Anatomy

Two essential organs of reproduction are located within the head of the animal. The
hypothalamus controls several body processes and behaviors along with reproductive processes.
Body temperature, concentration and components of body fluids and the drive to eat and drink
are just a few functions of the hypothalamus. It is classified as a neuroendocrine gland since it
sends and receives neural signals through the nervous system and hormonal messages through
the endocrine system.

The second organ, the pituitary gland, sits at the base of the brain. The pituitary is about half an
inch in diameter and weighs about 1 gram. Physiologically, the pituitary is divided into two
distinct regions: the anterior and posterior pituitaries. Each region secretes various hormones that
direct body processes. Some of these hormones are responsible for reproductive events, while
others control growth, metabolism and water balance.

The female reproductive organs consist of the ovary, uterus, cervix, vagina and vulva. A sketch
of the reproductive tract of the cow is shown in Figure 1. Female reproductive tracts of various
farm animals are similar to the cow, but differ primarily in the shape of the uterus and cervix.

The ovary, or female gonad, is responsible for two basic functions

 Production of the female gamete, the egg or ovum

 Production of two primary reproductive hormones, estrogen and progesterone. A cow has
two bean-shaped ovaries located within the abdominal cavity. Size of the ovaries varies
with stage of the reproductive cycle and age of the female, but generally are 1 to 1-1/2
inches long.

The oviduct begins as a funnel-shaped tube that engulfs the ovary. This funnel portion of the
oviduct is called the infundibulum. When ovulation occurs, the ovum is picked up by the
infundibulum and channeled into the oviduct (also known as the Fallopian tube), where
fertilization takes place if viable sperm are present. Here the ovum remains capable of
fertilization for only a short time. Thus it is essential that sperm be present in the oviduct near the
time of ovulation. The ovum moves through the oviduct into the uterine horn within the next 3 to
4 days. If the ovum is fertilized, it then begins embryological development; if not, it degenerates
and disappears and the next estrous cycle ensues.

The body of the uterus of the cow is short and poorly developed, while the uterine horns are
relatively long and well developed. The fertilized embryo moves from the oviduct into the
uterine horn, where fetal and maternal membrane development begins. This newly developing
fetus grows within a layer of membranes called the placenta, through which nourishment from
the dam diffuses. There is no direct blood connection between the fetus and the dam, but rather a
complex system that selectively allows certain molecules to pass from the maternal side of the
placenta to the fetal side and vice versa. It also provides nutrients and carries waste products
from the fetus.

The cervix is in effect the neck of the uterus. It has thick walls and a small opening which
softens and relaxes to allow a passageway for sperm at mating and expulsion of the fetus at the
time of birth. During pregnancy, the cervix is filled with a thick mucus secretion known as the
cervical plug, which protects the uterus from infections entering from the vagina. The cervical
plug is expelled and the cervical opening begins to dilate in the days prior to calving.

The vagina serves as a receptacle for the male's penis during service. In the cow, the semen is
deposited in the vagina near the cervix during natural mating with the bull. When artificial
insemination is used, an insemination instrument is threaded through the vagina and cervix and
semen is deposited at the uterine side of the cervix. Urine is discharged from the urinary bladder
through the urethra, which opens into the base of the vagina. The region behind the urethral
opening is called the vestibule and is a common passageway for both the urinary and
reproductive systems. The external opening of the vagina is called the vulva.

OVARY:

The ovary is the female Gonad homologous to the male Testes. It is usually a paired organ in
domestic species, but in the bird only the left Ovary is present. The structures found within the
ovary are undergoing constant changes throughout the oestrus cycle from the Follicles containing
Oocytes, to the formation of Corpus Haemorrhagicum, Corpus Luteum, and finally Corpus
Albicans. Ovaries are ellipsoidal in shape with an irregular surface due to the projection of
dominant follicles and corpora lutea. These irregularities are absent in the mare due to the cortex
and medulla being reversed with ovulation only occuring from the ovulation fossa. They are
greatest in Polytocious animals such as the sow due to many dominant follicles, and so corpora
lutea, developing at once.

