K. P.

Paudel
Sem VIII
VOG 421
Lecture 2

Role of various Hormones on Reproduction

After the differentiation during embryonic life, the development of Mullerian or Wolffian
duct into respective reproductive tracts is entirely under the influence of fetal steroid
hormones. In females, Mullerian duct develops into reproductive tract organs (refer to
Lecture 1) under the influence of estrogen produced by the fetus itself. So does Wolffian duct
in males under the influence of fetal testosterone.
Until several months prior to puberty, general growth and development of all reproductive
organs is under the influence of growth hormone as are other tissues of the body.
The reproductive hormonal system consist of three hierarchies as follows:
 Hypothalamic releasing hormones (GnRH) under the influence of CNS and steroidal
hormone feed back mechanism.
 The anterior pituitary sex hormones (gonadotropins) in response to the effect of
GnRH.
 The ovarian hormones, estrogen and progesterone.

The physiological functions of gonadotropins and steroid hormones are briefly discussed
here.

Gonadotropins:
Those hormones secreted from the anterior pituitary and acting mainly on the gonads
are termed as gonadotropins. Gonadotropic hormones (FSH and LH) are secreted from the
basophilic cells (gonadotropes) and prolactin from acidophilic cells of the anterior pituitary
gland.

FSH and LH hormone both consist of two distinct peptide chains, alpha (α) and beta (β)
chains. The α subunit is common to both FSH and LH within species whereas the β subunit is
different in each hormone and therefore confer the functional specificity.

Follicle stimulating Hormone (FSH):

FSH is a glycoprotein hormone with a molecular weight of about 32,000 Dalton. The
half-life of this hormone is 3- 4 hours. The pituitary output of FSH is under the hypothalamic
control, including feedback mechanism involving the steroid hormones of the gonads. Rising
estrogen level (ovarian hormone) from the matured follicle feeds back to depress GnRH
(gonadotropin releasing hormone) from the hypothalamus and then the FSH output. The other
probable control of FSH occurs through environmental stimuli. For example in certain
species visual stimuli is known to stimulate hypothalamus to release GnRH which then cause
release of FSH. Similarly, environmental conditions such as changing seasons and day light
mediated through some exteroreceptors such as eye to the hypothalamus thus, again
influencing GnRH output.
The physiologic effect of FSH in hypophysectomized (pituitary gland removed surgically)
animal is known to cause multiple follicular growth in the ovaries without estrogen
production and/or ovulation.
The main physiological function of FSH include
o Stimulation of the growth of ovarian follicle or the graffian follicle.
 o It stimulates maturation of oocyte and secretion of follicular fluid. Follicular fluid
provides nutrients and is the medium for the development of the ovum in the follicle.
o It also helps proliferate granulosa cells and thecal layer thus helping in overall
development of the follicle.
o Stimulation of estrogen production in the presence of LH. However, it alone cannot
cause secretion of estrogen.
o Acts on germinal cells in the seminiferous tubules of the testes and is responsible for
spermatogenesis up to the stage of secondary spermatocytes.
The main function of FSH is to favor protein synthesis by acting on primary
spermatocytes in male.
The clinical use of pure pituitary FSH is limited in veterinary medicine because of the cost of
the substance. An economical source of an FSH like hormone is available from the serum of
pregnant mares, which is called equine chorionic gonadotropin or also called pregnant mare
serum gonadotropinn (PMSG). The clinical use of FSH is limited to induction of super
ovulation for embryo production and transfer.

Luteinizing Hormone:
LH of female and interstitial cell-stimulating hormone (ICSH) in male is in fact the
same hormone and the difference in nomenclature only reflects the sex difference. This is
also a glycoprotein hormone with two peptide chains alpha and beta having a molecular
weight of about 30,000 Dalton in swine. Its half-life is just about 30 minute implying how
soon it acts on the target organ and then gets metabolized.
Regulation of LH output from the pituitary gland is dependent on the hypothalamic control
(GnRH). This regulation is again in part is under a feedback control mechanism involving the
gonadal steroid hormones. The functions of the LH have to be understood in two levels:
- Basal level (the level occurring all the time except at around
ovulation) acts in conjunction with FSH to induce estrogen
secretion from the large ovarion follicle
- Pre-ovulatory surge induces ovulation.
- The basal level is also responsible for luteinization of the
ovarian tissue to form corpus luteum
- In male LH stimulates androgen secretion in the leydig cells.

Ovarian Steroid Hormones

Steroid hormones are basically derived from cholesterol, a 27- carbon compound through
different enzymatic pathways in different endocrine organs.

