“Midterm Notes for ANS150”

Fertilization and Cleavage

I. General Concepts
 Mammalian reproduction evolved to internal fertilization & cleavage.
 Purpose: avoid accumulation of harmful mutations & adapt in changing
environments.
 Sperm & ova have equal nuclear genome BUT different behavior, lifespan,
maturation.
 Egg removes almost all sperm cytoplasmic/nonnuclear elements after fertilization.

II. OVUM MATURATION

 Ovum arrested at prophase I during folliculogenesis.
 Ovulation normally releases ovum at metaphase II.
 Species exception (still in 1st meiosis at ovulation):
 Horse
 Dog
 Fox
 Meiosis completes only after fertilization → zygote formed.

III. SPERM MATURATION

 Requires 10–15 days transport through epididymis.
 Epididymal secretions + transit = essential for fertilization ability.
 Requires sequential modifications:
 Maturation
 Capacitation
 Acrosome reaction
 Purpose: enable penetration + fusion with ovum.

IV. CAPACITATION
 Discovered by Chang & Austin.
 Occurs in female tract (mainly uterus & isthmus of oviduct).
 Changes:
 Removal/modification of surface components
 Cholesterol depletion
 GAG alterations
 Ion changes
 Membrane destabilization
 Function:
 Prevents premature acrosome reaction
 Prepares acrosomal activation at ovum
 Leads to TRUE acrosome reaction → fusion of sperm plasma membrane + outer
acrosomal membrane → vesiculation.

True vs False acrosome reaction:

 True = fertilization, enzyme release, membrane fusion
 False = degeneration/aging
Enzymes released:
 Hyaluronidase
 Acrosin → required for zona penetration.

V. Interaction of Sperm & Ovum

 Lifespan importance
 Successful conception requires overlap of:
  Sperm viability
 Ovum viability
 Species differences:
 Highest longevity in pig & horse → good conception even if
insemination occurs early relative to ovulation.
 Sperm numbers:
 Billions ejaculated → drastically reduced:
 1,000–10,000 reach isthmus
 Only 10–100 in ampulla (site of fertilization)
 Reduction NOT due to slow transport but controlled restriction by:
 Uterotubal junction
 Lower isthmus
 Cervix/uterus barriers (ruminants)
 Importance:
 Prevent polyspermy
 Maintain reservoir of capacitated sperm
 Hyperactivation:
 Occurs in ampulla
 Increases probability of ovum contact

VI. SPERM–OOCYTE ENCOUNTER (3 Key Events)

1. Migration through cumulus cells
2. Binding and penetration of zona pellucida
3. Fusion of sperm & ovum plasma membranes
 Cumulus cells may secrete motility-activating factors (e.g., rabbit).

VII. SPERM ATTACHMENT (Zona Binding):

 Based on receptor recognition on zona (mainly ZP3).
 Blocked by:
 Anti-zona antibodies
 Trypsin
 Antisperm antibodies

 Soluble zona glycoproteins → blocks zona receptor sites.

 Mechanism resembles enzyme–substrate lock-and-key.

VIII. SPERM PENETRATION

 Occurs 5–15 min after attachment.
 Acrosome-intact sperm required for initial attachment.
 Binding to ZP3 triggers acrosome reaction.
 Multiple enzymes act synergistically: NOT only acrosin
 Motility + enzymes both required.
 ZP2 acts as secondary receptor maintaining attachment during penetration.

IX. GAMETE FUSION

 Vitelline membrane (oolemma) less specific than zona, but prefers homologous
sperm.
 Acrosome reaction is REQUIRED before fusion.
 Zona-free ova cannot fuse with non-acrosome-reacted sperm (attachment only, no
fusion).

KEY TERMS
a. Capacitation: membrane changes in female tract enabling acrosome reaction.
b. Acrosome reaction: membrane fusion + enzyme release for zona penetration.
c. Polyspermy block: cortical granule reaction.
 d. Hyperactivation: increased motility in ampulla.
e. ZP3: primary receptor; ZP2: secondary receptor.
f. Hyaluronidase & acrosin: zona penetration enzymes.
g. Sperm reservoir: uterotubal junction/isthmus.

Vitelline Membrane & Sperm Fusion

 Vitelline membrane covered with dense microvilli, except an elevated area near 2nd polar body extrusion (rare
sperm attachment here).
 Initial sperm attachment occurs at equatorial segment of sperm head → contacts ovum microvilli.
 Fusion does NOT involve plasma membrane above equatorial or post-acrosomal region.
 Equatorial region of sperm becomes incorporated into egg membrane → identifiable until 8-cell stage.

Block to Polyspermy

 After fertilization, ovum membrane changes to prevent entry of additional sperm.

 Failure → polyspermy → polyploid embryo → embryonic death/abnormality
 Polyspermy:
o Mammals: 1–2%
o Pig highest (up to 15%, delayed insemination)
o Sheep: delayed breeding → 39% abnormal or polyspermic eggs

Pronuclei Formation & Syngamy

 Upon sperm penetration:

o Ovum completes meiosis → extrudes 1st and/or 2nd polar body
o Female haploid chromosomes form female pronucleus
 Male and female pronuclei migrate → center → cytoskeleton rearrangement → syngamy
 Both pronuclei are essential → normal development requires male + female genome

Interspecies Fertilization

Examples

 Successful:
o Snowshoe hare × rabbit
o Mink × ferret
o Sheep × goat
o Horse × donkey
o Bos taurus × Bos indicus
o Cattle × bison
 Hybrids often:
o ↑ disease resistance
o ↑ environmental tolerance

Failure causes

 mostly physiological gamete differences, not just genetic incompatibility

Hybrid development
  Normal until blastocyst, later failure due to post-blastocyst species-specific requirements

Reciprocal cross difference

 Success rate depends on direction (♂A × ♀B ≠ ♂B × ♀A)

Trophoblast importance

 maternal immune rejection prevented by placing trophoblast-origin cells externally in chimeric embryo

Cleavage

 Series of mitotic divisions of zygote

 No increase in cell mass
 Decrease in cell size → increases nuclear : cytoplasmic ratio
 Early growth = “negative growth”

Nutritional Support

 Early embryo has little yolk → depends on oviduct & uterine secretions (histotroph)

Cleavage Pattern

 First division: vertical, animal → vegetal pole

 Second: vertical, perpendicular to first
 Third: perpendicular to second
 Produces: 1 → 2 → 4 → 8 blastomeres

Totipotency

 Rabbit: 2–8 cell stage = totipotent

 Sheep:
o 4-cell: max 3/4
o 8-cell: max 1/8

Morula Formation

 Around 9–16 cells

 Compaction begins at 8-cell
o flattening
o cell polarization
 Leads to formation of:
o Inner Cell Mass (embryo proper)
o Trophectoderm (future chorion)

Division planes

4th cleavage division:

 Anticlinal → perpendicular
 Periclinal → parallel
 Oblique → intermediate

→ Determines inner vs outer cell allocation

Blastocyst Formation

 Tight junctions in trophoblast → fluid accumulation → blastocoele formation

 Gap junctions isolate embryo from maternal environment

Cleavage Rate + Species Differences

 Fertilization in lower ampulla

 Enters uterus at:
o 8–16 cell stage (most spp.)
o Pig → 4-cell

Important exception

 Horse: unfertilized ova remain and degenerate in oviduct

Hormonal Regulation of Transport

 Oviduct transport influenced by maternal factors

 Estrogen/progesterone affect isthmus muscle function
 PGF inhibits embryo entry into uterus

Genetics

 Cleavage rate influenced by embryo genetics (e.g., mouse MHC-linked genes)

Term Key Fact

1. Polyspermy  Highest in pigs
2. Pronuclei  Both necessary for development
3. Totipotency  Up to 8-cell
4. Compaction  Starts at 8-cell
5. Inner cell mass  Embryo proper
6. Trophectoderm  Chorion/trophoblast
7. Cleavage  Mitotic, no mass increase
8. Histotroph  Maternal nutrition source

1. Expression of the Genome

Before fertilization

 Ovulated oocytes already contain rRNA, mRNA, tRNA, ribosomes

 Can synthesize proteins without sperm

After fertilization

 Protein synthesis continues into cleavage stages

 Early development relies on maternal RNA

Zygotic gene activation

 New transcription begins after early cleavage divisions

  Embryonic genome activates
o by 2-cell stage
o strong after 1st cleavage

Sheep protein synthesis

 High at 1st & 2nd cleavage

 Drops 95% by 3rd cleavage
 Low at 4th (8–16 cells)
 Rises at 5th (16–32)
 Full transcription activation = 4th cleavage
 Similar findings in cows

Types of embryonic gene products

1. Constitutive → present in all cells (actin, tubulin, metabolic enzymes)

2. Temporal → appear only at specific stages (ex: plasminogen activator in peri-implantation)

Control mechanisms

 Peptide growth factors likely regulate early conceptus development

2. Parthenogenesis

 Development of embryo without sperm

 Common in invertebrates
 Mammalian parthenogenesis occurs but rarely develops beyond 1–2 cleavages

Experimental types

 Androgenones = 2 paternal genomes

 Gynogenones = 2 maternal genomes
→ both fail to develop fully

Reason

 Maternal and paternal genomes are not functionally equivalent

o due to genomic imprinting
 Development requires both parental contributions

Chimeras

 Parthenogenetic cells + normal cells can form viable animals

Twinning in Domestic Animal

Dizygotic twins

 Common type
 Two ova, two sperm
 Influenced by:
o breed
 o age
o environment

Cattle

 Dairy: ≈3.5%
 Beef: <1%
 One fetus per horn = normal viability

Freemartinism

 91% of female calves born with a male co-twin are sterile (shared placental blood supply)

Sheep

 Prolific breeds (Finnsheep, Booroola Merino)

 Booroola F gene → high ovulation (↑ FSH)

Horses

 Double ovulation common

 Twin survival low (placental insufficiency)

Increasing twinning

 exogenous gonadotropins (eCG) → variable

 bilateral embryo transfer ~60%
 Immunization against inhibin → ↑ ovulation

Monozygotic Twins

 Split from the same embryo

 Rare naturally
 Occur after implantation (ICM divides into two)

Nuclear Transplantation (Cloning)

 Nucleus transferred to mature oocyte

 Requires cytoplasmic “reprogramming”
 Embryonic nuclei lose totipotency as development advances
 Original success rates were low but improved (sheep, cattle, rabbits)

Note: mitochondrial DNA comes from recipient oocyte (not a perfect clone)

Blastocyst Formation

 After morula
 Formation of blastocoele
 Differentiation into:
o Trophoblast → placenta (chorion)
o Inner cell mass → embryo (3 germ layers)

Zona Hatching
  Occurs 4–8 days post-ovulation
 Triggered by:
o blastocyst expansion
o uterine enzymes
o prostaglandins (E series important)

Blastocyst Elongation

Ruminants (sheep/cow)

 Rapid elongation
 Day 13–18
 Up to 25 cm

Pigs

 Fastest elongation of all species

 2 mm → 20 cm
 Cellular remodeling + hyperplasia
 Important for uterine space competition

Horses

 Remain spherical
 Slow enlargement

Embryo Migration & Spacing

 Essential in pigs to avoid crowding

 Pig embryos migrate then space evenly by day 12
 Rare in cows, minimal in sheep unless many ovulations
 Horses show frequent migration until fixation
 IMPLANTATION
Early embryonic development

Blastocyst Formation

 Compaction of morula → tight junctions form.

