Animal Reproduction 7 - 1101
Animal Reproduction 7 - 1101
Reproductive Cyclicity
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Reproductive Cyclicity Estrous
and Menstrual Cycles
1. Estrous cycle
the combination of physiological events
which begin at one estrus and end at the
next estrus
all mammals display estrus except
humans, great apes and a few other
primates
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Estrus behavioral pattern, noun,
characterizes the first phase of the estrous
cycle, this is the time of sexual receptivity
to the male
Estrus is derived from Greek oestrus
(frenzy , gadfly )
Its adjective form is estrous
Referred to as heat
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three types of estrous cycles
1. Spontaneous ovulation of mature
follicles
Formation of functional CL for a
definite period of time regardless of
mating
cow, sow, ewe, mare, bitch, guinea pig
Group includes seasonal and nonseasonal
breeders
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2. Spontaneous ovulation of mature
follicles
Formation of a functional CL occurs only
after mating
This is in response to a neuroendocrine
reflex arc stimulated by mating
Prolactin release maintains the CL
Estrous cycles are 4-5 days long
Rat, mouse, hamster, gerbil , skunks
嚙⿒類動物動情週期:4~5天
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3. Induced ovulators or reflex ovulators
Maturation and ovulation of follicles and
formation of functional CL occurs only if
copulation/mating occurs
Queen, rabbit, ferrets , mink , voles ,
camels
Most of these species have distinct estrous
cycles
These animals come into heat every 14-21 days
and have an estrus of ~8 days, by this time the
ovaries will have a group of mature follicles
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Final maturation of follicle and ovum will
occur only if mated then the CL will form
Sterile mating
pseudopregnancy will occur lasting
30-50 days (gestation is about 63 days)
If not mated, the follicles undergo atresia
and female will come into heat in
another 14-21 days.
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Rabbit is an estrus exception atypical
estrus, continual waves of follicle
growth and receptivity to the male is not
confined to a distinct period
Doe is receptive continuously
Gestation is about 30 days
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Terminology
Estrus noun, part of the estrous cycle
Estrous adjective, describes the cycles
Estrual female adjective
Oestrus and Oestrous
Anestrus vs. anestrous
Polyestrus multiple estrous cycles
throughout the year
Seasonal polyestrus multiple
cycles during a season of the year (long versus
short day breeders)
Monoestrus single estrous cycle per
year
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Polyestrus
Seasonal polyestrus
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Monoestrus
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Follicular
Phase
20%
Luteal
Phase 80%
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4 Stages of the Estrous Cycle
Proestrus
Estrus
Metestrus
Diestrus
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Proestrus
Proestrus is the period immediately preceding
estrus.
Failing CL, release of PGF2 begins during
diestrus and peaks at Diestrus/Proestrus
transition
follicle enlarges and estrogen increases
vascularity of the female reproductive
tract increases
endometrial glands begin to grow
estrogen levels peak
Cow 3-4 days; ewe 2-3 days; sow 3-4 days;
mare- 2-3 days
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Estrus
Estrus is the period during which the female
allows male to mount (copulation) , sexual
receptivity
estrogen decreases
LH surge occurs (peak)
ovulation 24-48 hr after surge of LH
uterine motility high with contractions moving
toward oviduct, sperm transport is optimal
cervical mucus volume increases
Cow: 8-24 hr; ewe: 24-36 hr; sow: 48-72 hr
Mare: 4-8 days
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Metestrus
Metestrus is the transition from estrogen
dominance to progesterone dominance
estrogen low
corpus hemorrhagicum present
CL is forming
Ovulation in cow, goat
uterus contractions subside
endometrial glands continue to grow and
become coiled
3-4 days in cow, 2-3 days in ewe, sow, mare
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Diestrus
Diestrus is the period of maximum luteal function
Progesterone high
FSH low
Uterus secretes fluid but the volume of fluid
decreases over time, contractions stop
corpus luteum regresses at the end of this
period if female is not pregnant
10-14 days in cow; 10-12 days in ewe; 11-13
days in sow; 10-12 in mare
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Estrous cycle Dogs
The estrous cycle of the bitch varies from
patterns previously described.
