Oestrous Cycle - Anatomy & Physiology

Introduction

Ovarian cycles are from oestrus to oestrus. Oestrus is the easily identifiable external marker. It should be noted that Oestrus is the noun and oestrous is the adjective.

Classification of Oestrous Cycles

Based on Vaginal Cytology

Pro-Oestrus

• The phase immediately preceding oestrus.
• Lasts 2-5 days, depending on the species.
• Marked increase in reproductive activity.
• Endocrine transition from progesterone to oestrogen dominance under the influence of gonadotrophins LH and FSH.
• Follicular growth and regression of the corpus luteum of the previous cycle (in polyoestrous species).
• Uterus enlarges slightly
• Endometrium becomes congested and oedematous, glands show secretory activity.
• Vaginal mucous becomes hyperaemic.
  • Increase in cell numbers in the vaginal epithelium.
  • Superficial layers become cornified.
• The bitch shows external evidence:
  • Vulval oedema
  • Hyperaemia
  • Sanguinous vulval discharge

Oestrus

• The period where the female will accept the male for copulation.
• The onset and end of oestrus are the only stages of the oetrous cycle that can be accurately measured. They are therefore used to determine cycle length.
• Female seeks out the male and 'stands' to be mounted.
• Uterine, cervical and vaginal glands secrete increased amounts of mucus.
• Vaginal epithelium and uterine endometrium becomes hyperaemic and congested.
• Cervix is relaxed
• Ovulation occurs during this phase in all domestic species except the cow.
  • The cow ovulates ~12 hours after the end of oestrus.
• Ovulation is spontaneous in all domestic species except the cat, rabbit and camelids. In these species it is induced by the act of copulation.
• During pro-oestrous and oestrus there is follicular growth with no functional corpora lutea present on the ovary (aka follicular phase).
• The main ovarian hormone production is Oestrogen.

Metoestrus

• The phase after oestrus
• During early metoestrus, both oestrogen and progesterone levels are relatively low.
• Granulosa cells of the ovulated follicles give rise to lutein cells, which are responsible for the formation of the corpus luteum (luteinisation).
• Progesterone secretion is detectable soon after ovulation, with significant quantities produced in 2-5 days. This marks a transition from oestrogen to progesterone dominance.
• Reduction in secretion from uterine, cervical and vaginal glands.

Dioestrus

• Longest phase of the oestrous cycle, which usually laste 10-14 days in polyoestrous females.
• The corpus luteum is present on the ovary and secreting large amounts of progesterone. The duration of this phase is directly related to the time that the corpus luteum remains functional.
• High progesterone levels prompt the uterus to prepare a suitable environment for development of the embryo, and eventual attachment of the conceptus to the endometrium (implantation).
• Uterine glands undergo hyperplasia and hypertrophy.
• Cervix becomes constricted
• Secretions of the tract are scant and sticky.
• Vaginal mucosa becomes pale.
• Ends with the regression of the corpus luteum (luteolysis).

Anoestrus

• Genital system is quiescent
• Follicular development is minimal and corpora lutea have regressed and are non-functional.
• Secretions are scanty
• Cervix is constricted
• Vaginal Mucosa is pale

Based on Secretory Patterns of the Ovary

Follicular Phase

• Begins after luteolysis, which causes the decline in progesterone.
• Gonadotrophins LH and FSH are produced, which cause ovarian follicles to produce Oestrogen.
• Ends at ovulation.
• Oestrus is designated as day 0.

Luteal Phase

• Begins after ovulation
• Includes the development of corpora luta, which produce progesterone.
• Includes luteolysis, which is brought about by Prostaglandin F2α.

Based on Vaginal Cytology

Proliferative Phase

• Oestrogen dominant
• Includes proliferation of the endometrium.

Secretory Phase

• Progesterone dominant
• Includes secretions from the uterus, such as 'uterine milk' to sustain the fertilised ovum until implantaion.

Comparative Oestrous Cycles

Species	Mare	Cow	Sow	Ewe	Bitch	Queen
Length of Oestrous Cycle (days)	21	21	21	17	93	Variable
Duration of Oestrus	4-8 days	12-36 hours	48-96 hours	24-48 hours	7-13 days	4-10 days
Duration of Follicular Phase (days)	7 (variable)	4	7	3	18	Variable
Duration of Luteal Phase (days)	14	17	14	14	75	40

Mare

Oestrous Cycle

• Seasonal breeder (long day)
• Oestrous cyclicity from spring-autumn. During the winter, the mare is normally anoestrus.
• Winter anoestrus is follwed by a transition to regular cyclic activity. During this time, the duration of oestrus may be irregular or very long.
• Before the first ovulation, behavioural oestrus may not be accompanied by the presence of palpable follicles on the ovary.
• Some long oestrus periods in spring are anovulatory.

