Chapter3 Reproductive hormones of

female animals

1
 Chapter 3 Reproductive hormones of female animals

• Specific/learning Objectives
• At the end of this chapter, you will be able to
• clarify the major sources of the Primary reproductive hormones
of female animals

• identify the reproductive hormones of female and their major

functions

• discuss endocrine feedback [positive (stimulatory) and negative

(inhibitory) feedback]

• explain the role of secondary hormones such as thyroid hormones

and adrenal steroids in animal reproduction.

• elucidate synthetic hormones that are widely used in veterinary

medicine
2
 Introduction

Both the endocrine and nervous systems function to initiate,

coordinate, or regulate the functions of the reproductive
system.

Unlike the nervous system, which controls body function

through rapid, electric nerve impulses e.g., musculoskeletal
system, the endocrine system uses chemical messengers or
hormones to regulate slow body processes, e.g., growth and
reproduction.

3
 Introduction
The classic definition of a hormone is a physiologic, organic,
chemical substance synthesized and secreted by a ductless
endocrine gland, which passes into the circulatory system for
transport.

Hormones inhibit, stimulate, or regulate the functional

activity of the target organ or tissue.

However, organs like the uterus and the hypothalamus

produce hormones, which do not meet the classic definition
of a hormone.

4
 Endocrine glands
Before discussing the hormones
of reproduction, it is worthwhile
to review briefly the functional
anatomy of the hypothalamus, the
pituitary, and the gonads.
Hypothalamus

The hypothalamus occupies only

a very small portion of the brain,
and it consists of the region of the
third ventricle, extending from
the optic chiasma to the
mammillary bodies

There are neural connections

between the hypothalamus and
the posterior lobe through the
hypothalamic-hypophyseal tract
and vascular connections between
the hypothalamus and the anterior
pituitary lobe 5
Endocrine cont’d
Pituitary Gland
The pituitary gland is located in the
sella turcica, a bony depression at
the base of the brain.

The gland is subdivided into three

distinct anatomic parts: anterior,
intermediate, and posterior lobes.

There are remarkable species

variations in the anatomy of the FIGURE 3 ..3. Hypothalamus..pituitary gonadal
pituitary gland, and for example, complex. Hypothalamic nerve cells releasing
neurohormones intothe portal vessels for
the pars intermedia is well transport to the anterior pituitary via the
developed in the hypophysis of hypothalamopypophyseal vessels. Solid
particles in nerve cells represent
cattle and horses. neurohormones. Blood is transported by the
retrograde venous system back to the
hypothalamus. 6
 Pituitary Gland
The cell types in the anterior pituitary have traditionally been classified on
their staining characteristics into agranular and granular chromophils.

The chromophils are divided into acidophils and basophils. This

classification has been revised with the advent of immunochemistry and
electron microscopy.

The anterior pituitary has five different cell types secreting six hormones.

By cell type, the somatotropes secrete growth hormone, corticotropes

secrete adenocorticotropic hormone (ACTH), mammotropes secrete
prolactin, thyrotropes secrete thyroid stimulating hormone (TSH), and
gonadotropes which secrete follicle stimulating hormone (FSH) and
luteinizing hormone (LH).

7
Gonads
In both sexes, the gonads plays dual role: the production of germ
cells (gametogenesis) and the secretion of gonadal hormones.

The interstitial cells that are located among the seminiferous

tubules are named the Cells of Leydig.

The Leydig cells secrete testosterone in the male whereas the

theca interna cells of the Graafian follicle are the primary source
of circulating estrogens.

Following rupture of the follicle (ovulation), the granulosa and

thecal cells are replaced with the corpus luteum (CL) that
secretes progesterone.
8
Pineal Gland
The pineal gland (epiphysis) originates as a neuroepithelial
evagination from the roof of the third ventricle under the posterior
end of the corpus callosum.

The pineal gland of the amphibian is a photoreceptor that sends

information to the brain, whereas the mammalian pineal is an
endocrine gland.

9
 Hormones
Hormones may be classified according to either their biochemical
structure or mode of action.

The biochemical structure of hormones includes glycoproteins,

polypeptides, steroids, fatty acids, and amines.

