Pig

Rodney D Geisert and Matthew C Lucy, University of Missouri, Columbia, MO, United States
© 2018 Elsevier Inc. All rights reserved.

Glossary
Altrenogest A synthetic progestogen used to synchronize estrus in gilts.
Anestrous sow A sow that fails to come back into estrus after weaning.
Bicornuate uterus Uterus in which there are two long uterine horns (150–200 cm) connected by a common short uterine body
(1 cm).
Blastocyst Early stage of embryo development containing an outer trophoblast with inner cell mass and fluid filled center
called the blastocoel.
Conceptus Embryo and its extraembryonic placental membranes.
Corpus luteum (CL) Ovarian structure that is formed from the follicle after ovulation that synthesizes and secretes
progesterone.
Cystic follicle Large follicle on the ovary that is incapable of ovulation.
Dystocia Difficult birth caused by a large or badly positioned fetus.
Epitheliochorial placentation A noninvasive type of placental attachment in which microvilli of placenta (chorion) and
uterine surface epithelium interlock superficially at the maternal-fetal interface in the uterus.
Estrus Period of the estrous cycle when females are sexually active and receptive to mating by the male.
Farrowing rate Percentage of gilts or sows that become pregnant and farrow a litter after insemination.
Follicle Ovarian structure that contains the developing oocyte (egg) and also synthesizes and secretes estradiol to bring the pig
into estrus.
Gilt Young female pig.
Luteolysis Functional and structural regression of the corpus luteum causing a decrease in blood progesterone concentrations.
Maternal recognition of pregnancy Period of early pregnancy when a chemical signal(s) produced by the developing
conceptus prevents the regression of the corpus luteum to maintain the corpus luteum beyond the normal length of the estrous
cycle.
Mesometrial The region of the uterus where the broad ligament containing maternal blood vessels attaches and enters into the
uterus.
Morula Early embryo stage of development where multiple blastomeres form a solid ball of cells contained within the zona
pellucida.
P.G. 600 A combination of a pregnant mare serum gonadotropin (PMSG) and human chorionic gonadotropin (hCG) that is
used to bring prepubertal gilts and postpartum sows into estrus.
Polyestrous Females having multiple estrous cycles throughout the year.
Polytocous Females that give birth to multiple offspring at parturition.
Psuedopregnancy Condition of female where the corpus luteum lifespan is extended beyond the length of the estrous cycle
without the presence of a fetus.
Puberty Period of adolescence when a male or female is first able to release gametes.
Sow Adult female pig that has given birth.

Introduction

The domestic pig (Sus scrofa domesticus) is a polyestrous, polytocous farm animal species. Modern breeds of domestic pigs
typically breed year-round. The European wild boar and feral pigs, however, can display a seasonal (July to September) anestrus
related to photoperiod and nutrient availability in their natural (wild) environment. General characteristics of reproduction for
the domestic pig are presented in Table 1. Gilts (young female pigs before first farrowing) from commercial maternal lines reach
puberty at 5–7 months of age and farrow their first litters when they are approximately 1 year old. The length of estrous cycle for gilts
and sows (mature females that have given birth) is approximately 21 days (range 18–22 days) with estrus (standing heat) lasting
from 24 to 72 h. Ovulation occurs approximately 30–40 h after the initiation of estrus. Gilts and sows are typically inseminated by
either hand mating with a boar or by artificial insemination (AI) on the first and second day of estrus.
Gilts and sows ovulate from 20 to 30 ovarian follicles that are 8–10 mm in diameter when mature. Oocytes are released from the
ovulatory follicles and transported into the oviduct for fertilization. The cervix of the pig has thick alternating interdigitating pads
that lock around the spiral corkscrew shaped glans penis of the boar during mating (Fig. 1). Pipettes used for artificial insemination

Encyclopedia of Reproduction, 2nd edition, Volume 2 https://doi.org/10.1016/B978-0-12-809633-8.20520-7 641

