm that
fertilized the egg.
The development of either male or
female external genitalia is shown in
Figure 21.12. By six weeks an undifferen- Y chromosome Y chromosome
tiated urogenital groove has developed, present 6 weeks absent
topped by a bud and surrounded by
labioscrotal swellings. After six weeks, the
presence of a Y chromosome causes the em-
bryo to start developing male sexual charac-
teristics. The process begins when a gene on
the Y chromosome called SRY (for Sex-
determining Region Y) is switched on. SRY
encodes for a protein called testis-determining
factor that directs the initial development of
the testes internally. Shortly thereafter the
10 weeks
testes begin to secrete testosterone, which in
turn stimulates the development of the male
Development requires
internal ducts and external genitalia. (The testosterone from
Development does
embryonic testes also secrete a second hor- not require hormones.
developing testes.
mone called anti-Mullerian hormone that

Figure 21.12 Development of male or

female external genitalia.
Occasionally an embryo has two X chromo- Testes
Vaginal
somes and a Y chromosome (three sex chro- opening
mosomes in total). What will happen to this
embryo during sexual differentiation?
Explain your answer. Birth approaching
496 Chapter 21 Development and Aging

21.6 Fetal development: Nine weeks The heartbeat is now loud enough to be heard with a stetho-
scope. Skeletal hardening continues.
to birth The sixth month marks the first point at which the fetus
After the eighth week the developing human is called a could, with the best neonatal care available, survive outside
fetus. This is the time when fetal organ systems begin to the uterus. At this stage the fetus seems to respond to external
function and the bones start to calcify. The mother, too, un- sounds. Most importantly, the lungs begin to produce surfac-
dergoes obvious changes in physical shape. Next, we exam- tant, the substance that reduces surface tension in the lungs
ine each of these stages in detail. and permits the alveoli to fill with air at birth.
The end of the sixth month marks the end of the second
trimester. At this point the fetus weighs nearly 700 grams, or
Months three and four
approximately 11⁄2 pounds.
During the third month the kidneys develop sufficiently for
the fetus to begin eliminating some wastes as urine into the Months seven through nine
amniotic fluid. The limbs are well developed, the cartilaginous
The seventh through ninth months (the third trimester) are a
skeleton begins to be replaced with bone, and the teeth form.
period of continued rapid growth and maturation in prepa-
The spleen participates briefly in the production of red blood
ration for birth. The eyes open and close spontaneously and
cells, a job that will soon be taken over by the liver and bone
can be conditioned to respond to environmental sounds. Ac-
marrow. The liver begins to function, and the genitalia are well
tivity increases, as if the fetus is seeking a more comfortable
enough developed that sex can be determined. The end of the
position in the increasingly restrictive space of the uterus.
third month marks the end of the first trimester.
Usually the fetus moves to a position in which the head is
During the fourth month the liver and bone marrow
positioned downward, near the cervix. The skin begins to
begin producing blood cells. The face takes on nearly its final
lose some of its reddish color and its coat of downy hair. In
form as the eyes and ears become fixed in their permanent
the male the testes descend into the scrotal sac. Although
locations (Figure 21.13). In the female fetus, follicles are
neither the lungs nor the digestive systems have had a chance
forming in the ovaries.
to function yet, both are now ready.
Overall growth is rapid during this time. By the end of
By nine months the fetus is about 20 inches long and
the fourth month the fetus is already 6 inches long and
weighs approximately 6–71⁄2 pounds. Birth usually occurs at
weighs about 170 grams (6 ounces).
about 38 weeks of development.

Months five and six

Recap The period of fetal development extends from nine
By the fifth month the nervous system and skeletal muscles weeks to birth at 38 weeks. Growth is rapid, with the mature
are sufficiently mature for the fetus to begin moving. These fetus weighing approximately 6–71⁄2 pounds at birth. The fetus
movements are called quickening, and the mother may feel begins to move at about five months, and life outside the
them for the first time. The skin, which is well formed womb is at least possible by about six months when the lungs
although red and wrinkled, is covered by soft, downy hair. begin to produce surfactant.

21.7 Birth and the early postnatal period

Other than perhaps conception and death, there is no period
in our lives marked by greater developmental change than
the hours during and immediately after birth. Within minutes,
the newborn makes the shift from relying solely on the mater-
nal circulation for nutrient and gas exchange to depending on
its own circulation and lungs. The digestive tract, too, starts to
function with the first swallow of milk. Let’s take a quick look
at the changes that occur during birth and the postnatal (“after
birth”) period.

Labor ends in delivery

Birth involves a sequence of events that we call labor (it’s
hard work!), which ends in the delivery of the newborn
into the world.
Labor begins as a result of a series of events that is
Figure 21.13 The fetus and placenta at four months of devel- triggered by maturation of the fetal pituitary gland, which
opment. Note the amnion and the clear amniotic fluid surrounding serves as the timing device to indicate that the fetus is now
the fetus. Actual length is 6 inches. ready for birth.
 Chapter 21 Development and Aging 497

The mature fetal pituitary gland begins to release ACTH, ■ Stage 1—dilation. The first phase of labor can last 6–12
which stimulates the fetal adrenal gland to secrete steroid hours. During this time the rate, duration, and strength
hormones that cause the placenta to increase its production of contractions increase over time, pushing the head of
of estrogen and decrease the production of progesterone. Es- the fetus against the cervix. The cervix itself is drawn
trogen, in turn, increases the number of oxytocin receptors back toward the uterus, widening the cervical opening
and stimulates production of prostaglandins. Together, the and expelling the mucus plug. Continued pressure of the
increased estrogen, prostaglandins, and oxytocin receptors fetus against the cervix widens the cervical opening even
(along with the mother’s oxytocin) stimulate the uterus to further until eventually it is large enough to accommo-
contract rhythmically. date the baby’s head, or about 10 centimeters. At about
A positive feedback cycle begins: rhythmic contractions this time the pressure of the baby’s head ruptures the
of the uterus cause the release of still more oxytocin from the amnion, releasing the amniotic fluid. This “breaking of
maternal pituitary, which increases contractions still further, the water” is a normal sign that delivery is proceeding.
and so on. As labor progresses the periods of contraction get The physician keeps a close watch on the stage of dila-
closer together, last longer, and become stronger, creating tion by measuring the degree of cervical dilation at regu-
enough force to push the fetus toward the cervix and eventu- lar intervals.
ally through the vagina. ■ Stage 2—expulsion. The period of expulsion extends
The period of labor and delivery lasts about 24 hours for from full cervical dilation through actual delivery. During
the first birth and slightly less time for subsequent births. this time uterine contractions strengthen and the woman
Labor and delivery are divided into three phases: dilation, experiences an intense urge to assist the expulsion with
expulsion, and afterbirth (Figure 21.14). voluntary contractions as well. To make the birth easier,

