Sex Hormones & Differentiation
Sex Hormones & Differentiation
C H A P T E R
5
Sex Hormones and
Sexual Differentiation
CHAPTER HIGHLIGHTS
Sex Hormones Descent of the Testes and Ovaries
Sex Hormone Systems in the Male Brain Differentiation
Sex Hormone Systems in the Female Homologous Organs
Atypical Prenatal Gender Differentiation
Prenatal Sexual Differentiation
Sex Chromosomes Sexual Differentiation during Puberty
Gonads Changes in the Female
Prenatal Hormones and the Genitals Changes in the Male
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SEX HORMONES 97
Sex Hormones
T his Way
Hypothalamus
–
Hypothalamus
Pituitary gland
GnRH
Pituitary
– –
Testosterone
Spermatogenesis
secretion
Ovary Inhibin
Estrogen Testes
Progesterone
Endometrium
SEX HORMONES 99
off. The temperature in the room then gradually falls menstrual cycle (see Chapter 6)
Inhibin: A substance secreted by the
until it produces another change in the thermostat, and during various other stages testes and ovaries which regulates
which then turns on the furnace, and the cycle is such as pregnancy and meno- FSH levels.
repeated. This is a negative feedback loop because pause. The levels of estrogen and
rises in temperature turn off the furnace, whereas progesterone are regulated by the
decreases in temperature turn on the furnace. two pituitary hormones, FSH and LH. Thus the
The hypothalamus, pituitary, and testes form a levels of estrogen and progesterone are controlled
similar negative feedback loop, ensuring that testos- by a negative feedback loop of the hypothalamus,
terone is maintained at a fairly constant level, just as pituitary, and ovaries similar to the negative feed-
the temperature of a room is kept fairly constant. back loop in the male (see Figure 5.3). For exam-
The pituitary’s production of LH stimulates the ple, as shown on the right side of Figure 5.3,
testes to produce testosterone. But when testos- increases in the level of GnRH increase the level of
terone levels get high, the hypothalamus reduces its LH, and the increases in LH eventually produce
production of GnRH; the pituitary’s production of increases in the output of estrogen; finally, the
LH is then reduced, and the production of testos- increases in the level of estrogen inhibit
terone by the testes consequently decreases. When it (decrease) the production of GnRH and LH.
has fallen, the hypothalamus again increases the The pituitary produces two other hormones, pro-
production of GnRH and the process starts again. lactin and oxytocin. Prolactin stimulates secretion of
While the level of testosterone in men is fairly milk by the mammary glands after a woman has
constant, there is probably some cycling, with vari- given birth to a child. Oxytocin stimulates ejection of
ations according to the time of the day and possibly that milk from the nipples. Oxytocin also stimulates
according to the time of the month (see Chapter 6). contractions of the uterus during childbirth.
Although it has been clear for some time that
there is a negative feedback loop between testos-
terone levels and LH levels, it has not been clear
what regulates FSH levels. Inhibin is a substance Figure 5.3 Schematic diagram of
produced in the testes (by cells called the Sertoli hormonal control of estrogen secretion and
cells) that serves exactly that function—it acts to ovum production by the ovaries (during the
regulate FSH levels in a negative feedback loop follicular phase of the menstrual cycle).
(Plant et al., 1993). Note how similar the mechanism is to the
one in the male.
Interest in inhibin has been intense because it
shows great promise, at least theoretically, as a Hypothalamus
male contraceptive. That is, because inhibin sup- –
presses FSH production, sperm production in turn
is inhibited. Future developments in this field
should be interesting.
GnRH
2 Ovary Ovary
We really should say estrogens because they are a group of hor-
mones like the androgens. Estradiol is one of the estrogens. To
keep things simple, we will just use the term estrogen.
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The female sex hormone system functions gonad has two parts, an outer cortex and an inner
much like the male sex hormone system. The medulla), two sets of ducts (the Müllerian ducts
ovaries and testes produce many of the same hor- and the Wolffian ducts), and rudimentary external
mones, but in different amounts. The ovaries, like genitals (the genital tubercle, the urethral folds,
the testes, produce inhibin, which in turn forms a and the genital swelling) (see Figure 5.4, top).
negative feedback loop with FSH production
(Burger, 1993). The functioning of the female sex
hormone system and the menstrual cycle will be Gonads
considered in more detail in Chapter 6. In the seventh week after conception, the sex chro-
mosomes direct the gonads to begin differentia-
tion. In the male, the undifferentiated gonad
develops into a testis at about 7 weeks. In the
female, the process occurs somewhat later, with
Prenatal Sexual Differentiation the ovaries developing at around 13 or 14 weeks.