642 Comparative Mammalian Female Reproduction j Pig

Table 1 Characteristics of female reproduction in the domestic pig

Age at puberty 5–7 months

Estrous cycle length 18–22 days
Duration of estrus 24–72 h
Time of ovulation 30–40 h after the initiation of estrus;
two-thirds to three-quarters of the time through estrus
Number of follicles ovulated 20–30
Gestation length 114–117 days
Age at first farrowing 12 months
Farrowing rate >85%
Litter size 8–18
Still born rate <7%
Lactation length 21–24 days
Weaning percentage (weaned/ live born) >90%
Number of farrowings per sow per year 2.3–2.4
Number of piglets per sow per year 25–30
Number of litters before culling 3–4

(AI) lock into the spiraling cervical rings of the pig to replicate the anatomy of the boar penis. Compared with other farm species, the
boar produces an extremely large volume of semen. Semen is diluted and used fresh for AI in pigs. The semen is infused into the
cervix or uterus in a relatively large volume over a 5–10 min period.
A high percentage of pigs become pregnant after breeding. The uterus of the pig has a short uterine body (1 cm) with long uterine
horns (150–200 cm) that support the extensive placentation of the conceptus (Fig. 1). Gestation is approximately 114 to 117 days
and litter sizes are typically 8–18 piglets per litter with an average of approximately 12 piglets per litter. Sows are weaned 21–24 days
after farrowing. After weaning sows return to estrus within 1 week and they are AI to reestablish pregnancy. The production cycle for

Fig. 1 Picture of pig bicornuate reproductive tract. Drawing by Lydia Michel and Sarah Petty.
 Comparative Mammalian Female Reproduction j Pig 643

a sow, therefore, is approximately 142 days (gestationþlactationþreturn to estrus). A highly productive sow, therefore, can farrow
nearly 2.5 litter per year and produce over 30 piglets per year.

Puberty and the Estrous Cycle

Puberty
Gilts are born with an immature ovary that contains primordial, primary and secondary follicles. Ovarian follicular development
depends on gonadotropins (follicle stimulating hormone (FSH) and luteinizing hormone (LH)) that arise from the anterior
pituitary under the control of gonadotropin-releasing hormone (GnRH). Circulating concentrations of FSH are elevated
until 10 weeks of age and provide an initial stimulus for ovarian follicular development. There is a gradual increase in the
number of LH pulses per day with increasing age. The negative feedback effect of estradiol on LH secretion is also reduced
over time. The ovary grows and there are successively larger ovarian follicles that secrete greater amounts of estradiol that in
turn stimulate growth of the uterus. When ovarian follicles achieve a critical threshold of circulating estradiol then there is an
LH surge that triggers first ovulation (puberty) at 5–7 months of age (80–120 kg of body weight). The age at puberty can be
influenced by breed, season of the year, nutrition, social environment, and exposure to the boar. A delay in puberty has
been observed in the summer months and also when young pigs are confined and crowded in indoor pens. Mixing gilts from
different pens or transporting gilts in a truck or trailer can advance the age at puberty if gilts have adequate nutrition. Daily
exposure of gilts to boars has a direct effect on advancing puberty in gilts that are between 135 and 160 day of age. Stimulation of
puberty by the boar occurs through physical contact, vocalization, and secretion of musk-smelling compounds, 16-androstenes,
that give the typical odor of mature boars. Boar odor provides signals to the female for inducing cyclicity and also the
strong lordosis response during estrus. The first pubertal estrus in the gilt is generally less fertile with a lesser ovulation rate
compared with subsequent estrous cycles. To achieve larger litters, therefore, gilts are typically inseminated at their second or
third estrus.
Puberty can be advanced by treating gilts with a combination of pregnant mare serum gonadotropin (PMSG) and human
chorionic gonadotropin (hCG) which is marketed under the trade name P.G. 600 (Merck Animal Health; Summit, New Jersey,
United States). Prepubertal gilts should be older than 5.5 months of age before treatment. Gilts will show estrus 4–10 days after
treatment (average¼7 days). Gilts can be inseminated at their first estrus after P.G. 600 or can be inseminated at the next estrus
which will increase farrowing rate and litter size.