Structure of Hormones

According to their chemical structure, the hormones of

reproduction are divided into four groups:

Proteins: These are polypeptide hormones ranging from a

molecular weight of 300 up to 70,000 daltons, e.g., oxytocin,
FSH, and LH.
10
Structure of Hormones
Steroids: These are derived from cholesterol and have a
molecular weight of 300· to 400 daltons, e.g., testosterone.

Fatty acids: These are derived from arachidonic acid and have a
molecular weight of about 400 daltons, prostaglandins:

Amines: These compounds are derived from tyrosine or

tryptophan, e.g., melatonin.

11
 Hormones
Modes of Intercellular Communication
Cells communicate with each other via chemical messengers such
as amines, amino acids, steroids, and polypeptides.

Thus, there are four modes of intercellular communications:

Neural communication, in which neurotransmitters are

released at synaptic junctions from nerve cells and act across
narrow synaptic clefts between neurons as neurotransmitters

12
 Hormones
Modes of Intercellular Communication

Neurotransmitters are chemical messengers that transmit signals

across a synapse, which is the junction between two neurons or
between a neuron and a target cell (such as a muscle or gland cell).

These molecules play a crucial role in communication within the

nervous system, influencing a wide range of processes including
mood, cognition, and behavior.

Examples of neurotransmitters include: Acetylcholine (ACh),

Dopamine, Serotonin, Norepinephrine (noradrenaline),
Gamma-aminobutyric acid (GABA) and Glutamate

13
Modes of Intercellular Communication

Endocrine communication,
in which hormones
transported through blood
circulation, typical of most
hormones. FIGURE 4. Modes of intercellular communication. Locally
produced growth factors acting in an autocrine/paracrine
manner mediate endocrine action in target cells.

Paracrine communication, in which the products of the cells

diffuse through extracellular fluid to affect neighboring cells that
are at a distance., e.g., prostaglandins.

Autocrine communication, in which cells secrete chemical

messengers that bind to receptors on the same cell that secreted
the messenger
14
 Hormones
Endocrine Feedback

Gonad.
A target gland hormone (e.g., estrogen) can influence the secretion
of the tropic stimuli that caused its own release (e.g., FSH).

The feedback control occurs at the level of the hypothalamus and

the pituitary gland

Depending on their concentration in the blood, steroid hormones

may exert a stimulatory (positive) or an inhibitory (negative)
feedback.

15
 Hormones
Inhibitory or negative feedback.

This system involves reciprocal interrelationships between two or

more glands and target organs.

For example, as stimulation of the ovary increases estrogen

secretion, FSH levels decline.

Similarly, when pituitary hormones reach a certain level, some

hypothalamic nuclei respond by decreasing the production of their
particular releasing hormone, a decline in secretion of pituitary
tropic hormone, and a lower level of target gland function.

16
 Hormones
Stimulatory or positive feedback.

In this system, an increasing level of hormone(s) causes subsequent

increase of another hormone.

For example, increasing levels of estrogen during the preovulatory

phase trigger an abrupt release of pituitary LH.

These two events are precisely synchronized, because a LH surge is

necessary for the rupture of ovarian follicle.

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Endocrine Feedback

Generally, both negative and positive feedback mechanisms are

involved in the regulation of reproductive hormones in farm
animals.

Negative feedback helps maintain hormonal balance and regulate

the timing of reproductive events, while positive feedback
triggers specific events such as ovulation at critical times during
the reproductive cycle.

These feedback mechanisms ensure the proper functioning of the

reproductive system and contribute to reproductive success in
livestock production

18
 Hormones
Hypothalamic hormones.

Both pituitary and steroid hormones regulate the synthesis,

storage, and release of hypothalamic hormones through two
feedback mechanisms: a long and a short loop.

Long feedback involves interaction among the gonad,

pituitary, and hypothalamus while the short feedback system
permits pituitary gonadotropins to influence the secretory
activity of the releasing hormones without mediation of the
gonads (see Hypothalamus retrograde back flow in the
hypothalamus).

19
 Hormones
Neuroendocrine Reflex.

Apart from the feedback mechanisms mentioned above, the

nervous system may control release of hormones through
neural pathways, e.g., oxytocin in milk-let down and LH
release following copulation, as will be pointed out later.

