ELSEVIER Livestock Production Science 50 (1997) 75-87

The lactation cycle in the sow: physiological and

management contradictions
P.E. Hartmann *, N.A. Smith, M.J. Thompson, CM. Wakeford, P.G. Arthur
Department of Biochemistry, The Unir~ersityof Western Australia, Nedlands, WA6907, Australia

Abstract

Although lactation is an essential component of mammalian reproduction, each species has evolved unique lactation
strategies that optimise the growth and development of their young. However, these evolutionary strategies have imposed
physiological and environmental contradictions in commercial pig production. In the sow only extremes of either under- or
over-nutrition have been reported to influence mammogenesis and lactation, and the incompatibility between the zones of
thermal comfort for piglets and sows often results in ambient temperatures in commercial piggeries that impair either the
growth rate of piglets or milk production in sows. Of the factors investigated to date, litter size has the greatest positive
influence on total production of milk from sows. This observation strongly suggests that milk production is regulated at the
level of each mammary gland (e.g., autocrine control) rather than systemic metabolic processes (e.g., hormonal control).
While the short period of milk flow (10 to 20 s) during milk ejection could facilitate survival of larger litters, the restriction
of milk intake by the piglet due to this short period of milk flow, and the local inhibition of milk synthesis by autocrine
control mechanisms, may be important rate-limiting steps restricting potential milk production in domesticated sows.
Although the very tight control of milk flow imposed by the milk-ejection reflex and the ambient temperature requirements
of piglets could have provided an evolutionary advantage to larger litters, these factors may limit the expression of the
domesticated sow’s capacity to produce milk as well as her piglets’ potential for growth and development. It is important to
determine the significance of these subtle influences on pig production so that both experimental protocols and management
practices do not compromise expression of the full lactation potential of the sow. 0 1997 Elsevier Science B.V.

Keywords: Mammogenesis; Lactogenesis; Milk production; Involution

1. Introduction and develop into adults. In many species of verte-

brates, the adults can thrive in environments that lack
Sexual reproduction is fundamental to the evolu- suitable foods for their young. In contrast, mammals
tion of vertebrate species and depends, in part, upon nourish their young with milk that is synthesised
an adequate supply of food for the young to grow from the products of maternal digestion. Therefore,
they can reproduce successfully providing the mother
has adequate nourishment. Furthermore, because
* Corresponding author. Tel.: +61 8 9380 3327; fax: +61 8 mammals have the ability to deplete their body
9380 1148. reserves to supply the precursors for milk synthesis,

0301-6226/97/$17.00 0 1997 Elsevier Science B.V. All rights reserved,

PII SO301-6226(97)00076-6
76 P.E. Hartmann et al./ Livestock Production Science50 (1997) 75-87

an interruption in food intake by the mother does not It is now clear that the composition of milk is
immediately interrupt the food supply for her young. uniquely appropriate for the neonate at a time when
An extreme example of this ability is seen in the many of its systems such as the digestive, hepatic,
Southern Right whale (Bannister, 1995). These neural, renal, vascular, visual, skeletal and immune
whales filter small crustaceans from the water through systems are functionally immature. Milk is tailored
their large sieve-like baleen plates, and although they to augment the specific developmental defects of the
spend the summer months in their feeding grounds in young of each species and thereby facilitates optimal
the sub-Antarctic waters, every April/May they mi- growth and development during early postnatal life.
grate to Australia’s temperate southern coastline to Human milk appears to be at one extreme of milk
calve and mate before returning to their sub-Antarctic composition, with very low protein and sodium but
feeding grounds in October/November. The calves high lactose and oligosaccharides, whereas aquatic
are born mid-year with a birth weight of about one mammals, such as Blue whales and Harp seals, are at
tonne. The initial physiological drain of lactation on the other extreme with high protein and about 50%
the mother is enormous as food supplies do not of their milk consisting of fat (Table 1). The compo-
normally occur in coastal waters of Australia, so a sition of sow’s milk is closer to the average for the
Right whale fasts for 4 to 5 months while simultane- range of mammals studied to date. Furthermore, the
ously putting around 10 tonnes of weight onto her lactating mammary gland has a high synthetic capac-
calf. ity. For example, the energy output in milk repre-
Lactation has provided a niche for mammals in sents approximately 25% of the total energy intake
the evolutionary tree and, as a result, there are over in the maternal diet for women exclusively breast-
4000 species of mammals (Oftedal, 1984) living in feeding single infants and as much as 50% of the
an amazing variety of habitats. There is not only total energy intake for those breastfeeding twins
great variation in the environments that mammals (Saint et al., 1986). Lactation in the sow is even
live in, but there also is great variation in the matu- more demanding as 100% of maternal dietary energy
rity of mammals at birth. Whereas the guinea pig is intake in the lactating sow may be required for milk
relatively mature at birth, the Joey kangaroo is very synthesis (Pluske et al., 1995b). So to some extent
immature at birth. There also is great variation in the the sow is like the whale, normally using consider-
rate of postnatal development. For example, a l- able body reserves to support the energy demands of
month old piglet is much more mature than a 3-month lactation. Furthermore, it can be concluded that the
old wombat (Reardon, 1993). Given these environ- milk of each species is uniquely appropriate for their
mental and developmental differences, it is not sur- young and that lactation represents a very significant
prising that milk composition varies greatly from energy cost to the mother.
