Hormones and Behavior 52 (2007) 78 – 85

www.elsevier.com/locate/yhbeh

Maternal behavior in pigs

Bo Algers ⁎, Kerstin Uvnäs-Moberg
Department of Animal Environment and Health, Swedish University of Agricultural Sciences, PO Box 234, 532 23 Skara, Sweden
Received 27 March 2007; accepted 28 March 2007
Available online 1 April 2007

Abstract

When sows kept under commercial conditions were put into crates in the early 1960s, the neuro-endocrine regulation of the maternal behavior
in these domestic animals was disputed. Thus, the study of sow maternal behavior intensified and today a significant body of knowledge has
accumulated to support the hormonal regulation of sow maternal behavior. The onset of nest building is associated with a periparturient decline in
progesterone, an increase in prolactin and a major rise in plasma concentrations of PGF2α the day before parturition. Some nest building
behaviors, such as pawing and gathering straw, have been found to correlate with changes in the levels of progesterone, prolactin and somatostatin.
The duration of the birth process correlates negatively with peripheral oxytocin levels. During lactation, the stimuli from the piglets affect the
release of several hormones which not only regulate the let down of milk but also sow metabolism and mammary milk production. The sow's
nursing behavior ensures an even distribution of milk to her piglets. The piglets suckling behavior, in turn, is mainly a way to communicate their
individual nutritional needs.
© 2007 Elsevier Inc. All rights reserved.

Keywords: Pigs; Sows; Maternal; Behavior; Hormones; Nursing; Suckling; Lactation; Metabolism; Communication

Introduction
The nursing behavior of sows has attracted the interest of
researchers since the 1950s. Much of this research was aimed at
increasing the knowledge to better understand how piglets
could be provided with milk from the sow in order to increase
weight gain.
The pig (Sus scrofa) is an omnivore living in groups of about
8–10 adult sows, some young individuals and, in the periphery,
single males (Jensen, 1986). Under natural conditions, a day or
so before the onset of parturition or farrowing, the sow separates
herself from the group and seeks a suitable nest site with some
shelter from rain and wind, well-drained soil and with
possibilities to root a shallow hole in the ground (Grundlach,
1968; Frädrich, 1974; Jensen, 1986). Nest building occurs
during the last 24 h before the onset of farrowing and is most
intensive during 12 to 6 h before farrowing (e.g., Vestergaard
and Hansen, 1984; Jensen, 1986). Nest building is performed in
phases: first, the sow digs a shallow hole in the ground after
which she gathers branches of bushes etc. and arranges them
⁎ Corresponding author. Fax: +46 511 67204.
E-mail address: Bo.Algers@hmh.slu.se (B. Algers).
0018-506X/$ - see front matter © 2007 Elsevier Inc. All rights reserved.
doi:10.1016/j.yhbeh.2007.03.022
along the edges of the nest. The nest building ends with the sow
gathering softer material such as grass that she distributes
throughout the nest by nodding head movements and pawing
with front legs (Jensen, 1986). Thus, the nest can provide some
comfort, thermoregulation and shelter for the piglets. Nest
building can be divided into two distinct phases; an initial phase
with rooting and pawing in the ground and a subsequent phase
with collecting, carrying and arranging nest material (Jensen,
1993).
Under commercial conditions the sow is usually bred at 7–8
moths of age, is pregnant for 133 days, and transferred to
gestation and farrowing crates where she is kept individually
without possibilities to turn around. This practice was suggested
to reduce piglet mortality due to crushing by the sow. In some
countries, gestating sows are instead kept in groups. The sows
are moved to a facility where they give birth — a farrowing
unit. This can either be equipped with farrowing crates where
there is some space for the piglets outside the crate or the sow
can be kept loose in a pen with a creep area for the piglets where
they are protected from being crushed by the sow.