 Water enters forming blastocoele due to Na⁺/K⁺ pumps on trophectoderm → creates
solute gradient → draws water.
 Blastocyst contains:
o Trophoblast – forms placenta, adhesion to uterus
o Inner cell mass (ICM/embryoblast) – forms embryo
 ICM later forms 3 germ layers (ectoderm, mesoderm, endoderm) → gives rise to skin,
nervous system, muscle, organs, etc.
 Trophectoderm has microvilli → uterine attachment.

Zona Hatching

 Occurs 4–8 days post-ovulation

 Mechanisms:
o Blastocyst expansion (fluid + hyperplasia)
o Rhythmic contractions
o Enzymes (plasmin, trypsin) soften zona
 Prostaglandin E important for expansion and hatching.

Intrauterine Migration & Spacing

Purpose:

 Contact with endometrium

 Prevent luteolysis by blocking PGF2α
 Provide space for conceptuses

Pig

 Highly important
 Migration day 5–12
 Requires ≥2 embryos per horn for pregnancy maintenance

Cow/Ewe

 Limited migration
 Occurs only with double ovulation

Horse

 Most extreme migration

 Occurs 10–13 times/day (Days 10–16)
 Prevents luteolysis
 Fixes after Day 16
Conceptus Expansion

 Trophectoderm grows and elongates

 Rapid elongation ensures contact with uterine surface

Cow & Ewe

 Slow, by cell hyperplasia

 Day 13: sphere → day 17: long filament

Pig

 Most rapid
 Tubular → filamentous within 2–3 hours
 Length 20 cm by day 12
 Driven mainly by cell remodeling, not hyperplasia
 Retinoic acid possibly involved

Horse

 Remains spherical up to day 50

Implantation

 Uterus becomes receptive when Muc-1 decreases

 Muc-1 normally prevents attachment
 Progesterone exposure → downregulates receptors → ↓Muc-1 → receptive uterus
 Livestock = non-invasive implantation

Pig

 Attachment Day 13–24

 Microvilli interdigitation
 Areolae absorb uterine secretions

Ruminants (cow, ewe)

 Binucleate cells form → fuse with uterine epithelium

 Transfer placental lactogen
 Immunological protection

Horse

 Attachment Day 24–40

 Chorionic girdle → endometrial cups (produce eCG)

Placentation

 Extraembryonic mesoderm forms:

o Yolk sac
o Amnion (surrounds embryo)
o Allantois (joins chorion → chorioallantoic placenta)
  In mare, yolk sac persists longer

Maternal Recognition of Pregnancy

Conceptus must prevent luteolysis (PGF2α) → maintain CL → maintain progesterone.

General

 Without recognition → PGF2α released → luteolysis → return to estrus

Sheep and Cow

 Conceptus secretes proteins (Days 12–21)

 ↓ PGF2α pulses
 Main signal = interferon-tau (oIFN-τ)
 Action local in uterus, not systemic
 Blocks uterine synthesis of PGF2α

KEY CONCEPTS
Process Key Mechanism
Blastocoele  Na/K pump → water
formation influx
Germ layers  Gastrulation
Hatching  PGs + enzymes
Migration  Prevent PGF2α
Pig elongation  Rapid, remodeling
Ruminant signal  Interferon-tau
Implantation
 ↓Muc-1
control
 Mesoderm +
Placenta formation
trophectoderm

High-yield species differences

Pig

 Rapid elongation
 Migration essential
 Estrogen signal

Cow/Ewe

 Interferon-tau
 Limited migration

Horse

 Vesicle migration essential

 Endometrial cups
 eCG
 CATTLE AND BUFFALO

Family Bovidae but DIFFERENT genera and chromosomes

 Cattle: Bos taurus, Bos indicus

 Buffalo (water buffalo): Bubalus bubalis
 African wild buffalo: Syncerus caffer
 American “buffalo”: Bison bison
-> All Bovidae, not the same species, so reproduction features are not interchangeable.

PUBERTY

Female

 Puberty = first estrus followed by NORMAL luteal phase

 First ovulation ≠ puberty (may be silent ovulation, short cycles, anovulation)
 Puberty depends on physiologic age, NOT chronological age

Factors affecting puberty

 Nutrition
o ↑ nutrition → earlier puberty
o ↓ nutrition → delayed
 Body size measurements more useful
than age
(bodyweight, wither height, pelvic area)
 Breed
o Dairy: 10–12 months
o Beef: 11–15 months
o Zebu: 18–24 months
 Season
o Winter = delayed puberty

Male

Puberty = ejaculate containing ≥50 million sperm with ≥10% progressive motility

Mechanism

 ↑ LH pulses → testosterone → Sertoli differentiation → spermatogenesis

Breed
  Holstein earliest
 Charolais intermediate
 Angus/Hereford later
 Zebu latest
 Crossbreds earlier than purebreds

Testes keep increasing sperm output until 18–24 months

ESTROUS CYCLE

 Cattle are non-seasonal breeders due to domestication

 Beef cows may retain some seasonality (photoperiod sensitive)

Follicular Dynamics

 Follicles grow in waves (1–4 waves, usually 3)

 From each wave, one dominant follicle
 Large follicles shift dependence from FSH → LH
 A dominant follicle is always present

Ovulation

 10–12 hrs after END of estrus

(~30 hrs after onset)

Frequency

 Usually 1 oocyte
 10% = double ovulation
 Right ovary 60% of time

HORMONES

Estrogen actions

 ↓ cervical mucus viscosity

 ↑ uterine tone
 Cervix dilates
 Behavioral estrus
 Metestrus bleeding (not related to conception)

Progesterone actions

 Thick cervical mucus

 Closed cervix
 Relaxed uterus

Estrus Detection
  Observe 2× daily
 Best indicators:
o standing heat
o uterine tone (rectal exam)
o mucus
 Aids:
o vasectomized bulls
o chin-ball marker
o pressure-sensitive patches

Efficiency
= % of estrus periods detected

Accuracy
= % detected that are true estrus

SPERM PRODUCTION

 Full spermatogenesis = 61 days

 Testicular size correlates with sperm output
 Ejaculate volume: 4–10 mL
 Concentration: 0.8–2.0 billion/mL

Stress (heat, fever, transport) = temporary infertility for ~2 months

MATING

 Natural service mostly in beef herds

 Most dairy = artificial insemination
 Bulls per cows: 1:30–60
 Social hierarchy affects mating success

GESTATION

 ~276–295 days
 Longest in Brown Swiss & Brahman

PARTURITION Signs

 Relaxed tailhead ligaments

 Vulvar swelling + mucus
 Isolation behavior
 Udder distension

PUERPERIUM

Uterine involution
  Faster in:
o primiparous
o suckled cows

Postpartum ovarian function

 First ovulation usually without estrus

 Dairy cows:
o estrus ~35 days
o breed at ≥50 days
o should conceive by 80 days (to maintain 12-month calving interval)

Effect of Suckling

 Suckling → ↓ gonadotropins (LH/GnRH)

→ extended postpartum anestrus
→ major problem especially in beef

Undernutrition

 moderate deficiency → affects ovulation

 severe deficiency → affects dominant follicle development
 important mediators:
o glucose
o insulin-like peptides
o PGF2α

SUPER-SHORT MNEMONICS

Puberty Factors (heifer)

B-N-S

 Body size
 Nutrition
 Season

Estrogen Effects

MUT

 Mucus thin
 Uterus tone ↑
 Tubular tract patent (cervix open)

Progesterone Effects

RTL

 Relaxed uterus
 Thick mucus
 Locked cervix
Reproductive Performance

Measures of Reproductive Efficiency

1. 60–90 day non-return rate

 % cows not returning to estrus after AI

 Estimates bull fertility + inseminator efficiency

2. First service conception rate

 Pregnancy diagnosis 6–8 weeks after insemination

3. Calving to conception interval (Days open)

 Indicates estrus detection + fertility of male/female

4. % cows pregnant

 Used in beef herds, more meaningful with short breeding season

5. Calf crop

 Measures pregnancy loss + calf mortality

 % calves weaned = reflects breeding season efficiency, calving ease, mothering ability &
calf survival

Nonreturn vs Conception Rate

 Nonreturn overestimates conception by 10–15%

Due to:
 failure of estrus detection
 anestrus
 early embryonic death
 sale/death of cows

Calving interval & Days Open

 Gestation fixed → both influenced by same factors

 Days open = earlier detection of problem cows

General Fertility

Optimal calving interval = 12 months

BUT rarely achieved

Beef cattle:

 net calf crop = 70–75%

Dairy cattle:

 % cows calving to first service = only about 50%

IMPROVING REPRODUCTIVE PERFORMANCE (CATTLE)

Beef cattle

 Most bred by natural service → bull must have:

o fertile semen
o libido
o stamina
 Huge improvement by breeding soundness exam of bulls
 Early estrus expression is critical

Nutrition

 Puberty earlier in well-fed heifers

 Postpartum estrus earlier with proper feeding

Current reproductive technologies

 calf manipulation
 estrus synchronization
 targeted nutrition
 improved environment
→ reduces postpartum anestrus

Dairy: To maintain 12-month interval

Cows should:

 show estrus by 60 days postpartum

 conceive by 85 days postpartum

Best conception after 60 days postpartum

(breeding earlier → lower conception)

Estrus Detection (very exam-important)

Improved by:

 observation 2× daily
 heat detectors
 milk progesterone
 rectal palpation

↑ detection efficiency from 50% → 90%

Bull contact stimulates estrus postpartum

Hormonal management

 Early postpartum GnRH = variable

 hCG: helpful for repeat breeders, NOT first service
 GnRH NOT recommended for first service (may reduce fertility)

AI Practices Improving Fertility

 inseminate only cows in estrus

 correct semen thawing
 deposit semen in uterus
 massage uterus/clitoris
 avoid high temperature after AI

Nutrition & Fertility

Negative energy balance:

 ↓ IGF-1
 ↓ luteal function
 ↓ fertility

rBST before 1st ovulation → harmful

BUFFALO
REPRODUCTION

Types:

