ANS 431: Estrous Cycle



I.	Introduction

	A.	Estrous cycle
		1.	Period of reproductive cyclicity
	B.	Anestrus
		1.	Periods of no estrous cycles
			a.	Pregnancy
			b.	Nursing
			c.	Season of year
			d.	Poor Nutrition
			c.	Pathological Conditions
	C.	Other Terminology
		1.	Estrus
			a.	Noun
			b.	Period of sexual receptivity
			c.	Heat = Common name
			d.	Oestrus = British equivalent
		2.	Estrous
			a.	Adjective to describe events associated 
				with estrous cycle
			b.	Oestrous = British equivalent
		3.	Seasonal Breeders
			a.	Animals that have one or more estrous cycles 
				during certain periods of the year 
		4.	Polyestrous Animals
			a.	Estrous cycles throughout the year
			b.	Cattle, Pigs, Humans
			c.	Are sometimes only bred at certain periods 
				of the year, but will cycle all year
		5.	Seasonally Polyestrous Animals
			a.	Animals that have multiple estrous cycles 
				only during certain periods of the year
			b.	Short-Day Breeders
				i.	Cycle when days are getting shorter
					--Fall
				ii.	Sheep, Goats, Deer, and Elk
				iii.	Anestrus in spring and summer
			c.	Long-Day Breeders
				i.	Cycle when days are getting longer
					--Spring
				ii.	Horses and Hamsters
				iii.	Anestrus in fall and winter
		6.	Monestrous animals
			a.	Animals that have one estrous cycle per year
			b.	Dogs, wolves, foxes, and bears
			c.	Extended periods of estrus (days)
				i.	Increase probability of mating 
					and pregnancy	

	
II.	Length of Estrous Cycle in Different Animals

					Length

	Ewe				17 d (13-19)
	
	Goat				21 d (15-24)
	
	Sow				21 d (17-25)

	Cow				21 d (17-24)
	
	Mare				21 d (15-25)

	Woman				28 d (Menstrual Cycle)

	Rodents				4-6 d

III.	Divisions of the Estrous cycle

	A.	Luteal and Follicular Phase
		
		1.	Luteal Phase = Period of the Corpus Luteum
		2.	Follicular Phase = Period of the Follicle
		3.	Order

			a.	Non-menstruating animals

				Estrus to Luteal Phase 
				to Follicular Phase to Estrus
	
			b.	Menstruating animals

				Menses to Follicular Phase 
				to Luteal Phase to Menses
				
	B.	Proliferative and Secretory Phase 
	
		1.	Proliferative Phase = Thickening of endometrium
		
			a.	Due to estrogen secretion
			b.	Menses + Proliferative Phase = Follicular Phase
		
		2.	Secretory Phase = Endometrium highly secretory
		
			a.	Due to progesterone secretion
			b.	Luteal Phase

	C.	Metestrus, Diestrus, Proestrus, and Estrus

		1.	Metestrus + Diestrus = Luteal Phase
		2.	Proestrus + Estrus = Follicular Phase
		3.	Metestrus
			a.	Begins at cessation of estrus
			b.	Lasts for about 3-5 days
			c.	Period of the formation of the 
				Corpus Luteum
			d.	Some Bleeding in Farm Animals
				--Declining Estrogen causes 
					capillary vessels to break
				--Not an indicator of conception
				--Not the same as menstruation
		4.	Diestrus
			a.	Lasts between 10-14 days
			b.	Period of a functional C.L.
			c.	Progesterone increases
			d.	Some follicular growth
		5.	Proestrus
			a.	Lasts 2 to 4 days
			b.	C.L. regresses (apoptosis begins)
			c.	Rapid follicular growth
			d.	Estrogen increases
		6.	Estrus
			a.	Period of sexual receptivity
			b.	Usual period of ovulation