mammal to mammal (Table 1). From these considerations, it can be argued on
both evolutionary and physiological grounds that
lactation is an essential component of mammalian
reproduction. However, the direction of research
funding contradict the evolutionary and physio-
Table 1
Variation in milk composition between mammals
logical importance of lactation to pig reproduction
by directing most attention towards either early
Soecies Lactose (g/l) Protein (g/l) Fat (g/l)
weaning (21 days) or segregated early weaning at 10
Womana 68 8 41
to 14 days. The short-term expediency of segregated
Horsesa 69 19 13
Piga 50 56 83
early weaning may have considerable implications
Cow” 46 32 37 for the future of the pig industry as a greater under-
Rabbitb 18 103 152 standing of the importance of sow lactation emerges.
Blue whaleb 13 119 409 In this review, we propose to compare the physi-
Harp sealb 8 77 53.5
ology of lactation in the sow with that of other
aFrom Oftedal (1984). species. The consistencies and inconsistencies be-
bFrom Oftedal and Iverson (1995). tween the sow and other species will be considered
 P.E. Hartmann et al. /Lic,estock Production Science 50 (19971 75-87 77

in relation to the lactation strategy of the pig and the the least body reserves (low-low; 152 g/piglet/d).
factor(s) limiting the lactation potential of the sow. However, provided the sows had a high feed intake
during lactation, variation in body reserves at farrow-
ing did not affect piglet growth. Piglets from sows
with high-high, medium-high or low-high levels of
2. Lactation cycle
nutrition during pregnancy-lactation all had similar
growth rates: 2 10, 2 10 and 219 g/piglet/d, respec-
Conception marks the beginning of the lactation
tively. Thus, the work of Mullan and Williams (1989)
cycle (Fig. l), which includes: growth and develop-
suggests that reduced food intake during pregnancy
ment of the mammary gland (mammogenesis), initia-
does not compromise mammogenesis to the extent
tion of synthesis of unique milk constituents such as
that it affects subsequent milk production.
lactose, P-lactoglobulin, cu-lactalbumin and the ca-
Indeed, there appears to be a contradiction be-
seins during pregnancy (lactogenesis l), initiation of
tween nutrition in pregnancy and nutrition during
copious milk secretion at fan-owing (lactogenesis 21,
lactation. Recent research suggests that the more a
established lactation (lactation) and regression of the
sow eats during pregnancy the less she eats during
mammary gland after weaning (involution).
lactation (Mullan and Williams, 1989). Furthermore,
Head and Williams (1991) observed impaired mam-
mogenesis in gilts given a low protein-high energy
3. Mammogenesis diet during pregnancy. Although, this is not necessar-
ily a consistent finding (Smits et al., 19951, it is
Hormonal control of growth of the mammary probably not desirable to have pregnant sows on a
gland of the sow during pregnancy has not been diet which restricts lean tissue growth as this also
extensively investigated (Hartmann and Hughes, may compromise mammogenesis.
1996). The influence of nutrition of the pregnant sow
on mammary development during pregnancy and the
outcome of the subsequent lactation also has not 4. Lactogenesis 1
been completely resolved. Mullan and Williams
(1989) varied the body reserves of pregnant gilts at Although the timing of the first phase of lactogen-
farrowing by feeding either high, medium or low esis (lactogenesis 1) has not been extensively studied
diets during pregnancy. In addition, after far-rowing it appears to occur later in pregnancy in the sow
the sows received either high or low levels of feed. (Willcox et al., 1983; Dodd et al., 1994) than in
Severe restriction of feed intake during lactation other species, such as women (Kulski and Hartmann,
caused a marked reduction in litter growth during the 1981) and cows (Hartmann, 1973). Dodd et al. (1994)
fourth week of lactation, particularly in sows with reported that P-lactoglobulin is the earliest de-
tectable specific milk product found in sow’s blood
during pregnancy (5 weeks pre-parturn), with de-
tectable increases in most other milk constituents,
Birth such as a-lactalbumin (Dodd et al., 1994), lactose
Gestaticn/ Tb> and casein only occurring during the last few days of
pregnancy (Whitely et al., 1990). Furthermore, it is
Conception9 O”i..Gl”G0
very difficult to express any mammary secretion
%
n
Mammogenesis
from the sow until about 24 h before farrowing and
Lactogenesis 1 Involution
/I the earliest secretion is usually a clear straw colour.
The late initiation of lactogenesis 1 in the sow
(Willcox et al., 1983) may be, in part, attributed to
Lactation the fact that the sow lacks a placental lactogen
Fig. 1. Mammary development and the stages of the lactation (Forsyth, 1986); a h ormone that appears to be associ-
cycle in the sow. ated with growth and development of the mammary
78 P.E. Hartmann et al./ Livestock Production Science 50 (1997) 75-87
gland during pregnancy in a number of species,
including the cow (Byatt et al., 1994).