 Swamp (2n=48) =
draft
 River (2n=50) =
dairy

Puberty

Later than cattle

River = 15–18 mo
Swamp = 21–24 mo
First conception: 24–36 months

Estrous cycle

Signs milder than cattle

Estrus = 12–30 h
Ovulation = 15–18 h after estrus end
Best detection = acceptance of male
Homosexual behavior uncommon

Seasonal effects

Heat + long photoperiod → ↓ fertility

High temperature → ↓ libido

Male buffalo

 Smaller testes & glands

 Libido ↓ in hot weather
 Ejaculate small volume (5 mL max)
 Sperm motility lower than cattle

Gestation

River = 305–320 days

Swamp = 320–340 days
Longer than cattle

Postpartum

Uterine involution:

 Swamp suckled = 28 days

 River hand-milked = 45 days

Postpartum anestrus common

Reproductive Efficiency (Buffalo)

Conception rates:

 Natural: >60%
 Chilled semen: 50–60%
 Frozen semen: 25–45%

Calving interval:

 Range: 2 calves in 3yrs

 Well managed dairy: 14–15 months

Major problems in Buffalo

  delayed puberty
 poor estrus detection
 long days open
 low libido in heat

Improving Buffalo Reproduction

 Improve nutrition
 Early weaning
 Estrus induction (PGF2α)
 Synchronization
 Cooling systems
 Use frozen semen collected during cool season

KEY EXAM TAKEAWAYS

Cattle

 Optimal calving interval = 12 months

 First-service conception dairy = 50%
 Beef calf crop = 70–75%

Buffalo

 Lower fertility
 Later puberty
 Poor estrus signs
 Strong seasonal effect
 Frozen semen lower conception

SHEEP AND GOAT

Introduction
 Sheep (Ovis aries)
 Goats (Capra hircus)
 Both family Bovidae, but different species
 Early domestication
o Sheep → wool & meat
o Goats → milk, meat, fiber
 Goats important protein source in tropics
 World population: sheep > goats
SEXUAL SEASON
Females
 Temperate zone
o Both seasonally polyestrous
o Breed in fall
 o Parturition in spring
o Triggered by decreasing daylength (short days)
 Tropics
o Day length variation minimal
o Breed year-round
o Temperate breeds lose seasonality over time
o BUT high temp & low feed suppress activity during dry months
o Activity increases after rainy season

Factors affecting breeding season

 Photoperiod (main)
 Breed (genotype)
o Near-equator breeds = longer sexual season
 Nutrition
Breed examples
 Sheep:
o Long season: Dorset, Merino, Rambouillet
o Short season: British breeds (Southdown, Shropshire, Hampshire)
 Goat:
o Dairy goats restricted (Aug–Feb)
o Nubian = least restricted

Hormonal Regulation
 Sheep seasonality controlled by:
o photoperiod → pineal gland → melatonin
o High melatonin during dark → stimulates reproduction
 LH feedback sensitivity changes with season

MALES
Rams
 Have sexual activity all year but highest in fall
 Short days:
o ↑ FSH
o ↑ LH
o ↑ testosterone
Bucks (male goats)
 Also seasonal
 Peak activity & testosterone in fall

PUBERTY
Females
 Does: 5–7 months
 Ewes: 6–9 months
 Late maturity: Merino, Angora (18–20 months)
 Early maturity: Pygmy goat, Finn sheep (3–4 months)
 Requires:
o 50–70% adult bodyweight
o Photoperiod exposure (long → short)
o Good nutrition
Puberty delayed by season even if endocrine system mature

Males
 Rams & bucks: 4–6 months
 First viable ejaculation at 40–60% adult weight
 Testosterone increases strongly at sexual maturity
  Sexual maturity depends more on body weight than age

ESTROUS CYCLE
Specie
Length
s
~17
Sheep
days
~21
Goat
days
 Short cycles early in the season (regressing CL)

DURATION OF ESTRUS
Specie Hour
s s
24–
Ewe
36
24–
Doe
48
 Shorter at beginning and end of season
 Angora goat shortest duration

ESTRUS SIGNS
Doe (more obvious)
 restless, bleating, tail-wagging, decreased appetite & milk
Ewe (less obvious)
 silent estrus common
 needs presence of ram
 mild vulvar edema, mucus discharge

MALE EFFECT
 Introduction of a male causes:
o ovulation in 3–6 days
o estrus in ~17–24 days
 Due to pheromones
 Stronger in goats than sheep

OVULATION
 Both spontaneous ovulators
 Ewe: 24–27 hrs after onset of estrus
 Doe: 24–36 hrs (Nubian later)
Ovulation rate
 Often >1 egg
 ↑ with age
 ↑ with nutrition
 ↑ early in breeding season
 Right ovary ovulates more (≈53%) than left ovary (46.6%)
 The ovulation rate is 1.2 for Merinos and 3 for the Finnish Landrace breed. In both species,
the ovula.. tion rate increases with age and reaches a maximum at 3 to 6 years, then
declines gradually.

BREEDING
 Mostly natural mating
 AI limited (low conception, frozen semen difficulty)
Mating ratio
  1 ram : 30 ewes
 1 buck : 50 does
 Breeding males may serve does as much as 20 times a day.
Copulation
 Small ejaculate volume
 Very high sperm concentration
Sperm storage
 Cervix stores sperm for ~3 days
 Sperm survive in uterus ~30 hrs
 Ova viable 10–25 hrs
 In both species, eggs enter the uterus about 72 hours after ovulation.
 Compaction at 8-cell
 Blastocyst implants at Day 6

EXAM MNEMONICS
Seasonal breeders
Short days → breeding
S = sheep
S = short days
S = sexual activity increases
Male effect
“Male = Melatonin + Pheromone”
Puberty requirements
B-P-N
 Bodyweight
 Photoperiod
 Nutrition

SHEEP vs GOAT QUICK DIFFERENCE

Feature Sheep Goat
1. Estrus duration  Shorter  longer
2. Estrus signs  silent  Obvious
 not  tail
3. Tail behavior
obvious wagging
4. Seasonal  less
 strong
breeding strong
5. Male effect  weaker  strong

GESTATION

Gestation length

 Average ~150 days in both species

 Variation:
o Sheep: shorter in early maturing & prolific breeds
o Variation within breed: ±13 days
o Goats: usually within ±2 days of 150 days
o Black Bengal goat = ~144 days

Factors affecting gestation length

Fetal genotype = MOST IMPORTANT

 Accounts for ~⅔ of variation in sheep

 Male lambs → longer
 Spring-born → longer than fall-born
 Singles → longer than twins
 Older dams → longer gestation

Source of progesterone

Stage Sheep Goat

1st trimester CL CL
Late gestation Placenta (main) CL (main)
 Sheep = placenta-dependent
 Goat = CL-dependent
o Ovariectomy ANY time → abortion!!

KEY EXAM STATEMENT

in goats, CL persists throughout pregnancy as the major source of progesterone

Hydrometra (“pseudopregnancy” in goats)

 fluid accumulation in uterus

 no fetus
 progesterone remains high
 diagnosed by ultrasound
 common in goats vs sheep

PARTURITION

Initiation

 fetus initiates parturition

 In goats must also have CL regression before birth

Presentation

 Most born anterior presentation:

o head + forelimbs first

Duration

 Variable
 prolonged with:
o oversized single fetus
o twins stuck
o abnormal presentation
Twin births

 faster than singles

 but time between twins = few minutes to >1 hour

Goats detect and care for twins better than sheep.

Neonatal behaviors

 Lamb stands within ~15 minutes

 Seeks udder within 1–2 hours

Maternal behavior

 vigorous licking & ingestion of membranes

 attracted to fetal fluids → “lamb stealing”
 critical attachment period = first few hours
o if removed at birth → rejected after 6–12 hrs

Poor maternal behavior associated with:

 primiparous ewes
 twins
 certain breeds (Merino, Angora)

Species difference in mother–young bond

 Sheep = follower species

o lamb follows mother from birth
 Goats = hider species
o kid hides for first few days
o mother seeks isolation at kidding

PUERPERIUM

Uterine involution

 Sheep = complete at ~27 days

 No data for goat

Post-partum ovarian activity

 seasonal effect is MAJOR determinant

 PP estrus can be 5–27 weeks depending on season

If lambing occurs during breeding season:

  rapid return to cyclicity
 first ovulation may occur ~20 days
 BUT silent ovulation (no estrus)

REPRODUCTIVE PERFORMANCE

Measures

 Conception rate ≈ 85%

 Lambing rate ≈ 150%
 Fecundity = number of lambs born per ewe
 Prolificacy = number of offspring per year

Fertility decreases with

 near equator
 heat stress
 undernutrition OR obesity
 young or old animals
 high-estrogen forage
 parasitism/disease/stress

Ovulation rate increased by:

 flushing
o works early or late breeding season
o not midseason
 crossbreeding with high-ovulation breeds:
o Booroola, Finn, Romanov
o may increase lamb crop 250–400%

ADVANCING BREEDING SEASON

Methods (ewe)

 progesterone + eCG
 ram effect
 artificial light (8L:16D)
 melatonin implants

Melatonin:

 implanted for 30–40 days

 advances season
 DOES NOT extend season if used mid-season
 animals can become refractory

Goats
  melatonin less effective
 prenatal photoperiod treatment → delays puberty & lowers testosterone (Saanen study)

ARTIFICIAL REGULATION

Increase frequency of breeding

 goal: 3 lambings in 2 years (8 months interval)

 or twice yearly (6 month interval)

Light/melatonin useful in sheep, limited in goat

Increase litter size

 flushing (most practical)

 exogenous gonadotropins (variable, embryo loss)
 steroid immunization (estrone, androstenedione)
o raises LH/FSH
o increases ovulation by ~0.6 eggs/ewe
o raises lamb output by ~30–40%

SUPER EXAM POINTS (memorize)

 Gestation sheep & goat = ~150 days

 Black Bengal goat = 144 days
 Sheep = placenta progesterone
 Goat = CL progesterone
 Ovariectomy in goat → abortion ANY TIME
 Hydrometra = pseudopregnancy in goats
 Sheep = follower; goats = hider
 Uterine involution sheep = 27 days
 Flushing ↑ ovulation
 Finn & Romanov → prolific
 Melatonin advances season but doesn’t extend
 PIGS

GENETIC SEX → GONADAL SEX → PHENOTYPE

 Genetic sex determines formation of testis or ovary
 Testis determines:
o male internal & external genitalia
o male body sex characteristics
o male behavior
 Porcine karyotype = 2n = 38 chromosomes

ORIGIN & FORMATION OF GONADS

Gonads originate from:
 coelomic epithelium
 mesenchyme
 primordial germ cells (extra-embryonic origin)
Gonadal ridge develops at:
 Day 21–22
Primordial germ cells:
 round/elongated
 diameter 10–20 μm
 divide by mitosis