Since the development of farrowing crates to house sows in
confinement in the late 1950s, it has been argued that the
modern pig does not necessarily build nests when approaching
 B. Algers, K. Uvnäs-Moberg / Hormones and Behavior 52 (2007) 78–85
the time of farrowing due to a high degree of domestication and
that it, therefore, does not need an environment which allows it
to do so. However, increasing evidence has accumulated to
challenge that assumption. Studies on nest building of domestic
sows were first investigated under semi-natural conditions in
the 1980s (Stolba and Wood Gush, 1989; Jensen, 1986; Algers
and Jensen, 1990). They showed that, e.g., Swedish landrace
sows, with experience from four prior farrowings in confine-
ment crates, were perfectly able to locate suitable nest sites and
to build farrowing nests, a behavior more or less identical with
that of the wild boar (Gustafsson et al., 1999). Since then, the
study of maternal behavior has intensified, such that, to date a
significant body of knowledge has built on the hormonal
regulation of sow maternal behavior.
Nest building
The onset of nest building is associated with a periparturient
decline in progesterone from about 50 to 20 nmol/l and increase
in prolactin (Ash and Heap, 1975; Taverne et al., 1978/1979;
Widowski and Curtis, 1989; Meunier-Salaün et al., 1991;
Castrén et al., 1993a). Prolactin concentrations are highly
variable among sows on the day before farrowing and increase
from about 15 ± 4 nmol/l (n = 10) 3 days before farrowing to
reach maximum levels of about 72 ± 27 nmol/l (n = 10) at 8 to
12 h before the onset of parturition (Castrén et al., 1993a). A
major rise in plasma concentrations of PGF2α on the day before
parturition in sows has been demonstrated (Watts et al., 1988)
and PGF2α metabolite (measured as 13,14-dihydro-15-keto-
PGF2α) levels have been shown to increase from 1–2 nmol/l to
about 50 nmol/l (Gilbert et al., 2002). An injection of
prostaglandin (PGF2α) leads to an immediate increase in
prolactin concentration and a rapid onset of nest building
(Widowski et al., 1990). In contrast the prostaglandin analogue
Cloprostenol led to a delayed increase in prolactin and delayed
nest building which would suggest that it is the rise in prolactin,
rather than the changes in prolactin concentrations that is
important to trigger nest building behavior (Widowski et al.,
1990). Indomethacin treatment specifically inhibits nest build-
ing in sows by a mechanism that may involve an inhibition of
endogenous PGF2α synthesis, independently of circulating
oxytocin, cortisol and progesterone levels. This points to a
major role of PGF2α in prepartum nest building in the sow
(Gilbert et al., 2000). Indeed, PGF2α given as a single dose to
cyclic nulliparous sows (gilts) induces rooting, pawing at the
ground and gathering straw, behaviors characteristic of nest
building in the sow (Burne et al., 2000a). Pawing and gathering
straw have been shown to be dependent on the dose of PGF2α
(Burne et al., 2000a), pawing and duration of rooting are
modulated by the provision of straw (Burne et al., 2000b) and
rooting and lying are modulated by environmental temperature
(Burne et al., 2001).
Some nest building behaviors have been found to correlate
with levels of specific hormones. Thus, the time spent carrying
and depositing straw correlated significantly with prolactin
(r = 0.65, n = 9, p = 0.05), tended to be positively correlated with
progesterone (r = 0.60, n = 9, p = 0.08) and tended to correlate
79
negatively with somatostatin concentrations (r = − 0.65, n = 9,
p = 0.06). However, only progesterone and somatostatin were
independently correlated with the time spent carrying straw
(Castrén et al., 1993a). Damm et al. (2002) found a negative
correlation between plasma oxytocin concentrations and nosing
(r = − 0.8, p < 0.01) and arranging nest material (r = − 0.9,
p < 0.001).