• Average cycle length is 20-23 days. The cycles are longest in spring and shortest from June-September.
• Oestrus lasts 6 days.
• Dioestrus lasts 15 days.
• Ovulation occurs on the penultimate day or last day of oestrus.
• The diameter of a ripe follicle ready to ovulate is 3-7cm. The day before ovulation, the tension in the follicle is reduced. Palpation of a large, fluctuating follicle is a reliable sign of impending ovulation.

• The onset of oestrus after foaling is known as the 'foal heat'.
• Occurs on day 5-10 post-parturition.
• Sometimes shorter than normal, lasting 2-4 days.
• The first two post-parturient cycles are a few days longer than normal.

• During Oestrus, a single egg is usually released. Thus, the mare is monotocious.
• Ovulation seems to occur more frequently from the left ovary.
• All ovulations occur from the ovulation fossa.
• Due to the reversed structure of the ovary, corpora lutea may only be seen sometimes at the ovarian hilus. However, because the ovary is curved and mostly covered by fimbrae of the oviduct, corpora lutea cannot be identified by rectal palpation.
• Only fertilized eggs can pass into the uterus.
• Non-fertilized eggs remain in the Utero-Tubal junction for months, where they finally disintegrate.

Cyclic Changes in the Ovaries

• Just before the onset of oestrus, several follicles enlarge to 1-3cm.
• By the first day of oestrus, one dominant follicle is significantly larger than the others, with a diameter of 2.5-3.5 cm.
• During oestrus, this dominant follicle matures and ruptures once it has attained a diameter of 3-7 cm.
• Several hours before ovulation, the ripe follicle becomes much less tense and can be recognised as an indentation on the surface of the ovary.
• There is usually haemorrhage into the follicle and the coagulum hardens within the next 24 hours.
• After ovulation, the other follicles regress.
• During the first 4-9 days of dioestrus, there are no follicles over 1cm present on the ovary.
• For 3 days after ovulation, the leutinising mass can be felt, but later it normally has the same texture as the ovary.
• The corpus luteum reaches full size at 4-5 days after ovulation but does not protrude from the ovarian surface.
• On section of the ovary, the corpus luteum will appear brown and later yellow. It is triangular or conical in shape, with the narrow end on the ovulaiton fossa. The centre of the corpus luteum normally contains dark brown fibrin.
• The corpus luteum of the cycle (non-pregnant) starts to regress on about the 12th day of the cycle. At the time of regression, there is a fall in blood progesterone concentration.

• During winter anoestrus, both ovaries are usually small and bean-shaped. They normally measure:
  • 6cm pole-pole
  • 4cm from the hilus to the free border
  • 3 cm side-side
• During the cycle, the size of the ovary depends on the number and size of the follicles.
• During oestrus, the ovary may contain 2-3 follicles. These can each measure 4-7 cm. These, combined with other less developed follicles, give the ovary a huge size.
• During Dioestrus, there is an active corpus luteum and atretic follicles present. However, these only give the ovary a size a little larger than in Anoestrus.

Endocrine Changes During the Oestrous Cycle

• Biphasic secretion of FSH with surges every 10-12 days.
  • One surge after ovulation and a second in mid-late Dioestrus (~10 days before the next ovulation).
• This increase in FSH secretion is unique to the mare.
  • Primes the development of a new generation of follicles, one of which will ovulate during the next oestrus.
• The pattern of LH secretion is also unusual.
  • No dramatic LH surge prior to ovulation.
  • LH gradually increases and elevated levels then persist for 5-6 days either side of ovulation.
• Oestrogens reach peak values during oestrus.
• Progesterone concentrations follow the changes of the corpus luteum closely.

Uterine Cytology

This is often performed routinely before breeding commences on stud farms. Endometrial and inflammatory cells are examined. Evidence of bacteria in the absence of inflammatory cells is normally due to contamination.

• Oestrus/Active Cycling
  • A high number of columnar epithelial cells with oval, basal nuclei containing finely stippled chromatin.
  • Columnar, finely vacuolated cytoplasm.
  • There should be no inflammatory cells or infectious agents visible. Rarely neutrophils may be present if it is a foal heat or the first oestrus cycle in maiden mares.

• Anoestrus/Ovarian Inactivity
  • Clumps of epithelial cells with crowded or overlapping, oval, hypochromatic ‘bland’ nuclei without granular chromatin or nucleoli.
  • Unsubstantial cytoplasm with indistinct boundaries.

Cow

Oestrous Cycle

• Polyoestrous
• The first ovulation in heifers is usually without behavioural oestrus. This is termed 'silent heat'.
• Cyclical activity persists except during pregnancy, for 3-6 weeks post-calving and during high milk yield.
• In heifers, the average length of the oestrous cycle is 20 days (18-22).
• In cows, the average length of the oestrous cycle is 21 days (18-24).
• The average duration of oestrus is ~ 15 hours (2-30 hours).
• The body temperature of dairy cows falls by ~0.5◦C the day before oestrus. It then increases in oestrus before falling by ~0.3◦C at ovulation.
• Vaginal pH also fluctuates throughout the oestrous cycle, but is lowest (7.32) on the day of oestrus.
• Ovulation is spontaneous, occuring ~12 hours after the end of oestrus.
• Within 2 days of service, there is yellow-white vulval discharge of mucus. This contains neutrophil leucocytes from the uterus.
• At ~48 hours post-oestrus, irrespective of service, there is a bright red sanguineous discharge from the uterine caruncles.