Estrous Cycle
In domestic farm animals, the estrous cycle is defined as the number of days from initiation of behavior estrus expression to onset of
a subsequent estrus. Endocrine changes during the estrous cycle are presented in Fig. 2. There are four stages of the estrous cycle:
proestrus, estrus, metestrus, and diestrus. During proestrus, circulating concentrations of progesterone decline rapidly. The decrease
in progesterone (regression of the corpus luteum) is caused by the episodic release of prostaglandin F2a (PGF2a) from the
uterine endometrium. The regression of the CL and concomitant decline in circulating progesterone is associated with an increase
in the frequency of LH pulses that initiates a wave of follicular development. The development of large ovarian follicles increases the
circulating concentrations of estradiol and inhibin. The increases in both estradiol and inhibin cause a decrease in circulating FSH

Fig. 2 Diagram of circulating hormone concentrations during the estrous cycle of the pig.
644 Comparative Mammalian Female Reproduction j Pig

when the pig approaches estrus. High circulating concentrations of estradiol bring pigs into estrus; the next phase of the estrous cycle.
Pigs display lordosis behavior or “standing heat” when they are in estrus meaning that they will stand still with erect ears when
mounted by the boar or following the application of back pressure by a person. Other signs of estrus include swelling and reddening
of the vulva, vocalization (estrual grunts), mounting other female pigs and boar-seeking behavior. In response to estradiol, the
cervix becomes rigid and the uterine horns are tightly coiled.
The duration of estrus tends to be shorter in gilts (24–48 h) compared with mature sows (24–96 h). Circulating estradiol reaches
peak concentrations near the onset of estrus and triggers the LH surge. The LH surge may occur before, during or after the initiation
of estrus. High circulating LH concentrations last for about 10 h during the LH surge. Ovulation occurs during a 1–3 h period
approximately 30–35 h after the LH surge. In general, ovulation occurs when the estrus period is 70% complete. The fact that
the LH surge and ovulation is poorly timed with the onset of estrus makes the timing of AI difficult. Gilts and sows, therefore
are typically inseminated two times during estrus and may be inseminated a third time if they are in estrus on day 3. A new animal
drug named “OvuGel” (JBSUnited Animal Health; Sheridan, Indiana, United States) was recently introduced. OvuGel is a GnRH
agonist that causes an LH surge with ovulation 40–48 h after treatment. OvuGel can be used in a program for single fixed time
AI in sows after weaning.
Following ovulation, the granulosa cells and theca cells of the follicle wall become luteinized and form corpora lutea (CL).
Initially blood fills the lumen of the ruptured follicle to form the corpus hemmorrhagicum (CH) and the CH is eventually replaced
by the CL. The metestrus phase of the estrous cycle follows as the CL grows, fills the ruptured follicle cavity of the CH, and increases its
synthesis of progesterone. The presence of multiple CL (8–11 mm in diameter) on each ovary results in a steady increase in circu-
lating progesterone. Diestrus, the period of maximal circulating progesterone follows metestrus and continues for approximately
2 weeks during the estrus cycle. Concentrations of progesterone greater than 30–40 ng/mL are achieved during diestrus. The episodic
release of PGF2a from the uterine endometrium and the regression of the CL defines the end of diestrus and beginning of the proes-
trus phase which leads back to estrus.

Synchronization of Estrus
Reproductive management of farm species typically involves protocols that are used to synchronize estrus in a group of females.
Synchronization of estrus is done to reduce the time and labor associated with estrous detection and to facilitate the use of AI.
Although exogenous PGF2a is commonly used to regress the CL in cattle, it is not routinely used to synchronize estrus in gilts or
sows because the CL does not respond to exogenous PGF2a until after day 12 of the estrous cycle or pregnancy. Estrus in gilts
can be synchronized by feeding altrenogest (a progestin) marketed under the trade name MatrixÒ (Merck Animal Health, Intervet
Inc.; Summit, New Jersey, United States) at a dose of 15 mg/day for 14 day. The altrenogest suppresses follicular development so
that a synchronous wave of follicular growth occurs in a group of gilts after feeding is stopped. Gilts typically express estrus 4–9 day
after the last day of altrenogest feeding.