Anatomical Location
 In the dog and cat, ovaries do not migrate in development. They remain in the dorsal part
of the abdomen, caudal to the kidneys at the tip of the uterine horns.
 In other domestic species, the ovaries migrate during development.
 The greatest degree of migration occurs in ruminants. Here, the ovaries come to lie close
to the ventral abdominal wall, cranial to the pelvic inlet.
 In the pig, they descend to the middle of the abdomen.
 In the mare, they are located ~8-10 cm ventral to the dorsal wall of the abdomen.
 In all domestic species except the horse, ovaries are ellipsoidal in shape and their surface
has large follicles and corpora lutea.
 The ovaries of the mare are kidney shaped and the surface is smooth.
 The ovaries are suspended in the abdominal cavity by the Mesovarium of the Broad
Ligament.
Structure
Outer Tunica Albuginea
 Connective tissue layer covering the ovarian cortex.
 Overlying this structure is a single layered Germinal epithelium.
Ovarian Cortex
 Contains:
o Follicles in various stages of development
 o Corpus Luteum (includes Corpus Haemorrhagicum and Corpus Albicans)
Ovarian Medulla
 The Medulla is made up of dense connective tissue. This is where all of the lymphatics, nerves
and vasculature of the Ovary are found.

The Mare

The Ovaries of a Mare showing the Ovulation Fossa-

 The cortex is in the centre, follicles develop here.
 This layer is enclosed by a dense, richly vascularised connective tissue layer which is analagous
to the medulla of other domestic mammals.
 The cortex reaches the surface of the ovary at the ovulation fossa, a deep indentation at the
free margin. This is where mature follicles rupture in ovulation, as opposed to at various points
on the surface in other domestic mammals.
 Follicles can be identified by transrectal palpation, but Corpora Lutea cannot. Identification of
Corpora Lutea requires ultrasonography.

Histology

Histological Section of a Rabbit Ovary under Low Power- Courtesy of J.Bredl, Copyright RVC 2008
Stroma
 The body of the ovary (ovarian stroma) consists of:
o spindle-shaped cells
o fine collagen fibres
o ground substance
 Stromal cells resemble fibroblasts, but some contain lipid droplets.
 Bundles of smooth muscle cells are scattered throughout the stroma.
Cortex
 Follicles containing oocytes in various stages of development.
 Atretic Follicles
 Corpora lutea
 Corpora albicans
  The superficial cortex is more fibrotic than the deep, and is called the tunica albuginea.
 On the surface of the ovary is the germinal epithelium. This is a continuation of the peritoneum.

Medulla
 Highly vascular
 Contains hilus cells, which are similar to the Leydig cells of the testes.

Arterial Supply
 The ovarian artery (a branch of the Aorta) and ovarian branches of the Uterine artery form
anastomoses in the mesovarium and the broad ligament.
 From this arterial plexus ~10 coiled Helicine arteries enter the hilus of the ovary.
 Smaller branches form a plexus at the corticomedullary junction, giving rise to straight Cortical
arterioles, which radiate into the cortex.
 Here they branch and anastomose to form vascular arcades, which give rise to a rich capillary
network around follicles.
Venous Drainage
 Venous drainage follows the course of the arterial system.
 Medullary veins are large and tortuous.
 The Ovarian Artery is closely associated with the Uterine Vein. This is important for the transfer
of luteolytic PGF2α from the Uterus to the Ovary.
Lymphatics
 Arise in the perifollicular stroma.
 Drain into larger vessels, which coil around the medullary veins.
Innervation
 Sympathetic fibres of the autonomic nervous system supply blood vessels and terminate on
smooth muscle cells in the stroma around follicles.
 May play a role in follicular maturation and ovulation, but the main control is via the endocrine
system.
Function
It has two main functions:
 Producing the female gametes oocytes via Gametogenesis.
 Producing the reproductive hormones Oestrogens and Progesterone, an endocrine function.
 PHYSIOLOGY OF REPRODUCTIVE HORMONES; HYPOTHALAMIC AND PITUTARY
HORMONES

DEFINITION
1. Hormone = chemical substance secreted from tissues or ductless glands into surrounding tissue
or the bloodstream to evoke a specific physiologic response
2. The specific response occurs at a given tissue because of presence of receptors on those tissues.
Knowledge about receptors and binding guides clinical decision-making - e.g. oxytocin does not
dissociate readily from myometrial receptors so, at some point, continued dosing is not effective.