Sex Chromosomes An important gene that directs the differentia-
As noted above, at the time of conception the tion of the gonads, located on the Y chromosome,
future human being consists of only a single cell, is called SRY, for sex-determining region, Y chro-
the fertilized egg. The only difference between the mosome (Page et al., 1987; Skaletsky et al., 2003). If
fertilized egg that will become a female and the SRY is present, testes differentiate and male devel-
fertilized egg that will become a male is the sex opment occurs (see Figure 5.5 for a summary of all
chromosomes carried in that fertilized egg. If there the genes that regulate sexual differentiation).
are two X chromosomes, the result will typically be Researchers discovered this gene by studying cases
a female; if there is one X and one Y, the result will of abnormal development—for example, adult
typically be a male. Thus, while incredibly tiny, the women who had XY sex chromosomes and were
sex chromosomes carry a wealth of information infertile. The researchers found that these individ-
that they transmit to various organs throughout uals were missing a section of the Y chromosome,
the body, giving them instructions on how to dif- precisely the section containing the SRY. The X
ferentiate in the course of development. The Y chromosome carries a number of genes that con-
chromosome, because it is smaller, carries less trol normal functioning of the ovaries (Winter &
information than the X. Couch, 1995). Surprisingly, a number of genes on
Occasionally, individuals receive at conception the X chromosome affect cells in the testes that
a sex chromosome combination other than XX or manufacture sperm (Wang et al., 2001).3
XY. Such abnormal sex chromo-
SRY: Stands for sex-determining some complements may lead to a
region, Y chromosome. variety of clinical syndromes, such Prenatal Hormones and the Genitals
Müllerian ducts: Ducts found in both
male and female fetuses; in males
as Klinefelter’s syndrome. In this Once the ovaries and testes have differentiated, they
they degenerate and in females they syndrome, a genetic male has an begin to produce different sex hormones, which
develop into the fallopian tubes, the extra X chromosome (XXY). As a then direct the differentiation of the rest of the
uterus, and the upper part of the result, the testes are abnormal, no internal and external genital system (see Figure 5.4).
vagina. sperm are produced, and testos- In the female the Wolffian ducts degenerate,
Wolffian ducts: Ducts found in both
male and female fetuses; in females
terone levels are low (Winter & and the Müllerian ducts turn into the fallopian
they degenerate and in males they Couch, 1995). tubes, the uterus, and the upper part of the vagina.
develop into the epididymis, the vas The single cell divides repeat- The tubercle becomes the clitoris, the folds
deferens, and the ejaculatory duct edly, becoming a two-celled organ- become the inner lips, and the swelling develops
ism, then a four-celled organism, into the outer lips.
then an eight-celled organism, and so on. By 28 The testes secrete Müllerian inhibiting sub-
days after conception, the embryo is about 1 cen- stance (MIS) (Vilain, 2000). MIS causes the Müller-
timeter (less than 1/2 inch) long, but the male and ian ducts to degenerate, while the Wolffian ducts,
female embryo are still identical, save for the sex supported by testosterone, turn into the epi-
chromosomes; that is, the embryo is still in the didymis, the vas deferens, and the ejaculatory
undifferentiated state. However, by the seventh
week after conception, some basic structures have
been formed that will eventually become either a 3
Scientists got so excited by this finding of genes on the X
male or a female reproductive system. At this chromosome acting on the testes that they started calling
point, the embryo has a pair of gonads (each them transsexual genes. Maybe you had to be there.
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Figure 5.4 Development of the male and female external genitals from the undifferentiated
stage. This occurs during prenatal development. Note homologous organs in the female and male.