5. Farrowing
The normal length of gestation in the sow varies
from 110 to 120 days, and it is apparent that initia-
tion of farrowing is under hormonal control. With-
drawal of progesterone (Ash and Heap, 1975) cou-
pled to an increase in prostaglandins and relaxin
(Whitely et al., 1990) are central to the control of
farrowing. oxytocin also may be involved, but its
exact role remains a puzzle (Gilbert et al., 1994).
In established lactation, acute changes in intra-
mammary pressure reflect transient changes in con-
centration of oxytocin in the sow’s blood (Ellendorff
et al., 1982). Between sucklings in established lacta-
tion, blood oxytocin ranges from < 0.2 to 6.0
pU/ml (Forsling et al., 1979; Ellendorff et al.,
1982; Holmes, 1991). However, oxytocin increases
to maximum concentrations of 3.9 to 21.2 @J/ml
just prior to the intra-mammary pressure increase
and milk ejection (Ellendorff et al., 1982; Holmes,
1991).
During farrowing, the concentration of oxytocin
in blood (Ellendorff et al., 1979; Forsling et al.,
1979; Taverne et al., 1979) is much higher (IO-265
pU/ml) such that lowest concentrations of oxytocin
over farrowing are at least as high as the maximum
concentrations reported for established lactation. The
concentration of oxytocin varies considerably over
farrowing, although the association between changes
in concentration of oxytocin in blood and birth of
each piglet is not convincing (Gilbert et al., 1994).
It is difficult to relate changes in concentration of
oxytocin in blood to birth of the piglets as oxytocin
has a short half-life. As an alternate approach, Ellen-
dorff et al. (1982) introduced the measurement of
changes in intra-mammary pressure as a qualitative
in vivo ‘bio-assay’ for oxytocin in the SOW, and
subsequently Gilbert et al. (1994) investigated the
changes in intra-mammary pressure during farrowing
with puzzling results-an increase in intramammary
pressure occurred consistently in the minute after
delivery of each piglet.
We also have investigated variations in intra-
mammary pressure in sows over farrowing (Fig. 2).
Fig. 2. Intra-mammary pressure changes during the birth of a
piglet. The plot shows (a) intra-mammary pressure increases
coinciding with movement of the sow, (b) expulsion of the piglet,
and (c) post-delivery increase as the sow lay motionless (Thomp-
son, Wakeford and Hartmann, unpublished observations).
The mammary duct of unrestrained conscious sows
cannulated with a Venflon@ 2 i.v. (22 G/0.8
mm OD, luer-lock) cannula (Viggo Products, Swe-
den) attached to a disposable pressure transducer
(COBE, Lakewood, CO, USA) and the amplified
pressure signal was captured by either a chart recorder
or a data recording system (MacLab/4e, ADInstru-
ments, Sydney, Australia). The cannula was held in
position manually throughout the measurement of
intra-mammary pressure. Irma-mammary pressure
was recorded over 61 normal deliveries of piglets
(Smith, 1994). There was a pressure peak in the
minute prior to four of the deliveries and at the same
time as four of the deliveries (i.e., to within 5 s of
delivery). During these eight deliveries, movement
of the sows obscured the measurement of changes in
intra-mammary pressure after delivery. For the re-
maining 53 deliveries, the sows remained at rest and
all showed an increase in intra-mammary pressure
within a minute after delivery (Table 21, suggesting
that a release of oxytocin occurred after the expul-
sion of a piglet. In this connection, it is of interest
that Fraser (1984) observed that colostrum ejections
Table 2
Characteristics of the intra-mammary pressure peaks associated
with the birth of oielets
Peaks Delay a (seconds) Duration (seconds)
Before Delivery (n = 4) 32 f 5.1 28f7.0
At Delivery (n = 4) 23*5.5**
After Delivery (n = 53) 18 + 7.4 37f9.0
’ * P < 0.01 from duration of the peaks after delivery.
aDelay refers to the delay between the end of the peak and the
delivery (before delivery peaks) or between the delivery and the
start of the peak (after delivery peaks).
 P.E. Hartntann et al. / Lic,estock Production Science 50 (1997) 75-87
began lo-25 s after the birth of 10 out of 23 piglets.
However, in his hand-milking study, 11 births were
not accompanied by any noticeable increase in
colostrum yield and two births occurred during the
initial periods of high yield. These findings also
suggest that a release of oxytocin occurs immedi-
ately after the birth of the piglet.
The delay between expulsion of the piglet and
pressure peak (18 f 7.4 s) was similar to the delay
measured following injection of oxytocin (50 mu>
into the ear veins of sows either during farrowing
(20 + 9.3 s, II = 6) or at 7 days of lactation (21 + 4.1
s, n = 6) (Smith, 1994). Thus, we concluded that the
delay between delivery of the piglet and the increase
in intra-mammary pressure was due to the circulation
time from the ear (or the neurohypophysis) to the
mammary gland. These observations appear to con-
tradict the accepted action of oxytocin. If oxytocin
promotes uterine contractions, why is the effect of
oxytocin release observed immediately after the birth
of the piglet?