20
 Hormones
Hormone Receptors

Each hormone has a selective effect on one or more target

organs. This effect is achieved through two mechanisms:

Each target organ has a specific method of binding that

'hormone not found in other tissue.

The target organs have certain metabolic pathways capable

of responding to the hormone metabolic pathways not shared
by nontarget tissue

21
 Hormones
Hormone Receptors

Specific binding is the usual mechanism, for example, all target

tissues that respond to steroid hormones contain a receptor protein
within the cell, which specifically binds the activating hormone.

Within the target cell, the steroid hormone is found in the

cytoplasm, bound to a relatively large protein (molecular weight,
200,000 daltons).

Binding results in transformation or activation of the steroid protein

complex, allowing it to move (translocate) into the cell nucleus.

At the nuclear site the steroid complex binds to specific receptor

and causes a sequence of physiologic responses specific for that
cell
22
 Hormones
The target cells of the anterior pituitary possess cell membrane
receptors that recognize and selectively bind the protein
hormones, including gonadotropins.

The binding phenomenon triggers the synthesis and secretion of

the pituitary hormone via the cyclic AMP-protein kinase system
of the cell. Estrogen levels, in turn, influence the gonadotropin
receptors

23
 Hormones
Primary Hormones of Reproduction

Primary hormones regulate the various reproductive

processes, whereas secondary or metabolic hormones
indirectly influence reproduction, and the latter hormones are
not discussed in this chapter.

The primary hormones are involved in many aspects of

reproductive processes such as spermatogenesis, ovulation,
sexual behavior, fertilization, implantation, maintenance of
gestation, parturition, lactation, and maternal behavior.

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 Hormones
Primary Hormones of Reproduction

Reproductive hormones are derived primarily from four

major systems or organs:

✓ various areas of the hypothalamus,

✓ anterior and posterior lobes of the pituitary gland,
✓ gonads (testis and ovary including their interstitial
tissues and corpus luteum), and
✓ the uterus and placenta.

25
 Hormones
Hypothalamic Releasing/Inhibiting Hormones
The hormones of the hypothalamus that regulate
reproduction are gonadotropin-releasing hormone (GnRH or
LH-RH), ACTH, and prolactin-inhibiting factor (PIF).

The hypothalamus is also the source of oxytocin and

vasopressin, which are stored in the neurohypophysis
(posterior lobe of the pituitary gland).

The neuroendocrine control of hypothalamic hormones,

sensory pathways, and integrating areas are summarized in
Table 3-1.
26
 Hormones
Hypothalamic Releasing/Inhibiting Hormones

GnRH is a decapeptide (10 amino acids) with a molecular

weight of 1183 daltons.

It is synthesized and then stored in the medial basal

hypothalamus.

GnRH provides a humoral link between the neural and

endocrine systems.

In response to neural signals, pulses of GnRH are released

into the hypophyseal portal system for the release of LH and
FSH from the anterior pituitary.
27
 Hormones

Adenohypophyseal Hormones

The anterior pituitary gland secretes three gonadotropic

hormones: FSH, LH, and prolactin (Table 3-2) (see cell types
under Pituitary Gland).

LH and FSH are glycoprotein hormones with a molecular

weight of about 32,000 daltons.

Gonadotropes in the anterior pituitary secrete both hormones.

28
 Hormones
Adenohypophyseal Hormones
As stated above GnRH and the gonadal steroids regulate
secretion of gonadotropins.

Additionally, gonadal peptides regulate FSH secretion.

These either stimulate (activins) or inhibit (inhibins,

follistatin) FSH secretion as will be discussed later in this
chapter.

29
 Hormones
Follicle Stimulating Hormone.

Follicle stimulating hormone stimulates the growth and

maturation of the ovarian follicle or the Graafian follicle.

FSH does not cause secretion of estrogen from the ovary by

itself; instead, it needs the presence of LH to stimulate
estrogen production.

In the male, FSH acts on the germinal cells in the

seminiferous tubules of the testis and is responsible for
spermatogenesis up to the secondary spermatocyte stage;
later androgens from the testis support the final stages of
spermatogenesis.
30
 Hormones
Luteinizing Hormone.
Luteinizing hormone is a glycoprotein composed of an alpha
and a beta subunit with a molecular weight of 30,000 daltons
and a biologic half-life of 30 minutes.