UNDIFFERENTIATED
Glans area
Genital tubercle
Urethral fold
Urethral groove
Lateral buttress
Anal pit
Anal tubercle
PARTIALLY DEVELOPED
MALE FEMALE
Glans
Urethral fold
Urogenital groove
Labioscrotal swelling
Anal tubercle
Anus
FULLY DEVELOPED
Body of clitoris
Urethral meatus
Inner labia
Scrotum
Outer labia
Vagina
Perineal raphe
thalamic cells are relatively insensitive to estrogen are both homologous (they develop from an indiffer-
levels, whereas female hypothalamic cells are ent gonad) and analogous (they produce gametes and
highly sensitive to them. Male hypothalamic cells sex hormones).
have more androgen receptors (Donahue et al.,
2000).
New magnetic resonance imaging (MRI) studies Atypical Prenatal Gender Differentiation
are giving us a view into the brain of alive, awake Gender is not a simple matter, a fact that is appar-
humans, in contrast to earlier techniques that dis- ent from the preceding discussion. Most people,
sected the brains of, well, dead people and animals. however, assume that it is. That is, people typically
The trade-off, at least for now, is that the MRI meas- assume that if a person is female, she will be femi-
ures are relatively crude, simple assessments of the nine; will think of herself as a woman; will be sexu-
volume or size of certain regions. One of these stud- ally attracted to men; will have a clitoris, vagina,
ies found a larger volume of the hypothalamus and uterus, and ovaries; and will have sex chromo-
amygdala—both being brain regions with high den- somes XX. The parallel assumption is that all males
sities of estrogen and androgen receptors—in men are masculine; think of themselves as male; are
compared with women (Goldstein et al., 2001). sexually attracted to women; have a penis, testes,
Regions of the brain that have few estrogen and and scrotum; and have sex chro-
androgen receptors did not show these gender differ- mosomes XY. Homologous organs (huh-MOLL-
ences in size. Interestingly, a different MRI technique, A great deal of research over the uh-gus): Organs in the male and
female that develop from the same
the functional MRI, or fMRI, has detected increased last several decades challenges embryonic tissue.
activation in one region of the hypothalamus in men these assumptions and provides Analogous organs (an-AL-uh-gus):
who were sexually aroused (Arnow et al., 2002). much information about sexuality Organs in the male and female that
and gender and their develop- have similar functions.
ment. Before the results of this
Homologous Organs research are discussed, however, some back-
The preceding discussion of sexual differentiation ground information is necessary.
highlights the fact that although adult men and We can distinguish among the following eight
women appear to have very different reproductive variables of gender (adapted from Money, 1987):4
anatomies, their reproductive organs have similar
origins. When an organ in the male and an organ in 1. Chromosomal gender. XX in the female; XY in
the female both develop from the same embryonic the male.
tissue, the organs are said to be homologous. When 2. Gonadal gender. Ovaries in the female; testes
the two organs have similar functions, they are said in the male.
to be analogous. Table 5.1 summarizes the major
homologies and analogies of the male and female 4
The distinction between the terms gender and sex,
reproductive systems. For example, ovaries and testes discussed in Chapter 1, is being maintained here.
Table 5.1 Homologous and Analogous Organs of the Male and Female Reproductive Systems
Embryonic Source In the Adult Male In the Adult Female In the Adult Male In the Adult Female
exposed to a high dose of androgens. (Progestin three, not two, gender categories— Androgen-insensitivity syndrome
and androgens are quite similar biochemicals, and and the society in which the (AIS): A genetic condition in which the
in the body the progestin acted like androgen.) In Guevodoces grow up is a three- body is unresponsive to androgens so
genetic females this produced an abnormal, mas- gender society. The third gender is that a genetic male may be born with
culinized genital development similar to that the Guevodoces. Their gender a female-appearing body.
Puberty: The time during which
found in CAH. identity is not male or female but there is sudden enlargement and
The reverse case occurs in androgen-insensitivity Guevodoce. The 5-Alpha Reduc- maturation of the gonads, other geni-
syndrome (AIS) (Wisniewski et al., 2000). In this syn- tase Syndrome has also been found talia, and secondary sex characteris-
drome a genetic male produces normal levels of among the Sambia of New Guinea, tics, so that the individual becomes
testosterone; however, as a result of a genetic condi- who also have a three-gender cul- capable of reproduction.
tion the body tissues are insensitive to the testos- ture. Again we see the profound
terone, and prenatal development is feminized. Thus effect of culture on our most basic ideas about sex
the individual is born with the external appearance and gender.
of a female: a small vagina (but no uterus) and unde-
scended testes. The individual whose poem
appeared at the beginning of this chapter has AIS.
