Lecture15 - (161) Hormone in Reproduction

1/27/2015 Hormonal
Regulation in Reproduction | Principles of Biology from Nature Education

Principles of Biology contents

161 Hormonal Regulation in Reproduction
Piglets nursing.
These piglets are the result of the precise timing of reproductive events: sexual maturity of the sow, timing of
ovulation, preparation of the uterus, changes in behavior to allow for mating, and more. The reproductive
hormones are the chemical signals that are ultimately responsible for the successful timing of these multiple
events.
Courtesy of U.S. Department of Agriculture.
Topics Covered in this Module
Hormones Regulate Reproduction
Major Objectives of this Module
Explain how hormones govern human female reproductive cycles.
Explain how hormones govern spermatogenesis in human males.

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1/27/2015 Hormonal Regulation in Reproduction | Principles of Biology from Nature Education


Hormones are well suited for regulating reproductive events that require coordination of multiple organ systems
because they are chemical signals that circulate to most parts of the body. In most cases, the brain integrates
environmental cues and internal cues to assure proper timing. The brain then controls the release of the hormones that
coordinate reproductive activities such as hormone release, gonad maturation, production of gametes and behavior
that ensure successful production of offspring.
What are some of the ways that hormones regulate reproduction?
Reproductive hormone release is usually influenced by environmental stimuli and can change organ function and even
behavior. For example, it is evolutionarily beneficial to have a system to control reproduction based on day length since
day length is linked to the seasons and food availability. Many birds will begin to court mates, establish territories and
produce gametes as the days begin to grow longer due to an increase in neurohormonecontrolled pulses of
reproductive hormones like estrogen and testosterone. Similar daylength effects on reproduction are seen in other
animals. In another example, if the Colorado potato beetle (Leptinotarsa decemlineata) reaches adulthood during a
time of short days (~10 hrs), its level of juvenile hormone (JH) will be low, causing it to burrow underground in the fall to
await spring (Figure 1). If it reaches adulthood during a time of longer days, it will have high JH levels and will continue
to feed, produce gametes and reproduce. Long days detected by the brain cause the release of allatotropin, a
neurohormone that stimulates the endocrine gland that produces JH. Neurohormones are hormones produced and
released from specialized neurons, while hormones that target other endocrine glands, as in this example, are called
tropic hormones.
Figure 1: A newly emerged adult Colorado potato beetle (Leptinotarsa decemlineata).
If the day length is short, this animal will make no gametes and will burrow underground for the winter.
Scott Bauer/Courtesy of U.S. Department of Agriculture.
The behavior of the animals around them (as perceived by the brain) can also be a cue for timing of reproductive
hormone release. Courtship displays or various forms of contact can serve to stimulate reproductive hormone release.
The key is to be ready to reproduce when in the presence of a suitable mate. An example of courtship behavior affecting
reproductive hormone levels is seen in male ring doves (Streptopelia risoria). Imagine an isolated male, who neither
sees nor hears a female nearby, versus a male kept in a cage right next to a female. Which bird will have a higher
testosterone level in his blood? Experiments showed that the isolated male dove had significantly lower testosterone (a
male reproductive hormone) (Figure 2). A male dove taken from isolation will start courting a female dove right away
when first introduced to her, and his blood levels of testosterone begin to rise within 30 minute of meeting her. Similar to
what happens in doves and other birds, in quails (Coturnix japonica), testosterone levels are often the trigger for
male sexual behavior (Figure 3).
http://www.nature.com/principles/ebooks/principlesofbiology104015/29145865/1 1/8
Figure 2: Testosterone concentrations in male ring doves
(Streptopelia risoria) are affected by the presence of a
female.
Plasma testosterone concentrations in male ring doves 24 hrs
after they were kept in isolation (Isol) or paired with a female
(+F).
© 2011 Nature Education All rights reserved.
Figure 3: Hormones affect behavior.
Effect of castration and exposure to increasing doses of
testosterone on sexual behavior (mount attempts and
crowing) in male Japanese quail (Coturnix japonica).
Castration completely suppressed both behaviors; however,
the behaviors were restored with increasing dosage of
testosterone administered to the castrated males.
Placental animals and ovoviviparous animals need to temporarily halt egg production to retain their offspring until a live
birth. Ovoviviparous animals retain their eggs internally until hatching, resulting in the release of live young. In the
ovoviviparous Madeira cockroach (Leucophaea maderea), the presence of eggs will inhibit release of tropic
hormones, halting the release of juvenile hormone (JH) that stimulates egg production and the receptivity of the female
to courting males.
Test Yourself Graph.
Test Yourself
Observe the above Test Yourself Graph of the Leucophaea madera reproductive cycle
plotting the JH production and egg growth during two cycles. What would you expect to
happen if the developing embryos were removed at the point marked by the arrow?
Submit
This pattern of neurohormone control over endocrine hormone production is a common one in the reproduction of many
animal species. In mammals, the neurohormones and nongonadal hormones are often called tropic hormones, tropins,
or releasing hormones, named for their effect on other endocrine organs. A tropic hormone is one that stimulates the
synthesis and secretion of a hormone by another endocrine gland. The gonadal steroid hormones, the sex hormones,
are androgens (esp. testosterone) from the testes and estrogens (esp. estradiol) and progestins (esp. progesterone)
from the ovaries. The timing of reproductive events is usually controlled by the brain, which responds to environmental
and internal cues such as day length or fat stores. In addition to the timing of events by these cues, the effect of the
hormones may be influenced by the developmental state of the animal. That is to say, the hormones must be present
during certain "critical periods" of development in order for their effects to be fully realized.
In many mammals that are nonseasonal breeders (such as humans), there are two stages of reproductive hormone