Placenta

Urinary bladder Ruptured

Pubic bone amnion

Urethra

Vagina

Cervix
9-month-old fetus
Rectum

a) 9-month-old fetus. As birth approaches, b) Stage 1: dilation. The cervical opening widens.
the fetus usually is positioned with the head The amnion may break at this stage.
down and toward the cervix.

Placenta

Uterus
Umbilical
cord

c) Stage 2: expulsion. The fetus passes headfirst d) Stage 3: afterbirth. The placenta detaches
through the cervical canal and the vagina. from the uterus and is expelled along with the
remainder of the umbilical cord.

Figure 21.14 The stages of birth.

498 Chapter 21 Development and Aging

some women opt for a surgical incision called an But what causes the infant to take that first breath? During
episiotomy to enlarge the vaginal opening. The intense labor, the placental connection to the mother begins to separate,
contractions push the baby slowly through the cervix, reducing gas exchange with the fetus. Clamping the umbilical
vagina, and surrounding pelvic girdle. The appearance of cord stops gas exchange entirely. This situation is equivalent to
the baby’s head at the labia is called crowning. As soon as someone covering your mouth and nose completely. Within
the baby’s head appears fully, an attendant removes mucus seconds the carbon dioxide concentration in the fetus rises to
from the baby’s nose and mouth to facilitate breathing. the point that the respiratory centers in its brain stimulate
The rest of the body emerges rather quickly. When the respiration. With the enormous effort of one who is being
baby has fully emerged, the umbilical cord is clamped and smothered, the infant takes its first breath and begins to cry.
then cut. By this time the infant should be breathing on its
own. The entire expulsion phase lasts less than an hour.
Changes in the cardiovascular system The fetal cardiovascular
■ Stage 3—afterbirth. Contractions do not stop with
system is different from the cardiovascular system described in
birth. Strong postdelivery contractions serve to detach
Chapter 8. While the fetus is still in the womb the fetal lungs
the placenta from the uterus and expel the umbilical
are not yet useful for gas exchange. The fetus receives its
cord and placenta, collectively called the afterbirth. The
nutrients and exchanges gases via blood that travels to and
afterbirth stage usually lasts less than half an hour after
from the fetus in the umbilical cord. Immediately after birth
the birth of the infant.
the umbilical circulation ceases, and pulmonary circulation
and gas exchange becomes crucial to survival.
Cesarean delivery: Surgical delivery of a baby
Let us follow the circulation of blood between mother
Some deliveries are not allowed to proceed as described above. and fetus, and also within the fetus, starting at the point that
Surgical delivery of a baby is called a cesarean delivery, or C- oxygen-laden, nutrient-rich blood from the placenta enters
section. In a C-section, an incision is made through the abdom- the fetus via the umbilical vein (Figure 21.15a). Some of the
inal wall and uterus so the baby can be removed quickly. blood is carried to the fetal liver either directly or via a
A C-section is often performed when the position or size branch of the umbilical vein that joins the hepatic portal
of the fetus could make a vaginal delivery dangerous. Possi- vein. But the fetal liver is not yet fully developed and cannot
ble reasons include a fetus that is too large for the birth handle the entire umbilical blood flow. Most of the blood
canal, improper position of the fetus with the legs near the bypasses the liver and is carried directly to the inferior vena
cervix, maternal exhaustion from a long delivery, or signs of cava by the ductus venosus. In the inferior vena cava, the nu-
fetal distress such as an elevated fetal heart rate. trient-rich blood mixes with the venous blood of the fetus.
In the past, if a C-section was performed it was common Most of the blood that enters the fetal heart must bypass
practice to do all subsequent deliveries by C-section as well. the fetal lungs because they too are not fully developed. Some
Today most women who have a C-section can still opt for a of the blood passes from the right atrium to the left atrium
vaginal delivery with the next child if they wish. through the foramen ovale, and some is shunted from the pul-
monary artery directly to the aorta via the ductus arteriosus.
The transition from fetus to newborn Even though the fetal lungs are not yet functional and
Somewhere between the point when its head emerges and the the digestive tract is not receiving nutrients, the aortic blood
umbilical cord is clamped, the fetus must become capable of still has sufficient oxygen and nutrients (brought to the fetus
sustaining life on its own. The first necessity is for the newborn via the umbilical vein) to supply all the fetal tissues. Some of
(also called the neonate for the first 28 days) to start breathing, the fetal arterial blood returns to the placenta via the umbili-
preferably even before its umbilical connection to the mother is cal arteries. In the placenta the blood again picks up
severed. Second, over the first few days of life, its cardiovascular nutrients and oxygen and gets rid of carbon dioxide
system undergoes a series of remarkable anatomical changes generated by fetal metabolism.
that reflect the loss of the placental connection for gas and nu- The situation changes dramatically at birth, for the um-
trient exchange. The newborn is now entirely on its own. bilical circulation is immediately cut off. Over time the um-
bilical vein, umbilical arteries, and ductus venosus regress to
Taking the first breath When the infant emerges, the three become vestigial connective tissue. The foramen ovale and
hundred million alveoli in its lungs have never been inflated. ductus arteriosus close off in the first few days or weeks after
After all, the infant essentially has been living underwater birth as well, so that all the cardiac output passes through
(more correctly, under amniotic fluid). The first inflation is the lungs for efficient gas exchange (Figure 21.15b). Nutri-
critical and not all that easy. If you’ve ever tried to blow up a ents are now absorbed by the newborn’s digestive tract and
really stiff balloon, you know that it takes a lot of effort to pass through the liver via the hepatic portal vein. The only
start the inflation. The reason for this phenomenon is that reminder of the intimate connection between fetus and
the surface tension generated by any small sphere is greater mother is the umbilicus (navel).
than the surface tension generated by a larger sphere. The To summarize the unique features of the fetal circulation:
first inflation of the alveoli (representing small spheres) is
facilitated by substances called pulmonary surfactants that ■ Blood enters the fetus via the umbilical vein. Some
reduce alveolar surface tension. of the blood flows through the liver, but most of it
 Chapter 21 Development and Aging 499