Breeding

Estrous detection is typically done twice daily using a boar. Gilts or sows that show an interest in the boar and that stand with
erect ears when pressure is applied to the back are considered in estrus. Estrus lasts 2–3 days. The reproductive tract of the pig
lacks a fornix vagina (blind pocket around the cervical opening) that is present in a number of other farm species (cow, mare
and ewe) (Fig. 1). Hand mating of boars to sows or AI of gilts and sows is quite easily accomplished because the fornix vagina
is not present. Hand mating or AI is usually done 12–24 h after first detection of estrus and then repeated on the next day. Gilts or
sows that are in heat for a third day may be inseminated three times (once for each day in estrus). Artificial insemination is very
typically done on swine farms because it conserves semen (allowing one boar to breed many females) and enables producers to
purchase semen from superior boars for use on farm. When AI is used, semen is diluted to a volume of 50 mL in extender and
a total of 2–3 billion sperm are used for each breeding. A plastic AI pipette attached to a bag of liquid semen is inserted into the
vagina and threaded or inserted into the cervical rings. The semen is then slowly infused into the cervix over 5–10 min. Some
AI pipettes lock into the cervix and a second catheter is fed through the cervix and into the uterine body for insemination.
Intrauterine insemination catheters enable the use of one-half the number of sperm cells per insemination without compro-
mising farrowing rate or litter size. A high percentage of oocytes are fertilized and most gilts or sows become pregnant after
breeding (typically >85% of those inseminated). The expectation is that breeding will yield both a high percentage of pregnan-
cies and also a large litter size (>11 piglets per litter).
Commercially raised females can exhibit a summertime decline in breeding efficiency (seasonal infertility) which is explained by
the combined effect of heat stress and photoperiod. Typical signs of seasonal infertility are anestrus, weak or irregular expression of
estrus, irregular estrous cycles, reduced farrowing rates, increased abortion rates, and depressed litter size. There are methods to miti-
gate heat stress on farm. For example, evaporative cooling systems will decrease the ambient temperature by 5–7  C inside swine
barns, depending on the outside temperature and relative humidity. These systems will reduce but not entirely eliminate the impact
of seasonal infertility on the reproductive efficiency of swine farms.
 Comparative Mammalian Female Reproduction j Pig 645

Early Embryonic Development

Following ovulation, the sticky cumulus cells surrounding the oocytes assist adhesion and transport to the ampullary-isthmic
junction (site of fertilization) of the oviduct through contraction of cilia lining the fimbria and ampulla luminal epithelium.
Follicular growth and the release of estradiol stimulates oviductal contractions to move the oocytes to the site of fertilization.
Estradiol stimulates a physical constriction of the isthmus lumen which restrains further movement until the uterus becomes
receptive for embryo development. Fertilized ova undergo their first cleavage division approximately 24 h after fertilization.
Cleavage continues with 4–8 cell embryos being rapidly transported into the uterine horns following progesterone induced
relaxation and widening of the isthmus luminal diameter (Fig. 3). The embryos continue to divide near the tips of the uterine
horns, developing into morula by Day 5 of gestation. The loose outer cells of the morula undergo compaction which is defined as
the formation of tight cellular junctions. Polarization of the outer trophectoderm allows sodium pumps located on the basal cell
border to transport fluid into the center to form the blastocoel. The presence of the blastocoel is the defining characteristic of
the blastocyst embryo which is found on Day 6–7 of gestation. Blastocysts continue to grow within the zona pellucida until
production of enzymes from both the uterus and blastocyst and the physical expansion and contraction of the blastocyst
stimulate hatching; the process through which the embryo breaks through the surrounding zona pellucida. Hatching occurs
on Day 8 of gestation (Fig. 3).
From Days 8–12 of gestation, blastocysts begin to migrate down and between the uterine horns to equidistantly space
themselves throughout uterus. Early uterine migration of blastocysts from the tip of the uterine horns is stimulated by blastocyst
synthesis of prostaglandins that induce the release of histamine from the endometrium to stimulate local myometrial contractions.
The spherical conceptuses continue to grow, migrate and complete spacing within the uterine horns through the production of
estrogens (Fig. 4(A)). Conceptus migration and trophoblast expansion is essential because the pig has both a local and systemic
pathway for PGF2a-induced luteolysis (CL regression) which would reduce progesterone and terminate the pregnancy. The pig
requires at least two conceptuses in each uterine horn to prevent luteolysis and maintain pregnancy.
The spherical conceptuses (Fig. 4(A)) steadily grow and increase in diameter from Day 10 (2–5 mm diameter) to 12 (8–10 mm
diameter) of gestation. Once reaching an approximate 10 mm spherical diameter, conceptuses undergo a rapid transition in
morphology to a thin filamentous tread-like structure measuring 100 mm in length (Fig. 4(B)). The rapid morphological change
during conceptus elongation occurs within approximately 1 h and results from massive cellular remodeling and movement rather
than cellular growth and proliferation. Pig conceptuses synthesize a unique form of interleukin-1b (IL1B2) that is involved in
regulating the rapid elongation of the conceptus. Gene edited IL1B2 null embryos do not elongate. IL1B2 is a proinflammatory
cytokine that stimulates a localized increase in blood flow to the uterus and regulates many endometrial responses to stimulate
implantation and the maternal immune system.
Conceptus elongation provides the placental surface area for nutrient uptake that is essential for continued embryonic and fetal
development to term. The pig endometrial surface is not a flat, smooth surface but has a substantial amount of primary and
secondary folding (Fig. 4(C)). Conceptuses follow the folds and attach along the mesometrial border of the uterine horns.