Ferguson (1941) measured uterine motility in the
8 to 48 h post-partum rabbit. He found that dilation
of either the uterine horn or cervix increased the
motility of the exposed uterus when both the spinal
tract and neurohypophysis were intact, but not when
either had been destroyed. On the other hand, vagi-
nal stretch had a varied response. He attributed this
increase in uterine motility to be due to a reflex
release of oxytocin from the neurohypophysis in
response to the stretching of the uterus. This re-
sponse has become known as the Ferguson reflex
(Folley, 1969). The consistent observation of pres-
sure peaks after the piglet was fully expelled (Table
2; Gilbert et al., 1994) suggests that, if oxytocin was
released from the neurohypophysis in association
with birth of the piglet, it must have been released as
the piglet was fully expelled and not while the piglet
was stretching either the uterus or the cervix. It
follows that it was the release of the stretch stimulus
which caused the reflex release of oxytocin rather
than the application of this stimulus to the
cervix/vagina. Upon closer examination of Fergu-
son (1941), there is further evidence to support our
hypothesis. In several of his experiments, there was
increased uterine activity after deflation of either the
vaginal or uterine dilators.
Debackere and Peeters (1960) reported that milk
19
ejection in the cow occurred after a ‘lag period’ of
3-4 min in response to the l-2 min inflation of an
airbag within the vagina. Roberts and Share (1968)
rhythmically inflated and deflated a balloon for 2
min and then in the third minute, kept the balloon
inflated at 100 mm Hg in lactating ewes during an
oestrous cycle. They found that there was a signifi-
cant increase in the concentration of oxytocin in the
plasma in both the first and third minutes of stimula-
tion as compared to pre-stimulation concentrations,
although the increase was more pronounced in the
first minute. The deflation of the balloon may have
been the stimulus for oxytocin release in both stud-
ies. A recent report on parturition and vagino-cervi-
cal stimulation in the ewe showed that the reflex
release of oxytocin could be induced by the use of a
plastic probe that was rhythmically inserted into, and
withdrawn from, the vagina (Kendrick et al., 1991).
Thus, the results of earlier studies, including those of
Ferguson ( 194 l), also are consistent with the hypoth-
esis that the relaxation of a stretched cervix/uterus
provides the stimulus for the release of oxytocin.
Further support for this hypothesis was provided
by measurement of the intra-mammary pressure as-
sociated with internal examinations of farrowing
sows. Following nine internal examinations, pressure
peaks were only recorded in the minute after the
withdrawal of the animal attendant’s hand from the
reproductive tract of the sows. The mean time inter-
val from completion of the internal examinations to
the pressure peak was 21 * 13 s and was similar to
the time interval observed after the full expulsion of
the piglet at normal farrowing (Table 2) except that
on a number of occasions multiple peaks were
recorded, whereas after the birth of a piglet multiple
peaks were rarely recorded.
We concluded that release of oxytocin was not
inducing birth of the piglet but rather was the result
of birth of the piglet. That is, it is not distension of
the birth canal but rather contraction of a distended
birth canal that stimulates release of oxytocin. There-
fore, the physiological strategy for release of oxy-
tocin at birth is probably related to its release in the
hypothalamus and its central nervous system effects
on maternal behaviour rather than peripherally influ-
encing the birth process.
It is of interest that although this reflex was
originally described by Ferguson (1941), it has had
80 P.E. Hartmann et al. /Livestock Production Science 50 (1997) 75-87
practical significance for centuries. For example,
Kolben 1731 (cited by Cowie et al., 1980) observed
the practical application of this reflex in Southern
Africa N To make a refractory cow yield milk. His
description of the process was as follows:
But if the cow is too cunning to be cozen’d in that
way (by bringing a Counterfiet calf to the Teats),
their last Shift to make her yield Milk is, after tying
her Hind-Legs together to prevent her Kicking, blow
with all their Might into the Vulva. This is done by
Men as well as Women.
6. Lactogenesis 2
Parturition and the second stage of lactogenesis
(lactogenesis 2) are two closely coupled events, It
has been recognised for many years that initiation of
copious milk secretion at parturition is under en-
docrine control in mammals. Indeed, the pioneering
work of Kuhn (1969) established that progesterone
withdrawal was the trigger for lactogenesis 2 in the
rat. Subsequently, Nicholas and Hartmann, 1981a,b,
established the temporal relationship between with-
drawal of progesterone during late pregnancy and
lactogenesis 2 in the rat. They demonstrated that
when progesterone fell to low levels about 24 h
before parturition, there was an increase in lactose
synthesis at parturition.
Evidence supporting the progesterone withdrawal
theory in the sow has been reviewed recently (Hart-
mann and Holmes, 1989; Hartmann et al., 1994).