Tonic or basal levels of LH act in conjunction with FSH to

induce estrogen secretion from the large ovarian follicle.

The preovulatory surge of LH is responsible for rupture of

the follicle wall and ovulation, and LH stimulates the
interstitial cells of both the ovary and testis.

In the male, the interstitial cells (Leydig cells) produce

androgens after LH stimulation. 31
 Hormones
Prolactin.
Prolactin is a polypeptide hormone secreted by the
adenohypophysis. As stated earlier, it is not a glycoprotein
like other gonadotropins.

Ovine prolactin is a 198 amino acid protein with a molecular

weight of 24,000 daltons.

Prolactin molecules are similar in structure to growth

hormones, and in some species, these hormones have similar
biologic properties.

32
 Hormones
Prolactin initiates and maintains lactation. It is regarded as a
gonadotropichormone because of its luteotropic properties
(maintenance of corpus luteum) in rodents.

However, in domestic animals, LH is the main

luteotropichormone, with prolactin being of less importance
in the luteotropic complex.

Prolactin may mediate the seasonal and lactational effects on

reproduction in farm animals.

33
 Hormones
Prolactin Inhibiting Factor (PIF)

An inhibiting hormone termed regulates secretion of

prolactin.

PIF is probably the catecholamine, dopamine, that is an

amine of low molecular weight synthesized from L-tyrosine.

It is secreted from nerve terminals mostly in the arcuate

nucleus located in the median eminence and transported
through the hypophyseal portal system to the
adenohypophysis.
34
 Hormones
Neurohypophyseal Hormones
The hormones of the posterior pituitary (neurohypophysis)
differ from the other pituitary hormones in that they do not
originate from the pituitary, but are only stored there until
needed.

The two hormones, oxytocin (milk letdown hormone) and

vasopressin (antidiuretic hormone or ADH) are actually
produced in the hypothalamus.

These hormones are transferred from the hypothalamus to

the posterior pituitary not through the vascular system, but
along the axons of the nervous system.

35
 Hormones
Oxytocin.

Oxytocin is synthesized in the supraoptic nucleus of the

hypothalamus and is transported in small vesicles enclosed
by a membrane down the hypothalamic-hypophyseal nerve
axons.

They are stored at the nerve endings next to the capillary

beds in the neurohypophysis until their release into the
circulation.

As stated earlier, oxytocin is also produced in the corpus

luteum. Thus, oxytocin has two sites of origin, the ovary and
the hypothalamus. 36
 Hormones
Oxytocin also plays an important part in reproductive processes.

During the follicular phase of the estrous cycle and during the late
stages of gestation, oxytocin stimulates uterine contractions,
which facilitate sperm transport to the oviduct at estrus.

The stretching of the cervix at parturition caused by the passage of

the fetus stimulates a reflex release of oxytocin (Ferguson's
reflex).

However, the best known action of oxytocin is the reflex release

of milk.

37
Hormones
Oxytocin
In the lactating female, visual and
tactile stimuli associated with
suckling or milking induce the
release of oxytocin into the
circulation.

Oxytocin causes contraction of

myoepithelial cells (smooth muscle
cells) that surround the alveoli in the
mammary gland, resulting in milk
letdown (Fig. 3-7).
Ovarian oxytocin is involved in luteal
function. It acts on endometrium to
induce prostaglandin F2α (PGF2a )
release, which has a luteolytic action
(regression of the corpus luteum).

38
 Hormones
Melatonin

Melatonin (N-acetyl-5-methoxytrptamine) is synthesized in the

pineal gland. Pineal parenchymal cells take up the amino acid,
tryptophan, from the circulation and convert it to serotonin.

Two steps in the metabolism of serotonin are under neural

control.

The first is the conversion of serotonin to N-acetylserotonin,

which is followed by the conversion N-acetylserotonin to
melatonin.

The second step involves the melatonin-forming enzyme,

hydroxyindole-O-methyl-transferase (HIOMT) 39
 Hormones

Melatonin

Synthesis and secretion of melatonin is greatly elevated

during darkness.

Long daily periods of elevated secretion of melatonin are

probably responsible for the induction of ovarian cycles in
ewes and the .inhibition of cyclicity in mares.