CHEMICAL STRUCTURES
1. Proteins - e.g. LH, FSH. Unique structure in each species, requires unique assay in each species
2. Steroids - e.g. progesterone, estrogen, testosterone. Same structure in all species, common
assay - Canine progesterone can by assayed using a bovine or human assay.

CONTROL OF HORMONE SECRETION

1. Feedback systems - e.g. increased serum testosterone concentration has negative feedback on
the pituitary, decreasing release of LH
2. Homeostasis - e.g. increased blood glucose stimulates release of insulin
3. Environmental stimuli - e.g. Suckling of neonates stimulates oxytocin release from the
pituitary, daylength controls reproductive function in several species via the pineal gland

ENDOCRINE TISSUES AND THEIR PRODUCTS

1. HYPOTHALAMUS - The hypothalamus is connected directly to the pituitary by the

hypophysial portal system. Hormones released from the hypothalamus are never in the general
circulation.
 a. Gonadotropin releasing hormone (GnRH) - causes release of LH and FSH from the
pituitary
b. Thyrotropin releasing hormone (TRH) - causes release of TSH from the pituitary

2. PITUITARY
a. Posterior lobe = neurohypophysis - The actual cell bodies producing these hormones are
neurons in the hypothalamus, their axons lie in the posterior pituitary. Neural stimuli cause
release of these hormones.
i. Oxytocin - stimulates mammary gland to cause milk letdown,
gravid uterus at parturition to cause uterine contraction and cervical dilation
b. Anterior lobe = adenohypophysis - This is a true gland; hormones are
produced and stored here
i. Follicle stimulating hormone (FSH) - stimulates growth of
follicle on ovary (female) and production of androgen-binding protein to maintain high
intratesticular testosterone concentrations (male)
ii. Luteinizing hormone (LH) - causes ovulation of mature ovarian follicle, maintains CL on
ovary after ovulation (female), stimulates and sustains testosterone release and spermatogenesis in
testicle (male)
iii. Prolactin - stimulates mammary gland to promote milk
production, induces maternal behavior
iv. Thyroid stimulating hormone (TSH) - stimulates thyroid gland
to assemble T3 and T4
v. Adrenocorticotropic hormone (ACTH) - stimulates adrenal to
produce steroids

3. THYROID - thyroid hormones stimulate oxygen consumption and increased metabolism

in all body tissues. Either hypothyroidism or hyperthyroidism is associated with poor reproductive
function but exact cause-and-effect relationship is not reported
a. Tri-iodothyronine (T3)
b. Thyroxine (T4)

4. ADRENAL
a. Sex steroids
b. Glucocorticoids
c. Epinephrine, norepinephrine

5. OVARY
a. Follicle
i. Estrogen - stimulates female sexual development, breeding
behavior, physical changes of heat (vaginal cornification, vulvar swelling, serosanguinous vulvar
discharge)
b. Corpus luteum (CL)
i. Progesterone - responsible for pregnancy maintenance, uterine
and mammary growth, induction of maternal behavior
ii. Relaxin - relaxes pubic symphysis, contributes to cervical dilation late in gestation
6. PLACENTA/PREGNANCY-SPECIFIC TISSUES
a. Human - Human chorionic gonadotropin (hCG) produced by
fetal/placental unit as early as 6 days post-conception. This is what is used to identify pregnancy
in human early pregnancy tests. In dogs, has LH activity.
b. Horse - Equine chorionic gonadotropin (eCG = PMSG) produced by
endometrial cups. In dog, has FSH and LH activity.
c. Dog - No pregnancy-specific gonadotropins produced. Some relaxin may be produced at the
placenta.
d. Cat - No pregnancy-specific gonadotropins produced, progesterone
produced by CL and may be produced by the fetal/placental unit