Wild canines one breeding season per
year
Domestic canines
Monoestrous one estrous cycle per
breeding season may display two breeding
seasons per year
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Bitch Proestrus, estrus, diestrus,
anestrus
Proestrus is considered the start of the
cycle
characterized by bloody discharge
, does not allow mounting
Large follicles present, large increase in
estradiol and peak
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Estrus
mounting, other behavior changes
declining estradiol and rising progesterone
levels
Ovulation occurs 2-3 days after LH surge
Fertilization occurs ~2 days after ovulation
because the oocyte is still in meiosis I at
ovulation, completes meiosis I while in oviduct
Dog sperm survive up to 11 days longer than
other species
Oviduct transport time of egg/embryo is ~ 7days
in bitch
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No distinct metestrus sometimes merged
with estrus , sometimes with diestrus
Diestrus starts with bitch refuses to
be mounted
Pregnancy and diestrus last the same
length of time (~64-65 days)
Pseudopregnancy
Covert no external signs of
pregnancy, high progesterone
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Anestrus Sexual quiescence , ovary
still has low level of follicular development,
~120 days
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Estrous cycle Felines
Seasonally polyestrous (estrus every 17 d)
Recall induced ovulators, mating regulates
phases of estrous cycle
Rapid follicular growth during proestrus (1-3
days) with increase of estradiol.
Estrus (7 - 9 d) estradiol decreases, follicles
retain ability to ovulate during entire estrus
period
No mating follicles undergo atresia, estradiol
rapidly drops, postestrus (no metestrus or
diestrus)
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Mating occurs ovulation occurs 24-50
hrs later, number of ovulations (and LH
surge) depends upon day of estrus cycle
(day 2 is best) and number of matings
Can induce ovulation with LH or GnRH
injection during estrus
Sterile mating results in pseudopregnancy
(45 days) maintain CL but no mammary
development or behavioral changes
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Causes of Anestrus
Seasonal
Gestational (Pregnancy)
Lactation
Presence of Offspring
Stress
Pathology
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How intrinsic is seasonal reproduction?
Cannot completely over-ride [ ]
seasonal reproduction - refractory
Recall sheep must be exposed to long
photoperiod prior to short photoperiod for
puberty
Sheep kept in constant stimulatory
photoperiod will eventually stop cycling
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Seasonal Cyclicity in Mares
Long daylight breeder need a threshold
of light to breed
Breeding season mare displays overt
estrus behavior
Estrous cycle is ~21 days in length with
estrus lasting 5-7 days
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Fall transition (Autumn) and winter anestrus
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Alteration of photoperiod
Stimulates estrous cyclicity high levels of
melatonin associated with winter anestrus
can be lowered by exposure to artificial
light at minimum intensity of 10-12
footcandles for 14.5 hr/day. Response by
individuals may vary.
Same progression of follicular
development as occurs normally, just
shifted.
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First estrus may be prolonged (10-20 days),
ovulation and cyclicity.
Requires 2 months of exposure before breeding,
first ovulation of breeding season is less fertile.
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Seasonal Cyclicity in ewes
In contrast to mares, ewes are short day
length breeders
High levels of melatonin stimulate cyclicity
Fig 7-7
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-
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-
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Melatonin is thought the be a major
regulator of seasonal cyclicity
GnRH and/or Anterior Pituitary hormones
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Factors affecting seasonal anestrus
Mechanism for maximal reproductive
performance and offspring survival
Examples fertilization prior to hot
weather offspring when nutrient availability
is high to support lactation
Season photoperiod zeitgeber
refractory
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Gestational Anestrus
A corpus luteum present during pregnancy produces
high levels of progesterone that has negative
feedback effects on GnRH
LH and FSH do not surge and estrogen remains low
to moderate
The estrogen that is present has negative feedback
effects in the presence of progesterone
The corpus luteum regresses before parturition but
the hypothalamus still does not increase GnRH
pulse frequency and amplitude for a while
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The delay in hypothalamic response to
hormone feedback is possibly due to long
inhibition of hypothalamus during gestation
This delay is useful biologically to allow for
uterine involution prior to the first
postpartum estrous cycle
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Lactational Anestrus
Females suckling offspring experience
anestrus
Sow lactation completely suppresses
estrus, estrus 4-6 days after weaning
Cow - suppressed for up to 60 days
Potential mechanisms visual, olfactory or
possibly auditory encounter with calf
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Suckling interferes with hypothalamic
release of GnRH, provoking a marked
suppression in pulsatile LH release,
resulting in extended postpartum anestrus.