Briefly, there is a negative relationship between con-
centration of progesterone in blood and concentra-
tion of lactose in milk at both normal farrowing and
when farrowing is delayed by administration of pro-
gesterone. Furthermore, de Passille et al. (1993)
reported a reduced growth rate and increased mortal-
ity in piglets from sows that had higher concentra-
tions of progesterone in their blood immediately
after farrowing.
In other mammals such as women, lactogenesis 2
occurs whether or not the baby suckles at the breast
(Kulski et al., 1978). Therefore, milk withdrawal is
not required for lactogenesis 2 in these species.
Closer examination of initiation of lactation suggests
that an additional level of control may exist within
each sow. The fat content of colostrum increased
significantly in suckled compared to unsuckled
mammary glands (Atwood and Hartmann, 1993),
while concentration of lactose and metabolites asso-
ciated with lactose synthesis also increased in the
colostrum from suckled compared to unsuckled
mammary glands during the post-partum period
(Atwood et al., 1995). Thus it is probable that,
whereas the withdrawal of progesterone releases the
systemic hormonal block to lactogenesis 2, either
removal of colostrum by the piglets or the local
effect of sucking switches on increased milk (lactose
and fat) synthesis in the individual mammary glands.
Our investigations suggest that a relatively large
volume of colostrum is ingested initially by most
piglets within an hour of birth (Fig. 3) and that
piglets tend to develop a preference for the teat from
which they first obtained colostrum. Piglets are born
agammaglobulinanaemic and with virtually no fat
reserves. The strategy of teat preference and milk
removal triggering lactogenesis 2 would ensure that
the last born piglet obtained colostmm of similar
concentration of immunoglobulin to that available to
the first born piglet, while the rapid increase in fat
and lactose triggered by milk removal ensures that,
after birth, each piglet obtains a crucial supply of
energy as it establishes itself on its preferred teat.
80r
%
0 30 60 90 120 150 180 210
Time after first delivery (min)
Fig. 3. Changes in the weight, relative to the birth weight, of the
individual piglets of one litter for 3.5 h after delivery of the first
piglet (Smith, 1994). Each delivery time is indicated by a vertical
line at the bottom of the figure with the symbols indicating the
piglet.
 P.E. Hartmann et al. /Livestock
7. Lactation
The major changes in the composition of sows’
milk occur during the first 72 h after farrowing and
this is characterised by a major reduction in concen-
tration of IgG in milk (see reviews by Hartmann and
Holmes, 1989; Hartmann et al., 1995).
Based on studies in dairy animals, it would be
expected that the diet of the lactating sow would be
one of the most important determinants of milk
production. However, the sow does not respond as
might be expected.
Unlike dairy animals, there is not a consistent
relationship in sows between dietary energy intake
and milk production measured by the growth rate of
standard sized litters (Williams, 1995). Since the ad
libitum fed piglet has the capacity to grow at up to
twice the rate of the normal suckling piglet (Pluske
et al., 1995a), it is unlikely that this variable re-
sponse in milk production is due to piglet satiety. On
the other hand, milk production in the sow is linearly
related to litter size and this relationship is due to an
increase in the number of functional glands (Auldist
and King, 1995). Indeed, litter size has a greater
influence on milk production in the sow than any
other factor studied to date.
Recently, Auldist et al. (1995) demonstrated that
suckling demand can greatly increase milk produc-
tion from individual glands. They measured milk
production from 6 to 27 days of lactation in three
treatment groups of sows. Group 1 sows had six
functional teats and a litter size of six, group 2 sows
had 12 functional teats and a litter size of 12, and
group 3 sows had six functional teats and were cross
suckled with two litters of six piglets at 30 min
intervals. The six glands of the sows in the cross-
suckled group supported similar piglet growth (2 195
g/d) to that in the 12 glands of the group of sows
with litters of 12 piglets (2571 g/d). The additional
production from the glands of the cross-suckled group
was the result of hypertrophy of the glandular tissue
(mean gland weight 124.0 g DM and 64.9 g DM,
respectively). Since piglets have a strong teat prefer-
ence, it can be concluded that total milk production
of the sow is closely related to the number of
functional glands and to the suckling intensity and
resultant milk withdrawal from individual glands.
Thus, the growth potential of the piglet is limited by
Production Science 50 (19971 75-87 81
the intake of milk from their preferred gland. Indeed,
under increased suckling pressure, individual glands
within sows have the potential to produce almost
double their normal milk production. It can be con-
cluded that milk production in the sow is regulated,
in part, within individual glands rather than respond-
ing to systemic metabolic control mechanisms.
The possibility of local regulation within each
gland together with the teat preference of piglets
suggests that suckling behaviour and, in particular,
the milk-ejection reflex, might limit the transfer of
milk from the sow to the piglet. The sow has the
highest level of control over milk ejection of any
mammal studied to date. Piglets suckle once every
40 to 60 min and observational and intra-mammary
pressure measurements suggest that milk is only
available to the litter for a very short period of time
about 10 to 20 s at each suckling. Folley and Knaggs
(1966) as well as Ellendorff et al. (1982) demon-
strated that milk ejection in the sow, as in other
mammals, is triggered by the release of oxytocin.