Cyclic Changes in the Vagina

The main variations are in the epithelial cells of the anterior vagina and in the secretory function of the cervical glands.

• Oestrus: Anterior vaginal epithelium becomes greatly thickened due to cell division and the growth of the tall, columnar, mucus-secreting superficial cells.
• Dioestrus: Cells of the cranial vaginal epithelium vary from flat to low columnar. Invasion of the vaginal mucosa by leucocytes is maximal 2-5 days after oestrus.
• Copious secretion of mucus by the cervix and anterior vagina begins 1-2 days before oestrus.
• Secretions diminish to the 4th day after oestrus.
• Mucus is transparent and flows readily.
• Variations in cervical mucus:
  • During oestrus and for a few days after, crystals are disposed in a distinct aborisation pattern.
  • For the remainder of the cycle this pattern is absent.
  • This pattern, and the amount of cervical mucus depends on oestrogen concentration.
• Postoestrous vaginal mucus:
  • Floccules of leucocytes
  • Blood is frequently present
• Hyperaemia of vaginal and cervical mucosae is progressive during pro-oestrus and oestrus.
• At this time, the protrusion of the cervix into the cranial vagina is relaxed, so that 1-2 fingers can be inserted into the cervical os.
• During metoestrus, there is a rapid reduction in vascularity.
• From day 3-5 postoestrus:
  • Mucosa of the cervix is pale and quiescent.
  • The external os is constricted.
  • Mucus becomes scanty, sticky and pale yellow/brown.
• There are also cyclical changes in the vaginal pH:
  • pH falls from 7.0 to 6.72 one day before first signs of behavioural oestrus.
  • At the start of oestrus, pH falls again to 6.54.

Cyclic Changes in the Uterus

• Oestrus:
  • Uterus is congested
  • Endometrium contains oedematous fluid, which causes its surface to glisten.
  • Muscularis is contractile, so when palpated per rectum, the uterus feels turgid and the horns feel erect and coiled.
    • Tonicity is present the day before and the day after oestrus, but is at its maximum during heat.
  • Marked increase in vascularity
• 24-48 hours postoestrus:
  • Uterine caruncles show petechial haemorrhage. This gives rise to the discharge of blood from the vagina.
    • In heifers, there is also often perimetral subserous petechiae.
• Dioestrus:
  • Endometrium is covered with scanty secretion from the uterine glands.

Cyclic Changes in the Ovaries

• Usually one follicle ovulates, but twin ovulations occur in 4-5% of cows and triplet ovulations more rarely.
• In dairy cattle 60% of ovulations are from the right ovary, but in beef cattle, they are about the same.
• Size of the ovaries is dependent on the phase of the oestrous cycle.

Follicular Growth and Development

• There is follicular growth and atresia throughout the oestrous cycle.
• There are normally two waves of follicular growth, but 3 waves also commonly occur:
  • One begins on day 3-4
  • The second begins on day 12-14
• A normal follicle of 9-13mm is present between day 5-11 before becoming atretic.
• In the second wave, an ovulatory follicle develops, measuring 9-13mm between day 15-20.
• Ovulatory follicle is selected ~3 days prior to ovulation.
• Follicular growth is under the influence of FSH.
• One follicle normally obtains dominance and subsequently ovulates.
• Dominance does not involve inhibin. It is probably due to an intra-ovarian mechanism which does not suppress FSH secretion. Other hormones such as IGF-1 (Insulin-like growth factor-1) may also be involved in follicular growth patterns.
• During dioestrus, there are several large follicles on the ovary. These range in size 0.7-1.5cm in diameter. These follicles do not alter the contours of the ovaries, but cause variation in size.
• During Pro-oestrus and oestrus, the dominant follicle enlarged and ovulation occurs once it is at least 1.9cm.
• The ripening follicle is detectable by rectal palpation during heat.
  • It can be felt as a soft, bulging area on the surface of one ovary.
• Ovulation can occur from anywhere on the ovarian surface.
• The shape of the ovary when the corpus luteum develops will depend on the site of ovulation.
• The point of ovulation is usually an avascular area of the follicular wall. Therefore, haemorrage of the follicle is not seen with ovulation in the cow. However, after ovulation there is congestion around the point of follicular rupture and sometimes there is a small blood clot in the centre of the new corpus luteum.
• When the follicle ruptures, the ovum is expelled through a small breach in the surface.
• Most of the antral fluid escapes and the follicle collapses.
• The ovary feels flattened and soft on palpation.
• The surface from which ovulation has occurred is wrinkled and may be blood-stained.