High oxygenation Moderate oxygenation Low oxygenation Very low oxygenation

Aorta

Ductus arteriosus Pulmonary

Superior (becomes connective artery
vena cava tissue) Left atrium
Right atrium Left
ventricle
Foramen ovale
Right Lung
(closes shortly after
ventricle
birth)
Inferior
vena cava Hepatic
portal vein
(from
Ductus venosus digestive
Liver (becomes connective organs)
tissue)
Abdominal
aorta

Umbilical vein Inferior

(becomes connective vena cava
tissue)
Umbilical
cord Common
Umbilical arteries iliac artery
(become connective
tissue)

a) Fetal circulation. b) Circulation after birth.

Figure 21.15 The cardiovascular systems of the fetus and of the newborn. In the fetus the im-
mature lungs are not used for gas exchange, so most of the blood returning to the heart bypasses the
lungs by flowing through either the foramen ovale or the ductus arteriosus. Gas exchange occurs via
the placenta, so special fetal blood vessels connect the fetus to the placenta. These unique
pathways for blood circulation in the fetus close shortly after birth.

What would happen if either the ductus arteriosus or the foramen ovale did not close after birth?

bypasses the liver and joins the inferior vena cava of the Lactation produces milk to nourish the newborn
fetus via the ductus venosus (venous duct). The intimate relationship between mother and child does not
■ Blood leaves the fetus via two umbilical arteries that end at birth, for the human infant must rely entirely on its
originate from arteries in the lower extremities. mother (or someone else) for all its nutritional needs. During
■ In the fetus there is a hole between the two atrial cham- pregnancy, high concentrations of estrogen and progesterone
bers of the heart called the foramen ovale (oval open- cause the woman’s breasts to enlarge in preparation for
ing). The hole permits some blood to pass from right lactation (milk production). However, the breasts do not secrete
atrium to left atrium, bypassing the pulmonary milk before childbirth because estrogen and progesterone
circulation. The hole closes after birth. prevent the action of prolactin, the hormone that actually stim-
■ In the fetus there is a shunt, or shortcut, from the pul- ulates milk production.
monary artery directly to the aorta called the ductus ar- During the first day or so after birth, the breasts produce a
teriosus (arterial duct). Consequently, most of the watery milk called colostrum that is rich in antibodies but
blood pumped by the right ventricle into the pul- low in fat and lactose. The antibodies help to protect the new-
monary artery bypasses the lungs. The ductus arteriosis born until its own immune system matures. Later, the high fat
closes after birth. and lactose content of milk help the infant to grow rapidly.
A fourth hormone, oxytocin, is responsible for the con-
tractions that deliver colostrum or milk only when needed.
Quick Check Infants born prematurely often have not yet Oxytocin is released during childbirth and also as a result of
produced surfactants. What problems might this cause, and a neural reflex every time the infant nurses. The expulsion of
how could it be treated? milk from the breast during suckling is called milk ejection.
500 Chapter 21 Development and Aging

Recap The period of labor and delivery is composed of

three stages; dilation, expulsion, and afterbirth. The dila-
tion phase may last 6–12 hours. At birth a sharp
increase in carbon dioxide causes the newborn to take
the first breath. Shortly after birth, anatomical changes
in the newborn route all blood through the lungs. In the
mother, prolactin stimulates lactation, and oxytocin
stimulates the release of milk during suckling.

2 3 newborn 2 5 13 22 80
months months years years years years years
21.8 From birth to adulthood Figure 21.16 Changes in body form and relative proportion throughout
The long process of human development continues prenatal and postnatal growth and development.
after birth. Because development is a continuum,
experts do not always agree on how to categorize its
stages. Some categories may seem somewhat arbitrary, The neonatal period: A helpless time
such as the distinction (if there is one) between The human neonate (newborn) is fairly helpless. In particular,
adulthood and old age. At the risk of oversimplifying, we the nervous system and the muscular system are relatively im-
define the neonatal period as the first month, infancy as mature at birth. Consequently, movements are uncoordinated,
months 2–15, childhood as the period from infancy to weak, and governed primarily by reflexes rather than con-
adolescence, adolescence as the transitional period from scious control. The neonate hardly has enough strength to
childhood to adulthood (approximately 12 to 18 or 20 hold its head up, so its head must be constantly supported
years of age), and adulthood as 20 years and beyond. whenever it is carried. The eyes are open but often they seem
Table 21.2 summarizes the stages of human develop- to wander about, unable to focus. Neonates are aware of their
ment. Bear in mind that these ages are approximate, environment and can respond to it, but they are unable to re-
intended as a guide to orient you with regard to your tain any long-term memories. (What is your first memory?)
progression through life. We concentrate primarily on bio- They hear a narrower range of pitches than adults and are
physical development, leaving human psychosocial and more sensitive to higher-pitched tones.
cognitive (learning) development to the fields of psychol- What neonates do best is suckle, urinate, and defecate.
ogy and education. Figure 21.16 depicts the changes in The digestive system can absorb the constituents of milk but
body form that accompany human development to is not yet ready for solid foods. The stools are soft, and defe-
adulthood. cation and urination occur as reflexes.