Intersex persons provide good evidence of the
great complexity of sex and gender and their devel- Sexual Differentiation
opment. Many variables are involved in gender
and sex, and there are many steps in gender differ-
during Puberty
entiation, even before birth. Because the process is Puberty is not a point in time but rather a process
complex, it is vulnerable to disturbances, creating during which there is further sexual differentiation.
conditions such as intersex. Indeed, the research It is the stage in life during which the body changes
serves to question our basic notions of what it from that of a child into that of an adult, with sec-
means to be male or female. In CAH, is the genetic ondary sexual characteristics (such as breasts or a
female who is born with male external genitals beard) and the ability to reproduce sexually.
a male or a female? What makes a person male Puberty can be scientifically defined as the time
or female—chromosomal gender? External genital during which there is sudden enlargement and
appearance? Gender identity? maturation of the gonads, other genitalia, and sec-
A related phenomenon was first studied in a ondary sex characteristics, leading to reproductive
small community in the Dominican Republic capacity (Tanner, 1967). It is the second important
(Imperato-McGinley et al., 1974). Due to a genetic- period—the other being the prenatal period—dur-
endocrine problem, a large number of genetic ing which sexual differentiation takes place. Per-
males were born there who, at birth, appeared to be haps the most important single event in the
females. The syndrome is called 5-Alpha Reductase process is the first ejaculation for the male and the
Syndrome. They had a vaginal pouch instead of a first menstruation for the female, although the lat-
scrotum and a clitoris-sized penis. The uneducated ter is not necessarily a sign of reproductive capabil-
parents, according to the researchers, were unaware ity, since girls typically do not produce mature eggs
that there were any problems, and these genetic until a year or two after the first menstruation.
males were treated as typical females. At puberty, a The physiological process that underlies
spontaneous biological change causes a penis to puberty in both genders is a marked increase in
develop. Significantly, their psychological identity levels of sex hormones. Thus the hypothalamus,
also changed. Despite rearing as females, their gen- pituitary, and gonads control the changes.
der identity switched to male, and they developed Adolescence is a socially defined period of devel-
heterosexual interests. In their culture, these people opment that bears some relationship to puberty.
are called Guevodoces (“penis at 12”). Adolescence represents a psychological transition
Anthropologist Gilbert Herdt (1990) is critical from the behavior and attitudes of a child to the
of the research and interpretations about the behavior, attitudes, and responsibilities of an
Guevodoces. The major criticism is that the West- adult. In the United States it corresponds roughly
ern researchers assumed that this culture is a two- to the years from age 10 to age 20. Modern Ameri-
gender society, like the United States, and that can culture has an unusually long period of ado-
people have to fall into one of only two categories, lescence (Steinberg, 2002). A century ago,
either male or female. Anthropologists, however, adolescence was much shorter; the lengthening of
have documented the existence of three-gender the educational process has served to prolong ado-
societies—that is, societies in which there are lescence. In some cultures, in fact, adolescence
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Focus 5.1
The Debate over the Treatment of Intersex Individuals
hen Chris was born, her clitoris was 1.7 cm In the last decade, however, intersex individuals
W long. That’s about halfway between the
length of the average newborn clitoris and
have come out of the closet and formed an activist
organization, the Intersex Society of North Amer-
the average newborn penis. She had a scrotum but ica (ISNA).* Intersex activists argue that they have
no testes in it. The physician was unsure whether cases of genital variability, not genital abnormal-
she was a girl or a boy. A blood test revealed that her ity. The medical standard is that an infant’s organ
sex chromosomes were XY. After 24 hours of con- that is 0.9 cm or less is a clitoris and 2.5 cm or more
sultations, during which her parents were in agony, is a penis. Activists argue that these cutoffs are
the physician decided that Chris should be a girl arbitrary. What is wrong with a clitoris that is 1.7
because it would be impossible for her to function cm long? Perhaps the only thing wrong with it is
as a boy with such strange genitals. While a baby, that it makes doctors, and perhaps parents,
she had several surgeries, one to remove her testes, embarrassed. Issues of medical ethics are raised:
which were still in her abdomen. Her clitoris was Should essentially cosmetic surgery be performed
surgically reduced in size when she was 5, old on a baby who cannot give informed consent?
enough to remember it. Today she is 27 and angry Should parents be encouraged to lie to their child?