However, despite the short half-life of oxytocin in
blood, milk flow ceases in the sow while significant
concentrations of oxytocin remain in the circulation.
If a piglet is removed before a suckling, milk is not
accessible for expression from the unsuckled gland
once the flow of milk in the suckled glands has
ceased. Thus, compared to other mammals, milk
ejection in the sow may be more complex consisting
of both ‘on’ and ‘off’ mechanisms. The ‘on’ mecha-
nism is the classical neuro-hormonal reflex of oxy-
tocin triggering contraction of the myoepithelial cells.
Whereas, an opposing ‘off’ mechanism acutely stops
the flow of milk within a few seconds of oxytocin
triggering contraction of the myoepithelial cells and
regardless of whether or not milk has been removed
from the gland. In this context, we propose that the
term ‘milk ejection’ should be used to describe the
neuro-hormonal reflex, which stimulates the neuro-
hypophysis to release oxytocin and, in turn, causes
contraction of the myoepithelial cells surrounding
the alveoli and expulsion of milk. We also propose
the term milk ‘let down’ should be used to specifi-
cally describe the length of time that milk is avail-
able to the piglets following milk ejection.
We measured intra-mammary pressure in one
gland as the piglets suckled the other glands in 11
sows at 12- 14 days of lactation and at 19-24 days
82 P.E. Hartmann et al./Livestock Production Science 50 (1997) 75-87
IL
2Gfi-z
Fig. 4. Intra-mammary pressure changes during suckling in a sow
at 14 days of lactation (Thompson, Wakeford and Hartmann,
unpublished observations).
of lactation. Since the piglets stopped fast sucking (3
jaw movements per second) before the pressure re-
turned to basal levels, the duration of milk ejection
was arbitrarily defined as the time from the initial
sharp increase in pressure to the peak pressure re-
sponse (Fig. 4). On this basis, duration of milk
ejection was 14.0 s with a coefficient of variation of
27%. Therefore, it is possible that the short duration
of milk flow at each suckling restricts the ability of
the piglet to remove all of the available milk from
the gland and that variation in duration of milk flow
could explain a significant proportion of variation in
milk production. If this is so, then it is probable that
local (autocrine) inhibitory mechanisms (Peaker,
1995; Wilde et al., 1995) control the rate of milk
synthesis within individual glands in the sow de-
pending on how much milk the piglet has been able
to remove from the gland in this limited period of let
down. The autocrine control of milk synthesis would
explain both the linear relationship between litter
size and milk production, and increased production
from glands subjected to increased suckling demand
(Auldist et al., 1995). Furthermore, autocrine control
would restrict the influence of diet and exogenous
hormonal administration on milk production in the
sow.
The above observations coupled with the finding
that milk ejection only occurs if the majority of the
piglets are stimulating the teats means that it is very
difficult for dominant piglets to obtain milk from
more than one teat at a suckling. Thus, the restriction
imposed by the tight control over milk ejection in the
sow is a strategy that promotes survival of large
litters.
One of the most puzzling contradictions in lacta-
tion in swine relates to ambient temperature. The
lower temperature for the zone of thermal comfort of
the piglets is greater than 30°C whereas, the higher
temperature for the sow’s zone of thermal comfort is
less than 24°C. Thus, there is an incompatibility
between the ideal ambient temperature for sows and
piglets. A number of workers have shown that if
ambient temperature of the sow is increased to near
the zone of thermal comfort for the piglets, both food
intake by the sow and milk production decrease. For
example, Mullan et al. (1992) increased the tempera-
ture of the sow from 20°C to 30°C and food intake
decreased from 4.05 to 3.13 kg/day and milk pro-
duction decreased from 8.88 to 7.53 l/day. Black et
al. (1993) raised the ambient temperature of sows
from 18°C to 28°C and reduced voluntary feed intake
by 40% and milk production by 25%. The decline in
milk production was more than would be expected
from an equivalent decline in food intake for sows
housed within their zone of thermal comfort (Black
et al., 1993) and this has been attributed to heat
stress diverting blood flow to the skin with a con-
comitant decrease in blood flow through other tis-
sues including the mammary gland (Williams et al.,
1994).
Whatever the physiological explanation, it is clear
that the ideal ambient temperature for the sow (Black
et al., 1993) is not the ideal temperature for her
piglets (English and Edwards, 1996) and this im-
poses a contradiction for swine management in in-
tensive piggeries. It is difficult to provide a micro-
environment for piglets which provides appropriate
access to the sow yet has minimal air movement and
is at least 10°C higher than the ideal environmental
temperature for the lactating sow. Since newborn
piglets have very limited energy reserves and indi-
vidual piglets cannot survive cold stress for more
than a few hours, the selective advantage of this
environmental contradiction is not clear. However,
perhaps nest building by feral (and range managed)
sows more closely approximates the micro-environ-
mental requirements of both the sow and her piglets.