7. TESTICLE
a. Mullerian inhibiting factor (MIF) - inhibits female ductal development during
embryogenesis
b. Testosterone - Stimulates male sexual development and behavior, spermatogenesis

Reproductive Hormones

GnRH = gonadotropin-releasing hormone - A protein hormone produced by the hypothalamus

which stimulates (under negative feedback control by either estrogens, progesterone, and inhibin
in females or androgens and inhibin in males) the anterior pituitary gland to produce and begin
secreting luteinizing hormone = LH and follicle-stimulating hormone = FSH.

luteinizing hormone (LH) - The adenohypophyseal protein hormone which: (1) in the female:
(in cooperation with FSH), regulates the ovarian cycle, stimulates ovulation and development of
the corpus luteum and, thus, estrogen and progesterone production; (2) in the male: stimulates
production of androgens, especially testosterone from interstitial cells; in males, it is also known
as Interstitial Cell Stimulating Hormone (ICSH); LH secretion is regulated by hypothalamic
GnRH.

follicle-stimulating hormone (FSH) - The adenohypophyseal protein hormone which: (1) in the
female: (in cooperation with LH), regulates the ovarian cycle, stimulates maturation of the
ovarian follicle and estrogen production; (2) in the male: stimulates sperm production; FSH
secretion is regulated by hypothalamic GnRH.

testosterone - A steroid hormone, the major androgen of human males, produced primarily in
the interstitial cells of the testes in response to LH = ICSH and responsible for the development
and maintenance of male secondary sex characteristics, sperm production, virility and libido
(sexual desire); it is also produced synthetically for use in medical treatment. [Did you know
testosterone levels in utero influence male behavior later in life?

androgens - The general name given to the class of male sex hormones, e.g., testosterone or
androsterone, which are steroids produced by the interstitial cells of the testes in response to LH
= ICSH, and, in much lesser amounts, by the adrenal cortex; these hormones control the
development and maintenance of male secondary sex characteristics, sperm production, virility
and libido (sexual desire); they also increase nitrogen and water retention and stimulate skeletal
muscle growth.

anabolic steroids - Any of the group of synthetic analogs of testosterone, which encourage
growth, especially of bone and muscle mass, and have lesser androgenic properties, which are
used clinically to promote growth and repair in body tissues, to treat anemias, and for patients in
negative nitrogen balance; they are also widely misused and abused by athletes, body-builders,
and workers with strenuous jobs, e.g., soldiers, policemen, firemen, etc, who desire increased
muscular strength; these drugs have many serious negative side-effects with long term use.

estrogens - Any of several steroid (cholesterol-based) hormones produced chiefly by the ovaries,
and in minute quantities by the adrenal cortex, and responsible for promoting the development
and maintenance of female secondary sex characteristics as well as regulating the uterine cycle to
produce an environment suitable for the fertilization, implantation and nutrition of the
developing embryo; also produced by the placenta during a pregnancy; their output is regulated
primarily by FSH from the anterior pituitary.

progesterone - A steroid (cholesterol-based) hormone secreted by the corpus luteum of the

ovary and by the placenta, which acts to prepare the uterus for implantation of the fertilized
ovum, to maintain pregnancy, and to promote development of the mammary glands; it also
provides negative feedback to the ovary to prevent further follicular development; its output is
regulated by FSH and especially LH from the anterior pituitary.

relaxin - A female protein hormone secreted by the corpus luteum which helps to soften the
cervix and to relax the pelvic ligaments and pubic symphysis and causes appropriate smooth
muscle relaxation in childbirth; in males, it is secreted by the testes, and encourages the motility
of spermatozoa within semen.

inhibin - A polypeptide/protein hormone secreted by the follicular cells of the ovary and the
sustentacular = Sertoli cells of the testis which inhibits secretion of follicle stimulating hormone
(FSH) from the anterior pituitary; it exerts negative feedback control on the hypothalamus and
pituitary by increasing its blood levels as spermatogenesis or corpus luteum activity increase.
prolactin (PRL) - The adenohypophyseal protein hormone which stimulates the mammary
glands to produce milk; its secretion is regulated by hypothalamic PRH and aided by negative
feedback from the infant's suckling at the nipple.

