OESTROUS CYCLE

Dr.R.S.Abhilash,
Assistant Professor,
Dept. of ARGO

After puberty the female enters a period of reproductive cyclicity, which continues,
throughout most of her productive life. The word "Oestrous" is derived from a Greek word
"Oistros" meaning "Gad fly" - used to describe the behaviour of cows when attacked by such
flies. They continue throughout the adult female‟s life and are interrupted by pregnancy,
nursing and by season in some species. Oestrous cycles provide females with repeated
opportunities to copulate and become pregnant. If conception fails, another oestrous cycle
begins, providing the female with another opportunity to mate and conceive.
Oestrous (UK English)/ estrous (US English) : Noun
Oestrus (UKEnglish)/ estrus (US English): Adjective
Animals could be categorized based on the occurrence of oestrous cycle as:
Monoestrus: Only one oestrous cycle per year. eg. Wild animals.
Polyoestrus: Periodic oestrous cycles throughout the year. eg. Cow and sow.
Seasonally polyoestrus: Periodic oestrous cycles only during a particular season. eg. Sheep
and mare.

Depending on the ovarian activity again oestrous cycle is classified as:

Regular oestrous cycle: Characterised by ovulation and formation of corpus luteum. eg.
Cow, sheep, mare, bitch and sow

Spontaneous ovulators: Ovulations are spontaneous but the corpus luteum formed will not
be functional until mating has occurred. eg., rat or mouse

Induced ovulators: Ovulation and corpus luteum formation depends on whether mating has
occurred or not. eg. Cat, rabbit, mink, camel.

PHASES OF ESTROUS CYCLE

Follicular Phase

It is the period from the regression of corpora lutea to ovulation. The primary ovarian
structure is the Graafian follicle and the primary reproductive hormone is oestrogen.

Luteal Phase

The luteal phase is much longer than the follicular phase and extends from ovulation to
luteal regression. Predominant ovarian structure is the corpora lutea and the primary
reproductive hormone is progesterone. Even though the luteal phase is dominated by corpus
luteum, follicles continue to grow and regress during this phase. However, these follicles do
not produce sufficient amount of oestrogen to bring about oestrus and ovulation.

Stages of the cycle

Proestrus

Proestrus is the "building up period" and begins with luteal regression and ends with
onset of estrus. Duration is 2 to 5 days and is characterised by a major endocrine transition,
from a period of progesterone dominance to a period of oestrogen dominance brought about
by FSH and LH. Graafian follicle is growing under the influence of FSH and producing
increasing amounts of oestradiol. Regression of CL of previous cycle (in polycyclic animals).
Uterus enlarges, endometrium becomes congested and oedematous, and its glands show
evidence of increased secretory activity. Vaginal mucosa becomes hyperaemic; the number of
cell layers of the epithelium starts to increase, and the superficial layers become cornified. In
cattle the mucus changes to milky, viscid mucus to finally a clear, transparent, stringy mucus.
Late in this phase the female animal usually exhibits interest in the male. The shows external
evidence of prooestrus with vulval oedema, hyperaemia and a sanguineous vulval discharge.

Oestrus

Oestrus is the period of sexual receptivity and mating. The onset and end of the phase
are the only accurately measurable points in the oestrous cycle Oestradiol is the main
reproductive hormone responsible for oestrous behaviour and physiological changes in the
reproductive tract. Duration is 18 to 24 hrs. The animal exhibits sexual desire. The animal
usually seeks out the male and „stands‟ for him to mate her. The uterine, cervical and vaginal
glands secrete increased amounts of mucus; the vaginal epithelium and endometrium become
hyperaemic and congested; the cervix is relaxed. In most domestic animals, ovulation occurs
during oestrum, except in cow, where ovulation occurs about 12 hours after the end of
oestrus. Ovulation is a spontaneous process in all domestic species with the exception of the
cat, rabbit, camel and ferret – induced ovulators.

During pro-oestrus and oestrus there is follicular growth in the absence of functional
corpora lutea.

Metoestrus
 Metoestrus is the period from ovulation and formation of corpora lutea and poorly
defined period following oestrum. It is an ill defined phase. Duration is 3 to 5 days. CL grows
rapidly from granulosa cells of the ruptured follicles under the influence of LH. There is a
reduction in the amount of secretion from the uterine, cervical and vaginal glands.
Metoestrum is largely under the influence of P4 produced by CL – inhibits FSH secretion by
pituitary, prevents development of another oestrum.