release. The first is early in life either before or just after birth, and the second is later in life at the point of sexual
maturation. The timing between these two stages depends on the species. Release of gonadal steroids is stimulated
following a cascade of events that starts with activation of a part of the brain called the hypothalamus. The
hypothalamus releases the neurohormone gonadotropinreleasing hormone (GnRH) directly into blood vessels that
feed into the anterior pituitary. At the anterior pituitary, GnRH stimulates production and release of two hormones,
folliclestimulating hormone (FSH) and luteinizing hormone (LH).
GnRH is released by the neuroendocrine cells of the hypothalamus in pulses, in a manner similar to the way the
neurotransmitters are released by neurons — an action potential will release vesicles filled with hormone. During early
development, the GnRH neuroendocrine cells undergo a temporary increase in pulse frequency, which leads to an
increase in GnRH concentrations in blood flowing through the anterior pituitary. The increased GnRH causes an
increase in FSH and LH release from the anterior pituitary.
FSH and LH stimulate the ovary or testes to secrete estrogens or androgens. This early pulse establishes the sexual
differentiation of the animal into a male or female appearance and establishes male or femalespecific changes in the
brain. Between this period and sexual maturation, the secretion of GnRH slows to a less frequent pulse of release. The
GnRH pulse will not increase frequency again until the animal becomes sexually mature.
Multiple signals determine the timing of the transition to sexual maturation, or puberty. Most of these are factors that
would play a role in the successful production of offspring. Is the animal large enough? Are there enough fat stores?
What is the season, as represented by the length of day? All of these signals are integrated by the brain, causing GnRH
secretion pulses from the hypothalamus to increase (Figure 4). This second increase stimulates the gonadotropins FSH
and LH to complete gonadal maturation.
Figure 4: The hypothalamicpituitarygonadal axis.
The hypothalamus releases GnRH, which stimulates the synthesis and release of pituitary gonadotropins FSH and
LH from the pituitary. The inset shows the release of GnRH from hypothalamic neurons (top) and release of LH and
FSH from anterior pituitary cells (bottom), as well as the frequency of hormonal secretion pulses in immature and
mature individuals.
In addition to gonadal maturation at puberty, the sex hormones complete the changes that differentiate fully adult males
and females from immature males and females. Table 1 shows some of these characteristics in humans.
Type Boys Girls
Stage 1 (prepubertal) No sexual development No sexual development
Breast budding, first pubic hair
Testes enlarge
Stage 2 Body odor
Body odor
Height spurt
Penis enlarges Breasts enlarge
Stage 3 First pubic hair Pubic hair darkens, becomes curlier
Ejaculation (may include wet dreams) Vaginal discharge
Continued enlargement of testes and
penis Onset of menstruation
Penis and scrotum deepen in color Nipple becomes distinct from surrounding
Stage 4
Pubic hair curlier and coarser areola
Height spurt Pelvis begins to widen
Male breast development (temporary)
Pubic hair extends to inner thighs
Pubic hair extends to inner thighs
Stage 5 (Fully mature Increase in height slows, then stops
Increase in height slows, then stops
adult) Deposition of fat in hips, buttocks, thighs, and
Increased muscle mass
breasts
Table 1: Changes that take place in human males and females as puberty progresses.
Negative feedback mechanisms control sex hormone production in males.
After gonadal maturation, some animals enter into reproductive cycles tied to their environmental and internal cues. In
human males, there is no pronounced cycling of reproductive behavior or sperm production; sperm are produced
continually throughout life. GnRH pulses maintain a steady state and the subsequent release of FSH prepares the
Sertoli cells in the testes to respond to androgens, while LH stimulates testosterone synthesis from Leydig cells in the
testes. Testosterone maintains sperm production and male sexual behavior. As testosterone concentrations rise,
production of GnRH by the hypothalamus and of LH by the anterior pituitary is inhibited through a negative feedback
loop (Figure 5). In addition, the Sertoli cells release inhibin, which acts on the pituitary to reduce FSH secretion.
Figure 5: Negative feedback.
Testosterone and inhibin negatively feed back on the
hypothalamus and pituitary in males.
Test Yourself
A male birth control pill has long been sought after. How might inhibin or a synthetic inhibin
be used as a male contraceptive?
Submit
Cyclical changes in hormone concentration control human female reproductive cycles.
While males continuously produce sperm, egg production is cyclic. In mammals these cycles exist as estrous cycles or
menstrual cycles. In either type of cycle, ovulation is timed to occur when the lining of the uterus is prepared to support
an embryo. The difference between a menstrual and estrous cycle is whether or not the uterine lining is shed or
reabsorbed if pregnancy does not occur. Great apes, including humans, have a menstrual cycle and are receptive to
mating throughout the year. In mammals with an estrous cycle, mating takes place only at certain times of the year when
behavioral and physical changes lead to a period of sexual activity called estrus or "heat."
The human menstrual cycle takes an average of 28 days to complete. During this time, the uterus must be prepared to
receive a fertilized egg and the egg must be released when the timing is best suited for a successful pregnancy. The
uterine cycle refers to the changes that occur to the uterus. The ovarian cycle involves the development and release of
an egg from one of the ovaries. While separate, these two cycles are coordinated by the same hormones used in males
to control androgen production and sperm formation in the testes.
The menstrual cycle, the coordination of the ovarian and uterine cycles, starts on the day the uterine lining begins to
shed, the beginning of a menstruation or "a period" (Figure 6). At this point, in response to GnRH, a baseline level of
FSH and LH are secreted by the pituitary. The FSH induces growth of follicle cells around a developing egg in the ovary.
The follicle cells have few receptors for LH at this point.
Stimulated by FSH, the follicle cells synthesize estrogen hormones, most importantly estradiol. As the follicle cells
continue to develop, they release increasing amounts of estradiol. In response to the increasing estradiol, the lining of