Corpus Luteum of the Oestrous Cycle

• The corpus luteum develops by hypertrophy and luteinization of the follicular granulosa cells, this occurs rapidly.
• By 48 hours after ovulation, the corpus luteum is ~1.4cm in diameter. It feels soft and yields on palpation.
• It is dull cream in colour and the luteinized cells can be seen in loose pleats.
• It attains its maximum size by day 7-8 of dioestrus.
• Luteinized pleats are now compact.
• The body is a homogenous mass that is yellow/orange in colour.
• The shape varies, most are oval but they can be square or rectangular.
• Sometimes the centre is occupied by a cavity which is occupied by yellow fluid. This is known as a cyctic corpus luteum, but is rarely pathogenic and usually completely normal.
• If a cavity is present, it must be distinguished from a luteinisation of the walls of the follicle without ovulation (abnormal).
  • If it is a corpus luteum with a cavity, there will be a pin-head depression in the centre of the projection from the surface of the ovary. This depression indicates ovulation has occured.

Projection of the Corpus Luteum from the Surface of the Ovary

• As the corpus luteum enlarges, it pushes itself out of the ovary and so stretches the ovarian surface as it does so.
• By the time it reaches maximum development it often forms a disctinct projection from the surface of the ovary.
• Usually, this is a distinct bulge about 1cm in diameter with a clear-cut constriction where it joins the general contour of the ovary.
• Sometimes, it is nipple like.
• Another type of projection is indistinct and diffuse, occupying the greater part of the ovary.
• The type of protrusion depends on the extent of the ovarian surface that was occupied by the follicle just before ovulation.

Regressing Corpus Luteum

• The corpus luteum remains its maximum size until the onset of pro-oestrus (~24 hours before heat).
• When the cow comes into heat, the corpus luteum undergoes rapid reduction in size and changes colour and appearance.
• By the middle of oestrus, its diameter is reduced to 1.5cm and its protrusion from the surface of the ovary is less distinct. Also, its colour changes to bright yellow. (It is after this discovery in the cow that the corpus luteum was given its name. Corpus luteum is latin for 'yellow body').
• Its consistency is dense and it becomes invaded with scar tissue.
• By the second day of dioestrus, it is reduced to ~1cm and its outline is becoming irregular.
• The colour then changes from yellow to brown.
• By the middle of dioestrus, it has shrunk to 0.5cm and its protrusion from the ovarian surface is a bit larger than a pin-head.
• As it ages, its colour goes from red to scarlet.
• Small remnants can persist for several months.
• For more general information on regression of the corpus luteum, click here

Size of Ovaries

• Depends on the phase of the oestrous cycle and whether it contains an active corpus luteum.
• The presence of follicles does not alter the size of the ovary to anything like the extent of the corpus luteum.
• In the majority of heifers and young cows examined between day 6-18 of dioestrus, one ovary will be larger than the other.
• The larger ovary will be approximately:
  • 3.5cm pole-pole
  • 3cm from the attached side to the free border
  • 2.8cm from side to side
  • The corpus luteum will project from some point on the surface.
• The smaller ovary will be approximately:
  • 2.5cm pole-pole
  • 1.5cm from the attached side to the free border
  • 1.2 cm from side to side
• During the 4-5 days after oestrus, the developing corpus luteum has not attained sufficient bulk to influence the size of the ovary significantly and the regressing corpus luteum has lost significant bulk.
• During oestrus there will also be little difference in size.
  • If the ovary undergoing enlargement of a dominant follicle also contains a regressing corpus luteum, it will be a little larger than the other ovary but not strikingly so.

Ovaries in a Multiparous Cow

• Larger
• Partly due to progressive deposition of scar tissue from prolonged function.
• Can be due to large numbers of small, but visible follicles.
• Often, the ovary that does not contain a corpus luteum measures:
  • 4cm pole-pole
  • 3cm from the fixed edge to the free border
  • 2cm from side to side
• In mid-dioestrus it is still possible to detect the corpus luteum because, aside from its protrusion, the ovary appears plump. The ovary that does not contain a corpus luteum is flattened from side to side.
• Corpora lutea (active and regressing) and follicles approaching maturity are the same as those in the heifer.
• In addition, however, there are old scarred corpora lutea of previous pregnancies.
  • Generally white (corpus albicans - latin for white body), pin-head sized projections on the surface of the ovary.
  • Contain mainly scar tissue
  • Irregular outline
  • Maximum of 0.5 cm
• The corpus luteum of pregnancy does not atrophy as quickly after parturition as the corpus luteum of the cycle. It becomes invaded by scar tissue and remains throughout the cow's life.
• The fully developed corpus luteum is present by day 7 and persists unchanged until the onset of pro-oestrus at day 19-20.