40
 Hormones
Gonadal Steroid Hormones

The ovaries and testes primarily secrete gonadal sterok hormones.

Non gonadal organs such as the adrenals and the placenta also
secrete steroid hormones to some extent.

They are of four types: androgens, estrogens, progestins, and

relaxin. The first three types are steroids while the fourth is a
protein.

The ovaries produce two steroid hormones, estradiol and

progesterone, and a protein hormone, relaxin; the testis secretes a
single hormone testosterone
41
 Hormones
Steroid hormones secreted by the ovary, testes, placenta, and adrenal
cortex have a basic or common nucleus called the
cyclopentanoperhydrophenanthrene nucleus.

It consists of three, six...member fully hydrogenated (perhydro)

phenanthrene rings designated A, B, and C, and one five member
cyclopentane ring designated D.

An 18-carbon steroid has estrogen activity, a 19-carbon steroid has

androgen activity, and a 21-carbon steroid has progestogen
properties. Cholesterol, a 27-carbon steroid, becomes pregnenolone
(20-carbon) when its side chain is cleaved.

Pregnenolone is subsequently converted to progesterone, which is in

turn converted to an androgen and on to estrogens

42
 Hormones

In blood plasma, steroid hormone is mostly bound to albumin, a

plasma protein with low affinity and high capacity for steroids.

Another portion of the steroid hormone is bound to one or more

specific proteins with high affinity.

The half-life of naturally occurring steroids in the body is , very

short.

Therefore, several steroids with modified biochemical structure have

been synthesized for clinical use.

43
 Hormones
The secretory activity of steroid hormones by the gonads is under
endocrine control of the anterior pituitary.

Hypophysectomy or removal of the hypophysis before or after

puberty causes atrophy of the gonads, whereas injection of pituitary
preparation or implantation of pituitary tissue restores the secretory
activity.

Estrogens. Estradiol is the primary estrogen, with estrone and estriol

representing other metabolically active estrogens.

Several substances of estrogenic activity are found in both the

animal and plant kingdom.

44
 Hormones
Estradiol is the biologically active estrogen produced by the ovary
with smaller quantities of estrone.

Except for the possible secretion of small amounts of estriol in the

luteal phase of the cycle, most estriol and related urinary
estrogens are metabolic breakdown products of secreted
estradiol/estrone.

All ovarian estrogens are produced from androgenic precursors.

45
 Hormones
The biosynthetic pathways in all endocrine organs
that produce steroid hormones are similar, the
organs differing only in the enzyme systems they
contain.

The testis primarily synthesizes androgens, whereas

the ovaries synthesize two major types of steroids:
18-carbon estrogens and the 2-carbon progestins.

In blood plasma, steroid hormone is mostly bound

to albumin, a plasma protein with low affinity and
high capacity for steroids.

Another portion of the steroid hormone is bound to

one or more specific proteins with high affinity.

Figure 3 -10. The biosynthesis of steroid hormones from cholesterol. This

scheme provides a simplistic view of a highly organized and complicated
process that requires multiple enzyme systems.
46
 Hormones
Binding proteins in the circulation carry estrogens. Of all the
steroids, estrogens have the widest range of physiologic functions.
Some of these functions are:

Act on the CNS to induce behavioral estrus in the female; however,

small amounts of progesterone with estrogen are needed to induce
estrus in some species such as the ewe and cow.

Act on the uterus to increase both amplitude and frequency of

contractions by potentiating the effects of oxytocin and PGF2a•

Physical development of female secondary sexual characteristics.

Stimulate duct growth and cause the development of the mammary

gland.

47
 Hormones
Exert both negative and positive feedback controls on LH
and FSH release through the hypothalamus.

The negative effect is on the tonic center in the

hypothalamus, and the positive effect is on the preovulatory
center.

In ruminants, estrogens also have a protein anabolic effect

to increase body weight gain and growth.

The possible mechanism for increased growth may be due

to the ability of estrogens to stimulate the pituitary to
release more growth hormone.
48
 Hormones
Estrogens have been used to abort cows and sheep because of
their luteolytic properties (regression of CL) whereas in the sow,
estrogens have a luteotrophic action (helps to maintain CL).