Establishment of Pregnancy

Rapid conceptus elongation not only provides the surface area for later placental development and growth, but provides the
mechanism to deliver conceptus estrogen (maternal recognition signal) across the endometrium to block luteolysis and maintain
pregnancy. Two periods of conceptus estrogen production and release are involved in maintaining the CL throughout pregnancy in
the pig. Conceptuses release estrogen during the process of elongation on Day 12 of gestation. This short pulse of estrogen produc-
tion 1–2 days) extends the lifespan of the CL to 25–30 days. Although conceptus elongation normally occurs between Day 11–12,
the window for elongation and uterine estrogen stimulation for establishment of pregnancy in the pig extends from Day 11–14 of
gestation. A second sustained release of estrogen occurs during period of attachment and placental formation from Day 16–25 of
gestation. The short (Day 12) and prolonged (Day 16–25) release of estrogen is essential for CL maintenance to term. Treatment

Fig. 3 Cleavage and early development of pig embryos from fertilization until blastocyst hatching. Day 0¼time of ovulation.
646 Comparative Mammalian Female Reproduction j Pig

Fig. 4 Conceptus development from Day 11–12 of gestation. (A) Spherical conceptus (5 mm diameter) on Day 11 of gestation. (B) Elongated
conceptus (100 mm) on Day 12 of gestation. (C) Segment of uterine horn demonstrating the multiple folding of the endometrium during placentation
in the pig.

of cyclic gilts or sows with estrogen or environmental exposure to estrogenic compounds such as aflatoxins induces psuedopreg-
nancy (extended lifespan of CL without conceptuses present) which can range from 28 to 110 days.
Conceptus secreted estrogens do not function to prevent luteolysis through inhibition of endometrial PGF2a synthesis and
release as occurs in a number of large farm animal species. In the pregnant pig, estrogen alters endometrial PGF2a movement to
redirect the PGF2a away from the CL. The PGF2a is sequestered into the uterine lumen and metabolized rather than being trans-
ported into the uterine capillaries to induce luteolysis as occurs during the estrous cycle. Pig conceptuses secrete PGE which also
assists in sequestering PGF2a into the uterine lumen and stimulates CL progesterone production and maintenance during early
gestation.

Placental Attachment and Development

Evenly spaced, elongated conceptuses do not overlap and developing placenta do not fuse together during gestation. The
conceptuses continue to grow and expand throughout the uterine horns growing from 100 mm to 500 mm length from Day
15–30 of gestation. The placenta will continue to grow reaching lengths of approximately 800–1000 mm by Day 70 of gestation.
The pig has a noninvasive, epitheliochorial type of placentation which the chorion attaches to the surface epithelium along the
folds of endometrium (Fig. 5(A)). The multiple primary and secondary folds of the endometrial lining allow the meter-long
diffuse placenta (Fig. 5(B)) of each fetus to occupy a uterine area approximately one-third of its length (Fig. 5(A)). The endo-
metrium of the pig contains thousands of uterine glands (Fig. 5(B)) to supply the nutrients needed for continued development
to term. With regression of the yolk sac on Day 15 of gestation, complete attachment of the chorion to the endometrial surface
epithelium occurs through expansion of the allantois from the hindgut of the developing embryo on Day 15. The fluid filled
allantois expands throughout the chorion reaching a volume of 200–300 mL by Day 30 of gestation. Fluid in the allantois
declines from Day 32–45 but a second increase in volume to 300–400 mL occurs and peaks on approximately Day 60 of
gestation. The allantoic fluid declines from its peak volume on Day 60 to approximately 50–80 mL on Day 100 of gestation.
The allantois serves as a reservoir for nutrients (proteins, amino acids, carbohydrates and ions) absorbed from the uterine glands
by the overlying chorionic areola (Fig. 5(B)). The allantois may not expand to the extreme ends of the chorion when tips of
placental membrane of individual fetuses make contact and ends fold over each other. The lack of placental blood
supply from the allantois to the tip of the chorion results in the formation of brownish necrotic tips (Fig. 6). Placental length
(800–1000 mm) and weight (200–250 g) of pigs continue to increase until approximately 70 days of gestation where it plateaus
until parturition at 114–117 days of gestation.
 Comparative Mammalian Female Reproduction j Pig 647