about what she considers the mutilations of her Sex researcher Milton Diamond conducted long-
body. She now knows that she has androgen- term follow-ups on several individuals treated using
insensitivity syndrome. So much of her clitoris was Money’s standard protocol. He found that, contrary
removed that she is not able to have an orgasm. to the glowing picture of perfect adjustment
Chris (a composite of several case histories in painted by Money and others, these intersex indi-
the scientific literature) is an intersex individual; viduals had serious adjustment problems that they
that is, her genitals have combined male, female, traced directly to the medical “management’’ of
or ambiguous elements. She was treated according their condition. Diamond’s research has sparked a
to a protocol that became standard beginning in debate over the proper treatment of intersex indi-
the 1960s and persists to the present day. This pro- viduals. Diamond has proposed a protocol in which
tocol was based on the pioneering research of Dr. he urges physicians, in cases of intersex infants, (1)
John Money and others. According to him, individ- to make their most informed judgment about the
uals such as Chris, whom he called “pseudoher- child’s eventual gender identity (CAH girls, for
maphrodites,’’ could successfully be assigned to example, almost invariably have a female identity)
either gender, provided that it was done before 18 and counsel the parents to rear the child in that
months of age and that the necessary surgeries gender; (2) not to perform surgeries that might later
and follow-up medical treatments (such as hor- need to be reversed; and (3) to provide honest coun-
mone treatment) occurred. Money’s research indi- seling and education to the parents and child as he
cated that individuals treated with the standard or she grows up so that the child can eventually
protocol grew up to be healthy and well adjusted. make an informed decision regarding treatment.
does not exist; the child shifts to being an adult and females being out of step with each other at
directly, with only a rite de passage in between. this stage creates no small number of crises for the
Before describing the changes that take place adolescent. Girls are interested in boys long before
during puberty, we should note two points. First, boys are aware that girls exist. A girl may be stuck
the timing of the pubertal process differs consider- with a date who barely reaches her armpits, while a
ably for males and females. Girls begin the change boy may have to cope with someone who is better
around 8 to 12 years of age, while boys do so about qualified to be on the basketball team than he is.
two years later. Girls reach their full height by Second, there are large individual differences
about age 16, while boys continue growing until (differences from one person to the next) in the
about age 18 or later. The phenomenon of males age at which the processes of puberty take place.
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Thus there is no “normal’’ time to begin menstru- years of age (Herman-Giddens et al., 1997). The
ating or growing a beard. Accordingly, we give age ducts in the nipple area swell, and there is growth
ranges in describing the timing of the process. of fatty and connective tissue, causing the small,
conical buds to increase in size. These changes are
produced by increases in the levels of the sex hor-
Changes in the Female mones by a mechanism that will be described
A summary of the physical changes of puberty in below.
males and females is provided in Table 5.2. The As the growth of fatty and supporting tissue
first sign of puberty in the female is the beginning increases in the breasts, a similar increase takes
of breast development, generally at around 7 to 13 place at the hips and buttocks, leading to the
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Girls Boys
1. Growth of breasts 7–13 Pituitary growth hormone, 1. Growth of 10–13.5 Pituitary growth hormone,
estrogens, progesterone, testes, testosterone
thyroxine scrotal sac
2. Growth of pubic 8–14 Adrenal androgens 2. Growth of 10–15 Testosterone
hair pubic hair
3. Body growth 9.5–14.5 Pituitary growth hormone, 3. Body growth 10.5–16 Pituitary growth hormone,
adrenal androgens, testosterone
estrogens
4. Menarche 10–16.5 GnRH, FSH, LH, estrogens, 4. Growth of penis 11–14.5 Testosterone
progesterone
5. Change in About the Testosterone
voice (growth same time
of larynx) as penis
growth
5. Underarm hair About two Adrenal androgens 6. Facial and About two Testosterone
years after underarm hair years after
pubic hair pubic hair
6. Oil- and sweat- About the Adrenal androgens 7. Oil- and sweat- About the Testosterone
producing glands same time producing same time
(acne occurs when as under- glands, acne as under-
glands are clogged) arm hair arm hair
Source: Goldstein (1976), pp. 80–81.