Infancy: Rapid development and maturation

Table 21.2 A summary of the stages of human development
of organ systems
Stage Age Infancy is a time of rapid change. In just 15 months the
Prenatal (after fertilization) infant more than triples in weight to about 22–24 pounds.
The bones harden gradually, and there is a disproportionately
Pre-embryo From fertilization to 2 weeks
large increase in muscle mass and strength. The brain also
Zygote Days 1–3 grows rapidly, with nearly half of all brain growth after birth
Morula Days 4–7 occurring in infancy. Most of the growth occurs in the cere-
Blastocyst Days 8–14 bral cortex, the area associated with sensory perception,
Embryo Weeks 3–8 motor function, speech, and learning. Most of the myelina-
tion of nerves occurs in infancy as well. The first teeth appear
Fetus Weeks 9–40 (birth)
at about six months, and all 20 baby teeth are generally pres-
Postnatal (after birth) ent by about one year. Most infants begin eating solid foods
by about six months to a year.
Neonate The first month
By about 14 months the combined maturation of the
Infant Months 2–15 musculoskeletal and nervous systems allows the human infant
Child 16 months to 12 years to begin to walk on its own. A rapidly growing body needs lots
Adolescent 12–20 years of rest, and infants sleep a lot. Infants can easily sleep 10 hours
a night and still take several naps during the day.
Adult Beyond 20 years
The immune system consistently lags behind most other
Old age Aging accelerates after systems in its development. Most vaccinations are ineffective
40 years in infants because their immune systems cannot produce the
 Chapter 21 Development and Aging 501

appropriate antibodies. The schedule of vaccinations in infants emotion don’t fully mature until about 18 years of age. This
and children is designed so that each vaccine is given only after may be why teenagers, who may seem to be responsible in
the immune system becomes able to respond to that vaccine. most of their decisions, still sometimes make bad judgments
when they are under intense peer pressure.
Childhood: Continued development and growth
The long period of childhood, from about 15 months to 12 Recap Maturation of organ systems occurs at different rates.
years of age, involves continued growth of all systems. The At birth, the neonate is physically helpless and cannot form
brain grows to 95% of its final size during childhood. The memories. Infants begin to eat solid foods and to walk. Brain
immune system continues its slow process of maturation growth is nearly complete by the end of childhood. Muscle
throughout childhood and even into adolescence. Body strength and coordination continue to improve throughout
weight increases to an average of about 100 pounds in both childhood, and the body changes to a more adult form. Ado-
boys and girls as the result of periods of slow growth lescence is accompanied by a growth spurt and by maturation
interspersed with growth spurts. Both muscle strength and of the reproductive systems.
fine motor coordination improve.
In addition, body form alters as the long bones of the
arms and legs lengthen. A lumbar curve develops in the
21.9 Aging takes place over time
small of the back, and abdominal musculature strengthens, Physiologically, at least, humans reach their peak in early
so that the childhood “pot belly” disappears. adulthood at about 20 years of age. Most body systems are
By the end of childhood the body form is distinctly adult fairly stable from 20 to about 40, and then the process that
like, though still sexually immature. Most organ systems are we call aging seems to accelerate. No matter whether we exer-
fully functional or nearly so (although not necessarily their cise regularly, eat right, and in general take care of ourselves,
full adult size). The lone exceptions are the male and female there still seems to be at least some “natural” aging process,
reproductive systems, which have not yet matured. just like the carefully timed sequence of events that occurs in
human development. To be sure, exercise and a proper diet
Adolescence: The transition to adulthood do help, but in the end we all must face the fact that we will
One of the most challenging times of life is the transition not be the same at age 80 as we were at 20. The big question,
from childhood to adulthood. The final growth spurt, one of of course, is why we age and whether the aging process can
the largest, occurs in adolescence. The growth spurt begins at be halted or at least slowed.
different times in different individuals, so thereis a wide Aging is the process of change associated with the
range of normal body weights and degrees of sexual maturity passage of time (Figure 21.17). Although the word does not
in the 12- to 14-year-old age group. Rapid growth can lead to necessarily imply deterioration (after all, some wines get bet-
awkwardness and a temporary loss of coordination. ter as they age), human aging invariably leads in that
Both the skeletal and muscular systems experience their direction. Aging as we use the term in biology could be used
greatest rates of growth during adolescence. Other organs in- synonymously with senescence, the progressive deterioration
crease in size as well. Notably, the lungs more than double of multiple organs and organ systems over time. Human
in mass, and the kidneys and stomach increase by more than senescence (or aging) has no apparent or obvious cause—
50%. The brain, however, gains only about 5%. that is, it cannot be blamed on a particular disease or event.
Adolescence is marked by maturation of the reproductive
systems and the human sexual response, an event known as
puberty. Puberty is defined as the onset of menstrual periods
(menarche) in the female at some time between 91⁄2 and 151⁄2
years and by reflex discharge of semen during sleep (nocturnal
emissions) in the male sometime between 111⁄2 and 171⁄2 years.
The concentrations of all the reproductive hormones are
low in children, but they begin to rise during the period of
sexual maturation (known as the pubertal period). The puber-
tal period is initiated by maturation of certain neurons of the
hypothalamus, which begin to secrete a releasing hormone
called GnRH (Gonadotropin Releasing Hormone). The initia-
tion of GnRH secretion by the hypothalamus stimulates the
secretion of luteinizing hormone (LH) and follicle-stimulating
hormone (FSH) from the anterior pituitary, which in turn
stimulate the production of sex hormones and the maturation a) Rolling Stones’ Mick Jagger b) Jagger at age 60 on
at age 23, in 1967. December 12, 2003, the night
of sex organs and secondary sexual characteristics. he became ‘‘Sir Mick.’’
The brain is one of the last organs to reach full maturity.
The areas of the cerebral cortex that regulate impulse and Figure 21.17 Aging.
502 Chapter 21 Development and Aging

A common statistic is that the average life span in the out of telomeres, cell division begins to eat away at vital ge-
United States has increased from 47 years in 1900 to the netic information. Eventually enough vital genetic informa-
upper 70s today. Does that mean we are aging more slowly tion is lost that the cell stops dividing and dies.
now? Not at all. Longevity—how long a person lives—does However, cancer cells manufacture an enzyme called
not depend solely on the aging process. Most experts believe telomerase that builds new telomeres to replace the ones lost
that our increase in longevity is largely due to a reduction in during cell division. There has been a lot of excitement about
deaths by accident or disease, not to a slowdown in the aging telomerase lately as a result of these findings. Some researchers
process itself. have even suggested that telomerase may be the magic bullet
In fact, it appears that nothing we have done (so far!) has that could reverse or prevent the aging process, or that an
decelerated the aging process. If we could solve every known antitelomerase substance might offer the cure for cancer.
disease and prevent every accidental death, would we still These ideas remain far-fetched at the moment; under-
come face to face with a natural endpoint of life? Nobody standing why a single cell stops dividing does not tell us how
knows for sure, but so far the answer seems to be a qualified to arrest the aging process at the level of the whole body.
yes. No one has beaten the aging process yet, and there are After all, aging is not just a decline in cell division but in cell
very few individuals with documented ages of over 100 years. function as well.