Appearance of the Ovaries on Ultrasound

• Normal structures that can be identified:
  • Ovarian stroma: Mottled echotexture
  • Antral follicles: Anechoic (black) structures of variable size with a clear definition between the follicular wall and the antral cavity. They will not always be a regular, spherical shape.
  • Corpora lutea: Well defined border and a mottled echogenic appearance. They are less echogenic than the ovarian stroma. The fluid-filled lacuna can be identified as a dark, non-echogenic area in the centre.
  • Ovarian blood vessels: Black,non echogenic structures which are easily confused with antral follicles. Movevement of the transducer can be used to differentiate the two, as this shows their elongated structure.
• In addition, pathological structures like ovarian cysts can be seen.

Endocrine Changes during the Oestrous Cycle

• Hormones are secreted in a pulsatile manner and fluctuate considerably.
• Just before onset of behavioural oestrus, there is a sharp rise in plasma oestrogens, particularly oestradiol.
  • Peak values occur at the beginning of oestrus
  • Levels decline to basal concentrations at ovulation
• During the rest of the cycle, there are fluctuations but there is a discrete peak around day 6 of the cycle.
  • Related to the first wave of follicular growth
• Pre-oestrus rise in oestrogens stimulates the LH surge from the anterior pituitary.
  • This is necessary for follicular maturation, ovulation and maturation of the corpus luteum.
• A second, less discrete peak occurs 24 hours after the LH surge.
• Changes in progesterone concentrations mimic changes in the corpus luteum.
• Progesterone levels peak 7-8 days after ovulation and decline quickly from day 18.
• When progesterone levels fall to low basal levels, the negative feedback on the pituitary gland is removed. This allows a sudden release of gonadotrophins.

Ewe

Oestrous Cycle

• Short day breeders (oestrous cycles commence in response to decrease in day length).
• In Britain, sheep cycle from October-February, during which time there are 8-10 recurrent oestrous cycles.
• Ewe lambs and yearling ewes have shorter breeding seasons than older ewes.
• The length of the oestrous cycle is about 17 days.
• Duration of oestrus in most British breeds is ~30 hours and is at least 10 hours in immature ewes.
• In merino sheep, behavioural oestrus (heat) may last 48 hours.
• Ovulation occurs towards the end of oestrus.

Cyclic Changes in the Ovaries

• Ovaries are smaller than those of the cow and their shape is more spherical.
• During anoestrus they measure approximately:
  • 1.3 cm pole-pole
  • 1.1. cm from the fixed edge to the free margin
  • 0.8 cm from side to side
• Transrectal ultrasound is difficult, because the ovaries are difficult to access. Therefore, folliculogenesis is much harder to visualise than in the cow.
• The ewe is similar to the cow, with 3-4 follicular waves in each oestrous cycle.
  • If there are 3 follicular waves, two will occur during the luteal phase and one during the follicular phase.
• Even during anoestrus, dominant follicles will reach the same size as those present during cyclical activity.
• At the onset of oestrus, one or more follicles reach a size of 1cm.
  • Follicular walls are thin and transparent.
• Rupture of the follicle at ovulation is preceded by the elevation of a small papilla above the general surface.
• Ovulation occurs through rupture of this papilla ~24 hours after the onset of heat.
• There is rapid development of the corpus luteum.
  • Development is linear from day 2-12 after ovulation.
• By day 5 of dioestrus, the corpus luteum is 0.6cm in diameter.
• It then attains a maximum size of 0.9cm diameter and has a central cavity.
• Over the period of dioestrus, the colour changes from blood red to pale pink. The size stays constant until the onset of the next oestrus.
• At the onset of the next oestrus:
  • Regression of the corpus luteum of the previous cycle is rapid
  • Colour changes to yellow and then brownish-yellow.
• The luteolytic mechanism is similar to in the cow.
  • At the end of dioestrus, under the influence of oestradiol and progesterone, there is an increase in the number of uterine oxytocin receptors.
  • At the same time, the corpus luteum produces oxytocin, which stimulates PGF2α. PGF2α then acts to stimulate the release of oxytocin in a feedback loop mechanism.
• The first corpora lutea formed after the first ovulation at the start of the breeding season have a shorter lifespan than subsequent ones.
• In twin ovulations, the two corpora lutea can occupy the same or opposite ovaries.
• During pregnancy:
  • The corpus luteum is 0.7-0.9 cm in diameter.
  • It is pale pink in colour
  • The central cavity that is seen in cyclic corpora lutea has disappeared and is filled with white tissue.
• Ovulation with corpus luteum formation but without heat may occur in the anoestrus period. This is known as spurious ovulation.
• The number of ova shed at each oestrus is dependent on genetic and nutritional factors.