SEX DETERMINATION
 Begins once gonad starts differentiating
 Y chromosome contains a testis-determining gene
 Testicular organogenesis = Sertoli + Leydig cells

AMH (MIS – Müllerian inhibiting substance)

Secreted by Sertoli cells:
 appears Day 27
 before Leydig cells (Day 30)
Function:
 regress Müllerian ducts in males
Molecular weight:
 MIS = dimer
 70,000 daltons

MALE EMBRYO DEVELOPMENT

Seminiferous cords form → become tubules
Cell differentiation:
 Sertoli cells = sustentacular
 Leydig cells = interstitial
Wolffian duct → epididymis, vas deferens, vesicles, prostate, bulbourethral glands

Testicular Development Timeline

Development Age
 21–24
 Primitive gonad
days
 Sertoli begin MIS  27 days
 Leydig appear  30 days
 Wolffian duct  26–50
differentiation days
Testicular descent begins 60 days
Testes enter inguinal canal 85 days
 Development Age
in scrotum shortly after birth

Testicular growth phases

 prenatal → slow
 birth–3 weeks → rapid (Leydig hyperplasia)
 after 3 weeks → regress
 7 weeks → seminiferous growth ↑
 near completion by 25 weeks
Testes weight:
 6 g (40 days) → 120 g (250 days)

HORMONES (MALE)
Fetal testis produces testosterone
 regulates internal/external genital development
Serum gonadotropins
 peak 1 week after birth (LH, FSH, prolactin)
 decrease by week 7
Estradiol produced in boar
 synergistic with testosterone
 increases sexual behavior
 stimulates accessory glands

Spermatogenesis
 Seminiferous cycle = 8.6 days
 One full spermatogenic cycle = 34.4 days (4 seminiferous cycles)
 Epididymal transit = 10.2 days
Puberty:
 first mating behavior = 4 months
 first ejaculation = 5–8 months
 heritability h² = 0.32

FEMALE — OVARY DEVELOPMENT

Germ cell number
Age Germ Cells
20 days
5,000
embryo
50 days 5,000,000
embryo (peak)
birth 400,000
oogenesis
fetal day 100
ends

Meiosis
 begins Day 40
 diplotene stage begins Day 50
 all diplotene by 20 days after birth
Oocyte size:
 oogonia = 13 μm
 primordial follicle = 27 μm
 Graafian follicle = 120 μm
Zona pellucida thickness = 8.6 μm

GILTS – PUBERTY & HORMONES

Differentiation to ovary ≈ Day 35
FSH cells detectable = 70 days fetus
LH pulses start = 10 weeks after birth
Tertiary follicles = 8 weeks
Puberty = 25 weeks average

Environmental influence
Factor Effect
puberty delayed >40
restricted feed 50%
days
puberty ↓ from 232 →
boar exposure
191 days
group housing >
earlier puberty
individual

Estrous Cycle (Pig)

 Length = 21 days (19–23)
 Estrus duration = 40–60 hrs
 Ovulation = 38–42 hrs after estrus onset
 Ovulation earlier in mated = 4 hrs
Pig = polyestrous all year

Ovarian Follicle Sizes

Type Size
1. Small  2–5 mm
2. Medium  6–8 mm
3. Pre-  8–11
ovulatory mm

Corpus Luteum
 corpus hemorrhagicum immediately
 luteinization complete by Day 4
 max size Day 6–8 (8–11 mm)
 max weight 350–450 mg
 regression Day 16→18
 becomes corpus albicans

Hormones per cycle

Progesterone:
 low at estrus
 peak Day 8–12
 decline Day 18
Estrogen:
 peaks 2 days BEFORE estrus
 low during luteal

FSH
 peaks Day 2–3 after estrus

IMPORTANT KEY NUMBERS

 Chromosomes = 38
 MIS appears day 27
 Leydig day 30
 Testes descent day 60
  Germ cells peak 5 million - day
50
 Oogenesis finished day 100
fetus
 Seminiferous cycle: 8.6 days
 Total spermatogenesis 34.4
days
 Estrus 40–60 hr
 Ovulation 38–42 hr
 Cycle = 21 days

OVULATION RATE
Factors affecting ovulation rate
 Breed / genetic lines / crosses
 Amount of inbreeding
 Age at breeding
 Weight at breeding
 Parity (reproductive experience)
Effect of inbreeding
 Inbred lines: +1.1 ova from 1st → 3rd
estrus
 Little or no increase after 4th estrus
 Inbreeding reduces ovulation rate
 Crossing increases ovulation rate
Increasing ovulations with parity
 Ovulations increase up to 7 or more litters
Heritability
 Ovulation rate H² ≈ 0.40

METHODS TO INCREASE OVULATION RATE

Hormonal (superovulation)
🟠 PMSG
 Dose-dependent increase
 Given Day 15–16 of cycle
 Induces superovulation
 Can shorten estrous cycle
 Increases estrus duration
 May increase cystic follicles
 Ova still normally fertilizable
🟠 hCG
 500 IU IM during proestrus
 Ovulation occurs 44–46 hours later
 Induces ovulation
 BUT does NOT significantly increase ovulation rate
Nutrition ("flushing")
 High energy diets increase OR
 Only when short duration feeding
 Effect mainly genetic, not nutritional
Energy example:
 Restrict 3,000–5,000 kcal → follow with
 High energy 8,000–10,000 kcal
 Optimum feeding duration = 11–14 days before estrus
Protein ↑ = little effect
Drug treatments
 Insulin + high energy diet → ↑ ovulation + ↓ atresia
  Altrenogest fed 14 days → slight increase in OR, different effect from flushing

🔶 CONCEPTION RATE
 Fertilization rate usually > 90%
 Very high fertility
 High or low ovulation rate has little effect on fertilization
 Entire litter loss may be fertilization failure or total embryo death

🔶 EMBRYO SURVIVAL
Total embryonic loss
 ≥40% embryos lost before farrowing
Time-specific losses
 0–18 days → 17% loss
 By Day 25 → 33%
 By Day 50 → up to 40%
Gilts vs Sows
 Sows have:
o higher ovulation
o BUT more embryo loss during first 40 days
Genetic effects
Every 10% inbreeding causes:
 0.5–0.8 fewer ova
 0.5 fewer fertilized ova
 0.8 fewer embryos by Day 25
Crossing:

o 0.5 ova

o 0.3 fertilized ova

o 0.8 embryos
Heritability of embryo survival = 0.30

🔶 LITTER SIZE
Measured by:
 live births
 pigs farrowed per year
 total weaning weight
Effect of parity
 Max litter size at parity 4–5
 After parity 8:
o live births decrease
o stillborns increase
After 4.5 years of age reproductive performance declines
Genetic effect
 Heritability H² ≈ 0.17
 Mostly environmental
Heterosis examples
Purebreeds:
 Duroc & Yorkshire = 13.8 CL
Crosses:
 Duroc sow × other breed boar → +1.4 piglets
Three-breed crosses > two-breed crosses in:
 day-30 survival
 farrowing rate
  weaning survival

PREGNANCY DEVELOPMENT
Fertilization
 Occurs in ampulla
 Movement assisted by cilia
Embryo stages
Stage Day
 4-cell  Enter uterus
 Morula  Day 5
 Blastocyst  Day 6–8
 Day 6 (~150
 Hatching
cells)
 Elongation begins  Day 11
 Completed
 Day 13
elongation
 Blastocyst may exceed ->60 cm length
 Conceptuses grow exponentially ->Days 9–18

🔶 FETAL GROWTH
From Day 20–100:
 weight increases from 0.06 g → 1000 g
Placental size correlates with fetal weight:
 Placental length (r=.64)
 Placental surface area (r=.72)
 Areolae surface (r=.65)

🔶 HORMONE CONTROL OF PREGNANCY

Corpus luteum (CL)
 Required throughout pregnancy
 Maximum size Day 8
 Declines after Day 114 after parturition
Progesterone:
 Peak around Day 12
 Declines gradually to
o 20–25 ng/ml near Day 104
 Drops rapidly at farrowing
Estrogen
 Increases at Day 30 and Day 112
 Produced mainly by placenta
 Fetal maturity stimulates estrogen increase
Relaxin
 Low first 90 days (<2 ng/ml)
 Peaks 2 days before farrowing
 Needed for cervical dilation, mammary development
Prolactin
 Luteotropic
 High late pregnancy & early lactation

🔶 PARTURITION CONTROL
Induced by:
 Placental estrogens ↑
 Fetal cortisol ↑
 CL fails → abortion
Induce abortion experimentally:
  PGF₂α from Day 23
 Dexamethasone
 RU486 (Days 111–112 → induces parturition in 31 hours)

🔶 POSTPARTUM ESTRUS
 may occur 1–3 days postpartum
 BUT conception fails because follicle immature

🔶 LACTATION & ANESTRUS

 Estrus rare during lactation
 Ovary inactive (lack of pituitary gonadotropins)
 Follicles shrink from 4mm → 2mm
 Recover to >5mm by Week 5 of lactation
 Uterus involutes 21–28 days
Hormonal state:
 Low LH and FSH
 High prolactin (stimulated by suckling)

🔶 WEANING
At 3–5 weeks postpartum:
 Weaning → follicle development → estrus + ovulation in 4–8 days
 LH increases
 Estradiol rises until preovulatory LH surge

🔥 MOST IMPORTANT EXAM NUMBERS (CHEAT LIST)

Value Importance
1. Ovulation heritability  0.40
2. Embryo survival
 0.30
heredity
3. Litter size heredity  0.17
4. Energy restriction  3000–5000 kcal
 8000–10000
5. High energy feeding
kcal
6. High energy duration  11–14 days
7. PMSG injection  Days 15–16
8. hCG dose  500 IU
9. Ovulation after hCG  44–46 hrs
[Link] loss by Day
 33%
25
[Link] loss by Day
 40%
50
[Link] length  >60 cm
 2 days pre-
[Link] peak
partum
[Link] → estrus
 4–8 days
interval

KEY TERMINOLOGIES

Ovulation rate
Number of ova released at each estrus.

Inbreeding
Breeding between genetically related animals that increases homozygosity and reduces
reproductive performance.

Crossing (outcrossing)
Mating unrelated or inbred lines to increase heterozygosity and improve fertility traits.

PMSG
Pregnant Mare Serum Gonadotropin
FSH-like hormone used to stimulate follicular development.

hCG
Human Chorionic Gonadotropin
LH-like hormone used to induce ovulation.

Superovulation
Abnormally high number of ovulations induced hormonally.

Proestrus
Stage before estrus characterized by follicular growth.