Fig. 5 Histological section of the epitheliochorial placentation in the pig at Day 30. (A) Attachment of the chorion to the luminal epithelium of the
endometrium (note the intense folding). (B) Uterine glands that provide nutrients for fetal and placental growth throughout pregnancy. (Insert
provides the uterine location for magnification of A and B). Cap, capillary; Endo, endometrium; Myo, myometrium.

Endocrinology of Pregnancy and Parturition

Corpora lutea do not regress during pregnancy because the conceptus prevents the secretion of PGF2a into the uterine vasculature.
There is a slight decline in circulating progesterone concentrations after Day 14 until term. The decline in circulating progesterone is
explained by the steroid metabolism by the uterus and conceptus during pregnancy. Maintenance of circulating progesterone
concentrations throughout gestation is totally dependent on the CL. Regression of the CL during pregnancy leads to abortion of
the pregnancy.
Circulating concentrations of unconjugated and conjugated estrogens such as estrone sulfate increase from Day 16 to peak levels
between Day 23 and 30 of gestation. The circulating concentrations then decline to low levels by Day 45 before slowly increasing to
attain a second peak on the day before parturition. The fetoplacental unit is the primary source of circulating estrogen during
pregnancy. Increases in estrogen follow the steroidogenic activity of the pig placenta and change in placental allantoic volume
throughout gestation.
Parturition occurs on Day 114–117 of gestation in the pig. The release of glucocorticoids from the maturing fetal adrenal is asso-
ciated with the initiation of parturition in the pig. The glucocorticoids stimulate the release of PGF2a from the endometrium which
in turn causes a rapid decline in circulating progesterone concentrations 1–2 day before parturition. There is the release of prolactin
from the anterior pituitary and relaxin from the CL at this time as well. The release of prolactin stimulates mammary growth and
lactogenesis. Along with relaxin, placental estrogen softens and dilates the cervix. The PGF2a stimulates oxytocin release that
increases myometrial activity that is essential for delivery of the piglets through the softened cervical canal. Sows can be induced
648 Comparative Mammalian Female Reproduction j Pig

Fig. 6 Pregnant uterus (A) and fetal/placenta (B) collected on Day 60 of gestation.

to farrow after 110 of gestation. Before this time, induction of parturition results in a high percentage of stillborn piglets. Admin-
istration of exogenous PGF2a (LutalyseÒ, Zoetis Animal Health, Parsippany, New Jersey, United States) is an effective method to
induce parturition after day 110. Delivery typically occurs within 2 days after treatment.
Sows are moved into farrowing rooms that are equipped with specially designed “farrowing crates” a few days before
expected farrowing. The farrowing crates are designed to minimize the number of piglets that are crushed by the sow when she
lays down. Farrowing in the sow typically occurs over a 2–5 h period. A typical interval between the delivery of each individual piglet
is 10–15 min. Piglets are delivered randomly between the uterine horns. A typical delivery involves live born pigs, stillborn pigs, and
mummies. Mummified pigs are brown in color and died at some time during gestation after there was skeletal development.
Stillborn pigs appear completely normal but are dead at birth. Risk of stillbirth is greater for piglets born late in the farrowing
process, for sows farrowing large litters, and for piglets born >20 min after the last piglet. If piglets fail to be delivered on time
then it may indicate that there is a piglet lodged within the birth canal (dystocia). If his occurs then the piglet is removed manually
(gloved hand passed through the cervix and into the uterus) by the person monitoring the sow at farrowing. Oxytocin can be admin-
istered to sows that fail to deliver piglets on time if there appears to be weak myometrial activity and there is no blockage within the
uterus. The placenta are passed within 4 h of the delivery of the last piglet.