Endocrine Changes during the Oestrous Cycle

• Just before the onset of oestrus, there is a rise in oestrogens, particularly oestradiol 17β.
• This is follwed by an LH surge, which peaks at ~14 hours before ovulation.
• At the same time as LH peaks, there is a rise in FSH.
• There is also a second peak in FSH two days after ovulation.
• Progesterone concentrations closely follow changes in the corpora lutea.
  • Maximum progesterone concentrations are lower than those of the cow.
• Prolactin fluctuates throughout the oestrous cycle.
  • Concentrations rise during oestrus and ovulation
  • This pattern reflects the role of prolactin in formation of the corpus luteum.

Nanny Goat

Oestrous Cycle

• Breeding season in Britain is August-February with greatest activity in October, November and December. However, closer to the equator, does will cycle all year round.
• Polyoestrous
• Interoestrous interval of 20-21 days, but this is irregular at the start of the breeding season.
• Duration of oestrus is 30-40 hours.
• Ovulation occurs 12-36 hours after the onset of oestrus.

Cyclic Changes in the Ovaries

• Ovaries vary in shape, depending on the stuctures present.
  • The longest dimention is ~2.2cm.
• The largest follicles reach a maximum diameter of ~1.2cm.
• When these follicles protrude from the surface of the ovary, they often have a bluish tinge.
• There are four follicular waves of growth, with ovulation occuring during the fourth wave.
• Corpora lutea are pink in colour

Endocrine Changes during the Oestrous Cycle

• Endocrine changes are the same as in the ewe.

Sow

Oestrous Cycle

• The domestic sow is generally polyoestrous, but the wild pig is a seasonal breeder with the main breeding season occuring in autumn with another peak in April.
• There is some influence of photoperiod on reproduction in the domestic sow.
  • Anoestrus occurs more commonly in summer and less commonly in February and March.
  • Ovulation rate is lower in summer.
  • Artificially reducing the day length from May-August will decrease the interval from weaning to oestrus from 23.6 days on average to 5.7 days.
• The oestrous cycle normally lasts 21 days, interrupted only by pregnancy and lactation.
  • During lactation, the physical stimuli of suckling will suppress cyclic activity.
• Oestrus normally lasts ~53 hours.
• Ovulation occurs 38-42 hours after the onset of oestrus.
• Many sows show an anovulatory oestrus 2 days after farrowing.
• When weaning occurs at 5-6 weeks, oestrus occurs within 4-6 days.
• Earlier weaning results in a slightly longer interval before subsequent oestrus.

Cyclic Changes in the Ovaries

• The ovaries of a mature, cyclic sow are large and look like mullberries.
  • Surface lobulations are due to elevations of large follicles and corpora lutea.
• When mature, follicles reach a diameter of 0.8-1cm.
• When mature, corpora lutea reach a diameter of 1-1.3 cm.
• It is difficult to study follicular dynamics in a sow because it is hard to identify a single follicle due to the large number present.
• Except during the follicular phase, there is continuous proliferation and atresia of follicles.
• There is normally a pool of about 50 follicles 2-5mm in diameter at any one time.
• Between day 14-16 of the cycle,there is follicular recruitment under the influence of gonadotrophins.
  • This is possible due to a decline in progesterone, and subsequent withdrawal of negative feedback.
• A large number of these recruited follicles are destined for ovuation.
• At day 21-22 it is possible to identify the pre-ovulatory population.
• The growth of pre-ovulatory follicles during this follicular phase is associated with rapid atresia of small follicles. There is also a block to replacement of atretic follicles within the proliferating pool, so there must be some ovarian control mechanism. What exactly this mechanism is is currently unknown, but various substances have been proposed, including:
  • Steroids
  • Growth factors
  • Growth factor binding proteins
  • 'Follicle regulatory protein'
• The ripe follicle is a 'se-shell pink' colour. It has a fine network of surface blood vessels and one point which is very transparent. This transparent point indicates the site of imminent ovulation.
• Haemorrhagic follicles are common.
• After ovulation, a considerable number of follicles ~ 0.4cm remain on the ovary. Some of these enlarge gradually to 0.9cm by day 18.
• Immediately after ovulation, the ruptured follicle is seen as a congested depression on the ovary.
• Accumulation of blood clot soon gives the ruptured follicle a conical shape.
• By day 3 after ovulation, its cavity is filled with a dark red blood clot.
• By day 6, the blood clot is replaced by a connective tissue plug, or slightly yellow fliud.
  • Clots may persist up to day 12.
  • Fluid may persist up to day 18.
• Corpora lutea attain maximum size at 12-15 days post-ovulation.
• After this, they gradually regress to the next oestrus.
• They are dark red up to day 3.
• They then change colour and remain 'wine red' up to day 15.
• As the corpus luteum regresses between days 15-18, the colour rapidly changes to yellow, creamy yellow or buff.
• The mechanism of luteolysis is not fully understood in the sow.
  • Corpora lutea are unresponsive to PGF2α until 12-23 days after ovulation.
  • During luteolysis, corpora lutea are invaded by macrophages.
  • Macrophages produce Tumour Necrosis Factor (TNF).
  • TNF and PGF2α probably work together to cause luteolysis.
  • TNF also inhibits oestradiol production, thus eliminating the luteotrophic source.
• There is rapid regression of corpora lutea at the next oestrus, but throughout the next dioestrus the corpora lutea remain as distinct entities.
• After this they rapidly regress to grey pin-head foci on the ovary.
• During the luteal phase of the cycle, oestrogens are luteotrophic. This may prolong the lifespan of the corpora lutea for several weeks. The result of this is pseudopregnancy.