Cystic follicles
Follicles that fail to ovulate and become fluid-filled → can cause irregular estrus.

Flushing
Short-term feeding of high energy diet before mating to stimulate ovulation.

Atresia
Degeneration of follicles that fail to ovulate.

Altrenogest
Synthetic progestogen (17α-hydroxy-17-(2-propenyl)-estra-4,9,11-triene-3-one) given to delay
estrus or synchronize heat; sometimes increases ovulation rate.

Conception rate
Percentage of matings that result in fertilization.

Early embryonic death

Loss of embryos during the first days of pregnancy—conceptuses are reabsorbed.

Fetal mummification
Death of fetus after formation of bones resulting in dried, preserved fetus.

Heritability
Proportion of variation in a trait due to genetics.

Litter size
Total pigs born per farrowing including live and stillborn.

Ampulla (oviduct)
Upper part of oviduct where fertilization occurs.

Morula
Solid ball of cells formed after cleavage stage (≈ day 5).
Blastocyst
Fluid-filled embryo with inner cell mass (day 6-8).

Hatching
Escape of blastocyst from zona pellucida.

Zona pellucida
Glycoprotein shell around embryo.

Filamentous blastocyst
Elongated conceptus form (>60 cm) characteristic of pigs around day 11–13.

Conceptus
Embryo + associated membranes.

Maternal recognition of pregnancy

Signal by embryo to prevent luteolysis and maintain CL (estrogen in pigs).

Corpora lutea
Post-ovulation structures secreting progesterone and essential for pregnancy maintenance in
pigs.

Fetoplacental unit
Combined fetal + placental tissue responsible for steroid hormone production.

Relaxin
Hormone from corpora lutea causing cervical dilation and softening connective tissues for
parturition.

Progesterone
Hormone maintaining pregnancy and inhibiting uterine contractions.

Estradiol / Estrone
Estrogen hormones from fetoplacental unit increasing late in pregnancy.

Prostaglandin F2α (PGF2α)

Luteolytic hormone that induces luteal regression and can induce abortion.

Hysterectomy
Surgical removal of uterus.

Luteolytic
Causing regression of corpus luteum.

Postpartum estrus
Heat occurring 1–3 days after farrowing, usually not fertile.

Lactational anestrus
Period during lactation when estrus is suppressed due to high prolactin and low gonadotropins.

Sow at weaning
Physiological state after removal of litter—follicular development resumes and estrus occurs 4–8
days later.
 HORSE

TAXONOMY
 Species: Equus caballus
 Family: Equidae
 Order: Perissodactyla

Unique Reproductive Features of Horses

 Limited genetic selection for reproduction (vs cattle, swine, sheep)
 Major selection historically was ability to walk/run
 Natural breeding season affected strongly by photoperiod
 Horses used in racing are aged from January 1, so breeding tends to be early in the year
(to make foals older as 2-year-olds)

BREEDING SEASON
Stallion
 Semen can be collected all year
 BUT strong seasonal variation in:
o reaction time
o mounts per ejaculation
o gel-free semen volume
o total sperm number
o motility & agglutination
 Season affects seminal plasma more than sperm cells
 First ejaculate less affected by season than second

Mare
 Most variable reproductive cycle of all domestic animals
 Majority = seasonally polyestrous
 Behavioral estrus may occur Feb–Apr, BUT often no ovulation
 Best conception: May to July
 At equator, little seasonal variation
 Temperate areas = breeding in summer, anestrus in winter
Photoperiod
 Long days stimulate hypothalamic–pituitary–gonadal axis
 Artificial light advances season
 LH is photoperiodically entrained

Classification of mares by breeding pattern

1. Defined breeding season
o wild horses
o limited foaling season
2. Transitory breeding season
o estrus throughout year but ovulation only in breeding season
3. Year-round
o estrus and ovulation all year

Endocrine Manipulation to Advance Season

Used to advance breeding by 3–4 months
 PMSG
 hCG
 equine pituitary extract
 GnRH
 progesterone
 prostaglandins
REPRODUCTIVE PARAMETERS – STALLION
Testes
 Descent: 1–3 weeks after birth
 Begin rapid growth around 11 months
 Left testis develops earlier and grows faster

Semen Characteristics
 Ejaculate = 6–9 jets
 First jets contain ≥70% of sperm
 Gel fraction = secreted by seminal vesicles
 Gel = ~1/3 of ejaculate
 Gel DOES NOT affect motility or fertility
 Gel amount varies – not consistent stallion characteristic
 Sperm output, libido ↑ in spring & summer
 Testosterone also shows seasonal variation

Ejaculation Physiology
Two phases:
1. Emission
o transport into posterior urethra
o internal sphincter closes (prevents reflux into bladder)
2. Ejaculation
o rhythmic contractions of bulbocavernosus & perineal muscles
o high-pressure jets into uterus
o last jets deposited in vagina
Jet percentage
 Jets = 5–10 (average 8)
 First 3 jets contain 80% of sperm
 Later jets mostly mucous fraction

Sperm and Fertility Notes

 Hyperactivation required for zona penetration
 Motility defects = infertility
 zona drilling may allow fertilization in motility disorders
 Severe teratozoospermia treatable via micromanipulation
 Sperm antibodies in follicular fluid may prevent fertilization

ESTROUS CYCLE OF THE MARE

Average 21 days:
 14 days diestrus
 7 days estrus
Estrus characteristics:
 swollen vulva
 relaxed labia
 orange/wet mucosa
 clitoral “winking”
 Posture like urination

LH and FSH Patterns in Mare

 LH surge is prolonged and peaks day AFTER ovulation
 No sharp LH pre-ovulatory peak (unlike sheep)
 FSH surges every ~10 days
 Mid-diestrus FSH surge = prepares next ovulation
Ovulation and Corpus Luteum
 Most ovulations occur days 3–5 of estrus
 Ovulation ~24–48 hr before end of estrus
 Fertility peaks 2 days before estrus end
 CL reaches max size at day 14
 CL is ½–¾ size of ovulating follicle
CL persistence
 Often persists 2–3 months
 May be accompanied by follicular activity

Luteolysis
 Uterus causes luteolysis via systemic route
 Local uterine PGF2α effect minimal
 Systemic PGF2α effective beginning day 5 after ovulation
 PGF2α causes abortion
 Minimal effective systemic dose = 1.25 mg PGF2α

Inducing or Controlling Estrus

Saline infusion
 induces estrus in anestrus mares
 diestrus mares infused day 5–9 return to heat 4 days early
 prolonged diestrus → heat in 3–9 days
Prostaglandins
 induce luteolysis + estrus + ovulation
 artificial synchronization possible

OVA & OVIDUCT TRANSPORT

 At ovulation: ovum lacks corona radiata
 Surrounded by gelatinous mass that separates within 2 days
 Fertilized ova go to uterus
 Unfertilized ova trapped in oviduct for up to 7 months
 Embryo may “outrun” old ova
Species-unique:
 Embryo enclosed by acellular capsule (replaces zona pellucida)
 Fertilized ova reach uterus >144 hours
 Have more advanced cleavage stage than cattle
 50% transuterine migration

MUST-REMEMBER NUMBERS (for exam)

Parameter Number
 Length of estrous cycle  21 days
 Estrus duration  ~7 days
 Diestrus  ~14 days
 24–48 hrs before end of
 Ovulation timing
estrus
 First ejaculatory jets sperm
 70–80%
content
 Ovum arrival uterus  >144 hrs
 Testicular descent  1–3 weeks
 PGF2α minimal dose  1.25 mg
 CL maximal size  day 14
 CL persistence  2–3 months
 Unfertilized eggs retained  up to 7 months
GESTATION – MARE
Gestation Length
 Normal = 315–360 days
 Influenced by:
o Maternal size
o Fetal genotype
o Season of breeding
 Endometrial degeneration (cystic fibrosis, glandular atrophy) → placental insufficiency
→ fetal growth retardation → prolonged gestation

Pregnancy Diagnosis
Methods and Accuracy

Method When accurate Notes

 40–50  Detect uterus
a. Rectal palpation
days and ovaries
 Immature mouse
b. Mouse bioassay
 ≥35 days ovarian
(PMSG)
stimulation
c. Hemagglutination-  35–40  MIP commercial
inhibition test (PMSG) days test
 early
 Up to 15% false
d. Ultrasound pregnanc
negatives
y

Twinning
 Rare
 Mare almost never carries twins to term
 Most are dizygotic
 Monozygotic twins not reported in equids
 In 2,673 Thoroughbred mares → incidence 2%
 High abortion rate due to:
o Placental competition
o Placental insufficiency
 Outcomes:
o Usually one fetus dies mid-gestation → initiates abortion
o Stillbirth common
o Only ~50% of delivered twins survive
 Twins are usually aborted purposely early and mare remated
 Identical blood groups in surviving pairs → chimerism

Ovarian Function – 4 Stages

1. Early pregnancy
 One CL verum
 Previously thought to regress at day 40
 Now known persists > day 40
2. Day 40–150
 10–15 follicles may luteinize → accessory corpora lutea
 Usually 3–5 per ovary
 Some follicles ovulate but ova remain unfertilized
3. Month 5–7
 Both primary and accessory CL regress
 Mare remains non-estrus due to placental progesterone
4. Month 7 onwards
  Only vestiges present
 Near term → follicular activity resumes → postpartum estrus preparation

Placenta
 Type: Diffuse, microcotyledonary, epitheliochorial
 Microcotyledons formed by 5th month
 Chorionic villi fit into maternal crypts
 Close fetal–maternal apposition → microvillous junction

Uterine Defense After Breeding

 Breeding causes transient inflammation
 Sperm = faster inflammation than bacteria
 Normal mares clear bacteria quickly
 Persistent infection due to:
o Poor cervical dilation
o Weak myometrial contraction
o Poor lymphatic drainage
 Frozen/centrifuged semen → more inflammation

Placental Barrier Mechanisms

Movement controlled by:
1. Diffusion factors (permeability, area, distance)
2. Blood flow, shunts, exchange areas
3. Special transport mechanisms (Hb O₂ affinity differences fetal > maternal)

Endocrine Profile
Progesterone
 2 progesterone peaks:
o 3rd month = high PMSG → secondary CL
o 11th month = placental progesterone
 CL progesterone correlates with luteal morphology
PMSG
 Produced by endometrial cups
 Present day 40–130
 Stimulates accessory CL formation
 Accessory CL degenerate by day 180
 Regulated by season, maternal size, parity, fetal genotype
Fetal genotype effects
 Mare bred to donkey → PMSG only 10% of normal
 Due to fetal allantochorion influence

Foaling (Parturition)
 Mostly between 1900–0700 hours
 Peak 2200–2300
 Signs:
o Mammary hypertrophy
o Waxing of teats
o Milk leakage
o Patchy sweating (flanks, elbow)