Several factors may influence the motivational systems
regulating nest building in the sow. The onset of nest building is
suggested to be regulated by the increase in prolactin levels
(e.g., Castrén et al., 1993a) although in sows that gave birth for
the first time (i.e., gilts) there was either no increase or only
moderate rise in prolactin levels (Lawrence et al., 1994). The
authors suggested that with an increasing parity there is a closer
temporal relationship between rise in prolactin and nest
building. However, older parity sows tended to have a lower
prolactin concentration 2 days before farrowing (Castrén et al.,
1993a).
The gathering of nest material appears to be regulated by
external stimuli such as temperature or udder comfort, whereas
a softer texture, such as abundance of grass, would inhibit
further gathering of soft material (Algers and Jensen, 1990;
Jensen, 1993). This is supported by the findings that changes in
the sow farrowing environment had greater effects on the
gathering of straw than on rooting and pawing behavior (Jensen,
1993; Arey et al., 1991) and that these behaviors are not
correlated (Castrén et al., 1993a). In addition, the finding that
the time spent carrying and depositing straw was independently
correlated with high progesterone and low somatostatin
concentrations 2 days before farrowing (Castrén et al., 1993a)
suggests that not only sensory cues from the environment but
also endocrine mechanisms take part in the control of these
aspects of nest building behavior.
Somatostatin is a gastrointestinal hormone which exerts
inhibitory effects on gastrointestinal and metabolic functions.
Somatostatin levels are under inhibitory vagal nerve control,
i.e., somatostatin levels are decreased by vagal nerve activity.
When vagal nerve activity is increased, as e.g. during lactation
(Uvnäs-Moberg, 1994) somatostatin levels fall and conse-
quently gastrointestinal and metabolic functions are enhanced
leading to increased nutrient assimilation and growth (Uvnäs-
Moberg, 1994) (for a more detailed description see paragraph
on integration between maternal behavior, physiology and
metabolism during lactation). The findings by Castrén et al.
(1993a) indicate that somatostatin plays a role also in maternal
behavior, decreasing from approximately 15 nmol/l on day 3 to
10 nmol/l 1 day before farrowing. It is, in fact, possible that the
relationship between low levels of somatostatin and the time for
carrying and depositing straw is a reflection of a variant of
ingestive behavior occurring during nest building. The sows
take nest building material into their mouths but instead of
swallowing it they place the material in their nests. However, it
cannot be ruled out that the presence of straw in the oral cavity
per se leads to an activation of the vagal nerves and a
consequent decrease of somatostatin levels.
Estrogen levels (estradiol and estrone) remain low during the
first half of pregnancy (< 100 pg/ml levels) but rise rapidly (2–
80 B. Algers, K. Uvnäs-Moberg / Hormones and Behavior 52 (2007) 78–85
4000 pg/ml) during the second half of pregnancy (K'nchev et
al., 1975; Velle, 1976). No relationships between estradiol
levels and nest building have been demonstrated although levels
are extremely high in the periparturient period. Most of the
production of estrogens occurs in the placenta and therefore
estrogen levels drop abruptly after birth to reach the same levels
as in the beginning of pregnancy. Estradiol and estrone levels
remain low during location but increase slightly before estrus
(Hultén et al., 2006).
The cessation of nest building has been suggested to occur
when udder comfort is sufficient (Baxter, 1983). The provision
of a preconstructed nest led to a quicker cessation of nest
building but straw carrying was not eliminated: instead there
was an increase in the amount of rooting behavior (Arey et al.,
1991).
In contrast, the removal of nest material every 4th h until
parturition has been shown to increase plasma cortisol and heart
rate in gilts in comparison to sham removal whereas plasma
oxytocin was unaffected by such a procedure indicating a more
complex regulation of oxytocin release (Damm et al., 2003).
However, moving sows in early parturition from a crate to a pen
has been shown to decrease oxytocin concentrations (Lawrence
et al., 1992).