What causes aging? Cells become damaged beyond repair The second hypothesis
There are three hypotheses regarding the causes of aging. The holds that over time, accumulated damage to DNA and
first is that there is some sort of internal genetically determined errors in DNA replication become so great that they can no
program that dictates the timing of cell death. The second is longer be countered by the cell’s normal repair mechanisms.
that accumulated cell damage or errors eventually limit cells’ The damage is thought to be caused by certain toxic by-
ability to repair themselves. A third hypothesis is that the products of metabolism, such as oxygen free radicals.
decline in function of a critical body system may lead to the Eventually, like a ticking time bomb, the damage becomes
parallel senescence of other systems. There is evidence to sup- too great to allow survival.
port each hypothesis. According to proponents of this hypothesis, how long
cells live is a function of the balance between how often
An internal cellular program counts cell divisions It has long damage occurs and how much of it can be repaired. Cells
been known that even when cells are grown under laboratory that undergo frequent injury and are slow to repair the dam-
conditions that are ideal for cell growth and division, most age have shorter lives than cells that are rarely damaged and
cells will not continue to divide indefinitely. Most cells go repair themselves efficiently.
through approximately 50–90 successive divisions and then This hypothesis is supported by the finding that severe
just stop. Cancer cells are the exception, for they will con- caloric restriction (reduced food intake) in certain animals
tinue to divide indefinitely. prolongs life. One possibility is that by lowering body tem-
Why do normal cells stop dividing? Apparently every perature just a little, caloric restriction may slow metabolism
strand of DNA ends with a long tail of disposable bits of and reduce the rate at which damaging metabolic by-prod-
DNA called telomeres that do not code for any genes. The ucts form. Some people believe that taking antioxidant vita-
telomeres are like a roll of tickets, and the idea is that every mins may slow the aging process and prolong life, but there
time the DNA is replicated prior to cell division a ticket is not yet enough long-term evidence to know for sure if this
(telomere) is removed (Figure 21.18). When the cell runs is true.

Useful gene Telomere section Aging is a whole-body process Why does aging seem to
section of DNA of DNA happen to virtually all body systems at once, instead of just
one or two systems? The third hypothesis of aging is that
because all the organ systems in a complex organism are
interdependent, a decline in the function of any one of them
eventually impairs the function of the others.
For example, a decline in the secretion of growth hormone
Telomere by the endocrine system could well tip the balance away from
growth and replacement and toward a net loss of cells or organ
Newly forming Old (template) function. Impaired function of the cardiovascular system
strand strand would affect nutrient delivery to and waste removal from
Figure 21.18 Telomeres and aging. Short terminal segments every cell in the body. A critical system in terms of aging may
of DNA called telomeres are removed each time the DNA is repli- turn out to be the immune system. Indeed, the elderly have a
cated. As a result, the new strand will be shorter than the original higher incidence of cancer than the young, which may indi-
strand. When all telomeres are gone, further cell divisions erode cate that the immune system is not recognizing damaged
the useful genes. cells as efficiently as it once did.
 Chapter 21 Development and Aging 503

in muscle mass is a combination of a loss of muscle cells and

a decrease in the diameter of remaining cells. By age 80 the
average person has lost 30% of his or her muscle mass, and
strength declines accordingly. Often the joints become stiff
MJ’s Human Biology Blog and painful because they secrete less joint fluid. Ligaments
and tendons become less elastic.
Caloric Restriction and Longevity The skin becomes thinner, less elastic, and wrinkled.
There are fewer sweat glands, so it becomes harder to adjust
It has been known for some time that severe caloric re- to warm temperatures. Pigmentation may change as well,
striction can retard the aging process and prolong the with the appearance of pigmented blotches called liver spots.
life of a variety of species, from worms to mice. But skep-
Cardiovascular and respiratory systems Lung tissue becomes
tics of the idea that caloric restriction might also slow
less elastic, decreasing lung capacity. The heart walls become
the aging process in humans point out that the metabo-
slightly stiffer due to an increase in the ratio of collagen to
lisms of worms and mice are quite different from that of
humans.
muscle, with the result that an aging heart generates less force.
Now a new study reports that caloric restriction Blood vessels also lose elasticity, so the systolic pressure and
slows the aging process in primates, too. Macaque pulse pressure may rise in elderly people. Overall there is a
monkeys that have been on a calorie-restricted diet reduced capacity for aerobic exercise as aging progresses.
(by 30%) for the past 20 years are living longer and Immune system The thymus decreases to only 10% of its
are healthier than their age-matched control counter- maximum size by age 60, so there are fewer T cells. The acti-
parts. Excluding animals that died of non-age-related vity, but not the number, of B cells declines as well. Wound
causes (accidents, for example), 50% of the animals
healing slows, and susceptibility to infections increases—
on a normal diet have died of age-related causes, com-
in the elderly, even the flu can become life-threatening. In
pared to only 20% in the restricted-diet group. The
addition the immune system fails to recognize “self” as well as
calorie-restricted animals also have fewer age-associated
it once did, leading to higher rates of autoimmune disorders.
diseases such as diabetes, cancer, and cardiovascular
disease. The study is still ongoing. Nervous and sensory systems Neurons in most areas of the
Reference: Coleman, R.J. et al. (2009). Caloric Restriction Delays Dis-
brain do not divide in adults, so neurons that are lost
ease Onset and Mortality in Rhesus Monkeys. Science 325: 201–204. throughout life generally are not replaced. The brain shrinks
only slightly, however, because the neurons that remain tend
to increase their connections with each other. Nevertheless,
brain and sensory function do decline, in part because of
degenerative changes that occur within the cell bodies of the
It is also possible that aging is due to a specific type of
remaining neurons. Virtually every neural activity is affected,
tissue change that affects many organ systems at once. One
including balance, motor skills, hearing, sight, smell, and
hallmark of aging is that a certain type of protein cross-
taste. In addition to neural changes, sensory organs change
linking increases with age. This cross-linking could con-
as well. For example, hair cells in the inner ear stiffen,
tribute to multiple aging processes, ranging from decreased
reducing hearing, and the lens of the eye stiffens, making it
elasticity of tendons and ligaments to functional changes in
more difficult to focus on near objects. As a result of these
lungs, heart, and blood vessels. Of course, this hypothesis
many changes, elderly people become prone to injury. In
still leaves unanswered the question of what initiates the
addition, they may fail to eat properly because food seems
aging process at the cellular level.
less appealing.
Reproductive and endocrine systems Female reproductive
Quick Check What might be one negative consequence of capacity ceases when a woman reaches menopause (the
increasing the amount of telomerase in cells? (Hint: What cessation of ovulation and menstruation) in her late 40s or
cells often have high levels of telomerase?) early 50s. Menopause occurs because the ovaries lose their
responsiveness to LH and FSH and slowly reduce their
secretion of estrogen. The onset of menopause may be
Body systems age at different rates gradual, with menstrual periods becoming irregular over
several years.
What actually happens during aging is fairly well documented,
The symptoms of menopause are variable. As a result
even if we don’t know exactly why. Next we describe important
of the decline in estrogen, some women experience “hot
developments in each organ system.
flashes” (periods of sweating and uncomfortable warmth).
Musculoskeletal system and skin Bone and muscle mass begin There is a tendency for vaginal lubrication to decline after
to decrease in early adulthood. Bone loss is more marked in menopause, making intercourse potentially less comfortable.
women than in men, especially after menopause. The decline Mood swings are not uncommon. However, female sexual
504 Chapter 21 Development and Aging