Endocrine Changes during the Oestrous Cycle

• Oestrogens start to rise at the time of corpora lutea regression.
• Oestrogens reach a peak about 48 hours before the onset of oestrus.
• The ovulatory LH surge occurs at the start of oestrus, 8-15 hours after the peak in oestrogen.
• LH concentrations remain low and fluctuate throughout the rest of the cycle.
• FSH value vary considerably, but there are two surges.
  • One surge is concurrent with the LH peak.
  • A larger surge occurs on day 23 of the cycle.
• Progesterone concentrations closely follow changes of the corpora lutea.
  • In the first 8 days post-ovulation, there is good correlation between progesterone concentrations and the number of corpora lutea present.
  • By day 12 post-ovulation, the correlation is less obvious.
• There are two surges of prolactin.
  • One surge occurs at the same time as the LH and oestrogen surge.
  • A second surge occurs during oestrus.

Bitch

Oestrous Cycle

• Monoestrous, with no seasonal influence.
• A single very long cycle, follwed by a long period of anoestrus.
• Cycle about 3 times every 2 years.
• The luteal phase is about the same length as gestation (~60 days), so pregnancy does not affect the length of the oestrous cycle. Thus, no luteolytic mechanism is required.

Pro-oestrus

• Characterised by vulval oedema, swelling and sanguinous discharge.
  • May not always be visible because some bitches will continually lick the perineum.
• Bitch is attractive to males, but will not accept them.
• Duration is ~9 days

Oestrus

• Accepts the male
• Vulva becomes less oedematous
• Vulval discharge lessens and becomes more clear.
• Duration is ~9 days
• Ovulation usually occurs 1-2 days after the onset of oestrus.
  • Some follicles continue to ovulate 14 days later.

Metoestrus

• Starts when the bitch will no longer accept the dog.
• There is dispute over the duration of this phase, with two main theories:
  • Ends when corpora lutea have regressed at 70-80 days.
  • Ends when endometrial repair is complete at 130-140 days.

Anoestrus

• Transitions from metoestrus to anoestrus with no external signs.
• This is also true after parturition.
• Lasts about 3 months before return to pro-oestrus.

Cyclic Changes in the Uterus

• The endometrium shows considerable change during the oestrous cycle.
• In pro-oestrus and oestrus, endometrial glands are loosely coiled with obvious lumina and a deep epithelial lining.
• During metoestrus, the glands become larger. Lumina become smaller and the basal layer of the endometrium becomes more tortuous.
• As the bitch reaches anoestrus, there is a reduction in the number of glands and the amount that they coil.
• In metoestrus (~98 days after the onset of oestrus), there is desquamation of the endometrial epithelium.
• By day 120-130, the endometrium is restored by proliferation of cells from crypts of the endometrial glands.

Cyclic Changes in the Ovaries

Anoestrus

• In anoestrus, the ovaries are oval and slightly flattened.
• In a medium sized breed, they measure approximately:
  • 1.4cm pole-pole
  • 0.8 cm from the attached end to the free border
• No follicles can be seen.
• On section, tiny remnants of the corpora lutea of previous cycles are visible as yellow/brown spots.

Pro-oestrus

• Developing follicles are about 0.5cm in diameter.
• They enlarge until ovulation, where their size is 0.6-1cm.

Oestrus

• By the time of ovulation, the ovary is enlarged.
• The size of the ovary depends on the number of ripe follicles present.
• The shape of the ovary becomes irregular due to the projection of follicles from its surface.
• The follicle wall is thick, so it is hard to distinguish between follicles and corpora lutea.
• Prior to ovulation, the surface of follicles shows a slightly raised papule. This is about the size of a pin-head. Also, the epithelium covering this papule is brown, whereas the rest of the follicle is flesh-coloured.
• A unique feature of the bitch follicle is the thickness of its wall.
  • Due to hypertrophy and folding of granulosa cells which is visible to the naked eye.
  • This is evidence of pre-ovulatory luteinization.
• Spontaneous ovulation usually occurs 1-2 days after oestrus onset.
• Most follicles rupture over a period of 48 hours.
• The oocyte can be fertilized for up to 108 hours after ovulation.