Nest building stops at about 4 h before the onset of
parturition and a dramatic elevation of oxytocin levels starts at
about 6 h before the onset of parturition, a change that seemed to
end the nest building phase (Castrén et al., 1993a). At that time,
uterine contractions increase in frequency (Taverne et al., 1979).
Electric activity of the uterine muscles coincides with oxytocin
release (Forsling et al., 1979) and spontaneous contractions
occur only when oxytocin concentrations are elevated (Taverne
et al., 1979). Sows with high oxytocin concentrations 8 h
prepartum stopped nest building earlier than other sows
(Castrén et al., 1993a). Whether the peripheral effect of
oxytocin in generating uterine contractions is the cause of the
cessation of nest building or whether oxytocin acts to stop nest
building at a central level remains to be investigated. A direct
effect on the central nervous system as a result of the increase in
oxytocin release cannot be ruled out.
In summary, it can be concluded that there is strong evidence
to suggest that the onset of nest building in sows is triggered by
a rise in prolactin levels, which is itself related to decreased
progesterone and increased prostaglandin concentrations.
Further, some nest building activities such as carrying and
depositing straw seem to be related to changes in somatostatin
and progesterone levels. Nest building ends when oxytocin
levels begin to rise.
Farrowing
Farrowing involves the expulsion of fetuses which takes
place when the sow lays down in the farrowing nest. It has been
suggested that oxytocin release is of importance for effective
contractions of the uterus and rapid birth of the piglets (Forsling
et al., 1979; Taverne et al., 1979; Lawrence et al., 1997; Gilbert
et al., 2000). Sows give birth to 4–7 offspring (wild boar)
(Harris et al., 2001) and, under commercial conditions, typically
to 10–12 (domestic sow, van Dijk et al., 2005). The duration of
parturition is 166 min (26–505) with some variability (130–245
min) among strains (van Dijk et al., 2005). During farrowing,
oxytocin plasma baseline concentrations are elevated and
oxytocin is further released in a pulsatile pattern (e.g., Forsling
et al., 1979). Baseline levels of oxytocin before farrowing are
around 5–6 fmol/l, increasing to about 10 fmol/l just before
farrowing and to 45 fmol/l at 3 h after the start of farrowing,
whereafter it decreases to approximately 5 fmol/l (Castrén et al.,
1993b).
Research on several species such as miniature pigs, rabbits,
rats, goats, and horses (Forsling et al., 1979; Haldar, 1970;
McNeilly et al., 1972; Allen et al., 1973; Higuchi et al., 1986)
suggests that the expulsion of each fetus is related to a peak
release of oxytocin. Studying pigs and the pulsatile release of
oxytocin in detail, however, Castrén et al. (1993b) were unable
to relate the expulsion of fetuses to a peak release of oxytocin.
Only about 50% of piglets were born during an oxytocin peak
and the first piglets in the litter were born at much lower
oxytocin levels than those born later when increased oxytocin
peaks were observed. It may well be that the vaginal distension,
occurring as a result of the first pigs being born may, in turn,
increase the circulating levels of oxytocin as suggested by
Ellendorff et al. (1979). The release of oxytocin during
farrowing in sows may also be stimulated by the udder massage
performed by already born littermates since it has been shown
that udder stimulation during established lactation causes
release of oxytocin (e.g., Algers et al., 1990). Moreover, as
oxytocin also is released into the brain, it stimulates its own
release (Hatton and Tweedle, 1982; Theodosis, 2002). This
positive feed back may explain the increase of oxytocin over
time during farrowing.
Cortisol concentrations rise at farrowing (Taverne et al.,
1979; Lawrence et al., 1994, 1997; Jarvis et al., 1998), probably
as a result of physical strain or as a response to a psychological
factor(s).