efficiently. Nutritional requirements decline somewhat due

to slower cellular metabolism, smaller muscle mass, and,
often, less physical activity. Adequate nutrition can be a
problem if the elderly fail to eat a balanced diet. Tooth loss
MJ’s Human Biology Blog varies widely among individuals, but people who have lost
teeth may find it harder to eat fruit and other healthy but
Hormone Replacement Therapy Revisited fibrous foods.
The liver’s ability to detoxify and remove drugs from the
Back in the 1990s the federal government launched the body declines significantly. As a result, elderly people often
Women’s Health Initiative to investigate the health of require smaller doses of medications to achieve the appropri-
older women. One of its goals was to determine the risks ate therapeutic blood concentration.
and benefits of hormone replacement therapy (HRT) in
Urinary system Kidney mass declines, and both blood flow
postmenopausal women. The HRT study was stopped
and filtration rate may fall by as much as 50% by age 80.
early (in 2002) when it was discovered that HRT led to an
increased risk of breast cancer, heart attacks, and strokes.
However, there is normally such a tremendous reserve of
What about the women who were on HRT during the renal functional capacity that the decline is usually of little
study—do they remain at higher risk even after discon- consequence. The elderly do lose some capacity to respond
tinuing HRT? The good news is apparently most of the to extreme dehydration or overhydration, but otherwise the
increased risk goes away with time. The first follow-up kidneys carry out their functions rather well.
study indicates that three years after HRT is discontin- The primary problem in men is usually prostate hypertro-
ued the increased risk of heart attacks disappears, and phy. The enlarged (hypertrophic) prostate presses against
the increased risk of cancer declines significantly. the ureter, making urination difficult. In women the most
Future reports will show whether any increased risk of common problem is incontinence due to a weakened urethral
cancer remains as time passes. sphincter.
Current recommendations for HRT remain unchanged;
women should consider HRT only if they have moderate to Quick Check Which of the above aging-related declines do
severe postmenopausal symptoms, and for the shortest
you think could be minimized, or even temporarily reversed,
time and lowest dose that is effective.
with regular exercise? Which specific type of exercise might
Reference: Heiss, Gerardo et al. (2008). Health Risks and Benefits benefit specific age-related declines?
3 Years After Stopping Randomized Treatment with Estrogen and
Progestin. JAMA 299: 1036–1045, Mar. 5.
Aging well
It would be easy to get depressed after reading the long list of
declines in function that accompany aging. Bear in mind that
not all of these developments occur to the same degree in
everyone, nor do they all occur at the same age. The effects of
desire (libido) may be undiminished because the source of
aging are normal events in our long and exciting journey
female androgens is the adrenal glands, not the ovaries. After
through life.
menopause, women are at increased risk for cardiovascular
Will a healthy lifestyle that includes exercise and a
disease and osteoporosis. Hormone replacement therapy
proper diet slow the aging process? The answer is far from
with estrogen and progesterone is sometimes prescribed for
certain. However, regular exercise and healthy nutrition can
postmenopausal women to reduce menopausal symptoms
certainly help us age better. Throughout human history,
and other health problems, although the therapy itself adds
longevity has been determined primarily by the prevalence
an additional health risk.
of disease, not by the “natural” length of a human life span
Men exhibit a long, slow decline in the number of viable
(if indeed there is one). Exercise and a healthy diet improve
sperm as they age. In addition, after age 50 men may take
cardiovascular and skeletomuscular fitness and reduce your
longer to achieve an erection. Nevertheless, both sexes can
risk of major killers such as cancer and cardiovascular
and do remain sexually active into old age.
disease. A healthy lifestyle also improves the quality of life,
The rate of decline of various components of the
providing a heightened sense of well-being and increased
endocrine system varies widely. Notably, the secretion of
energy and vigor (Figure 21.19).
growth hormone decreases substantially. However, there is
With these benefits it really doesn’t matter whether a
not an appreciable decline in renin, TSH, ACTH, or ADH.
healthy lifestyle can extend the “natural” human life span
Digestion and nutrition The digestive system continues to from 100 to, say, 105. If you can’t live forever, you can at least
function well except that some vitamins are not absorbed as age well and gracefully.
 Chapter 21 Development and Aging 505

or digestive system, does not necessarily kill immediately.