the uterus, the endometrium, thickens and proliferates. While low levels of estrogens can inhibit the release of FSH and
LH through negative feedback on the hypothalamus, the large amounts of estradiol produced by growing follicle
simulates additional GnRH release, and estradiol increases the sensitivity of the LHreleasing cells to GnRH.
As the levels of FSH and LH continue to increase, the follicle cells become more receptive to LH (by increasing the
number of LH receptors). The peak in LH induces final maturation of the follicle, and ovulation occurs when the follicle
ruptures and releases the egg. The ruptured follicle remains in the ovary to become the corpus luteum. LH stimulates
the corpus luteum to secrete estradiol and progesterone. The combination of estradiol and progesterone now inhibits
the secretion of GnRH, diminishing the release of LH and FSH by the pituitary.
Without LH, and in the absence of fertilization, the corpus luteum eventually degenerates and the levels of estradiol and
progesterone decrease rapidly. The decrease in progesterone levels causes the uterine lining to be shed
(menstruation), and a new cycle will begin. With the level of estrogen low, the inhibition of GnRH secretion is lost, and
additional FSH and LH are secreted as new follicles in the ovary begin to mature.
If, instead, the egg is fertilized and implantation (pregnancy) occurs, an embryonic hormone called human chorionic
gonadotropin (hCG) takes over the role of LH and rescues the corpus luteum from degeneration. Progesterone and
estradiol continue to be produced and the thickened lining of the uterus will be retained. The lining will become the
decidua, part of the placenta, providing support for the developing embryo.
The number of cycles that a human female undergoes is limited to 500 or so. The end of reproductive cycling in women
is called menopause. During menopause, the ovaries stop responding to FSH and LH, resulting in a decline in estradiol
and progesterone.
The specific timing of hormone release can be used for medical purposes. hCG detection is the basis for most
pregnancy tests, because it is unique to and necessary for pregnancy. Failure of hCG to rescue the corpus luteum is
often a reason for loss of a pregnancy. The basis for birth control pills exploits the sensitivity of the hypothalamus to
estrogens and progesterone. The most commonly used type of pill contains a combination of estradiol and
progesterone. The two hormones negatively feed back on the hypothalamus to slow the release of GnRH, reducing the
levels of LH and FSH released from the pituitary gland. Without the peak in LH and FSH, follicle production and
ovulation will not occur.
Figure 6: Hormonal regulation of ovarian and uterine cycles.
Levels of hormones from the anterior pituitary and the ovary affect the ovarian and uterine cycles.
Test Yourself
Would you expect a pill that contains only estrogen to act as a contraceptive? Why or why
not?
Submit
Future perspectives.
Several reports of animals with malformed genitalia and girls entering puberty early have implicated environmental
exposure to substances that mimic or disrupt the action of sex hormones. These substances are called environmental
endocrine disruptors (Table 2). The proper function of the hypothalamuspituitarygonad axis requires precisely timed
production and response to reproductive hormones; therefore, endocrine disruptors may disrupt endocrine function at
several stages of reproduction, from development of sexual characteristics to proper sexual behaviors. Diethylstilbestrol
(DES), a synthetic estrogen, is a wellstudied endocrine disruptor. Exposure to this chemical during critical periods of
development can result in malformed reproductive organs, and increased incidence of cancers of the reproductive
organs. Evidence suggests that the adverse effects of DES can be transmitted to subsequent generations.
Daidzein, genistein, formononetin, biochaninA, prunetin, pratensein, glycitein, equol,
Phytoestrogens
desmetilangolestin, enterolactone, enterodiol, matairesinol, zearalenone
Organohalogens Dioxins, furans, polychlorinated biphenyls, hexachlorobenzene, pentachlorophenol
Dichlordiphenyltrichloroethane (DDT), methoxychlor, endosulfan, 2,4dichlorophenoxyacetic
acid, alachlor, aldicarb, amitrol, atrazine, benomyl, dibromo chloropropane, carbaryl, chlordane,
Pesticides
ethyl parathion, heptachlor, kepone, ketoconazole, lindane, methomyl, permethrin, malathion,
trifluralin, vinclozolin
Diethylhexyl phthalate, butyl benzyl phthalate, diNbutyl phthalate, dihexyl phthalate, dipropyl
Phthaltes
phthalate, dichloro hexyl phthalate, diethyl phthalate
Heavy metals Arsenic, cadmium, uranium, lead, mercury
Drugs Oral contraceptives, diethylstilbestrol, cimetidine
Industrial
Bisphenol A (BPA), polybrominated biphenyls
products
Table 2: Environmental endocrine disruptors.
Some of the major endocrine disruptors are listed here. These compounds disrupt the function of mammalian
endocrine systems.
BIOSKILL
What Level of Endocrine Disruptors in the Environment is Acceptable?
Will increasing levels of a hormone always cause increased effects on a target organ? On the basis of earlier studies
using high doses of estrogens, exposure to estrogen during fetal life is known to inhibit prostate development of
developing male fetuses. However, other studies of hormone function have shown that low concentrations of a hormone
can have one effect, while high concentrations can have the opposite effect.
One study measured the effects of prenatal exposure to DES in mice. Researchers measured prostate weight in 8
monthold male mice born of females fed different doses of DES from day 11 to day 17 of pregnancy, the time during
which the important accessory organ, the prostate is developing (Figure 7).
Figure 7: Dose response curve of the effect of prenatal diethylstilbestrol (DES) exposure.
Prostate size in 8monthold male mice varies with prenatal DES dose.
When low doses of DES (0.02, 0.2, and 2.0 ng/g of body weight per day) were fed to pregnant mice, prostate
enlargement was observed in their adult male offspring, with greatest enlargement at a dose of 0.2 ng/g body weight.
Prostate weight then decreased with increasing dose, resulting in an invertedU doseresponse relationship. This type
of invertedU response is not uncommon in endocrinology; however, current toxicology tests that help determine the
safety of drugs and chemicals assume that there will be a simple doseresponse relationship, where the response is
assumed to increase or reach maximum response as the dose increases. Depending on the starting doses selected, the
doses reached in testing may be too low or too high to show their adverse effects.
BIOSKILL
IN THIS MODULE
Summary
Test Your Knowledge