During metoestrum in some species the epithelium of the vaginal loses most of its new
growth. In the cow during early part of metoestrum epithelium over caruncles of the uterus is
very hyperaemic and some capillary haemorrhage occurs – metoestrual bleeding – occurs due
to withdrawal of oestrogen. Metoestrual bleeding is seen in cows about 15-36 hrs after
ovulation or 24-48 hrs after oestrum. It is observed in about 75% of heifers and 48% of cows.
This does not indicate either a failure or a successful pregnancy. (74 % of the heifers do not
show oestrus signs in the first oestrum).

“Menstruation” in primates – occurs at the time of P4 withdrawal – associated with a loss of

superficial layers of endometrium. In case of animals having diestrum, metoestrum is about
equal to the time it takes for the ova to reach the uterus about 3-4 days.

Dioestrus

Diestrus is the longest phase of the oestrous cycle characterised by a fully functional
corpus luteum and high levels of progesterone. Duration is 10-12 days. Females do not exhibit
sexual receptivity. The CL has matured and is fully functional, effects of P4 on reproductive
tract are marked. Endometrium becomes thicker and glands hypertrophy. Cervix is
constricted, vaginal mucus is scant and sticky. Mucus membrane of vaginal is pale. Uterine
muscles are relaxed. Late in the period the CL shows retrogressive changes. Endometrium
and its glands atrophy or regress in size. This leads to initiation of development of primary
and secondary follicles.

REGULATION OF OESTROUS CYCLE

Regulation of cyclical activity is under the control of the hypothalamic–pituitary–
ovarian axis. The pineal gland plays an important role in controlling reproduction in seasonal
breeding species by influencing the release of FSH, LH and prolactin.
The hypothalamus releases GnRH, which is responsible for the control of release of
gonadotrophins from the anterior pituitary. Noradrenaline stimulates the release of FSH and
LH; serotonin inhibits the basal secretion of LH. In the domestic species the secretion of FSH
and LH is controlled by two functionally separate, but superimposable, systems:
1. Episodic/tonic system - continuous basal secretion of gonadotrophin and stimulates the
growth of both germinal and endocrine components of the ovary,
2. Surge system, which controls the short-lived massive secretion of gonadotrophin,
particularly LH, responsible for ovulation. The positive-feedback effect of oestradiol
leads to preovulatory surge.
Cats, rabbit, camel, and mink are some animals which are induced ovulators.
The episodic/tonic hypothalamic release centre is influenced by the negative-feedback
effect of oestradiol and progesterone. FSH secretion is also controlled by a number of
ovarian-derived peptide hormones. Inhibin (produced by the granulosa cells of large antral
follicles) and oestradiol act in concert in suppressing FSH secretion. Two other peptide
hormones isolated from ovarian follicular fluid: activin, which stimulates, and follistatin,
which suppresses, FSH secretion.
The negative feedback of progesterone is mediated via a reduction in pulse frequency
of gonadotrophin release, whereas oestradiol exerts its negative feedback effect via a reduced
pulse amplitude.
Progesterone is the main regulatory hormone which controls the oestrous cycle of
most of the domestic animals.
Follicular growth
Throughout the oestrous cycle, during pregnancy and other reproductive stages, there
is dynamic follicular activity with growth and atresia; only about 1% of antral follicles
subsequently ovulate. In cattle, the growth of the follicles during oestrous cycle occurs in a
wave like pattern and majority of the cycle comprised of two to three waves. There are
individual variations in follicular dynamics among cattle and three wave patterns are found to
be most common.

Follicular growth involves three different stages viz.

Recruitment – gonadotrophin stimulation of a pool of rapidly growing follicles. Recruitment

defines the transition period when the primordial follicles leave the non-growing phase and
enter into the growing pool of follicular population. In cattle, a follicle becomes antral when it
reaches a diameter of 0.2 to 0.4 mm diameter and they are independent of gonadotrophin for
its growth. In cattle, there will be recruitment of 10- 40 follicles which gain the ability to
respond to gonadotropin later and require gonadotropin for its continued growth.
Selection – After recruitment, few follicles from this group will be selected for
continued growth while the rest will undergo atresia. During selection of follicles, declining
concentrations of FSH are required.
Dominance – Those follicles greater than 5mm diameter have the ability to suppress the level
of FSH by means of intra- ovarian as well as feedback inhibitions by inhibin. A dominant
follicle (DF) will acquire more number of LH receptors in granulosa cell layer and will
synthesize more oestradiol. So even in FSH decline, the dominant follicle can shift its
gonadotropin dependence to LH and can continue to grow while subordinates regress. DF will
suppress the growth of subordinate follicles as well as new follicular wave emergence by
systemic control through endocrine pathways. The DF will have larger diameter, more LH
receptors and oestradiol level than the second largest follicle (SF) and these differences are
called as deviation. The DF will be morphologically and functionally dominant as compared
to the SF.