III.	Duration of Estrus and Time of Ovulation

	A.	Ewe

		1.	Estrus Duration--30 h (18-48)
		2.	Ovulation--24-30 h from beginning of estrus
		3.	LH surge to Ovulation--26 h

	B.	Sow

		1.	Estrus Duration--50 h (12-96)
		2.	Ovulation--36-44 h from beginning of estrus
		3.	LH surge to Ovulation--40 h

	C.	Cow

		1.	Estrus Duration--15 h (6-24)
		2.	Ovulation
			--24-32 h from beginning of estrus
			--10-12 h after end of estrus
		3.	LH surge to Ovulation--28 h

	D.	Mare

		1.	Estrus Duration--7 d (2-12)
		2. 	Ovulation
			--5 d from beginning of estrus
			--1-2 d before end of estrus
		3.	LH surge to Ovulation--2 d


V.	Anestrus

	A.	Gestational Anestrus
	
		1.	Inhibition of GnRH by Progesterone
			a.	From CL verum (early pregnancy)
			b.	From Placenta (later pregnancy)

	B.	Postpartum Anestrus 

		1.	Reproductive System needs a Period of Recovery
		2.	Uterus need to undergo involution
			a.	Time Required
				i.	Cow--35-40 d
				ii.	Sheep--25-30 d
				iii.	Pig--25-28 d
		3.	Postpartum Ovulations
			a.	Sows
				i.	Estrus w/o ovulation
					--few days after parturition
			b.	Horses
				i.	Fertile Estrus
					--1-3 weeks after parturition
			c.	Ruminants
				i.	Silent Ovulation
					--2-3 weeks after parturition
					--Not very fertile

		4.	Fertility Maximum after Parturition
			a.	Cows--60-90 d
			b.	Sows--After Weaning
			c.	Sheep--Next Season
			d.	Horse--Weeks after parturition

	C.	Lactational Anestrus

		1.	Prevents a new pregnancy before young are weaned
			a.	Weaning induces Estrus
		2.	Sow
			a.	No cycles during lactation
			b.	Estrus and ovulation 4-8 days after weaning
		3.	Cow
			a.	Suckling can delay cycles up to 60 days
				i.	Very variable
			b.	Needs to be at least 2 encounters per day
				i.	Effect not increased 
					with more sucklings
					--Threshold effect
		4.	LH pulsatility is decreased
			a.	Weaning causes increased episodic pulses
		5.	Mammary stimulation is not totally responsible 
			for lactational anestrus
			a.	Weaning causes same effect in denervated cows
			b.	Encounter with offspring appears 
				to be necessary
				i.	Visual, Olfactory, and/or Auditory
				ii.	Dairy cows that have offspring 
					removed early don't encounter 
					lactational anestrus
				iii.	Needs to be own offspring
			c.	Negative energy balance related to lactation 
				can also cause anestrus	
				i.	Especially in primiparous 
					(first time birth) females
				ii.	Still growing

	D.	Other factors associated with anestrus
		
		1.	Nutrition
			a.	Poor nutrition reduces reproduction
		2.	Pathology
			a.	Uterine infection	
			b.	Persistent corpus luteum
			c.	Mummified fetus
			
VI.	Dogs and Cats

	A.	Dogs
	
		1.	Mono- or Biestrus depending on breed.
			a.	3-9 month estrous cycle
				i.	5 months anestrus (major variant)
					--large increases in LH pulsatility before proestrus
				ii.	9 day proestus
				iii.	9 day estrus (4-21 days)
				iv.	2 month diestrus
		2.	Ovulation
			--4-24 days from beginning of estrus
		3.	LH surge to Ovulation--2-3 days
		
	B.	Cats
	
		1.	Induced Ovulators
			a.	Only ovulate if mated
		2.	Estrous cycle length--17 days (4-30)
		3.	Duration of estrus--9 days
		4.	LH surge to Ovulation--30-40 hours