not capable of becoming pregnant until ovulation nisms that control the menstrual cycle shut down
begins, typically about two years after the menar- menstruation.6
che. The first menstruation is not only an impor- Before leaving the topic of running, we should
tant biological event but also a significant note that there is some evidence that serious exer-
psychological one. Various cultures have cere- cise also affects the male reproductive system. One
monies recognizing its importance. In some fami- study of male distance runners found that their
lies, it is a piece of news that spreads quickly to the testosterone levels were only about 68 percent as
relatives. Girls themselves display a wide range of high, on the average, as a control group’s testos-
reactions to the event, ranging from negative terone levels (Wheeler et al., 1984). There are some
ones, such as fear, shame, or disgust, to positive reports of male long-distance runners complain-
ones, such as pride and a sense of maturity and ing of a loss of sex drive, but it is unclear whether
womanliness. this results from reduced testosterone levels or
Some of the most negative reactions occur from the perpetual feelings of fatigue that such
when the girl has not been prepared for the menar- runners have from their intensive training
che, which is still the case surprisingly often. Par- (Wheeler et al., 1984).
ents who are concerned about preparing their Other body changes in girls during puberty
daughters for the first menstruation should include a development of the blood supply to the
remember that there is a wide range in the age at clitoris, a thickening of the walls of the vagina, and
which it occurs. It is not unusual for a girl to start a rapid growth of the uterus, which doubles in size
menstruating in the fifth grade, and instances of between the tenth and the eighteenth years. The
the menarche during the fourth grade, while rare, pelvic bone structure grows and widens, con-
do occur. tributing to the rounded shape of the female and
What determines the age at which a girl first creating a passageway large enough for an infant
menstruates? One explanation is the percent body to move through during birth.
fat hypothesis (Frisch & McArthur, 1974; Hopwood The dramatic changes that occur during
et al., 1990). During puberty, deposits of body fat puberty are produced, basically, by the endocrine
increase in females. According to the percent body system and its upsurge in sex hormone production
fat hypothesis, the percentage of body weight that during puberty. The process begins with an
is fat must rise to a certain level for menstruation increase in secretion of FSH by the pituitary gland.
to occur for the first time and for it to be main- FSH in turn stimulates the ovaries
tained. Thus very skinny adolescent girls would to produce estrogen. Estrogen is Leptin: A protein produced in the
tend to be late in the timing of first menstruation. responsible for many of the body that is related to the onset of
Leptin, a protein manufactured in the body, seems changes that occur; it stimulates puberty.
Adrenal glands (uh-DREE-nul):
to be related to the onset of puberty in girls and in breast growth and the growth of
Endocrine glands located just above
boys as well, although scientists have not yet the uterus and vagina. the kidneys; in the female they are
sorted out the details (Apter, 2003; Phillip & Lazar, Also involved in puberty are the the major producers of androgens.
2003; Wilson et al., 2003). In prepubertal girls and paired adrenal glands, which are Adrenarche (AD-ren-ar-key): A time
boys, leptin levels rise as body fat increases. Leptin located just above the kidneys. In of increased secretion of adrenal
androgens, usually just before age
stimulates the growth of skeletal bone and the the female, the adrenal glands are
eight.
release of LH. the major producer of androgens,
The percent body fat hypothesis also helps to which exist at low levels in females.
make sense of two related phenomena: the cessa- Adrenal androgens stimulate the growth of pubic
tion of menstruation in anorexics and the cessa- and axillary hair and are related to the female sex
tion of menstruation in women distance runners. drive. Adrenarche—the time of increasing secre-
Anorexia nervosa is a condition in which the per- tion of adrenal androgens—generally begins
son—most commonly an adolescent girl— slightly before age 8 (Grumbach & Styne, 1998).
engages in compulsive, extreme dieting, perhaps
to the point of starving herself to death. As
anorexia progresses, the percentage of body fat Changes in the Male
declines and menstruation ceases. It is also fairly As noted above, puberty begins at about 10 or 11
common for women who are runners, and all years of age in boys, about two years later than it
women who exercise seriously to the point where does in girls. The physical causes of puberty in
their body fat is substantially reduced, to cease
menstruating. For both anorexics and female run- 6
On the other hand, programs of moderate, regular aerobic
ners, it seems that when the percentage of body fat exercise have been shown to reduce menstrual problems
falls below a critical value, the biological mecha- such as cramps (Golub, 1992).