After ovulation the corpus luteum (CL) is rapidly formed from the granulosa and the
thecal cells. Although CL develops as a result of ovulation, in some species, like the bitch,
there are early signs of luteinisation of the follicle before it has ovulated. The hormones which
are involved in the formation of CL are prolactin and LH. Endogenous peptides with opioid
activity like β-endorphin and met-enkephalin inhibits the secretion of FSH and LH whilst
stimulating the secretion of prolactin.
The presence of a functional CL, due to its production of progesterone, inhibits the return to
oestrus by exerting a negative feedback effect upon the anterior pituitary
 Mechanism of ovulation

CHANGES FOLLOWING OVULATION

Following ovulation the theca interna and the granulosa cells of the follicle undergo
luteinisation. Immediately after ovulation the walls of the follicle collapse into many folds.
These folds begin to interdigitate, allowing thecal cells and granulosa cells to mix. The luteal
tissue consists of large and small luteal cells. The large cells originate from granulosa cells
and secrete oxytocin and progesterone. The small cells originate from cells of theca interna
and secrete progesterone only.

The functions of progesterone are

 1. stimulates hypertrophy of endometrial glands,
2. increases uterine milk secretions,
3. inhibits contractions of uterus
4. prevents neutrophilic infiltration,
5. plays an important role in endocrine control of reproduction because it exerts a
strong negative feedback on the hypothalamus thereby preventing development
of
a. preovulatory follicles (follicles develop during luteal phase but
do not reach preovulatory stage unless progesterone decreases),
b. production of estrogen,
c. behavioural estrus,
d. preovulatory surge of GnRH and LH.

If the animal is not pregnant, the CL undergoes luteolysis. Luteal oxytocin and PGF2
alpha from endometrium cause luteolysis. Oestrogen from the follicular waves causes
formation of oxytocin receptors in the uterus. The oxytocin from the CL binds to oxytocin
receptors in the endometrium and activates the enzyme system necessary for formation of
PGF2 alpha. The PGF2 alpha reaches the ovaries by a vascular counter current exchange
system.

Luteolysis
Prostaglandin appears to be the principle agent involved in luteolysis in most of the
domestic species. The luteolytic PGF2 is produced from the endometrium. In many species
like cow, mare, ewe, goat and sow, removal of part or the entire uterus will result in the
prolongation of the life span of the CL. In the human, dog and cat the normal life span of the
CL is unaltered in the absence of the uterus. In the cow, ewe and goat the „luteolytic‟ action of
the uterine horn is directed exclusively to the CL on the adjacent ovary. Thus, if one of the
uterine horns is surgically removed on the side adjacent to the ovary with a CL then the latter
will persist. If the contralateral horn is removed, then the CL will regress at the normal time.
In these species the PGF2 is transported directly from the uterus to the ovary. In the ewe the
most likely route for transport of the substance is the middle uterine vein. In the mare the
luteolysin is transported throughout the systemic circulation.
In the pig the luteolytic substance is transported locally but not exclusively to the
adjacent ovary. Following surgical ablation of parts of the uterine horns, provided at least the
cranial quarter of the uterine horn is left, regression of the CLs occurs in both ovaries. If more
than three-quarters of the horn is excised, then regression of the CLs occurs only in the ovary
adjacent to the intact horn.

The close proximity of the ovarian artery and uteroovarian vein is allows the leakage
of the luteolytic substance from the uterine vein into the ovarian artery and thus to the ovary,
by a form of counter-current exchange through the walls of the vessels.
Mechanism of Luteolysis

In ruminants, luteal regression is caused by episodic release of PGF2α from the uterus
at intervals of about 6 hours. This is induced by oxytocin secreted by the CL; thus, each
episode of PGF2α release is accompanied by an episode of oxytocin release. Furthermore,
PGF2α stimulates further secretion of oxytocin from the ovary. The sensitivity of the uterus to
oxytocin is determined by the concentration of endometrial oxytocin receptors. At the time of
luteal regression in sheep they rise approximately 500-fold. Their concentration is determined
by the effects of progesterone and oestradiol. Thus, the high concentrations of progesterone
which occur after the formation of the CL reduce the number of receptors, so that in the
normal oestrous cycle of the ewe they start to increase from about day 12. Exogenous
oestradiol causes premature induction of oxytocin receptors, resulting in premature luteolysis.
The CL becomes more sensitive to the leuteolytic effect of PGF2α as it ages. The early CL is
unresponsive to PGF2α. In case of cattle CL would not be responsive to exogenous PGF2α till
about day 6 of the cycle, and further from about day 16 of the cycle, when luteolysis would be
initiated by the endogenous release of PGF2α.