In this connection, Bruce Mullan (Personal Commu-
nication, 1996) in Western Australia has demon-
strated that optimising the ambient temperatures for
 P.E. Hartmann et al. /Livestock Production Science 50 (1997) 75-87
both the sow and her piglets in an intensive piggery
can have profound effects on improving the growth
rates of piglets.
If the short period of milk flow at a suckling, and
the different ambient temperature requirements of
sows and piglets are currently important rate-limiting
factors restricting milk production in the sow, both
endocrine manipulation and nutritional supplementa-
tion would be unlikely to greatly improve the lacta-
tional performance of sows.
8. Involution
In commercial piggeries, weaning is an acute
event that has occurred at progressively earlier stages
of lactation as the industry moves from 3-week
weaning to segregated early weaning at around 10
days of lactation.
Within an hour or so of removal of the litter,
concentration of prolactin in the sow’s blood begins
to fall and reaches low levels within 4 to 6 h. During
the first 24 h after weaning, there are significant
changes in concentration of a number of the metabo-
lites associated with synthesis of lactose including
glucose, glucose 6-phosphate, glucose 1-phosphate,
uridine S-diphosphate, orthophosphate and adeno-
sine S-phosphate (Atwood and Hartmann, 1995).
These changes occur 24 to 48 h before the detection
of increased apoptosis (programmed cell death) after
weaning even in rats (Bielke et al., 1995). It must be
emphasised that despite the current interest in apop-
tosis after abrupt weaning, metabolic changes still
provide first indications of a change in cell function
and that these changes occur before apoptotic
changes. Thus, the link between altered metabolic
activity of the cell and apoptosis should provide a
fertile area for investigation.
9. Conclusion
The apparent physiological and environmental
contradictions associated with the lactation cycle in
sows reflect strategies favoured during the evolution
of the lactation phase of the reproductive cycle in the
feral sow. Although the very tight control of milk
flow imposed by the milk-ejection reflex and the
83
ambient temperature requirements of piglets could
have provided an evolutionary advantage to larger
litters, these factors may limit expression of the
domesticated sow’s capacity to produce milk as well
as her piglet’s potential for growth and development.
It is important to determine the significance of these
subtle influences on pig production so that both
experimental protocols and management practices do
not compromise expression of the full lactation po-
tential of the sow.
10. Discussion
Ron Kensinger: Does the first intake of colostrum
by each newborn coincide with the presence of the
next pig in the cervix? I wonder if you would
comment on the role of oxytocin to make sure that
colostrum is available for that pig?
Peter Hurtmann: In sows, colostrum is generally
available. Oxytocin levels are high and considerable
amounts of colostrum can be expressed at any time.
So the firstborn piglet really does not necessarily
need milk ejection to get colostrum. We could not
associate the increase in the weight of the piglet with
actual milk ejection or increase in mammary pres-
sure for the initial piglets, but once you got halfway
through birth of the litter, you start to get some sows
having a definite milk ejection and increase in mam-
mary pressure. This was a suckling effect. So oxy-
tocin release at suckling could be superimposed on
the normal release of oxytocin associated with the
birth of each piglet. The interesting thing is that if
you take the amount of oxytocin that is required to
elicit an increase in mammary pressure during estab-
lished lactation, and you inject that small amount of
oxytocin intravenously into a sow during farrowing
you will get an increase in mammary pressure, de-
spite the fact that there are extremely high levels of
oxytocin there in the first place. So it really is a bit
of a puzzle. In summary, I think colostrum is avail-
able for piglets from the start of farrowing but as the
farrowing proceeds you start to get definite milk
ejection occurring in response to the piglets suckling.
Martin Tang S@rensen: You told us that piglets
have the capacity to grow much more than they do
when they are with the sow, and also that the sow
has much more capacity for milk production, than
84 P.E. Hartmnn et al. /Livestock Production Science 50 (1997) 75-87
she shows when she has her litter. So why is it that
the litter doesn’t grow more and the sow give more
milk at the same time when they both have the
capacity to do so?
Peter Hartmann: I think that is interesting. Allen
Tucker said yesterday that speculation was a good
thing. So let us speculate: What I think is occurring
is that the sows have evolved to produce large litters.
Now, the short period of milk availability at a suck-
ling switches off consistently in all glands at the
same time, so that the more dominant piglet really
does not have enough time to consume the milk
available to it and then go and attack its neighbour
and get its milk as well. So it is actually giving the
less dominant piglet a chance to have an equal
suckling period. Now in doing that it is actually
restricting the milk production capacity of the sow. If
you do not take milk from the gland, by stopping the
piglet from suckling, at the end of that 12 s you
cannot get any more than a couple of drips of milk
out of the gland, but during the 12 s you could squirt
it a couple of meters. It just shuts off whether or not
the milk is being removed. So that may be regulating
the amount of milk availability rather than the actual
synthetic capacity of the sow. The benefit of this
may be to allow the less dominant piglets to survive
against the dominant piglets. Just a suggestion!