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( ) LH
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LH
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Complete mammary denervation did not
prevent suckling induced inhibition of LH
LH ?
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Nutritional Anestrus
Negative energy balance can cause anestrus
Similar to pre-puberty in that inhibit GnRH pulse
frequency because insufficient energy available
Recall - effects of 2-deoxyglucose infusion in
that it inhibits GnRH release
There are problems with primiparous
females with combined requirements of lactation
and growth
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Menstrual Cycle in Human Females
Average cycle length is 28 days from start of
menstruation until start of next menstruation
(menses )
Menarche first menstrual period
Mensis (Latin) = month
Mene (Greek) = moon
Cycles are more regular in humans in the
absence of artificial light at night
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Ovarian events
Follicular wave starts day 1 -2 with
recruitment followed by selection and
dominance
Ovulation occurs on day 14
CL forms and lasts 12- 14 days
Luteolysis is not dependent on the uterus
CL lasts only 12 - 14 days and regresses if
the female is not pregnant
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Comparison of Menstrual and Estrous cycle
1. Higher primates exhibit a bloody vaginal discharge
associated with shedding of the uterine lining
2. sexual receptivity is not associated with a
particular stage of menstrual cycle ( near ovulation)
3. follicular phase of the menstrual cycle is prolonged (no
secondary surge of FSH has been detected).
4. The human CL produces substantial quantities of
estradiol - unlike luteal tissues of lower mammals and
anthropoid (human-like) primates.
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Menstrual Cycle - Gonadotropins
FSH increases initiate the follicular wave
at days 1-2 and LH is low at this time.
FSH - declines
Positive feedback (E2)
LH rise ~d10
LH surge d13 with ovulation on day 14.
Both LH and FSH then decline and remain
low through the remainder of the cycle
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Ovarian steroids
As follicles begin to grow during selection and
dominance, estradiol is secreted by tertiary follicles.
Estradiol has negative feedback effects initially on the
hypothalamus and anterior pituitary to inhibit GnRH
release and its effects on the anterior pituitary.
Around day 10 - estradiol has positive feedback on the
hypothalamus and anterior pituitary - surge.
The CL - estradiol and progesterone
Luteolysis causes a drop in both estradiol and
progesterone and initiates menses
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Uterine events
Day 1, menses begins and involves loss of
uterine endometrial tissue
Uterine endometrium begins to proliferate under
the influence of estradiol from growing follicles
After ovulation and formation of the CL, estradiol
and progesterone from the CL cause the uterine
endometrium to further proliferate
Luteolysis on day 25-27 results in a drop of both
estradiol and progesterone.
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(Menses) PGF2
Endometrial PGF2 is highest before the onset of
menses.
Vasoconstriction PGF2 causes ischemia ,
toxic catabolites, tissue necrosis , and desquamation
.
Coiled arteries dilate and bleed into the uterine
lumen as levels of PGF2 subside .
Lack of oxygen and nutrients from the blood vessels
tissue death, blood vessels bleed
Blood clots and then liquefies
Menstrual flow consists of both serum and endometrial
tissue debris .
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Menstrual Cycle
Portions of the cycle, for an average cycle
Menses - 5 days
Follicular phase - 9 days
Luteal phase - 14 days
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Variation in cycle length
Menses does not change, approx. 5 days
Luteal phase does not change, approx. 14
days
Follicular phase can be variable
6 days for a 25 day cycle
9 days for a 28 day cycle
16 days for a 35 day cycle
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Mammary tissue tenderness occurs 5 -
7 days before menses
Progesterone and estradiol
Lactation does not occur due to low
prolactin levels
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Effect of age on Menstrual Cycle
Length (average)
35 days for 15 - 19 year olds
30 days for 30 year olds
28 days for 35 year olds
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Amenorrhea Nutritional and Lactational
Absence of menses for an extended period of
time in women of reproductive ages
Lactational - high levels of prolactin suppresses
GnRH pulse frequency and amplitude
Length depends upon nutrition varies different
locations in the world.
Is viewed as a form of birth control in certain
cultures(Fig 7-11)
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7
1.
2. 3 (Causes of
Anestrus)
3. 7-7
4. (follicular phase,
ovulation, fertile period, luteolysis
sexual reception)
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7
5. (stages)
LH
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7
6.
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7
7.
90