Luteal regression involves two aspects:

1. Functional regression - secretion of progesterone declines rapidly.

2. Structural regression - CL is reduced in size, process takes more time than the
functional regression

Luteolysis is brought about by

1. Reduction in blood flow to the CL by vasoconstriction.

 2. PGF2 alpha binds to specific receptors on large luteal cells causing influx of Ca
ions. High intracellular Ca ions are thought to cause apoptotic effects
(programmed cell death).
3. PGF2 alpha receptor complex also activates protein kinase C (PK-C) that
inhibits progesterone synthesis.

Cattle

Under conditions of domestication, normal and well-cared-for cattle are polyoestrous

throughout the year. The age at first oestrus, or puberty, is affected by nutrition and season of
birth. A small proportion of heifers do not ovulate at the first heat, and in a majority of young
cattle the oestrus associated with the first ovulation is „silent‟. Poor feeding and calfhood
disease delay puberty. Once puberty has been reached, cyclical activity should persist, except
during pregnancy, for 3–6 weeks after calving, during high milk yield (especially if there is
some evidence of dietary insufficiency), and with a number of pathological conditions. Some
cows and heifers also fail to show overt signs of oestrus yet have normal cyclical activity, a
condition referred to as „silent heat‟ or suboestrus.
In heifers, the average length of the oestrous cycle is 20 days, and in cows 21 days, the
normal ranges being 18–22 and 18–24 days, respectively. The average duration of oestrus is
about 15 hours with a wide range of 2–30 hours. There are a number of factors which can
influence the duration: breed of animal, season of year, presence of a bull, nutrition, milk
yield, lactation number and, perhaps most important, the number of cows that are in oestrus at
the same time. There is also good evidence that more signs of oestrus are observed during the
hours of night, perhaps when the animals are least. Ovulation is spontaneous, and occurs on
average 12 hours after the end of oestrus.
Signs of oestrus
Accurate detection of oestrus is paramount in ensuring optimum fertility wherever AI
is used. There are great variations amongst individual cattle in the intensity of heat signs; the
manifestations tend to be more marked in heifers than in cows. However, it is generally
agreed that the most reliable criterion that a cow or heifer is in oestrus is that she will stand to
be mounted by another. The oestrous animal is restless and more active; an average increase
in activity of 393% is observed. There tends to be grouping of sexually active individuals;
there is a reduction in the time spent eating, resting and ruminating, and frequently a
reduction in milk yield. – which is a reliable indicator of the onset of oestrus. As the cow
approaches oestrus she tends to search for other cows in oestrus, and there is licking and
sniffing of the perineum. During this period, during oestrus and just afterwards, the cow will
attempt to mount other cows. A positive mounting response lasts about 5 seconds; however, if
both cows are in oestrus it will be increased to about 7.5 seconds.
Sometimes there are signs of a vulval discharge of transparent mucus whose elasticity
causes it to hang in complete clear strands from the vulva to the ground; it also adheres to the
tail and flanks. The vulva may be slightly swollen and congested, and there is a small
elevation of temperature. The tail may be slightly raised. The hair of the tail-head is often
ruffled and the skin sometimes becomes excoriated due to mounting by other cows. When she
is put with a bull, the two animals lick each other and the cow often mounts the bull before
standing to be mounted by him. For a short time after service, the cow stands with raised tail
and arched back, and where actual service has not been seen this posture indicates that mating
has occurred.
Within 2 days of service, there is an occasional yellowish-white vulval discharge of
mucus containing neutrophil leucocytes from the uterus. At about 48 hours after heat,
irrespective of service, heifers and many cows show a bright red sanguineous discharge, the
blood coming mainly from the uterine caruncles – metoestrual bleeding. The body
temperature of dairy cows falls about 0.5°C the day before oestrus, increases during oestrus
and falls by about 0.3°C at ovulation. The vaginal temperature, of 37.74°C, was lowest on the
day before oestrus, increased by 0.1°C on the day of oestrus, and increased for the next 6 days
until a plateau was reached. This was followed by a gradual decline from 7 days before
oestrus.

SYMPTOMS OF OESTRUS IN COW

 Standing to be mounted by other cows

 Attempt to mount other cows
 Stringy mucous hanging from vulva
 Mucus smeared on buttocks
 Increased restlessness
 Drop in milk yield
 Reduced feed intake
 Bellowing
 Chin resting on cow's rump by other cows, tail raising
 Frequent urination
 Vulval edema

Buffalo

 Overt signs of estrus are not pronounced as in cattle

 Heterosexual behaviour, particularly standing to be mounted by a bull is the most
reliable sign
 Homosexual behaviour, such as standing to be mounted by other females, is observed
only occasionally
 Signs such as vulval swelling, clear mucoid vulval discharge, spontaneous milk
letdown, bellowing, restlessness, frequent urination and raised tail vary in occurrence
and intensity from animal to animal, and in relation to standing oestrus.