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boys parallel those in girls. They are initiated by at puberty, had them castrated so that they remained
increased production of FSH and LH by the pitu- permanent sopranos. Contrary to popular belief,
itary. At the beginning of puberty, the increase in castration in adulthood will not produce a high
LH stimulates the testes to produce testosterone, voice, because the larynx has already grown.
which is responsible for most of the changes of A great spurt of body growth begins in males at
puberty in the male. around 11 to 16 years of age (Figure 5.9). Height
The first noticeable pubertal change in males is increases rapidly. Body contours also change. While
the growth of the testes and scrotal sac, which the changes in girls involve mainly the increase in
begins at around 10 to 13 years of age as a result of fatty tissue in the breasts and hips, the changes in
testosterone stimulation. The growth of pubic hair boys involve mainly an increase in muscle mass.
begins at about the same time. About a year later Eventually testosterone brings the growth process
the penis begins to enlarge, first thickening and to an end, although it permits the growth period to
then lengthening. This change also results from continue longer than it does in females.
testosterone stimulation. As the testes enlarge, Puberty brings changes and also problems. One
their production of testosterone increases even is acne, which is stimulated by androgens and
more; thus there is rapid growth of the penis, affects boys more frequently than girls. Acne is a dis-
testes, and pubic hair at ages 13 and 14. tressing skin condition that is caused by a clogging
The growth of facial and axillary hair begins of the sebaceous (oil-producing) glands, resulting
about two years after the beginning of pubic-hair in pustules, blackheads, and redness on the face
growth. The growth of facial hair begins with the and possibly the chest and back. Generally, acne is
appearance of fuzz on the upper lip; adult beards do not severe enough to be a medical problem,
not appear until two or three years later. Indeed, by although its psychological impact may be great. In
age 17, 50 percent of American males have not yet order to avoid scarring, severe cases should be
shaved. These changes also result from testosterone treated by a physician, the treatment typically being
stimulation, which continues to produce growth of ultraviolet light, the drug Retin-A, and/or antibi-
facial and chest hair beyond 20 years of age. otics. The drug Accutane is highly effective for
Erections increase in frequency. The organs that severe cases. However, it must be used cautiously,
produce the fluid of semen, particularly the because it may have serious side effects, including
prostate, enlarge considerably at about the same birth defects if taken by a pregnant woman.
time the other organs are growing. By age 13 or 14 Gynecomastia (breast enlargement) may occur
the boy is capable of ejaculation.7 By about age 15, temporarily in boys, creating considerable embar-
the ejaculate contains mature sperm and the male rassment. About 80 percent of boys in puberty
is now fertile. The pituitary hormone FSH is experience this growth, which is probably caused
responsible for initiating and maintaining the pro- by small amounts of female sex hormones being
duction of mature sperm. produced by the testes. Obesity may also be a tem-
Beginning about a year after the first ejaculation, porary problem, although it is more frequent in
many boys begin having nocturnal emissions, or girls than boys.
“wet dreams.’’ For the boy who has never mastur- In various cultures around the world, puberty
bated, a wet dream may be his first ejaculation. rites are performed to signify the adolescent’s pas-
At about the same time penis growth occurs, the sage to adulthood. In the United States the only
larynx (“voice box’’) also begins to grow in response remaining vestiges of such ceremonies are the Jew-
to testosterone. As the larynx enlarges, the boy’s ish bar mitzvah for boys and bat mitzvah for girls
voice drops, or “changes.’’ Typically the transition and, in Christian churches, confirmation. In a
occurs at around age 13 or 14. Because testosterone sense, it is unfortunate that we do not give more
is necessary to produce the change in voice, castra- formal recognition to puberty. Puberty rites proba-
tion before puberty results in a male with a perma- bly serve an important psychological function in
nently high voice. This principle was used to that they are a formal, public announcement of
produce the castrati, who sang in the great choirs of the fact that the boy or girl is passing through an
Europe during the eighteenth century. They began as important and difficult period of change. In the
lovely boy sopranos, and their parents or the choir- absence of such rituals, the young person may
master, hating to see their beautiful voices destroyed think that his or her body is doing strange things
and may feel very much alone. This may be partic-
7 ularly problematic for boys, who lack an obvious
Note that orgasm and ejaculation are two separate
processes, even though they generally occur together, at
sign of puberty like the first menstruation (the first
least in males after puberty. But orgasm may occur without ejaculation is probably the closest analogy) to help
ejaculation, and ejaculation may occur without orgasm. them identify the stage they are in.