Metoestrus

• At first, the corpus luteum contains a central cavity.
• By day 10 after ovulation, the cavity becomes filled by compact luteinised cells.
• At this time, the corpus luteum reaches its full size, measuring 0.6-1.0 cm.
• Corpora lutea comprise the greater mass of the ovary.
• Generally, equal amounts of corpora lutea are found on each ovary.
  • The number of foetuses in the uterine horns frequently differ from the number of corpora lutea on the ovaries of their respective sides.
  • Embryonic migration to the uterine horn of the opposite side is common.
• On section:
  • Corpus luteum is yellow-pink
  • It remains unchanged in the non-pregnant bitch until about day 30 after ovulation.
  • It then slowly atrophies
  • During pregnancy, corpora lutea persist at their maximum size thoughout the pregnancy, but regress rapidly after parturition.

Appearance of the Ovaries on Ultrasound

• Developing antral follicles can be seen at the beginning of pro-oestrus.
  • Appear circular and anechoic
• In oestrus, antral follicles are increased in size, reaching a maximum diameter of 4-13mm on day 13 (day 0 is the onset of pro-oestrus).
• From day 10 onwards, walls of the follicles become thickened due to pre-ovulatory luteinisation.
• No evidence of follicular collapse associated with ovulation.
• 25-30 days after the onset of pro-oestrus, the ovaries are difficult to identify.

Endocrine Changes during the Oestrous Cycle

• The luteal phase is prolonged compared to other species, this is shown by persisting levels of progesterone in the bloodstream.
• Progesterone levels begin to rise 60-70 hours before ovulation has occured. This is evidence of pre-ovulatory luteinisation of mature follicles.
• The pre-ovulatory rise in progesterone may signal the bitch to accept the male.
• Oestrogens rise rapidly before the onset of standing oestrus.
• The LH surge rapidly follows and lasts longer than in other species.
• Ovulation occurs 24-96 hours after the LH surge.
• FSH levels peak at the same time as the LH surge.
• As progesterone levels fall towards the end of metoestrus or pregnancy, prolactin increases.
• Prolactin is a major luteotrophic hormone in the bitch.

Pseudopregnancy

• Almost all bitches show some evidence of pseudopregnancy during metoestrus.
  • Covert pseudopregnancy: where the bitch is in metoestrus, but shows little or no signs.
  • Overt pseudopregnancy: where the bitch shows signs ranging from slight mammary development and lactogenesis to mock-parturition with nesting, loss of appetite, straining, emotional attachment to inanimate objects and heavy lactation.
• There is no difference in progesterone concentrations in bitches with or without pseudopregnancy.
• Prolactin is probably responsible for initiating the changes.

Queen

Oestrous Cycle

• Seasonally polyoestrous
• Long day breeders with a period of anoestrus beginning in late autumn.
• Increasing daylight is the most important factor for inducing the resumption of reproductive activity. The first oestrus will usually occur soon after the shortest day of the year.
• If constant 14 hours of daylight is provided, the queen will continue to cycle all year round.
• Some non-pedigrees have regular oestrous cycles lasting about 3 weeks, but others show no regular pattern.
• Duration of oestrus is 7-10 days regardless of mating.

• Oestrogen concentrations increase dramatically at the time of oestrus and may double over 24 hours.
• The principle oestrogen is Oestradiol 17β.
• The rise in Oestradiol 17β corresponds with behavioural oestrus, there are no external signs of pro-oestrus.

Mating and Ovulation

• The queen is an induced ovulator, so mating is important in triggering ovulation.
• There are receptors in the vulva, which are stimualated by copulation.
• This results in release of LH fromt he anterior pituitary gland.
• Only 50% of queens ovulate after a single mating, because multiple ovulations are usually required for adequate LH release.
• The ovulatory LH surge begins within minutes of coitus.
• LH peaks within 2 hours, then returns to basal levels within 8 hours.
• Further matings before the peak in LH will result in further LH release.
• After multiple matings over 4 hours or more, further matings will not result in any further LH release.
  • Due to depletion of the pituitary pool of LH, or the pituitary becoming refractory to further GnRH stimulation.
• Once significant quantities of LH are reached, all ripe follicles will rupture (all or nothing response).
• Occasionally, ovulation will occur in the absence of contact with an entire tom.
  • Receptors similar to those in the vulva are found in the lumbar area.
  • These can be stimulated if the queen is mounted by other females or castrated cats.

Pseudopregnancy

• Sterile matings induce ovulation and result in pseudopregnancy.
• Progesterone concentrations are similar to those of pregnancy for the first 3 weeks.
• After this, progesterone levels gradually fall and reach baseline at 7 weeks.
• Oestrus will occur shortly afterwards.
• Nesting behaviour and lactation are rarely seen, but hyperaemia of the nipples is as evident as in pregnancy.
• Appetite may increase, and fat may distribute to create an increased abdominal size.

Links

• Pharmacological Manipulation of Oestrous Cycles
• Oestrus Behaviour and Methods of Oestrus Detection