OESTROUS CYCLE IN THE MARE

Mare
The mare is normally a seasonal breeder, with cyclic activity occurring from spring to
autumn; and showing anestrous during the winter. However if provided sufficient feed and
photoperiod the seasonality can be influenced. Seasonality is mediated by the release of
melatonis from the pineal glands in response to photoperiodism. The winter anoestrus is
followed by a period of transition to regular cyclic activity. During this transition, the duration
of oestrus may be irregular or very long, sometimes more than a month. The manifestations of
heat during the transitional phase are often atypical and make it difficult for the observer to be
certain of the mare‟s reproductive status. It is common for the early heats to be lacking in
follicular growth and being anovulatory. However, once ovulation has occurred, regular
cycles usually follow.

The average length of the equine cycle is 20–23 days; the cycles are longer in spring
and shortest from June to September. Typically, oestrus lasts 6 days and dioestrus 15 days.
Ovulation occurs on the penultimate or last day of heat, and this relationship to the end of heat
is fairly constant and irrespective of the duration of the cycle or the length of oestrus. Manual
rupture of the ripe follicle results in termination of oestrus within 24 hours. The diameter of
the ripe follicle is 3–7 cm. During the last day before ovulation, the tension in the follicle
usually subsides, and the palpable presence of a large fluctuating follicle is a sure sign of
imminent ovulation. The onset of heat after foaling occurs on the fifth to 10th day, called as
foal heat. This foal heat is sometimes rather short, 2–4 days. It is traditional to cover a mare
on the ninth day after foaling.

During oestrus, a single egg is usually released, around 52.2 % of the ovulations occur
from the left ovary. Twin ovulation commonly occurs in mares. In mares only fertilised eggs
pass into the uterus; non-fertilised eggs remain for months in the uterine tubes, where they
slowly disintegrate. All equine ovulations occur from the ovulation fossa; only at the ovarian
hilus may occasional protrusions of corpora lutea be seen, but because of the curvature of the
ovary and the presence of the adjacent substantial fimbriae these protusions cannot be
identified by rectal palpation. For ovulation to occur the enzymes released from the cells of
the germinal epithelia are needed, and in the case of ovary of the mare, the germinal epithelia
is restricted to the ovulation fossa, hence ovulation can occur only from the ovulation fossa.
During oestrus, this follicle matures and ruptures when it has attained a diameter of 3–7 cm.
Several hours before ovulation the ripe follicle becomes much less tense. The collapsed
follicle is recognised by an indentation on the ovarian surface; there is usually some
haemorrhage into the follicle, and the coagulum hardens within the next 24 hours. Quite
frequently the mare shows evidence of discomfort when the ovary is palpated soon after
ovulation. Unless sequential transrectal palpation or ultrasonic examinations are performed, it
is sometimes possible to confuse a mature follicle with the early corpus haemorrhagicum,
since before ovulation the follicular antrum is filled with follicular fluid and then soon after
ovulation it becomes filled with blood. For this reason mares are sometimes incorrectly
diagnosed as having failed to ovulate. For the next 3 days the luteinising mass can be felt as a
resilient focus, but later it tends to have the same texture as the remainder of the ovary. The
cyclical CL begins to regress at about the 12th day of the cycle, when there is a parallel fall in
the blood progesterone concentration.

Ovulation, with the subsequent formation of a CL, does not always occur; the follicle
may regress or sometimes undergo luteinisation. During winter anoestrus, both ovaries are
typically small and bean-shaped, common dimensions being 6 cm from pole to pole, 4 cm
from the hilus to the free border and 3 cm from side to side.

By visual examination of the vagina and the cervix using an illuminated speculum, it
is possible to detect the preovulation period. In dioestrus, the cervix is small, constricted and
firm; it and the vagina are pale pink, while mucus is scanty and sticky. During oestrus, there is
a gradual increase in the vascularity of the genital tract and relaxation of the cervix with
dilatation of the os. As oestrus advances and ovulation time approaches, the cervix becomes
very relaxed and its protrusion can be seen lying on the vaginal floor, with its folds
oedematous; the vaginal walls are glistening with clear lubricant mucus. After ovulation there
is a gradual reversion to the dioestrous appearance. During anoestrus, as in pregnancy, both
the vagina and cervix are blanched; the cervix is constricted and generally turned away from
the midline, the external os being filled with tenacious mucus.
On palpating the uterus per rectum, cyclic changes can be detected. With the
development of the CL the uterus increases in tone and thickness, but these features diminish
when the CL regresses. At oestrus there is no increase of tone. During anoestrus and for the
first few days after ovulation the uterus is flaccid. During dioestrus, pregnancy and
pseudopregnancy the cervix is identified on rectal palpation as a narrow firm tubular
structure; at oestrus it is soft and broad.