oxytocin (OT) - A neurohypophyseal peptide hormone which stimulates the contraction of

smooth muscle of the uterus during labor and delivery and facilitates ejection of milk from the
breast during nursing; its output is aided by negative feedback from the infant's suckling at the
nipple.

human chorionic gonadotropin (hCG) - The protein hormone produced only by the placenta
after successful implantation of a developing embryo which signals to the hypothalamus,
pituitary gland, and ovary that a pregnancy has begun; this positive feedback regulation
maintains the output of estrogens and progesterone by the corpus luteum during pregnancy; this
is also the hormone detected in over-the-counter urine pregnancy tests.

prostaglandins - Any of the group of ~20 potent local hormones which are produced in various
tissues, are derived primarily from arachidonic acid (a cell membrane phospholipid) via the
cyclooxygenase pathway, and mediate a wide range of physiological functions, e.g., control of
blood pressure, contraction of smooth muscle, and modulation of inflammatory and immune
responses; their synthesis is inhibited by aspirin and other NSAIDs.

GROWTH, PUBERTY AND ESTROUS

CYCLE:

Puberty is the time in an animal's life cycle when the animal reaches a developmental state in
which it is capable of reproducing. By defination, it is the time in adolescence when male and
female gonads are capable of releasing gamates. This happens several weeks before sexual
maturity; the first gamates are usually incapable of fertilization. In a production system that relies
on reproduction to be profitable, the sooner an animal reaches puberty, the sooner the manager
will be able to realize a profit from the animal.

A number of factors influence the age at which an animal reaches puberty. Among the factors
are:

o Animal species - smaller species usually reach puberty before larger species
o Breed - some breeds within a species reach puberty earlier than other breeds
 in cattle, Hereford and Angus reach puberty before Brahma and Zebu
 in swine, Landrace reach puberty earlier than other breeds
o Climate - animal generally have an earlier onset of puberty in warmer climates,
however excessive heat may actually delay the onset of puberty
o Season - many breeds and species of animals are seasonal breeders
 Long-day breeders begin to cycle and breed during increasing day length
 Horses, chickens, swine are examples of long-day breeders. While
this trait has been bred out of many of our domesticated swine and
chickens, a decrease in fertility (fall infertility syndrome, or fall
abortion syndrome in swine) is thought to be due, at least in part, to
the effect of decreasing day length.
 Short-day breeders begin to cycle and breed and are most fertile during
decreasing day length
 Sheep, goats, deer, and elk are examples of species who are
normally fertile and actively breeding in the fall of the year during
decreasing day length. Exceptions to this in the Ovine species are
the Dorset and Ramboullet breeds of sheep, who have been selected
to remain fertile and capable of breeding at other times of the year.
o Nutrition - an adequate, well-balanced diet is necessary for animals to reach
puberty at an optimal time. Severely malnurished animals will have a delayed onset
of puberty.
o Sex - the female of the species or breed usually reaches puberty earlier than male
animal.
o Management system - the type of housing, pen arrangement, lighting, and the
presence or absence of mature animals will affect the onset of puberty.
 the sight, smell and sound of the opposite sex enhances the onset of puberty
in swine
o Stress - recent studies indicate that low levels of stress results in early onset of
puberty, excess stress, however, may delay the onset of puberty

Once animals reach puberty, we tend to concentrate our attention on the female, since she is
producing the desired product. It is not necessary to keep as many males as females for breeding
purposes, and when artificial insemination (AI) is used, there may be no intact male animals in a
livestock production system. Through use of AI, new genetics can be introduced into the herd or
flock without running the risk of inroducing disease agents that may be carried by an apparently
healthy animal. In addition, male animals tend to be aggressive and unpredictable, often fighting
among themselves. However, until or unless cloning of animals from somatic cells becomes
economically feasible and socially acceptable the male will continue to play an important role in
reproduction.

The Estrous Cycle -- A Female Thing