1/27/2015 Summary of Hormonal Regulation in Reproduction | Principles of Biology from Nature Education


Summary
OBJECTIVE Explain how hormones govern spermatogenesis in human males.
Gonadotropinreleasing hormone (GnRH), which is produced by the hypothalamus, stimulates release of follicle
stimulating hormone (FSH) and luteinizing hormone (LH) from the anterior pituitary. In males, FSH and LH act on two
different types of cells in the testes. FSH stimulates growth of Sertoli cells, which nourish sperm. LH stimulates Leydig
cells to produce testosterone. Testosterone stimulates spermatogenesis. Through a negative feedback loop,
testosterone inhibits GnRH production by the hypothalamus and LH production by the anterior pituitary. The Sertoli cells
produce the hormone inhibin, which inhibits production of FSH by the anterior pituitary.
OBJECTIVE Explain how hormones govern human female reproductive cycles.
LH and FSH regulate the menstrual cycle. In the ovaries, FSH stimulates the maturation of follicles, which consist of an
oocyte and support cells. Maturing follicles produce estradiol, which causes the uterine lining to grow. As the follicle
matures, it produces more estradiol. Low levels of estradiol inhibit FSH and LH production, but high levels stimulate
production of these hormones. Thus, as the follicle matures, increased estradiol levels begin to stimulate FSH and LH
production. A surge in FSH and LH coincides with ovulation. The remnants of the ruptured follicle differentiate into the
corpus luteum, an endocrine structure that secretes estrogen and progesterone, another female sex steroid hormone.
Together, estrogen and progesterone inhibit the secretion of FSH and LH from the anterior pituitary, preventing the
maturation of another follicle.
Key Terms
androgen
Type of steroid hormone that influences male characteristics; for example, testosterone.
estrogen
Type of sex hormone produced in large amounts by ovaries; also produced in small amounts by testes; present in
male and female organisms.
folliclestimulating hormone (FSH)
Stimulates growth of follicles in females and activates the spermforming cells in males.
gonadotropinreleasing hormone (GnRH)
Tropic neurohormone produced in the hypothalamus; triggers the anterior pituitary to release folliclestimulating
hormone (FSH) and luteinizing hormone (LH).
inhibin
Triggers the pituitary to reduce folliclestimulating hormone secretions.
luteinizing hormone (LH)
Stimulates ovulation and development of the corpus luteum in the female and production of testosterone in the
Leydig cells of the male.
menstrual cycle
Hormonal cycle in mammalian females to prepare the body for pregnancy.
menstruation
Shedding of the uterine lining if pregnancy does not occur.
ovarian cycle
Maturation and release of eggs from the ovary.
sex hormone
Type of hormone that affects the reproductive organs, such as estrogens or androgens.
testosterone
Type of androgen; a male sex hormone produced in the testes that maintains sperm production and male sexual
behavior.
tropic hormone
Triggers the synthesis and/or release of another hormone by another endocrine gland.
uterine cycle
Changes to the uterus that occur during the menstrual cycle.
IN THIS MODULE
Summary
Test Your Knowledge