Signs of oestrus
The mare becomes restless and irritable; she frequently adopts the micturition posture
and voids urine with repeated exposure of the clitoris – winking of the clitoris. When
introduced to a stallion or teaser, these postures are accentuated; the mare raises the tail to one
side and leans her hindquarters. The vulva is slightly oedematous, and there is a variable
amount of mucoid discharge. A mare which is not in oestrus will usually violently oppose the
advances of a stallion, and for this reason when „trying‟ mares at stud it should be done over a
gate, box-door or stout fence. If the mare is in oestrus the stallion usually exhibits „flehmen‟.

1. The normal oestrous cycle in mare is 21-22 days.

 2. Oestrus, the period when the mare displays behavioural signs of sexual receptivity to
the stallion, lasts for 5 to 7 days
3. During diestrus, the luteal phase that lasts for 14-15 days, the mare displays sexual
rejection to the stallion.
4. Duration of oestrus varies among individuals and also among oestrous cycles of the
same mare.
5. The intensity of behavioural oestrus varies both throughout the oestrous period and
among individual mares at comparable stages of the period.
6. Old mares, mares underfed during the early part of the breeding season and during
twin ovulations have a longer duration of oestrus.

SYMPTOMS OF OESTRUS IN MARE

 Restless and irritable.

 Frequently adopts the micturition posture and voids urine with repeated exposure of
the clitoris by prolonged rhythmic contractions. This is known as winking of clitoris.

 Introduction of a stallion or teaser, these postures are accentuated.

 Raises the tail to one side and leans her hind quarters.
 Vulva becomes large and swollen.
 Labial folds are loose and readily open for examination.
 Vulva becomes scarlet or orange, wet, glossy and covered with a film of transparent
mucus.
 Variable amount of mucoid discharge.
 If the mare is in oestrus, the stallion will usually exhibit “Flehmen”.
 Vaginal mucosa is highly vascular, and thin watery mucus may accumulate in the
vagina.
 During oestrus cervix dilates enough to admit 2-4 fingers, during diestrus only one
finger can be inserted.

OESTROUS CYCLE IN SOW

Puberty in gilts is reached at about 7 months, but diet, breed (including the degree of
in-breeding) and season of birth influence its onset. At the first oestrus the number of
ovulations is low, but it increases thereafter so that if mating is delayed until the third heat a
larger litter will result. The provision of a high-energy diet for 11–14 days before the expected
oestrus increases the ovulation rate: Flushing. Continuing such a diet after mating, however,
increases embryonic loss. Fecundity is best from the fourth to seventh gestations. Although
the domestic sow is generally considered to be polyoestrous, the wild pig is a seasonal
breeder.
The length of oestrous cycle is 21 days and the average length of oestrus is 53 hours,
with spontaneous ovulation occurring between 38 and 42 hours from the beginning. During
lactation, the physical stimuli of suckling suppresses cyclical activity, but many sows show an
anovulatory oestrus 2 days after farrowing. When weaning occurs at 5 or 6 weeks, oestrus can
be expected in 4–6 days.
Signs of oestrus
Beginning 3 days before oestrus, the vulva becomes progressively swollen and congested;
these features persist throughout oestrus and gradually subside during the 3 days afterwards.
Restlessness is an unfailing sign of the approach of heat, and a peculiar repetitive grunt is
emitted. With other sows the pro-oestrous animal sniffs their vulvae and may try to ride them,
or will be the recipient of such attentions. At the height of oestrus the sow assumes a
stationary, rigid attitude with her ears cocked, and she appears to be quite oblivious to her
environment. oestrus can readily be determined by firmly pressing the loin of the sow with the
palms of both hands; the oestrous sow will stand motionless with cocked ears whereas sows
not in heat will object to this approach: lordosis reflex. The same immobilisation response can
be elicited if the attendant sits astride the sow, and it can also be obtained in the absence of a
boar by reproducing the voice or odour of the boar. The substance responsible for boar odour
has been identified as 5α-andost-16-ene-3-one and it is secreted by the salivary glands. In the
form of an aerosol it can be sprayed in the vicinity of sows to promote the standing reaction of
oestrus. „Silent heats‟ occur in about 2% of porcine cycles.

