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Michael D. Johnson - Human Biology - Concepts and Current Issues, 6th Edition - Benjamin Cummings (2011) Bab 21
Michael D. Johnson - Human Biology - Concepts and Current Issues, 6th Edition - Benjamin Cummings (2011) Bab 21
Development
and Aging
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
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.
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
Second
meiotic
division
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.
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.
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
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.
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
Yolk sac
Amnion
Uterus
Umbilical cord
Amnion
Connection
to yolk sac
Cavity of uterus
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
Somites
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
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
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.
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
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.
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
Aorta
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
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.
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