49
 Hormones
Progestogens.
Progesterone is the most prevalent, naturally occurring
progestogen and is secreted by luteal cells of the corpus luteum,
the placenta, and adrenal gland.
Progesterone is transported in blood by a binding globulin as
for androgens and estrogens. LH primarily stimulates
progesterone secretion.
Progesterone performs the following functions:

Prepares the endometrium for implantation and maintenance of

pregnancy by increasing activity of secretory glands in the
endometrium and by inhibiting the motility of the myometrium.
50
 Hormones
Progestogens
Acts synergistically with estrogens to induce behavioral
estrus.

Develops the secretory tissue (alveoli) of the mammary

glands.

Inhibits estrus and the ovulatory surge of LH at high levels.

Thus, progesterone is important in the hormonal regulation

of the estrous cycle.

Inhibits uterine motility.

51
 Hormones
Progestogens

Synthetic progestogens are available to synchronize the estrous

cycles of ruminants. The progestogens act by inhibiting LH secretion
from the pituitary.

The hormone is either fed or inserted into the vagina as an

intravaginal device for a period of one estrous cycle length. On
cessation of the treatment, animals will display estrus and ovulate 48
to 72 hours later.

52
 Hormones
Androgens.

Androgens are 19-carbon steroids with a hydroxyl or oxygen

at positions 3 and 17 and a double bond at position 4.

The androgens are called 17-ketosteroids when oxygen is

found at position 17 (Fig. 3-10).

Testosterone is an androgen produced by the interstitial cells

(Leydig cells) of the testes, with a limited amount produced
by the adrenal cortex.

53
 Hormones
The functions of testosterone are:

Stimulate late stages of spermatogenesis and prolong the

life span of epididymal sperm.

Promote growth, development, and secretory activity of

the accessory sex organs of the male.

Maintain secondary sex characteristics and sexual

behavior or libido of the male

54
Relaxin
Relaxin is a polypeptide hormone consisting of alpha and beta
subunits that are connected by two disulfide bonds, and it has a
molecular weight of 5700 daltons.

Relaxin is secreted primarily by the corpus luteum during

pregnancy.

In some species, the placenta and uterus also secrete relaxin.

The main biologic action of relaxin is dilation of the cervix

and vagina before parturition.

It also inhibits uterine contractions and causes in creased

growth of the mammary gland if given in conjunction with
estradiol. 55
 Inhibins and Activins

Inhibins and activins were isolated from gonadal fluids because

of their effects on the production of FSH .

Inhibins and activins are paracrine regulators whereby they

modulate the endocrine LH signal.

Inhibins

The gonads are the main source of inhibin and related proteins,
which contribute to the endocrine regulation of the
reproductive system.

Sertoli cells in the male and the granulosa cells in the female
produce inhibins. 56
Inhibins.

Inhibins are not steroids but proteins comprising two disulfide

bridged subunits called a and β.

In the male, inhibins are secreted via the lymph and not by
venous blood as in the female.

Inhibins play an important role in the hormonal regulation of

ovarian folliculogenesis during the estrous cycle.

Inhibins act as chemical signals to the pituitary gland on the

number of growing follicles in the ovary.

57
Inhibins

Inhibins reduce the secretion of FSH to a level, which

maintains the species specific number of ovulation in both
single and litter bearing species.

By inhibiting FSH release without altering LH release,

inhibins may be partly responsible for the differential release
of LH and FSH from the pituitary.

Besides the regulation of pituitary FSH, inhibin related

proteins regulate Leydig cell function.

58
Activins.

Follicular fluid contains a fraction that stimulates rather than

inhibits the secretion of FSH.

The proteins responsible for this activity were characterized as

activins.

Activins are potent FSH-releasing dimers (dimers of inhibin-

subunits, (3) and are present in gonadal fluids, e.g., follicular
fluid and rete testis fluid.

These heterodimeric hormones are composed of a a-subunit

and one of two β-subunits (βA or, βB).

Activin is a fully functional member of the growth factors.

59
Follistatin.
Follistatin is another protein isolated from follicular fluid.

Follistatin not only inhibits the secretion of FSH similar to

that of inhibins but also binds activing and neutralizes its
biological activity, and thus, it modulates the secretion of
FSH.