VII.	Seasonal Breeders

	A.	Introduction

		1.	Wild species have a breeding season that 
			is initiated at a time when the environment 
			will allow for the best survival of the young.
			--Spring is usually the optimal season 
				for parturition
		2.	Time of breeding is then dependent 
			on Gestation length.
		3.	Domestication has stopped seasonal breeding 
			in some species
			a.	Cattle & Swine
		4.	Of the farm species, Sheep, Goats and Horses 
			are still seasonal breeders

	B.	Short Day Breeders:  Sheep and Goats

		1.	Most sheep and goats are seasonal breeders

			a.	Breeds native to the tropics may cycle 
				throughout the year
			b.	Arctic natives shorter season 
				than temperate natives

		2.	Ewes are short-day or fall breeders

			a.	Cyclicity dependent on photoperiod
				i.	Starts Daylight to Darkness ratio 
					decreases
				ii.	Ends Daylight = Darkness again
				iii.	Between Autumn and Spring Equinox
				iv.	Time of change called Transition
				v.	Some breeds have extended breeding
					--Horned Dorset, Merino, Rambouillet
			b.	Silent ovulations occur more frequently at 
				beginning and end of season
			c.	Introduction of Ram during fall transition 
				causes synchrony of estrus
				i.	1st estrus 15 to 20 days 
					after introduction of male

		3.	Does are also short-day breeders
			a.	Cycle between Late June and April
			b.	Peak Breeding between September and January
			c.	Introduction of Buck during transition also 
				causes synchrony of estrus
				i.	1st estrus 5-10 days 
					after introduction
				ii.	Lactation independent
	
		4.	Rams and Bucks
			a.	Also affected by photoperiod 
					(Less than females)
			b.	Highest breeding activity and fertility 
				in fall
				i.    	As photoperiod increases 
					(Spring & Early Summer)
				ii.	Sperm production decreases
				iii.	Abnormal Spermatozoa increases	
				iv.	Fertility decreases
			c.	Rams have some sexual activity year around
			d.	Bucks usually become sexually inactive 
				during anestrus
		5.	Other influencers
			a.	Season can be reversed by changing animals 
				from Northern to Southern hemisphere
			b.	Control of Light artificially has also been 
				used to reverse anestrus
				i.	16 hrs dark/8 hr light will induce 
					early transition
				ii.	Both sexes most be exposed to changes

	C.	Long Day Breeders:  Horses

		1.	Mares are long-day breeders
			a.	Cyclicity dependent on photoperiod 
				(opposite sheep & goats)
				i.	Starts when Daylight 
					to Darkness ratio increases
				ii.	Ends when Daylight Decreases
				iii.	Ponies: May to October
				iv.	Horses: February to November
				v.	Peak fertility: May to June
					--a lot of variation
				vi.	Artificial light will increase 
					earlier transition
					--16 hrs on/8 hrs off
		2.	Stallion not as sensitive as Mare
			a.	Fertile year around
			b.	Sexual activity and Semen production 
				decreases during mare anestrus period

	D.	Endocrinology
	
		1.	Pineal Gland
			a.	Responds to environmental lighting 
					(Photoperiod)
				i.	Retina receptors are stimulated
				ii.	Signals transmitted 
					to Suprachiasmatic nuclei 
					in Hypothalamus
				iii.	Signal then transmitted 
					via sympathetic nerves to Pineal
			b.	Secretes Melatonin
				i.	Tryptophan Derivative (Amine)
				ii.	Darkness stimulates synthesis 
					& release
				iii.	Inhibits reproduction 
					in long day breeders
					--Horses, Hamsters
				iv.	Stimulates reproduction 
					in short day breeders
					--Sheep and Goats
				v.	Probably acts on Hypothalamus
				vi.	Affects GnRH synthesis and release

		2.	Pituitary
			a.	LH increases in pulsatility just before 
				breeding season
			b.	Similar to puberty



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~~~~~Revised 2/25/04~~~~~ TAW