 Sow is polyestrus.
 The average length of the cycle is 21 days and may vary from 18-23 days.
 The estrogenic phase of the cycle is 6-7 days.
 The luteal phase is 14 days with corpus luteum regression beginning on day 16.
 The duration of estrum is 1-4 days, with an average of 2-3 days.
 Silent heats occur in about 2% of porcine cycles.
 Senility (after 6-10 years) may affect estrous cycle.

SYMPTOMS OF OESTRUS IN PIG

Oestrus symptoms are definite and marked and include

o Reduced appetite
o Restless and nervous
o Often pace back and forth by the fence
o Salivation
o Champing of the jaws
o Sow, if suckling, may ignore her piglets, may try to escape or if free will seek out the
boar and stand for service
o Vulval lips swollen and congested
o Mucous membrane pinkish-red
o Mucous discharge
o Mounting other animals
o Grunting and standing motionless for long periods in a position for service is the most
noticeable feature. This peculiar immobilizing reflex or stance (lordosis response) can
be produced by applying pressure of the hands on the sow‟s back or by straddling the
sow.

OESTROUS CYCLE IN SHEEP AND GOAT

The normal oestrous cycle length in sheep is 17 days and in goats is 21 days.
Photoperiodism is a characteristic of the reproductive cycle or sexual receptivity in sheep and
goats. A reduction in daylight to about 10 or 11 h, whether it occurs naturally or whether it
is induced artificially by penning the ewes in a darkened area during part of the day, will
induce the onset of oestrous cycle in anoestrous ewes. During early breeding season,
premature regression of corpus luteum or anovulation results in abnormally short cycles.
Oestrus lasts for 24-36 h in ewe and 24-48 h in the doe.

SYMPTOMS OF OESTRUS IN SHEEP AND GOAT

Doe

Near the equator, there is no evidence of seasonality but continuous cyclic activity.
The doe is polyoestrous, with an interoestrus interval of 20–21 days, although it is rather
irregular at the beginning of the breeding season. The duration of oestrus is 30–40 hours, with
ovulation occurring 12–36 hours after the onset.

When in oestrum the vulva shows some evidence of oedema and hyperaemia; the tail is
wagged from side to side and up and down and is the most reliable sign. The doe is restless
and more vocal, has a reduced appetite and milk yield, and shows mounting behaviour. The
presence of the pheromones from the male goat, which can be transferred from the scent
gland on to a cloth, will often intensify the signs

 Oestrus symptoms are more conspicuous in does and include

o Restless
o Frequent bleating
o Tail wagging from side to side and up and down (most reliable sign)
o Reduced appetite
o Decreased milk production
o Vulva oedematous and congested
o Clear mucous discharge from the vagina
o Occasionally does exhibit homosexual behaviour

Ewe

The stimulus for the annual onset of sexual activity is declining length of daylight.
The seasonal onset of sexual activity can be advanced by artifical manipulation of the
photoperiod and by the use of hormonal agents. The average duration of oestrus in mature
ewes is about 30 hours. Ovulation occurs towards the end of oestrus, and the length of the
oestrous cycle averages 17 days.
Oestrus ewes are restless. They seek the ram, and together form a following „harem‟.
When in full heat, ewe stands to be mounted by the ram, waggles her tail and moves it
laterally. The vulva is slightly swollen and congested, and there is often a slight discharge of
clear mucus.

 Oestrous symptoms are relatively inconspicuous, and are not evident in the absence of
ram.
o Ewes seek the ram, and together form a following “harem”
o Vulva oedematous and congested
o Clear mucous discharge from the vagina
 o Waggles her tail and moves it laterally

OESTROUS CYCLE IN BITCHES

Dogs experience ovarian cycles twice yearly. Breeding seasons depend on both
genetic and management factors. Breeds enter their first heat between 6 and 10 months of
age. Some dogs may not begin to cycle until 18 to 24 months of age. The inter-estrus
interval on an average is 7 months and ranges from 3.5 months to 13 months. However, the
Basenji breed cycles once a year only.

Phases of Oestrous Cycle

 There are four phases namely

o Proestrus
o Oestrus
o Diestrus
o Anestrus

Proestrus

Begins with the appearance of vaginal bleeding (spotting) and ends when the bitch allows
a male dog to mount and breed. Duration is 9 days (average and ranges from 2-3 to 25 days).