You can live without either of these systems for days as long
as their loss does not cause one of the critical systems to fail.
The challenge of defining the precise moment of death,
then, is that all of the organ systems don’t necessarily die at
the same time. Death is a process that begins with the failure
of certain life-critical organ systems and then proceeds
through the death of other organ systems, organs, and cells
until the very last of the body’s 100 trillion cells is no longer
alive. That could take hours, days, or (with the assistance of
life-support systems) even years.
Although death may occur gradually, our society requires
a practical way to define the moment at which an individual
dies. We need a definition of death so the living can begin to
grieve and make plans, and so health care workers can stop
the medical treatments that sustain life. We need a legal defi-
Figure 21.19 Aging well. Although it is not known for certain nition so that the police and courts can make a distinction
whether exercise and proper diet actually delay the aging process,
between assault and murder, and so that estates can be
they do improve performance and may provide an enhanced sense
of wellness throughout life. settled. A precise definition of death is especially important
because organs that might still be alive can be transplanted
into another person and save a life.
Recap Aging is a complex process that is still poorly Legal and medical criteria for declaring a person
understood. The number of times a cell can divide may have dead vary according to state law. Basically, all such criteria
an upper limit. In addition, damage may accumulate in cells attempt to define the moment when, to the best of our
until they no longer function properly. All organs and organ knowledge and abilities, the person no longer has even a
systems decline in function with advancing age, though not remote possibility of sustaining life. In general they define
necessarily at the same rate. Even if the aging process cannot death as either (1) irreversible cessation of circulatory and res-
be stopped, it is certainly possible to improve human health piratory functions, or (2) irreversible cessation of all functions of
and wellness throughout life. Regular exercise and healthy the entire brain, including the brain stem. Note that the defini-
nutrition are important factors in aging well. tions focus on critical systems that we cannot live without
for more than a few minutes.
The key, of course, is the word “irreversible.” Usually
21.10 Death is the final transition it is easy to tell when a person has had no respiration or
The final transition in the journey of life is death. In terms of blood flow for 10 minutes or so, long enough to cause
human biology, it is as natural as life itself. brain death. Brain death alone, especially in a person on
Death is defined as the cessation of life. The death of a cardiopulmonary life support, is harder to define. Medical
complex organism such as a human being is not necessarily professionals generally run down a checklist of criteria
accompanied by the immediate death of all its organs or that must be met in order to declare a person’s brain non-
cells. Death occurs when an organ system that is essential for functional, and even then they usually repeat the tests
life in the short term fails to function, making it impossible 6–12 hours later before they are willing to declare irre-
to sustain the life of other systems. versible brain death. Even for the medical profession, the
The critical organ systems whose failures lead to death final transition in life can be difficult to define.
very quickly are the brain (because it controls respiration),
the respiratory system (without oxygen, life cannot be Recap Death is the termination of life. Death starts with the
sustained), and the cardiovascular system (without oxygen failure of one or more critical organ systems and ends ultimately
and nutrient delivery to all cells, life ends). Failure of other with the inability to maintain an internal environment consistent
organs with more long-term functions, such as the kidneys with cellular life.
506 Chapter 21 Development and Aging

From birth to adulthood p. 500

Chapter Summary ■ Human neonates are relatively helpless because the nervous and
muscular systems are not yet mature.
Fertilization begins when sperm and egg unite p. 486 ■ There is a disproportionate increase in the development of the
musculoskeletal and nervous system during infancy. The
■ Sperm and egg meet (and fertilization occurs) in the upper third immune system remains relatively immature.
of the oviduct. ■ Body shape changes in childhood. The brain achieves 95% of its
■ Enzymes in the head of the sperm create a pathway through the adult size.
corona radiata and the zona pellucida to the egg cell membrane. ■ Adolescence is marked by the last spurt of rapid growth and the
■ Entry of the sperm nucleus triggers the completion of meiosis II attainment of sexual maturity.
by the secondary oocyte. Thereafter the nuclei of sperm and egg
fuse, creating a single diploid cell, the zygote. Aging takes place over time p. 501
Development: Cleavage, morphogenesis, differentiation, and ■ Most cells have an internal mechanism that limits the number of
growth p. 489 times they can divide.
■ Cumulative unrepaired cellular damage as a consequence of
■ Development begins by a process called cleavage and then metabolic activity may limit the life of cells.
proceeds with morphogenesis, differentiation, and growth. ■ Aging may affect all organ systems because a decline in one sys-
■ The periods of development prior to birth are known as pre- tem will affect other systems.
embryonic (the first two weeks), embryonic (weeks three
through eight), and fetal (weeks nine to birth). Death is the final transition p. 505
Pre-embryonic development: The first two weeks p. 490 ■ Death is a process. Death begins with the failure of one or more
critical organ systems, leading to the failure of other organ
■ During the first two weeks of development, the single cell devel- systems and eventually to the death of all cells.
ops into a hollow ball with an embryonic disk in the center that
will eventually become the embryo.
■ At about one week the pre-embryo (now called a blastocyst) begins
to burrow into the uterine wall, a process called implantation. Terms You Should Know
Embryonic development: Weeks three to eight p. 491 amnion, 491 fraternal twins, 488
■ Embryonic development is marked by the presence of three pri- amniotic fluid, 491 human chorionic
mary germ layers (ectoderm, mesoderm, and endoderm) in the blastocyst, 490 gonadotropin (hCG), 492
embryonic disk. cesarean delivery, 498 identical twins, 488
■ During development the embryo is completely surrounded chorion, 492 menopause, 503
by two membranes. The amnion contains amniotic fluid, and colostrum, 499 miscarriage, 494
the chorion develops into fetal placental tissue. Two other ectopic pregnancy, 490 morphogenesis, 489
membranes (the allantois and the yolk sac) have temporary embryo, 491 placenta, 493
functions only. embryonic disk, 490 puberty, 501
■ By the fifth week the embryo begins to take on distinctly human endoderm, 491 umbilical cord, 493
features. fertilization, 488 zygote, 486
■ By the eighth week the placenta and umbilical cord circulation
are fully functional, and male or female gonads have begun to
develop. Concept Review
Gender development begins at six weeks p. 494
■ Until six weeks an embryo is “sexually indifferent.” Answers can be found at the Human Biology Place.
■ The presence or absence of a Y chromosome determines whether www.humanbiology.com
the embryo will become phenotypically male or female. 1. Why does only one sperm fertilize the egg, and why is this
important?
Fetal development: Nine weeks to birth p. 496
2. What determines whether an embryo will develop into a male
■ Months 3–9 are marked by rapid growth and development of or a female?
the organ systems. By the fifth month the fetus begins to move. 3. Describe the function of the placenta and the hormones that it
■ By the sixth month, life outside the womb is possible with good secretes.
medical care. 4. Describe the stages of labor and delivery.
5. What causes the newborn to take its first breath immediately
Birth and the early postnatal period p. 496 after birth?
■ Birth occurs at about 38 weeks of development (nine months). 6. Why aren’t all of a child’s vaccinations given in the first year?
■ The three phases of labor are dilation, expulsion, and afterbirth. 7. What causes the onset of puberty (sexual maturation)?
■ Shortly after the newborn takes its first breath, the newborn’s 8. Summarize the three main hypotheses of the causes of aging.
cardiovascular system undergoes substantial changes. Within 9. What has caused the increase in longevity of the U.S.
hours or a few days the umbilical vessels regress and the ductus population during the past 100 years?
arteriosus and foramen ovale close, rerouting all blood through 10. Why is it sometimes so hard to define the moment when an
the pulmonary circulation. individual dies?
 Chapter 21 Development and Aging 507