Cholesterol (27 - C) Pregnenolone (20 - C) Progesterone (21 - C)

Estradiol (18 - C) Testosterone (19 - C)

Estrogens are synthesized in theca interna cells, and progestins in granulosa cells of the
ovarian follicles. However, this generalization is still debatable.
- Estrogens are 18 - carbon steroids, mainly three compounds namely, estradiol,
estriol and estrone are identified in domestic animals. Estradiol - 17 β is the major
ovarian estrogen in the cow, mare, pig, bitch and human. Estrone is present in
small amounts. In most species the estrogenic secretion from placenta and testes is
similar to that from the ovary. However, estrogen isolated from sheep and goat
placenta appears to be estradiol - 17α while that isolated from swine placenta
 consists of estrone. Of these different estrogen compounds estradiol is the most
potent one in action. All ovarian estrogens are produced from androgenic
precursors and are transported in blood in bound form with globulin.
- Progestagens are 21 - carbon steroid compounds, progesterone being the main
one. The other similar compounds are 20 α and 20 β - hydroxy - 4 pregnene.
Pregnenolone is the precursor of progestagins. Progesterone is also transported in
blood in bound from with globulin.
- Androgens are 19 - carbon steroid compounds, testosterone being the most
important one.
Liver is the main organ that inactivates steroid hormones. The role of kidney and other organs
is also significant. The function of the liver is inactivation of potentially toxic compounds and
in case of steroid hormones it involves metabolism and chemical degradation and/or
esterification of steroids to form sulphates and conjugation to form glucuronides. The liver
handles estrogens and progestins most effectively and androgens and corticosteroids to a
lesser extent. These metabolites are probably excreted through bile to the gut and removed
from the feces.

Functions of Estrogens:
The predominance of estrogen secretion at higher than the basal level during proestrus and
estrus in domestic animals seems to be meant for the preparation of female genital passage
for copulation and successful union of the male and female gamete. However, the estrogen
hormone has wide spectrum of its function, which can be summarized as follows:
- Estrogens act on CNS to induce behavioral estrus in female farm animals. However, a
small amount of progesterone with estrogen is needed to induce estrus in some
species of farm animals such as the ewe and cow. Thus it is generally accepted that
estrogen and progesterone act synergistically in a defined ratio at different stages of
the estrous cycle.
- Acts on uterus to increase both amplitude and frequency of contractions by
potentiating the effects of oxytocin and PGF2 α. Uterine contractions are the result of
contractions of the smooth muscle (myometrium) fibers present in the wall of the
uterus.
- Estrogen acts on related organs responsible for the physical development of female
secondary sexual characteristics.
- Estrogen stimulates ductal growth and cause development of mammary glands.
- Exerts both negative (reduced GnRH) and positive ( increased GnRH for LH surge
some 12 hours before ovulation) feedback control through hypothalamus on LH and
FSH release from the pituitary gland.
- Estrogen exerts protein anabolic effect, therefore is a growth promotor.
- Estrogen also exerts luteolytic effect in many of the species.( regression of corpus
luteum). However, estrogen is luteotropic in sows
- Estrogen induces quick union of the epiphysis with the shaft of the long bones. Thus
estrogen is responsible for preventing unlimited elongation of long bones, i.e. stops
height growth of an individual animal after some times (few months to few years
depending on the species) of attainment of puberty.
Functions of progestagins
- It brings about secretory changes in the fallopian tube. The fluid secreted in the
fallopian tube is responsible for smooth transport of the ovum from ovary down to the
site of fertilization. This action is the result of the basal level of progesterone. Further
secretary action is promoted after ovulation till day four until the fertilized ovum is
there in the fallopian tube. This secretion supports the life of the fertilized ovum
(zygote). The changed ratio of estrogen and progesterone (increased progesterone
secreted from newly formed CL and reduced level of estrogen after ovulation) make
these secretory changes.
- Progesterone prepares endometrium for the implantation and maintenance of
pregnancy by increasing the activity of secretory glands in the endometrium and by
diminishing the contractility of the myometrium. Maintenance of CL is essential (in
many species) through out pregnancy. In some species in later stage of pregnancy (for
example in mare), placenta also secretes progesterone. Even the adrenal glands
synthesize progesterone in need.
- Acts synergistically with estrogens to induce behavioral estrus.
- Develops secretory tissue (alveoli i.e. Glandular tissue) of the mammary glands.
- Inhibits estrus and ovulatory surge of LH at high level.
- Inhibits uterine motility.
- Exerts protein catabolic effect.
Figure 6. Schematic representation of the changes in plasma hormone concentrations
during the bovine oestrous cycle

Day O = day cow first shows estrus