A short duration of the farrowing process is of importance
for piglet survival and a delay can result in anoxia or stillbirth
(Randall, 1972; Bille et al., 1974). Low oxytocin levels may
cause a prolongation of the farrowing time (Castrén et al.,
1993b) and a negative correlation between plasma oxytocin
levels and duration of parturition (r = − 0.7, p < 0.01) has been
shown (Damm et al., 2002). Parturition is prolonged in rats if
oxytocin secretion is inhibited by opioids (Russell et al., 1989)
and an environmental disturbance could cause the same effect in
pigs (Lawrence et al., 1992). Sows with long farrowing times
had lower basal as well as lower peak levels of oxytocin during
farrowing in comparison to sows with short farrowing times
(Castrén et al., 1993b). However, in that study, long farrowing
times could not be predicted from prepartum oxytocin,
prolactin, progesterone or somatostatin levels or from the
prepartum behavior of the sows.
Lactation
The sow is rather unique in that she gives birth to a large
number of behaviorally well-developed young. Each piglet is
 B. Algers, K. Uvnäs-Moberg / Hormones and Behavior 52 (2007) 78–85
very capable suckling on its own, but for the fitness of the sow it
is important to distribute her bodily resources through milk to
all of her piglets. Therefore, an array of different behavioral
elements has evolved allowing for the piglets to communicate
their needs of energy and the sow to distribute her resources
evenly to all her piglets (Algers, 1993). Unlike cows, sows have
no teat cisternae, which is why a piglet cannot obtain milk
without there being an increase in the intramammary pressure
mediated by oxytocin release. The increase in pressure lasts for
approximately 15 s. Piglets are called to the udder by the sow
milk grunt (Castrén et al., 1989) and the increase in grunting
frequency causes the piglets to switch from massaging behavior
to sucking on a teat (Algers and Jensen, 1985).
Early lactation
The first milk ejections can appear even before farrowing is
complete. Although it has been suggested that during the first
hours, colostrum would be available continuously (Lewis and
Hurnik, 1986), a further release of colostrum release requires a
discrete release of oxytocin (Fraser, 1984). Basal levels of
oxytocin increase during 7–8 h after the onset of farrowing to
about 45 fmol/l (Castrén et al., 1993b). Some, but not all, of the
early milk ejections are associated with a pulsatile oxytocin
secretion with peaks reaching about 100 and sometimes
200 fmol/l. During the first 8 h, discrete milk ejections appear
at shorter intervals and with a higher variability, 5–40 min., than
during established lactation, when they approach 1 milk
ejection per hour (Fraser, 1984; Castrén et al., 1993b; Fraser
and Rushen, 1992).
Piglets compete with their littermates for access to teats
during the first day but gradually the suckling rhythm
synchronizes within the litter and a “teat order” is established
(McBride, 1963; Hemsworth et al., 1976; Lewis and Hurnik,
1985; De Passillé and Rushen, 1989). The sow recognizes her
own piglets by olfactory cues (Frädrich, 1974) and she is often
aggressive towards intruding piglets although cross suckling
occurs frequently in the domestic pig (Newberry and Wood-
Gush, 1986).
Established lactation
Initiation
Either the piglets initiate the nursing by approaching the
sow's udder and emitting a “high/deep grunt” (Jensen and
Algers, 1984) or, as is more common during the first week, the
sow lays down, exposes her udder and emits a typical nursing
grunt (Algers and Jensen, 1985; Jensen et al., 1991). The sound
of the sow grunting attracts the piglet to the udder (Jeppesen,
1982; Castrén et al., 1989) which ensures that all piglets, each
having the capacity to produce a let down, assemble at the udder
to be prepared for the short time that milk is available to them.