Placental Hormones

The placenta secretes several hormones either identical to, or

with biologic activity similar to, hormones of mammalian
reproduction: equine chorionic gonadotropin (eCG), human
chorionic gonadotropin (hCG), placental lactogen (PL), and
protein B
60
Equine Chorionic Gonadotropin. The hormone eCG was
discovered when blood from pregnant mares produced sexual
maturity in immature rat.

eCG is a glycoprotein with a and f3 subunits similar to LH

and FSH but with a higher carbohydrate content, especially
sialic acid.

The higher sialic acid content appears to account for the long
half-life of several days for eCG.

Thus, a single injection of eCG has biologic effects on the

target gland for more than a week.

61
Equine Chorionic Gonadotropin
The equine uterus secretes this placental gonadotropin.

The endometrial cups are the source for the eCG, and the cups
that are formed at about day 40 of pregnancy persist until day
85 of pregnancy.

eCG has both FSH and LH biologic actions, with the FSH
actions being dominant, and eCG circulates in the blood of
pregnant mares and is not excreted in urine.

The secretion of eCG stimulates development of ovarian

follicles.

Some follicles ovulate, but most become luteinized follicles,

due to the LH like action of the eCG.
62
Equine Chorionic Gonadotropin

These accessory corpora lutea produce progestogens, which

maintain pregnancy in the mare. eCG was one of the first
commercially available gonadotropins used to induce
superovulation in farm animals.

Placental Lactogen.
Placental lactogen is a protein with chemical properties similar
to prolactin and growth hormone.

Its molecular weight is 22,000 to 23,000 daltons in the ovine

with 192 amino acids.

Placental lactogen is isolated from placental tissue but cannot

be detected in the serum of the pregnant animal until the last
63
trimester of pregnancy
Placental lactogen
Placental lactogen is more important for its growth hormone
properties than its prolactin properties.

It is important in regulating maternal nutrients to the fetus

and possibly is important for fetal growth.

Placental lactogen may play a role in milk production

because the level is higher in dairy cows (high milk
producers) than in beef cows (low milk producers).

64
Protein B.

The bovine conceptus produces numerous signals during

early pregnancy.

Currently only one protein from placental tissue has been

partially purified-pregnancy specific protein B (bPSPB).

The physiologic action of protein B may be involved in

preventing destruction of the corpus luteum in early
pregnancy of the cow or ewe.

This placental hormone has the potential to be the first

reliable hormonal pregnancy test for cattle.
65
Prostaglandins
Prostaglandins, first isolated from accessory sex gland fluids,
were termed prostaglandins because of their association with
the prostate gland.

Almost all body tissues secrete them. All prostaglandins are

20..carbon unsaturated hydroxy fatty acids with a cyclopentene
ring.

Arachidonic acid, an essential fatty acid, is the precursor for

prostaglandins most closely associated with reproduction,
mainly PGF2a and prostaglandin E2 (PGE2)

66
Prostaglandins

Most prostaglandins act locally at the site of their production

on a cell-to-cell interaction and therefore do not conform
exactly to the classic definition of a hormone.

Unlike other humoral agents, prostaglandins are not localized

in any particular tissue.

They are transported in the blood to act on a target tissue

away from the site of production.

Some forms never appear in the blood, whereas others are

degraded after they circulate throughout the liver and lungs.
67
Prostaglandins

PGF2a is the natural luteolytic agent that ends the luteal phase
(corpus luteum)of the estrous cycle and allows for the initiation of
a new estrous cycle in the absence of fertilization. It is particularly
potent in ending early pregnancy.

Prostaglandins may be considered hormones, which regulate

several physiologic and pharmacologic phenomena, such as
contraction of smooth muscles in the reproductive and
gastrointestinal tracts, erection, ejaculation, sperm transport,
ovulation, formation of the corpus luteum, partuntion, and milk
ejection.

Prostaglandins are involved in ovulation. For example, in the ewe

and cow, ovulation is blocked by the administration of
indomethacin, an inhibitor of prostaglandin synthesis. 68
Prostaglandins

Since LH release is unaffected in these animals, the action at the

level of the ovarian follicle involves either or both PGF2αand
PGE2

An increase in estrogen, which promotes myometrium growth in

the uterus, stimulates PGF2a synthesis and release.