Hormonal changes

Proestrus is under the influence of oestrogen, which is synthesized by the developing

follicles. Early proestrus is associated with oestradiol concentration of 25 pg/ ml which may
increase to 60 –70 pg/ml in late proestrus. Thereafter oestrogen levels start declining to reach
basal levels (5 – 15 pg/ml) at the onset of diestrus. Progesterone concentration start increasing
from basal levels (< 1ng/ml) at late proestrus to reach levels of 3 ng/ml at the onset of oestrus
and 5 ng/ml at the start of ovulation. This progesterone is secreted by follicle which
becomes partially luteinised prior to ovulation and developing CL.

Oestrus

Begins with the first acceptance of the male and ends with the refusal. The duration is 9
days but ranges from 1-2 days to 18-20 days.

Hormonal changes

The bitch is unique in that standing oestrus is exhibited when oestrogen levels have
started to decline and progesterone levels are increasing. This triggers two events namely

1. Exhibition of maximal oestrus behaviour

2. Initiation of LH surge in the bitch

Ovulation

Ovulation occurs 24 to 48 h after the LH surge and occurs over a span of 72 to 96 h.

The eggs are released as primary oocytes and require 24 to 72 h to mature and become
fertile. Once mature, their lifespan is 2 to 4 days. At the time of ovulation the serum
progesterone concentration is typically in the range of 4 to 10 ng/ml.

Diestrus

Begins with the cessation of standing heat and ends when blood progesterone levels fall to
basal levels. Progesterone levels continue to rise to reach levels of 50 to 60 ng/ml 20 to 30
days after ovulation. The bitch is unique in that whether the bitch is pregnant or not, the CL
continues to remain functional for a particular period of time. There is no pregnancy
recognition system in dogs. Therefore CL functions throughout normal gestational period
regardless of the presence or absence of foetus.

In a pregnant bitch the luteal phase ends with the onset of parturition (approximately 65
days) while in a non pregnant bitch the CL functions for a longer period. ie., 75 to 100 days
and regression is brought about by aging of CL. Thus the uterus is not involved in the
regulation of the lifespan of the CL. Apart form LH; prolactin is the major luteotrophic factor
especially in the second half of the luteal phase.

Anoestrus

Anoestrus is the phase in which uterus involutes. Average duration is 4.5 months but may
be variable.

SYMPTOMS OF OESTRUS IN BITCH

Proestrus

During proestrus the bitch is playful, appears sexually attractive to the male but refuses
mounting attempts by the male by moving away, bearing of the teeth and snapping. The bitch
may keep her tail tight against the perineum and cover her vulva. This behaviour changes as
proestrus progresses. The female becomes more receptive, seeks males, playing and teasing
but still refuses the male by crouching or lying down. In late proestrus, the bitch may sit or
stand passively when mounted. The bloody discharge fades and becomes transparent to straw
coloured and the vulva which is swollen and hard during proestrus, now becomes small and
soft as estrus approaches.

Oestrus

The bitch may crouch and elevate the perineum toward the male. Te bitch attracts males
over long distances due to the presence of potent pheromones. The vulva is soft and flaccid
and the vaginal discharge is often straw coloured or pink.

Length of Various Periods of Oestrous Cycle In Domestic Species

 FACTORS AFFECTING ESTROUS CYCLE

Nutrition

 Inanition or starvation caused by lack of energy or nutritional deficiency impairs or

prevents secretions of gonadotropic hormones from pituitary gland thereby preventing
the occurance of cycles.

Seasonal Influences and Light

 Seasonal breeding is regulated by light through the medium of eyes or the

hypothalamus and pituitary gland. The total daily amount of light is important in
control of the onset of estrous cycle. This is called as sexual photoperiodicity.
 In sheep and goat decreased day light influences onset of estrus.
 In mares increased total daily hours of light will hasten the onset of estrous cycle.

Temperature

 Excessive heat during summer months causes decreased thyroid activity which
directly reduces reproductive efficiency.
 In cattle under hot conditions length of the estrous cycle is increased to 25 days as
compared to 20-22 days in cool weather.

Age

 In cattle and swine the young female usually has a slightly shorter length of estrous
cycle than the adult animals.
Systemic Diseases

 Severe chronic wasting diseases cause debility and emaciation resulting in cessation of
estrous cycle.

Pathology of Uterus and Cervix

 In cattle, conditions like pyometra, foetal maceration, mummification causes

persistence of corpus luteum and cessation of estrous cycle.

Endocrine Disturbances

 Endocrine disturbances leading to cystic ovaries or pituitary, ovarian and adrenal

tumors affect the length of the estrous cycle.

Miscellaneous Causes

 Pregnancy causes physiological cessation of estrous cycle.

 The presence of males hastens onset of estrus.
 Infusion of iodine compounds into the uterus shortens the estrous cycle.

Ovulation and fertilization

Transportation of gametes and implantation