Induced Parturition
 Hormones used: Estrogen, PGF2α, Oxytocin
 Large breeds: 100 mg dexamethasone daily × 4 days
 Indications:
o Delayed parturition
 o Uterine atony
o Prolonged gestation
o Pre-parturient colic
o Injury
o Obtain colostrum-deprived foals

Postpartum Estrus (“Foaling Heat”)

 5–15 days after foaling
 Can occur up to 45 days
 Breeding at this time:
o ↑ abortion
o ↑ dystocia
o ↑ stillbirth
o ↑ retained placenta
 Due to contamination before uterine involution complete

Equine Hybrids
Cross Result
Horse ♂ × Donkey
= Hinny
♀
Donkey ♂ × Horse
= Mule
♀
=
Zebra × Donkey ♀
Zebronkey
=
Zebra × Horse ♀
Zebrorse
 Hybrids infertile
 Cause:
o Unequal chromosome sets
o Failure of chromosome pairing
o Block in meiosis
o Mitotic division continues normally

IMPORTANT TERMS
 Gestation: 315–360 d
 PMSG: 40–130 d
 Accessory CL: 40–150 d
 PLACENTA: diffuse – microcotyledonary – epitheliochorial
 Twins = 2% Thoroughbreds
 Accessory CL degenerate: ~180 d
 Foaling peak: 2200–2300
 Foaling heat: 5–15 d

Conception & Foaling

 Conception and foaling % vary widely
 Some pony studs = 100% conception & foaling
 Most studs → foaling rate is LOWER than conception rate
 Conception rate affected by:
o breed
o nutrition
o age
o management
 Thoroughbreds = lower conception rate
  “Foal heat breeding” → lower pregnancy rate than breeding after first post-partum
estrus

Estrous Irregularities
Caused by:
 seasonal photoperiod
 nutrition
 climate
Common patterns:
 variable cycle length
 irregular estrous behavior
 failure of ovulation
 failure of follicular development
 spontaneous prolonged CL
Named types:
 quiet ovulation
 anovulatory estrus
 split estrus
 prolonged estrus

Anestrus – 3 types
1. Winter anestrus
 lasts 11 weeks
 follicles rarely reach 35 mm
 progesterone ~1 ng/mL
 regress → no ovulation
2. Persistent CL
 lasts 5–13 weeks
 high progestogen
 usually in November–December
3. Cyclic ovulation w/out estrus
→ ovarian cycles occur but no behavioral estrus

Hormonal induction of cycles

 Single GnRH injection → FSH ↑ 3.7-fold
 LH increase small compared to cyclic LH peak
 GnRH + progesterone near end of non-breeding season can induce normal
ovulation

Prenatal Mortality
Higher in:
 lactating mares early season
 breeding after foal heat
 some horse families
Yearlings:
 higher prenatal mortality due to:
o low progesterone
o immaturity
o high nutritional requirement
o husbandry stress

Excessive Umbilical Cord Length

Not related to:
 gestational age
 body weight
  sex
 dam parity
Associated with:
 weight of allantochorion
 length of non-pregnant horn
Pathology results:
1. strangulation of fetus
2. excessive twisting → vascular obstruction / urinary retention
3. necrosis of chorioallantois at cervix
Findings:
 grooves around fetus
 edema
 urachal dilation
 failure to close bladder apex
 risk of abortion

Abortion in Mares
 Average abortion rate = 10%
 Lowest between 3–6 years of age
 Infection abortions → late pregnancy
 High abortion susceptibility at:
o 5th month
o 10th month
→ due to endocrine vulnerability
Post-abortion conception rate LOW
Older mares more affected
Cause in older mares:
 uterine inadequacy
 endometrial gland loss
 collagen deposition
 chronic inflammation

Neonatal Issues
Microphthalmos (“button eye”)
 congenital
 common in Thoroughbreds
 variable severity
 corneal distortion
 bilateral or unilateral
Entropion
 congenital + hereditary
 causes corneal opacity/ulcer

Barker Syndrome
(Convulsive Foal Syndrome / Neonatal Maladjustment Syndrome)
 occurs hours after birth
 esp in easy deliveries
 clinical signs:
o jerking movements
o hyperexcitability
o convulsions
o opisthotonus
o barking respiration
 50% recover
Pathology
  necrosis of cerebral cortex, diencephalon, brainstem
 hemorrhage in cortex & cerebellum

Ocular Changes
 asymmetrical pupils
 retinal petechiae present day 1–2
 resolve within few days

Perinatology Features (Important!)

Equine pregnancy unique:
 fetus matures very late
 maternal progestins rise near term
 fetal gonads enlarge → produce estrogen precursors
 fetal adrenal & thyroid axis crucial for maturation
 abnormal maternal progestins/estrogens signal fetal compromise
 hormones regulate parturition

Neonatal Mortality
Causes
 weak dam/foal
 umbilical infection
Prevention
 clean foaling area
 sanitation

Breeding Management
 estrus detection
 vaginal exam
 rectal palpation
 predict ovulation
 conception best when insemination timed properly
After mating:
 mare may strain & expel semen
 prevent by walking mare after breeding
Mare prone to endometritis postpartum because cervix poorly closes
LH deficiency:
 treat with exogenous LH
 hCG 1500–3000 IU IV induces ovulation if follicle ≥ 3 cm

Assisted Reproduction
Useful:
 AI
 MOET
 in-vitro embryo production
Limited value:
 semen sexing
 embryo sexing
 cloning (only one-time genetic boost)

ICSI (Intracytoplasmic Sperm Injection)

 micromanipulation technique
 works WITHOUT exogenous oocyte activation
 normal fetal development possible
  sex-sorted sperm can be used
 zona receptors:
o ZP3 = induces acrosome reaction
o ZP2 = inner binding
 acrosome organizes plasma membrane proteins
 necessary for fertilization

Stallions – Reproductive Failure

Types:
 abnormal sexual behavior
 ejaculatory dysfunction
 poor semen quality

Abnormal Sexual Behavior

 failure to maintain erection
 incomplete intromission
 pain during copulation
 poor libido
 failure to ejaculate
 masturbation
Treatment: behavioral retraining

Ejaculatory Disturbances
Signs:
 absent ejaculation despite erection
 may need several mount attempts
 count urethral waves (normal ~5)
Causes:
 neural inhibition
 fatigue
 stress
 exhaustion
 failure of smooth muscle contraction
 norepinephrine depletion

Poor Semen Characteristics

Terms:
 azoospermia = no sperm
 oligozoospermia = low sperm concentration
 asthenospermia = low motility
 teratospermia = abnormal morphology
 hemospermia = blood in semen
Hemospermia:
 urethritis
 painful ejaculation
Semen microflora commonly include:
 Pseudomonas, E. coli, Klebsiella, Proteus, Staph, Strep
→ usually do NOT affect fertility

Management Note
Frequent ejaculation ↓ sperm per ejaculate → ↓ fertility
Artificial insemination can prevent overuse of stallion
 MALE POULTRY

Definition of male fertility

Male fertility depends not only on sperm production, but on the successful completion of
multiple sequential reproductive steps:
Required steps (memorize!)
a. sperm production & maturation in testes
b. initiation of sperm motility at ejaculation
c. storage in sperm storage tubules (SST)
d. passive transport through oviduct (above vaginal sphincter)
e. acrosome reaction at perivitelline layer
f. perforation of perivitelline layer
g. delivery of condensed DNA into oocyte
h. male pronucleus fuses with female pronucleus
➡ therefore fertility ≠ just sperm count
➡ sperm function and transport are equally critical

Sperm motility characteristics

 poultry sperm are immotile before ejaculation
 become motile only after ejaculation (major difference from mammals)

significance:
motility determines:
 ability to ascend vagina
 ability to enter SST
 duration of fertility in hens

Sperm Storage Tubules (SST)

 located in female oviduct (uterovaginal junction)
 store viable sperm for multiple days
Different species = different duration:
Days
Species
stored
Japanese
longest
quail
Broiler intermedia
breeder te
White
moderate
Leghorn
Turkey long
(Duration influenced by sperm physiology and SST function)

Why do sperm LEAVE SST?

Hypothesis:
 sperm enter SST with acrosome toward blind end
 require ATP to maintain position
 if ATP ↓ → sperm cannot hold position → drift out (fluid current)
 sperm then re-enter oviduct

🔹 Male reproductive tract anatomy

Structures paired, lie on dorsal body wall.
Components:
 Testes (attached by mesorchium)
 Seminiferous tubules
 Rete testis
 Efferent ducts
 Epididymal duct
 Deferent duct → storage + ejaculation
Important: the deferent duct (not epididymis) stores semen prior to ejaculation
→ storage area is called receptaculum

Accessory reproductive structures

Paracloacal vascular body
 produces lymph
 causes tumescent tissue (not blood)
 enables eversion of non-intromittent phallus
 ejaculation occurs during phallus eversion

SPERMATOGENESIS (Galliform birds)

Key points:
 occurs at body temperature 40–41°C (not 32–35°C like mammals)
 circadian fluctuation: 1–1.5°C
 most studied in Japanese quail
Duration (IMPORTANT number!)
 one seminiferous cycle = 2.69 days
 full spermatogenesis = 12.8 days
  produces 32 sperm from each spermatogonial stem cell

Sperm production yield

 quail: ≈92.5 × 10⁶ sperm / gram testis / day
 rooster: 80–120 × 10⁶ sperm / gram / day

Cytoplasmic reduction
 ≈ 97% cytoplasm lost when spermatids become sperm
 results in vermiform shape

Extragonadal sperm maturation and transport

 occurs in excurrent ducts
 sperm released immotile
 become motile during transit
 90% stored in deferent duct
Mechanisms uncertain:
 likely hormonal (testosterone)
 possibly estrogen dependent
 may involve reproductive tract proteins

Fertilization Features (VERY UNIQUE TO POULTRY)

Fertilization site:
 infundibulum
Two major differences from mammals
1️⃣sperm do NOT undergo capacitation
2️⃣polyspermy = normal

Acrosome interaction
 perivitelline layer induces acrosome reaction
 sperm perforate perivitelline at multiple sites
Sperm surface glycoproteins and perivitelline receptors determine:
 binding
 fertilization success
 male fertility differences among chicken lines

Key Mechanistic Concepts to Memorize

✔ sperm immotile pre-ejaculation
✔ SST storage
✔ sperm efflux depends on ATP
✔ spermatogenesis at 40–41°C
✔ polyspermy = normal
✔ fertilization in infundibulum
✔ deferent duct stores sperm (not epididymis)
✔ phallus is lymphatic, non-intromittent