Pre let down
When lactation is established, let down and suckling occur at
regular intervals of 45–60 min when all piglets in the litter
suckle at the same time (Barber et al., 1955; Fraser, 1977). This
81
is triggered by a release of oxytocin from the posterior pituitary
(Folley and Knaggs, 1966; Ellendorff et al., 1982). As the sow
does not have a teat cistern, milk is only available during milk
ejection, which is stimulated by the piglets massaging each
udder segment (Whittemore and Fraser, 1974) for 1–3 min
(Fraser, 1980; Algers and Jensen, 1985; Algers et al., 1991). In
most mammals, such as humans and cattle, this period is much
shorter and milk ejection can be conditioned by other stimuli
that do not involve mechanical stimulation of the udder
(Schmidt, 1971). Fraser (1980) suggested that the function of
the pre let-down phase is an adaptation that ensures all the
members of a large litter are present at nursing. Indeed, when
fewer piglets are massaging the udder, it takes longer for
oxytocin release to occur (Algers et al., 1990).
Let-down phase
The grunting of the sow increases in frequency just before let
down which Fraser (1980) suggested to be a signal for the
piglets to change from massaging the udder segment to sucking
on the teat. If high-level fan noise is used to mask the vocal
signals from the sow in early lactation (days 1–3), suckling
behavior is much less synchronized and the time when piglets
switch from massaging to sucking is more variable, which
results in less milk ingestion by the piglets (Algers and Jensen,
1985). The fast grunting starts at about 23–25 s before the rise
of the intramammary pressure (Ellendorff et al., 1982; Fraser,
1975). There is an increase of oxytocin in the blood starting at
about 15 s after the increase of the grunt rate (Algers et al.,
1990). A latency of 15 to 25 s for the released oxytocin to reach
the udder can be expected (Whittlestone, 1953, 1954a,b), which
may be why it is the sharp increase, rather than the peak of the
grunt rate that seems to occur simultaneously with oxytocin
release (Algers, 1989).
Ellendorff et al. (1982) suggested that oxytocin facilitates
grunting. Oxytocinergic fibers, which originate in the para-
ventricular nucleus, project to the dorsal and dorsomedial vagal
nucleus (Swanson and Sawchenko, 1980) which also innervates
larynx. It could be hypothesized that there is a facilitatory effect
of oxytocin on grunting mediated either by the activation of
oxytocinergic fibers or by an increase of oxytocin in plasma. It
has been shown that the peak grunt rate level is influenced by
the time and number of piglets massaging but no such effects
could be demonstrated in response to the integrated grunt rate
response (Algers et al., 1990). Thus, the volume of grunting
seems to be rather constant but distributed differently depending
on the number of piglets stimulating.
Post let down
After the milk is consumed, pigs resume massaging the
udder of the sow, an activity that can go on for several minutes.
The longer time spent by the litter massaging the udder, the
more milk was produced by the sow (Gill and Thomson, 1956).
Indeed, piglets showed a decrease in weight gain following
periods where the post let-down massage was prevented
experimentally (Barber et al., 1955). As piglets keep to their
“own” udder segment through lactation, Algers and Jensen
(1985) presented the so-called “restaurant hypothesis” postu-
82 B. Algers, K. Uvnäs-Moberg / Hormones and Behavior 52 (2007) 78–85
lating that the piglets in a litter individually “order” the size of
their future meals by performing more or less post let-down
massage onto their “own” udder segment. The idea was that the
final massage acts as a regulator of sow milk production which
allows her to adjust this parameter to the number of vital piglets
suckling.
Tactile stimulation and hormonal release
Several studies have shown that piglet udder massage results
in the release of prolactin (e.g., van Landeghem and van de
Wiel, 1978; Algers et al., 1991) which reaches its highest
concentrations in plasma 10–20 min after nursing (Einarsson et
al., 1992; Rojkittikhun et al., 1993b; Spinka et al., 1997).
An increase in plasma prolactin concentration is associated
with an increase in lactational performance of the sow (Smith
and Wagner, 1980). Prolactin increases the number of insulin
receptors in the mammary gland (Uvnäs-Moberg, 1989) and
decreases these receptors in maternal fat stores. This directs
energy to the mammary glands rather than to fat deposits,
thereby promoting lactation.