1/27/2015 Summary of Hormonal Regulation in Reproduction | Principles of Biology from Nature Education



IN THIS MODULE

Test Your Knowledge Summary
Test Your Knowledge
1. Complete the following sentence: The period during which a sexually immature individual becomes sexually
mature is called...
menopause.
menstruation.
endocrine disruption.
puberty.
inhibin.
2. Complete the following sentence: The cessation of ovulation after about 500 cycles is called...
inhibin.
menstruation.
puberty.
endocrine disruption.
menopause.
3. In males, which of the following hormones exhibits negative feedback on GnRH secretion?
FSH
testosterone
LH
inhibin
LH and inhibin
4. Which best describes the relationship between estradiol levels and GnRH production during a female's
reproductive cycle?
Increasing estradiol levels always leads to increased GnRH production.
Increasing estradiol levels always leads to decreased GnRH production.
High levels of estradiol inhibit GnRH release, whereas low levels of estradiol stimulate GnRH release.
Increasing estradiol levels lead to increased inhibin levels, which shuts down GnRH production in the
hypothalamus.
Low estradiol levels inhibit GnRH release, whereas high estradiol levels stimulate GnRH release.
5. Compare the levels of estradiol and progesterone seven days after ovulation to the respective levels
immediately before ovulation.
The estradiol level is slightly lower than it was before ovulation, but the progesterone level is much
higher.
The progesterone level is slightly lower than it was before ovulation, but the estradiol level is much
higher.
Both levels are higher than before ovulation.
Both levels are lower than before ovulation.
The estradiol and progesterone levels are the same as they were immediately before ovulation.
6. Scientists discover a new chemical that is similar in structure to estrogen. What would be the most likely
consequences of excessive exposure to this drug?
irregular menstrual cycles in women
premature onset of puberty in girls
abnormal prostate gland development in males
irregular menstrual cycles in women; premature onset of puberty in girls; and abnormal prostate gland
development in males
None of the answers are correct.
Submit

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