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Figure 5.9 There is great variability in the onset of puberty and its growth spurt. Both girls are the same age.
All boys are the same age.
(a) (b)
Figure 5.10 Most cultures celebrate puberty, but cultures vary widely in the nature of the
celebration. (a) American Jewish youth celebrate a bar mitzvah (for boys) or bat mitzvah (for
girls). (b) The Samburu youth of Kenya celebrate a male circumcision ritual.
(a) (b)
111
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SUMMARY
The major sex hormones are testosterone, which is same embryonic tissue are said to be homologous
produced in the male by the testes, and estrogen and to each other.
progesterone, which are produced in the female by Intersex conditions are generally the result of
the ovaries. Levels of the sex hormones are regulated various syndromes (such as CAH) and accidents
by two hormones secreted by the pituitary: FSH that occur during the course of prenatal sexual dif-
(follicle-stimulating hormone) and LH (luteinizing ferentiation. Currently there is a debate over the
hormone). The gonads, pituitary, and hypothalamus best medical treatment of these individuals. The
regulate one another’s output through a negative Guevodoces provide an interesting case of gender
feedback loop. Inhibin regulates FSH levels. change at puberty.
At conception males and females differ only in Puberty is initiated and characterized by a great
the sex chromosomes (XX in females and XY in increase in the production of sex hormones.
males). As the fetus grows, the SRY gene on the Y Pubertal changes in both males and females
chromosome directs the gonads to differentiate include body growth, the development of pubic
into the testes. In the absence of the SRY gene, and axillary hair, and increased output from the
ovaries develop. Different hormones are then pro- oil-producing glands. Changes in the female
duced by the gonads, and these stimulate further include breast development and the beginning of
differentiation of the internal and external repro- menstruation. Changes in the male include growth
ductive structures of males and females. A male of the penis and testes, the beginning of ejacula-
organ and a female organ that derive from the tion, and a deepening of the voice.
1. Of the physical changes of puberty, which are fourth gender? (For further information, see
the most difficult to cope with? Herdt, 1990.)
2. The society in the Dominican Republic in 3. Teresa has just given birth to her first baby. The
which the Guevodoces are born (see page 105) doctor approaches her with a worried expres-
is a three-gender society, unlike the two- sion and says that her baby’s genitals are
gender society of the dominant U.S. culture. unusual and some decisions will have to be
What would the United States be like if it was a made. The phallus is too big for a clitoris, but
three-gender society? Who would be classified too small for a penis. What should Teresa do?
in the third gender? Would their lives be better What other information should she obtain
or worse as a result? Could we have a four- from the doctor before making a decision?
gender society? Who would be classified in the
Fausto-Sterling, Anne (2000). Sexing the body. New the medical and psychological professionals
York: Basic Books. The author, a developmental who treat them, and proposes new approaches
geneticist, has written a provocative book that in dealing with the condition.
calls into question our most basic understand- Larsen, P. Reed et al. (2003). Williams textbook of
ings of differentiation of the sexes, both physi- endocrinology. 10th ed. Philadelphia: Saunders.
cally and psychologically. An outstanding endocrinology text, with a par-
Kessler, Suzanne J. (1998). Lessons from the ticularly good chapter on sexual differentiation.
intersexed. New Brunswick, NJ: Rutgers Univer- Steinberg, Laurence (2002). Adolescence. 6th ed. New
sity Press. Kessler, a psychologist, reports on her York: McGraw-Hill. This is the definitive textbook
years of research with intersex individuals and on adolescence, written by a leading researcher.
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WEB RESOURCES
http://www.isna.org http://www.oxytocin.org/oxytoc
The Intersex Society of North America home Information and articles about oxytocin and its
page. roles in orgasm, childbirth, and bonding.
http://www.teenpuberty.com http://www.cah.org.uk
Teen Puberty; includes health information about Congenital Adrenal Hyperplasia Support
pubertal changes; intended for adolescents. Group.
http://www.urologyhealth.org/pediatric http://www.medhelp.org/www/ais
American Urological Association; discussion of Androgen Insensitivity Syndrome Support
pediatric abnormalities contains information Group.
regarding abnormal sexual differentiation.