c. the ductus venosus closes

Test Yourself d. both (a) and (b)
11. stimulates the production of milk while
causes ejection of milk from the breast.
Answers can be found in Appendix A.
a. Estrogen…prolactin
1. Which of the following lists the correct order of structures b. Prolactin…oxytocin
through which a sperm must pass to accomplish fertilization? c. Estrogen…oxytocin
a. zona pellucida, corona radiata, secondary oocyte plasma d. Oxytocin…prolactin
membrane 12. Possible activities of telomerase could be:
b. zona pellucida, secondary ooctye plasma membrane, corona a. stopping the aging process by rebuilding the telomeres
radiata b. contributing to cancer by allowing tumor cells to continue
c. ovum plasma membrane, corona radiata, zona pellucida dividing indefinitely
d. corona radiata, zona pellucida, secondary oocyte plasma c. blocking cancer by restoring telomeres after cell divisions
membrane d. both (a) and (b)
2. Which of the following will result in fraternal twins? 13. The aging of which organ or organ system is least problematic
a. the fertilization of one egg by two different sperm during the normal aging process?
b. the division of a fertilized egg into two masses after differen- a. bones b. immune system
tiation has begun c. nervous system d. kidneys
c. the fertilization of two eggs by two different sperm
14. All of the following are theories regarding causes of aging except:
d. the division of a fertilized egg into two masses before differ-
a. Cells run out of telomeres and stop dividing.
entiation has begun
b. Cortisol production increases and the stress response
3. Which of the following processes is the first to occur as a fertil- contributes to aging.
ized egg begins its journey down a uterine tube toward the c. Cells can no longer repair themselves properly, and damage
uterus? accumulates.
a. differentiation b. cleavage d. A decline in one critical body system induces parallel
c. growth d. all of these choices declines in others.
4. A sperm with a defective acrosome would not be able to: 15. Which hormone plays a critical role both in childbirth as well
a. swim b. penetrate the egg as breast-feeding?
c. penetrate through the cervical mucus a. oxytocin b. prolactin
d. provide the energy for the sperm to swim c. estrogen d. progesterone
5. Which of the following indicates the correct order of pre-
embryonic developmental stages?
a. morula, blastocyst, embryonic disk
b. blastocyst, morula, embryonic disk Apply What You Know
c. trophoblast, blastocyst, morula embryonic disk
d. embyronic disk, blastocyst, morula
Answers can be found at the Human Biology Place.
6. Which of the following is a correct statement?
www.humanbiology.com
a. The placenta is formed entirely from embryonic tissue.
b. The umbilical cord brings maternal blood to the embryo. 1. Why is it biologically impossible for humans and gorillas to
c. The embryonic/fetal blood and the maternal blood mix mate and produce offspring?
within the placenta. 2. Conjoined twins are twins that are attached to each other. They
d. The placenta secretes hormones including HCG and may even share limbs and organs. From what you know of early
progesterone. human development, how do you think conjoined twins come
7. Which of the following is mismatched? about?
a. neural groove: spinal cord 3. Why are many sperm but only one egg produced in the process
b. mesoderm: gastrointestinal tract lining of fertilization? Would humans be able to reproduce if the cell
c. urogenital groove: urethra numbers were inverted?
d. placenta: chorion and endometrium 4. Each year it is estimated that over 40,000 newborn babies are
impacted by the effects of alcohol consumed by the mother
8. Which of the following events is correctly matched with the
during pregnancy. Fetal alcohol syndrome is one of the leading
month of prenatal development?
causes of preventable birth defects. Why do you think alcohol is
a. kidneys producing urine: month 3
more likely to damage the fetus than the mother?
b. heart pumps to circulate blood: month 4
5. What exactly do home pregnancy tests test for as an indication
c. gender development is completed: month 5
that a woman is pregnant?
d. bone marrow takes over blood cell production: month 7
6. Babies born before a full nine months of gestation (premature
9. Which fetal hormone plays a role in initiating labor? babies) are often at a high risk of death. What is the earliest a
a. estrogen b. ACTH baby can be born and stand a good chance of survival, and
c. progesterone d. oxytocin which organ system is generally the limiting one?
10. Which of the following would be detrimental to the health 7. How is it possible that one’s life could be prolonged without
of a neonate? slowing the aging process?
a. the ductus arteriosus stays open
b. the foramen ovale closes