Nathalie Trottier: A question about the anterior
and posterior glands. Do you have words of wisdom
as far as capacity of the glands to produce more milk
in the anterior versus the posterior glands, or is that a
behavioural relationship for the perhaps more domi-
nant pigs migrating in the cranial direction of the
gland at birth?
Peter Hartmann: There has been quite a lot of
work done on actual production along the glands.
One of the real problems is the difficulty in measur-
ing milk production in the sow. Weigh-suckle-weigh
is quite difficult. Deterium oxide estimates are based
on a 3 or 4 day duration. So there is a lot of noise in
any milk production measurements that have been
done on the sow. We do not necessarily see a
gradation from teats 1 and 2 to 5 and 6 that other
people have reported, so I am not really sure whether
that is a real thing or not. Some suggest that it is, but
I do not know why this might occur. We certainly
have not seen the firstborn pigs getting out and
getting on to the front two glands. They tend to
sample a lot of glands and then they stop at the gland
where they seem to first get colostrum-not the first
gland they get to. But I am not sure about all of this,
Nathalie Trottier: It has been my observation and
many other people’s observation that with smaller
litter size the posterior part of the mammary gland
involutes before the anterior part. I think it deserves
further investigation.
Peter Hartmann: That is true, yes.
Denis Petitclerc: If I understand right, you have
shown a decrease in milk production during heat
stress while there was an increase in lactose concen-
tration in milk?
Peter Hartmann: We used deterium oxide to mea-
sure milk production, so it was the average over a
3-day period, whereas the concentration of lactose
went up and down during that period. We could not
get day by day estimates of milk production.
Denis Petitclerc: You do not have data that shows
when volume decreased concentration of nutrients
increased. But nutrients may have increased in milk.
Could that be a way for the sow to preserve water
while trying to maintain nutrient supply to the piglets
as much as possible?
Peter Hartmann: That could be. We did notice
that the sow’s suckling behaviour also was impaired
at that time, so we do not know whether she was
giving the piglets as good access to the gland as an
unstressed animal.
Denis Petitclerc: The sow regulates suckling time
for the piglet, but do you have any evidence whether
she also regulates the amount of milk given to the
piglet, which regulates their growth rate? And can
she purposely limit growth rate of the piglet?
Peter Hartmann: I think that is a good point
because we really do need to understand more about
milk ejection in the sow. If it increased from 12 to
15 s of flow it could make a big difference in milk
intake by the piglet. I think this is an area that really
needs a lot more study to see whether that is a
potential regulatory system for the sow.
Chantal Farmer: You mentioned that the ejection
reflex stops even though oxytocin levels are still
high. Do you have a suggestion as to what might
stop this ejection reflex, and what might be done to
prolong the reflex?
Peter Hartmann: Sows have made me rethink the
whole milk-ejection reflex because we have always
 P.E. Hartmann et al. / Lioestock Production Science 50 (1997) 75-87
just thought of it as a one-way thing. Namely, it
contracts the myoepithelial cells and then in some
way they relax and that is the reflex. But in the sow;
the relaxation is so precise and it occurs in all glands
at one time makes me think that there may be an
‘off’ mechanism. 1 really have no idea what that
‘off’ mechanism might be, but I think it is something
that needs further investigation.
Malcolm Peaker: You looked at relaxin in suck-
ling, and we have a manuscript in press now where
we injected relaxin interarterially in goats and looked
at responsiveness to oxytocin. The data show huge
variation in accordance with the reproductive cycle. I
just wondered whether your cut-off mechanism in-
volves relaxin? The second question: Do you know
whether the period of milk ejection increases with
increasing age of the sow because according to
parental investment theory it should (you know
younger litters being in competition with the litter-
mates as yet unborn). In a very old sow she should
be willing to invest far more in her later litters.
Peter Hartmann: We thought that relaxin might
be the off mechanism. We looked at relaxin, and
more relaxin is released at suckling than between
sucklings. However, the release at suckling is not
precisely related to the switch-off of the actual ejec-
tion of milk. The lack of a relationship could be due
to our sampling situation because the time is amaz-
ingly short when you have only got a few seconds of
let-down. I believe we should distinguish between
milk ejection and milk let-down, with milk ejection
being the natural priming process and milk let-down
being the duration and availability of milk. We are
now just starting to do experiments on milk let-down.
We have got a better apparatus for measuring mam-
mary pressure, and we want to look at comparisons
within and between sows in the let-down period.
Dieter Schams: Have you studied mechanisms
around the sphincter and whether the teat itself is
closing after 12 s or what?
Peter Hartmann: If you put cannula up into the
gland about 2.5 mm, pressure certainly decreases at
the time milk let-down switches off. So it is not just
local around the teat, and certainly you can go back
in and try to manually express milk from the sub-
stance of the gland, and you can only get a couple of
drops. I do not know how the shut-off mechanism
works.
85
Acknowledgements
We thank the Pig Research and Development
Corporation and the Pig Industry Compensation Fund
(WA) for financial support.
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