HIGH-YIELD NUMBERS & TERMS

Term Value
Temperature
40–41°C
spermatogenesis
Circadian temp +1–1.5°C
Cycle stage duration 2.69 days
Total spermatogenesis 12.8 days
Spermatogenic efficiency 80–120M
Term Value
sperm/g/day
Stored in deferent duct >90%
Cytoplasm reduced 97%
Sperm length 75–90 µm
Sperm width 0.6 µm

FEMALE POULTRY

Why poultry reproduction is unique

Domestic poultry reproduction is different from mammals because:
 Must ovulate single ova frequently
 Produces an egg that contains all nutrients for development
 No gestation—embryo develops outside the body

Reproductive Efficiency
 Poultry have short generation interval
 High reproductive capacity
 High egg and meat output due to:
o genetic selection
o controlled breeding
o improved nutrition
o artificial incubation

Composition of the Egg

Components
1. Yolk
2. Albumen
3. Shell

Yolk (Ovum)
  ⅓ of egg mass
 Contains female pronucleus
 Major nutrients (lipids + proteins)
 Contains maternal immunoglobulin G
 Blastodisc = site with maternal chromosomes
 After fertilization becomes blastoderm
White yolk
 protein rich, surrounds blastodisc
 Includes:
o Nucleus of Pander
o Latibra

Albumen (Egg white)

 58% of egg
 40+ unique proteins
 Functions:
o prevents dehydration
o water source
o antibacterial proteins
o mineral source

Shell
 composed of:
o shell membranes
o calcium carbonate layer
o cuticle
 Functions:
o protection
o gas exchange
o prevents water loss
o calcium source for embryo
Cuticle
 protects vs bacteria
 seals pores

Female Reproductive System

Ovary
 Only left ovary functional
 Produces:
o ova
o steroids
o estrogen
 Contains:
o medulla (blood supply)
o cortex (oogonia/oocytes)

Follicular development
 Thousands of follicles exist, only few mature
 Follicular hierarchy = follicles arranged by maturity
 High estrogen = liver produces yolk precursors
 Broilers have more follicles → double yolks → lower fertility
 Turkeys grow follicles slower → lower egg rate

Oviduct (5 parts)
1. Infundibulum
 o receives ovum
o fertilization site
o chalaza formation starts
o sperm storage site
2. Magnum
o secretes albumen (egg white)
3. Isthmus
o forms shell membrane
4. Shell gland (Uterus)
o shell formed
o calcium deposition
o pigment + cuticle added
5. Vagina
o expels egg
o sperm storage
o sperm selection

Egg Transport & Oviposition

Total time in oviduct ≈ 26 hours

 Infundibulum – 15 mins
 Magnum – 2–3 hrs
 Isthmus – 1–1.5 hrs
 Shell gland – 20 hrs
Controlled by:
 vasotocin
 prostaglandins
 neural reflex

Sperm Storage & Fertilization

 Sperm stored up to:
o 32 days (chicken)
o 70 days (turkey)
 Storage location:
o uterovaginal junction
o infundibulum
 Polyspermy common, but only one sperm fertilizes

Photoperiod Control
Egg laying triggered by increasing daylength
 Chickens start at 18–20 weeks
 Turkeys at 28–30 weeks
Hormones:
 LH
 FSH
 Estrogen increases yolk production

Important Terminologies

Blastodisc – circular disc on yolk containing female pronucleus

Blastoderm – expanded blastodisc in fertilized egg
Chalazae – twisted protein strands that suspend yolk
Cuticle – protein coating sealing shell pores
Medullary bone – temporary calcium storage for shell formation
Sperm storage tubules – structures storing sperm long-term
Photoperiod – amount of daylight controlling reproduction
Oviposition – laying of the egg
Infundibulum – site of fertilization
Magnum – albumen secretion region
Isthmus – shell membrane formation
Shell gland (uterus) – calcification + pigmentation
Vagina – sperm transport + egg passage
Polyspermy – multiple sperm penetrate perivitelline layer
Aromatase enzyme – converts androgens to estrogen
Anti-Müllerian hormone / MIS – causes regression of ducts
Nucleus of Pander – white yolk area under blastodisc
Latibra – white yolk core

REPRODUCTIVE FAILURE IN FEMALES

Reproductive failure: inability to produce viable offspring within a species-specific timeframe.

 Sterility – permanent inability to reproduce
 Infertility – temporary inability to produce viable young

1. OVARIAN DYSFUNCTION
Ovary functions:
 1. Production of ova
2. Secretion of ovarian hormones (estrogen, progesterone)
Abnormalities can affect estrus, ovulation, and uterine function.

A. Anestrus – absence of estrus

Physiologic causes:
 Prepuberty
 Pregnancy
 Lactation
 Seasonal breeders
Pathologic causes:
 Nutritional deficiency (energy/protein imbalance)
 Stress (lactation, heat)
 Aging
 Ovarian/uterine disease

B. Ovarian Abnormalities
1. Ovarian hypoplasia
 Underdeveloped ovary → never exhibits estrus
 Species: Swedish Mountain cattle, some mares
 May involve XO chromosomes, low estrogen, high LH
 Freemartins: heifer twins of bulls → poorly developed ovaries
2. Persistent corpus luteum (CL)
 Associated with uterine pathology: pyometra, mucometra, fetal mummification/maceration
 Pseudopregnancy: functional CL + uterine fluid accumulation
o Species: sow, mare, doe
o Diagnosis: elevated progesterone, ultrasound (no placentomes)
o Treatment: PGF₂α, oxytocin, progesterone devices
 Prolonged diestrus in mares → failure of PGF₂α release

C. Ovulatory Failure
 Failure of preovulatory follicle to rupture → anovulatory estrus
 Common in swine and horses
 Can lead to cystic ovarian disease (COD)
Cystic Ovarian Disease
 Dairy cattle > high-producing cows; also swine
 Follicular cyst: thin-walled, estrogenic → nymphomania or silent heat
 Luteal cyst: thick-walled, progesterone-producing → persistent anestrus
 Causes: LH release failure, postpartum uterine infection, high milk yield, seasonal
changes, heredity, pituitary dysfunction
 Diagnosis: transrectal ultrasonography
 Treatment:
1. GnRH / hCG (follicular cysts)
2. PGF₂α (luteal cysts)
3. Progesterone injections / intravaginal devices
4. Manual rupture (older method)

2. DISORDERS OF FERTILIZATION
A. Fertilization Failure
Causes:
1. Abnormal eggs – giant, oval, lentil-shaped, ruptured zona pellucida; environmental
stress (heat)
2. Abnormal sperm – acrosomal defects, DNA abnormalities, aging → reduced fertilization
3. Structural barriers – congenital (Müllerian duct defects, white heifer disease), acquired
adhesions, missing segments
 4. Phytoestrogens – e.g., red/subterranean clover → reduced ovarian function, impaired
sperm transport
5. Ovulatory failure – no ovulation → no fertilization
B. Atypical Fertilization
 Egg aging → loss of viability, polyspermy
 Polyspermy – entry of >1 sperm, leading to triploid embryos → nonviable
 Critical in horses and swine: timing of insemination relative to ovulation

3. PREGNANCY WASTAGE
Types:
1. Embryonic mortality – death before implantation
2. Fetal mortality – death after implantation
Embryonic Mortality
 Normal elimination of unfit genotypes
 25–40% in farm species
 Most deaths in early embryonic period
o Cow: days 8–16
o Pig: days 8–18 (≥4 embryos required by day 10 to maintain pregnancy)
 Causes: maternal factors, embryonic factors, maternal-embryo interactions
Maternal factors:
 Nutrition, infection, endocrine imbalance, uterine environment
Embryonic factors:
 Chromosomal abnormalities, genetic defects, gamete aging
Detection:
1. Fertilization rate / embryo recovery
2. Ultrasonography (≥20 days)
3. Progesterone measurement
4. Laparoscopy in sheep

Critical points
 Early embryonic death → often unnoticed, may return to normal estrus
 Late embryonic death → cycle length may be extended
 Fetal loss → recognizable by abortion, mummification, or stillbirth

4. HIGH-YIELD FACTS
Topic Key Points
Sterility vs
 Sterility = permanent; Infertility = temporary
Infertility
Ovarian hypoplasia  Swedish Mountain cattle, Freemartins
Persistent CL  Pseudopregnancy; treat with PGF₂α / oxytocin
Cystic Ovarian  Follicular vs luteal cyst; high milk yield; post-partum uterine
Disease infection
Fertilization failure  Egg/sperm abnormalities, structural barriers, phytoestrogens
Embryonic  Cow: 8–16 days; Pig: 8–18 days; ≥4 embryos in pigs for
mortality pregnancy maintenance
Polyspermy  Delayed insemination → triploid embryos → nonviable

LACTATION
 Embryonic mortality can occur during lactation in cattle, sheep, and horses.
 Characterized by prolonged estrous cycles.
 Causes:
o Stress of lactation
o Reduced uterine defense mechanisms
 o Incomplete endometrial regeneration
 Sows bred 7 days after weaning show high embryonic losses between days 9–20 of
pregnancy.

NUTRITION OF THE DAM

 Nutrition affects ovulation, fertilization, and embryo survival.
 High caloric or protein intake may increase embryonic mortality before implantation
(dairy cows, swine, sheep).
 Poor body condition increases embryo loss; moderate feed restriction may be less
harmful.
 Critical periods:
o Mare: 25–31 days post-ovulation
o Dairy cow: high rumen-degradable protein → lower uterine pH → embryo death

AGE OF DAM
 Young (gilts, ewe lambs) or old females (>5 years in swine, >6 years in sheep) show
higher embryonic mortality.
 Cause mostly embryo-associated rather than uterine.

OVERCROWDING IN UTERO
 High number of implantations → reduced vascular supply per embryo → increased
mortality.
 Twin/multiple ovulations: survival typically limited to 2–3 embryos in cattle/sheep.
 Transuterine migration (swine) ensures equal embryo distribution; its absence
increases mortality.

THERMAL STRESS
 Heat stress during early pregnancy increases embryonic mortality:
o Critical periods: Sheep/cattle: implantation; Pig: 8–16 days postbreeding.
 Heat alters uterine environment and conceptus development.
 Mitigation: environmental cooling during critical gestation periods.

SEMINAL FACTORS
 Male genetics, sperm quality, and semen storage affect embryonic survival.
 Aged sperm → reduced DNA integrity → early embryo death.

INCOMPATIBILITY
 Immunologic/genetic incompatibility between:
o Sperm & mother
o Sperm & egg
o Zygote & mother
 Example: Certain bovine blood group homozygosity → increased embryonic loss.

REPEAT BREEDERS
 Females failing to conceive after >3 services despite normal estrus.
 Early embryonic losses often occur at oviduct/blastocyst stage.
 Higher incidence in AI herds and older cows.
 Causes: uterine environment, chromosomal abnormalities, infections (e.g., contagious
equine metritis).