In pregnant animals, the developing embryo sends a signal to the

uterus (maternal recognition of pregnancy), preventing luteolytic
effects of PGF2a .

The capacity of PGF2a to induce luteolysis has been exploited for

manipulating the estrous cycle and the induction of parturition.

69
Hormonal· Regulation Of Reproduction

The hypothalamus functions as an interface between the

nervous and endocrine systems and plays an important role in
the hormonal regulation of reproduction.

Edqvist has reviewed the endogenous control of reproductive

processes in domestic animals.

As stated earlier, the gonadal hormones inhibit the release of

LH and FSH in both sexes (negative feedback) whereas they
enhance the release of LH and FSH only in the female
(positive feedback).

70
Female Cow Hormones: Description and Function

71
72
Reproductive functions of secondary hormones

Thyroid hormones play important roles in regulating

reproductive functions in farm animals, influencing various
aspects of reproductive physiology and fertility.

Here are some examples of how thyroid hormones affect

reproductive functions in farm animals:

Puberty and Sexual Development

Thyroid hormones are involved in the onset of puberty and
sexual maturation in farm animals.

Adequate thyroid hormone levels are necessary for the

proper development of reproductive organs and the initiation
of reproductive behavior. 73
 Reproductive functions … Cont’d

Puberty and Sexual Development

For example, in female cattle, thyroid hormones influence the
development of the ovaries and the onset of estrous cycles.
In male pigs, thyroid hormones are involved in the development
of the testes and the initiation of spermatogenesis.
Estrous Cycle Regulation

Thyroid hormones play a role in regulating the estrous cycle in

female farm animals.

They can influence the timing and regularity of estrus (heat)

cycles, which are essential for successful breeding and
reproduction.
74
 Reproductive functions … Cont’d

Estrous Cycle Regulation

For instance, thyroid hormones can affect the production and

secretion of reproductive hormones such as follicle-
stimulating hormone (FSH) and luteinizing hormone (LH),

which control follicular development, ovulation, and the

formation of the corpus luteum in the ovaries.

75
 Reproductive functions … Cont’d

Ovulation and Fertility

Thyroid hormones can influence ovulation and fertility in

female farm animals.

They affect the development and maturation of ovarian

follicles, the timing of ovulation, and the quality of oocytes
(eggs).

In dairy cows, for example, thyroid hormones have been

shown to impact ovarian function and fertility.

Suboptimal thyroid hormone levels may lead to irregular

estrous cycles, reduced ovulation rates, and decreased
conception rates. 76
 Reproductive functions … Cont’d

Spermatogenesis and Sperm Quality

Thyroid hormones also play a role in male reproductive

function, including spermatogenesis (the production of
sperm) and sperm quality.

In male livestock species such as pigs and cattle, thyroid

hormones influence testicular development, testosterone
production, and sperm production.

Suboptimal thyroid hormone levels may impair

spermatogenesis and decrease sperm quality, affecting
fertility.

77
 Reproductive functions … Cont’d
Pregnancy and Parturition
Thyroid hormones can affect pregnancy outcomes and
parturition in farm animals.

They are involved in maintaining pregnancy, regulating

maternal metabolism, and supporting fetal growth and
development.

In sheep, for example, thyroid hormones play a role in

maintaining pregnancy and influencing fetal growth and
development.

Thyroid dysfunction during pregnancy may lead to

complications such as abortion, stillbirth, or dystocia (difficult
78
birth).
 Reproductive functions … Cont’d

Overall, thyroid hormones are important regulators of

reproductive functions in farm animals, influencing various
aspects of reproductive physiology, including puberty,
estrous cycle regulation, ovulation, fertility, and pregnancy.

Maintaining optimal thyroid hormone levels is essential for

ensuring reproductive health and fertility in livestock
production.

79
 Reproductive functions … Cont’d

Adrenal steroids

Adrenal steroids are a class of steroid hormones produced by

the adrenal glands, which are small glands located on top of
each kidney.

These hormones play diverse and crucial roles in regulating

various physiological processes in the body.

The adrenal glands also produce small amounts of other steroid

hormones, including androgens (e.g., dehydroepiandrosterone,
DHEA) and small amounts of estrogen and progesterone
precursors.

80