Lactation

The increase in prolactin, glucocorticoids, growth hormone, and relaxin during the days prior to parturition stimulates lactogenesis
in the pig. Milk let down occurs following oxytocin release when piglets suckle. Piglets nurse as a group when allowed to suckle by
the sow. The group nursing ensures that milk from all of the teats is consumed at the same time. Nursing interval during early
 Comparative Mammalian Female Reproduction j Pig 649

lactation is less than 1 h but this interval increases over time. Milk production in the sow increases during the third and fourth weeks
of lactation.
Greater than 90% of live born pigs should survive until weaning. A common cause of death before weaning is crushing where
the mother accidently lays on top of a piglet and kills it. Runt piglets (extremely small piglets at birth) may also die before
weaning because they are either crushed or fail to compete for milk. Piglets are typically cross-fostered at one to two days after
farrowing by moving piglets from a sow with a large litter to a sow with a small litter. Cross-fostering is most-successful for
sows farrowing within 1 or 2 days of one another.
Most sows are weaned at approximately 21 days (3 week) of lactation. Sows rarely come into estrus while they are lactating
because pulsative LH release is suppressed by suckling. The development of large estrogenic ovarian follicles depends on LH
and therefore the ovary has only small follicles when the sow is lactating. After weaning, however, there is a rapid increase in
LH pulsatility that initiates the development of preovulatory follicles on the ovary.
Sows are moved out of farrowing crates into breeding and gestation barns immediately after weaning. Most sows will be in estrus
within 3–7 days after weaning and are inseminated at that time. The endocrine and follicular changes after weaning are very similar
to what was previously described for proestrus. Some sows do not return to estrus after weaning. These sows may be cystic
(have large non-ovulatory follicular cysts on the ovary), may have ovulated during lactation (relatively rare) or may be anestrus
(fail to develop follicles after weaning). There are no effective treatments for follicular cysts in the sow. Sows that are anestrus
can be treated with P.G. 600 to induce estrus and then can be AI.
Sows remain in the herd until they either become subfertile (reduction in litter size or farrowing rate), become injured, or
experience skeletal problems generally associated with feet and legs. A sow will typically be productive for 3–4 parities. The
production cycle for a sow is approximately 142 days (gestation (114 days)þlactation (21 days)þreturn to estrus (7 days); assumes
100% efficiency). A sow, therefore, can farrow approximately 2.5 litters per years if she is healthy and highly fertile. If she weans 12
piglets per litter then 30 piglets per sow per year are produced. This level of productivity is achieved by approximately 10% of sows
today. A more typical target for swine farms is 25 piglets weaned per sow per year.

Further Reading

Anderson, L. L. (2000). Pigs. In E. S. E. Hafez, & B. Hafez (Eds.), Reproduction in Farm Animals (pp. 182–191). London: Lippincott Williams & Wilkins.
Geisert, R. D., Whyte, J. J., Meyer, A. E., et al. (2017). Rapid conceptus elongation in the pig: An interleukin 1 beta 2 and estrogen regulated phenomenon. Molecular Reproduction
and Development, 84(9), 760–774. https://doi.org/10.1002/mrd.22813.
Geisert, R. D., Johnson, G. A., & Burghardt, R. C. (2015). Implantation and establishment of pregnancy in the pig. In regulation of implantation and establishment of pregnancy in
mammals: Tribute to 45 Year Anniversary of Roger V. Short’s “Maternal Recognition of Pregnancy.”. In R. D. Geisert, & F. W. Bazer (Eds.), Advances in Anatomy, Embryology
and Cell Biology (pp. 137–164). Springer.
Mauget, R. (1982). Seasonality of reproduction in the wild boar. In D. J. A. Cole, & G. R. Foxcroft (Eds.), Control of Pig Reproduction (pp. 509–526). London: Butterworth Scientific.
Perry, J. S. (1981). The mammalian fetal membranes. Journal of Reproduction and Fertility, 62, 321–335.
Senger, P. L. (2012). Pathways to Pregnancy & Parturition. Redmon, OR: Current Conceptions, Inc.