Daily ergocryptine treatment of sows 2 days before estimated
farrowing date significantly reduced mean plasma levels of
prolactin before and after farrowing, resulting in an absence of
milk production (Whitacre and Threlfall, 1981). Bromocriptine
treatment of sows from day 25 of lactation until weaning 9–17
days later suppressed prolactin serum levels but did not affect
piglet weight gains until 6 weeks after birth (Benjaminsen,
1981). Further, a similar treatment for 5 days, starting on day 21
in lactation did not affect litter body weight (Mattiolo and Seren,
1985) indicating that prolactin seems to be most important for
milk production in the early stages of lactation.
Integration between behavior, physiology and metabolism
during lactation
In addition to releasing oxytocin and prolactin, piglet udder
stimulation promotes the release of several gut hormones such
as gastrin, somatostatin and vasoactive intestinal polypeptide
(VIP) and of pancreatic hormones such as insulin and glucagon
in several species including pigs (Lindén et al., 1987; Uvnäs-
Moberg et al., 1984; Algers et al., 1991). The suckling/massage-
Table 1
Hormonal regulation of specific maternal behavior activities
Initiation Nest Termination
of nest building of nest
building building
Estradiol +? +? +?
Oxytocin 0 0 + +
Prolactin + + 0 0
Progesteron + + 0 0
PGF2α +
Somatostatin 0 + 0 +
Glucagon
Insulin
Gastrin
+ = Significant activity, 0 = no activity.
related release of hormones is due to an activation of the vagal
nerves. These hormones play an important role in adapting
maternal physiology and metabolism to the demands of
lactation.
The release of gastrin during nursing leads to growth of the
intestinal mucosa (Lankisch, 1980). This release is facilitated by
the vagally induced decrease of somatostatin levels, since this
hormone inhibits gastrointestinal functions (Efendic et al.,
1980). Thus the digestive tract adapts to a larger feed intake and
the digestive processes are optimized to meet the increased need
of energy during lactation.
VIP is produced in the gastrointestinal tract and in blood
vessels of the udder. Since VIP acts locally to dilate blood
vessels it helps allocate nutrient and hormone-rich blood to the
udder. In addition, udder skin temperature is increased as a
consequence of the increased blood flow. Stimulation of an
udder segment increases the skin temperature at the base of this
segment by more than 0.5 °C in comparison to the unstimulated
udder segments (Welch and Baxter, 1986; Algers, 1989). The
higher temperature caused by the increased blood flow may also
serve to increase sucking behavior.
The massage/suckling-induced release of insulin and glucagon
may serve to facilitate metabolic processes and transfer of nutrients
from maternal stores to the mammary gland. Glucagon seems to be
of particular importance as a catabolic agent during lactation, since
the activity of the hypothalamic–pituitary–adrenal axis and the
sympatho-adrenal system, normally associated with catabolic
processes, is partially repressed during lactation (Uvnäs-Moberg,
1996).
The finding of a quantitative relation between teat stimula-
tion and the release of prolactin, glucagon, VIP and (inverted)
somatostatin reported by Algers et al. (1991) supports the
concept of the restaurant hypothesis.
Measuring the duration and intensity of teat stimulation of
each piglet on each udder segment showed that both of these
parameters influenced the milk production during the first 3
days of lactation (Algers and Jensen, 1991). Further, nursing
frequency is shown to influence piglet weight gain (Spinka et
al., 1997; Auldist et al., 2000; Valros et al., 2002). Valros et al.
(2002) found that the duration (200–250 min/day) and
frequency (20–24 times/day) of nursing was rather stable for
Initiation Farrowing Lactation, Lactation,
of farrowing nursing not during
nursing
+? +? 0 0
+ + 0
0 + 0
0 0 0
+
+ +
+ +
 B. Algers, K. Uvnäs-Moberg / Hormones and Behavior 52 (2007) 78–85
each sow throughout lactation indicating the existence of a
stable individual nursing behavior pattern. This is probably due
to the amount of bodily resources that the sow can access.