FETAL MORTALITY
ABORTION
 Expulsion of fetus before viability:
o Cattle: <260 days
 oHorses: <290 days
oSwine: <110 days
 Causes:
o Noninfectious: genetics, chromosomal, hormonal, nutritional, twin pregnancy
o Infectious: bacteria, viruses, protozoa (Tables 17…5–8)
 Examples: Neospora caninum (cattle), EAE (sheep), equine herpesvirus I
(horses), SMEDI viruses (swine)
FETAL MUMMIFICATION
 Fetus dies but is retained → dehydration & resorption of fluids.
 Common in cattle & swine; less in sheep & horses.
 Causes persistence of corpus luteum, preventing abortion.
 Swine: mummified fetuses often occur in large litters, older sows, susceptible
breeds.

PERINATAL MORTALITY
 Definition: Death of offspring shortly before, during, or within 48–72 hours after birth.
 Includes stillbirths (born dead).
 Incidence:
o Cattle & horses: 5–15%
o Sheep & pigs: 20–30%
 Major causes:
o Asphyxia, starvation, chilling, congenital malformations
o Species differences:
 Cattle: trauma during prolonged/assisted birth
 Pigs: asphyxia due to premature umbilical rupture
 Induced parturition (e.g., PGF₂α in sows, corticosteroids in cows) can increase mortality.
 Swine specifics:
o Two types of stillbirth:
1. Prepartum (usually infectious)
2. Intrapartum (noninfectious, associated with meconium staining)
o Risk factors: uterine inertia, prolonged farrowing, extreme litter sizes, piglets in last
third of litter

NEONATAL MORTALITY
 Definition: Death of neonate during first few weeks of life.
 Causes: Heredity, environment, nutrition, infection.
 Key factors:
o Respiratory Distress Syndrome (RDS): Surfactant deficiency → unstable alveoli
→ death
o Poor maternal nutrition, weak dam, prolonged labor, delayed lactation
o Temperature extremes → hypothermia or heat stress
o Predation

DISORDERS OF GESTATION, PARTURITION, AND PUERPERIUM

DYSTOCIA (difficult birth)
1. Fetal dystocia: abnormal fetal presentation, oversized fetus, or monstrosities
o Common in dairy cattle, multiple pregnancies, sows with small litters
o Examples: head flexion, breech, twins
2. Maternal dystocia: inadequate uterine contractions (primary or secondary inertia)
o Primary: over-stretching from multiples
o Secondary: exhaustion due to obstructive dystocia
o Cervical spasm may occur
 3. Fetopelvic disproportion: mismatch between fetal size and maternal pelvis
o Common in cows, ewes (single lamb), sows (small litters)
o Prevention: planned mating, weight-based heifer mating, select bulls for smaller
calves

RETENTION OF FETAL MEMBRANES (RFM)

 Failure to expel fetal membranes within 12 hours (pathologic in cattle)
 Causes: uterine inertia, placental inflammation, infection, twins, induced parturition,
dystocia
 Consequences: metritis, endometritis, delayed uterine involution → reduced fertility
 Management:
1. Uncomplicated: usually no treatment
2. Manual removal: risk of uterine damage
3. Ecbolics: limited effectiveness; best within 1 hour of parturition
4. Antibiotics, hormones: inconsistent results
 Less common in buffalo, sheep, goats; mare: treated with oxytocin, manual removal if
>24h

HYDRAMNIOS & HYDRALLANTOIS

 Hydramnios: Excess amniotic fluid → often fetal cranial abnormalities → impaired
swallowing
 Hydrallantois: Excess allantoic fluid → massive abdominal enlargement, often twin
pregnancies → placental dysfunction
 Species: Mostly cattle; also reported in horses

MULTIPLE PREGNANCY
 Higher incidence of fetal loss than multiple births due to abortion/resorption
 Cow sequelae of twinning: shortened gestation, abortion, stillbirth, dystocia, RFM
 Economic impact: decreased fertility, neonatal mortality, lower birth weight, longer
calving interval, lower milk fat, freemartins (90% of females cotwin to male are sterile)
 Sheep: twins → higher neonatal mortality & twin-lamb disease
 Mares: high abortion rate for twin fetuses

PROLONGED GESTATION
 Causes: genetic & non-genetic factors
1. Fetal gigantism (Holstein, Ayrshire)
o Gestation prolonged 3 weeks–3 months
o Large calves → cesarean; die 6–8h postpartum due to hypoglycemia
o Adenohypophysis & adrenal cortex hypoplasia
2. Craniofacial abnormality (Guernsey, Jersey)
o Small fetuses, facial abnormalities, hydramnios, no adenohypophysis
o Survive only a few minutes post-delivery

UTERINE INFECTIONS
 Common postpartum sequelae of RFM & dystocia
 Endometritis: inflammation of endometrium
 Metritis: full-thickness uterine infection
 Pyometra: purulent uterine accumulation
 Common bacteria: Actinomyces pyogenes
 Infection delays cyclicity → suppresses follicular growth → reduced fertility
 Ovarian activity critical:
o Estrogenic phase → resistance
o Progesterone phase → susceptibility
 Avoid early GnRH to induce cyclicity in postpartum cows → risk of pyometra
 REPRODUCTIVE FAILURE OF FEMALE

Fertility of a male depends on:

1. Sperm production
2. Viability & fertilizing ability of sperm
3. Sexual desire (libido)
4. Ability to mate
Consequences:
 Sterile males → obvious
 Subfertile males → economic losses, especially in AI programs
Major causes:
 Gonadotropin deficiency, chromosomal/genetic disorders
 Excurrent duct obstruction
 Systemic or genital disease
 Environmental toxins, stress (heat, poor housing, high density, social changes)
 Neurologic or autoimmune disorders
Key focus areas for evaluation:
 Endocrine function
 Semen quality and sperm morphology
 Ejaculatory behavior
 Biochemical markers
 Therapeutic interventions

CONGENITAL MALFORMATIONS
Segmental Aplasia of Wolffian Ducts
 Missing segments of epididymis, vas deferens, or ampulla
 Unilateral: often fertile
 Bilateral: sterile
 May cause sperm accumulation (spermatocele)

Cryptorchidism
  Failure of one/both testes to descend into scrotum
 Higher incidence: swine, horses
 Hereditary: dominant in horses, recessive in other species
 Bilateral: sterile (spermatogenesis suppressed)
 Unilateral: fertile but reduced sperm count; testosterone mostly normal
 Should not be used for breeding
Testicular Hypoplasia
 Underdeveloped testes → reduced or absent spermatogenesis
 Genetic basis: recessive autosomal (e.g., Swedish Highland cattle)
 Signs: small testes, low sperm count, watery semen, abnormal sperm
 Diagnosis: scrotal measurements, semen evaluation, karyotyping

EJACULATORY DISTURBANCES
1. Lack of Libido
 Causes: hereditary, endocrine, psychogenic, environmental
 Species examples:
o Bulls: influenced by genetics, stress, management; good semen may be collected
via electroejaculation, but low-libido bulls should not be used in AI
o Stallions: overuse, pain, mismanagement, seasonal variations affect libido
o Rams: low service frequency due to poor libido, dexterity, or competition;
influenced by light and temperature
o Boars: obesity, heat stress, high nutrition, mismanagement
2. Inability to Copulate
 Failure to mount: hindlimb/joint problems, locomotor issues, arthritis, vertebral
exostoses
 Failure to achieve intromission: preputial/penile deformities, phimosis, hematomas,
congenital penile deviations, tumors, persistent frenulum
 Failure to ejaculate: may occur even with normal intromission; causes include nerve
dysfunction, poor AI techniques, stress, exhaustion, obesity

FERTILIZATION FAILURE
 Occurs despite normal libido and mating ability
 Causes: defective semen, abnormal sperm function, errors in breeding techniques

DISEASES OF TESTES AND ACCESSORY GLANDS

 Pathologies in testes, epididymis, seminal vesicles affect:
o Spermatogenesis
o Sperm maturation
o Semen quality
o Passage of sperm to urethra

ENVIRONMENTAL FACTORS
Heat Stress
 Elevated body temperature → testicular degeneration → reduced % of normal sperm
 Seasonal effects on semen quality and fertility:
o Bulls: semen quality decreases in hot months
o Rams: fertility may drop during hot months; conception failure often due to
fertilization failure
o Boars: semen volume higher in cool weather; heat exposure (≥35°C) → ↓ sperm
concentration & motility, ↑ abnormalities
 Epididymis in boars is highly sensitive to temperature → cytoplasmic droplets in sperm
 Infectious febrile diseases can reduce fertility for 5–6 weeks

BREEDING TECHNIQUES & MANAGEMENT

  Fertilization failure may result from: poor AI technique, stress, estrus synchronization,
ingestion of estrogenic pasture
 Natural mating considerations: species, age, libido, fertility, nutrition, mating season,
management system, pasture size
 Frequent ejaculations → temporary reduction in semen quality and libido
 Seasonal breeding species (stallions, rams): daylight changes affect semen quality

IMMUNOLOGICAL FACTORS
 Sperm can induce antibodies → may cause infertility
 Mechanisms of antisperm antibodies:
1. Macrophage activation → sperm destruction
2. Sperm immobilization (cytotoxicity)
3. Block sperm penetration of cervical mucus
4. Interfere with capacitation/hyperactivation
5. Block zona pellucida penetration
6. Affect sperm selection in female tract
 Antigens originate from testis, epididymis, vas deferens, accessory glands
 Breach of blood-testis barrier → autoimmune response → azoospermia
 Monoclonal antibodies against sperm surface → experimental inhibition of fertilization
 Semen extenders (egg-yolk, milk) can also trigger antibodies in repeatedly inseminated
cows

NUTRITIONAL FACTORS
 Underfeeding: delayed puberty, reduced libido & testosterone in young males; less
effect in mature males
 Obesity/Overfeeding: reduced libido, especially in hot weather
 Protein deficiency: affects young more than mature males
 Vitamin deficiencies:
o Vitamin A: testicular degeneration, azoospermia; reversible if damage not
permanent
o Vitamin E: essential but role unclear in fertility
 Minerals: iodine, copper, cobalt, zinc, manganese may improve semen quality
 Toxic agents: plant estrogens, chemicals, radiation → interfere with spermatogenesis

CHROMOSOMAL & GENETIC FACTORS

 Sex determination: Y chromosome → maleness (TDF/H-Y antigen)
 Spermatogenesis regulated by Y chromosome genes
 Chromosomal aberrations cause ~20% of male infertility
 Disorders include: Y chromosome translocations, numerical anomalies, sex reversal,
ambiguous genitalia
 Selection against chromosomal abnormalities in breeding males is important