Under commercial conditions, weaning usually takes place
when the piglets are 3–6 weeks of age depending on the
management practices applied, but under natural conditions
weaning is gradual and finishes at 14–20 weeks after birth
(Jensen and Recén, 1989).
Maternal metabolism during lactation
To allow for an increased energy demand during milk
production, the sow mobilizes her body reserves (see Quesnel
and Prunier, 1995). After parturition, the sow has to switch
from an anabolic to a catabolic state which prioritizes the
mammary gland energy output (Collier et al., 1984). This, of
course, is also related to feeding level and quantity of milk
output, but individual variations have been reported (Rojkitti-
khun et al., 1993b). Peak milk production occurs at 2–3
weeks post partum (Toner et al., 1996) when blood glucose
levels are low due to milk production (Rojkittikhun et al.,
1993a; Kraetzl et al., 1998). Measuring body weight loss,
however, Hultén et al. (1993) found a peak during the first
week of lactation but Rojkittikhun et al. (1993a) were unable
to demonstrate a correlation between prefeeding glucose levels
and body weight loss in the sow. However, sows are normally
fed diets rich in starch which is why glucose levels usually do
not reach low values (Kraetzl et al., 1998). As non-esterified
fatty acids (NEFA) are a product of fat metabolism and only
small amounts of them can be derived from feed intake, they
can be used as a sign of catabolic state (Armstrong et al.,
1986; Baidoo et al., 1992; Hultén et al., 1993). NEFA levels
in sows rise at the end of gestation and are highest during
mid- and late lactation (Kraetzl et al., 1998) although studies
are inconsistent (see e.g. Hultén et al., 1993; Rojkittikhun et
al., 1993b; Le Cozler et al., 1998). Valros et al. (2003) found
that sows differ in their metabolic state and milk production.
Sows with low piglet mortality were found to switch to a
catabolic state early in lactation in comparison to those with
high mortality and low litter weight gain, in which high NEFA
concentrations were found in the third week, when the
demand for milk production is at its maximum. Further, it was
shown that high baseline plasma oxytocin concentrations as
well as a high nursing-induced release of oxytocin were
associated with greater mobilization of bodily reserves of the
sow and faster piglet growth (Valros et al., 2004).
Interestingly, the milk ejecting hormone oxytocin has been
shown to stimulate several aspects of maternal behavior such as
caring behavior and bonding in several mammalian species (for
review, see Lim and Young, 2006). Oxytocin also adapts
maternal physiology and metabolism to the demands of
lactation. These effects are exerted in the brain and are caused
by a release of oxytocin from oxytocinergic neurons originating
in the paraventricular nucleus of the hypothalamus.
In sows the oxytocin-related increase in grunt rate occurring
before milk ejection is an expression of the oxytocin-related
maternal behavior. An oxytocin mediated increased secretion of
VIP from blood vessels in the udder, may dilate blood vessels in
83
the udder, thus increasing udder temperature and also facilitating
transportation of oxygen and nutrients to the mammary glands. In
addition, the vagally mediated increased activity of the gastro-
intestinal tract as well as the increased release of glucagon and
increased catabolic activity are expressions of oxytocin-mediated
effects aiming at integrating maternal behavior, physiology and
metabolism in the lactating sow.
The relations between the individual sow, its genetic
background and capacity to release hormones that influence
metabolism and milk output is complex and not completely
understood. Moreover, additional research is needed to better
understand all aspects of maternal behavior in pigs.
Conclusions
This review shows that much of the maternal behavior in sows is
governed by the release of specific hormones (see Table 1), but
mechanisms seem complex and most likely involve many other
nervous and hormonal systems. The sow's nursing behavior
ensures an even distribution of milk to her piglets. The piglets'
suckling behavior, however, is mainly a way to communicate their
individual nutritional needs. This communication is mediated
through tactile stimulation of hormonal release.
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