Brill Behaviour: This Content Downloaded From 175.45.191.251 On Thu, 04 May 2017 08:57:32 UTC

Maternal Behaviour in Domestic Sheep (Ovis aries): Constancy and Change with Maternal
Experience
Author(s): Catherine M. Dwyer and Alistair B. Lawrence
Source: Behaviour, Vol. 137, No. 10 (Oct., 2000), pp. 1391-1413
Published by: Brill
Stable URL: http://www.jstor.org/stable/4535781
Accessed: 04-05-2017 08:57 UTC
REFERENCES
Linked references are available on JSTOR for this article:
http://www.jstor.org/stable/4535781?seq=1&cid=pdf-reference#references_tab_contents
You may need to log in to JSTOR to access the linked references.
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted
digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about
JSTOR, please contact support@jstor.org.
Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at
http://about.jstor.org/terms
Brill is collaborating with JSTOR to digitize, preserve and extend access to Behaviour
This content downloaded from 175.45.191.251 on Thu, 04 May 2017 08:57:32 UTC
All use subject to http://about.jstor.org/terms
MATERNAL BEHAVIOUR IN DOMESTIC SHEEP (OVIS ARIES):
CONSTANCY AND CHANGE WITH MATERNAL EXPERIENCE
by
CAXIHERINE M. DWYERI) and ALISTAIR B. LAWRENCE2)

(Behavioural Sciences Department, Animal Biology Division, SAC, King's Buildings, West
Mains Road, Edinburgh, EH9 3JG, UK)
(Acc. 9-VI-2000)
Summary
Individual differences in the way ewes behave towards their lambs may help to identify
the sources of variation in maternal care. In this study we investigated how the pattern of
maternal behaviour shown by an individual animal changed across parities in domestic sheep.
The maternal behaviour of individual animals (Scottish Blackface and Suffolk sheep), in the
first 2 hours after lambing, was compared when lambing as primiparous and multiparous
ewes. More rejecting behaviours (e.g. withdrawal, aggression, lack of co-operation with
lamb sucking attempts) were expressed by ewes in their first parity than in subsequent
parities. Behaviours associated with affiliation and bonding (e.g. grooming attention) were
not, however, significantly influenced by parity within individual ewes. Principal Components
Analysis of maternal behaviours was carried out on behavioural data from primiparous and
multiparous ewes. In common with studies in primates, behaviours in both primiparous
and multiparous ewes were clustered on two main axes or Factors, labelled 'maternal
rejection' and 'maternal care/warmth'. However, in multiparous ewes, a third Factor, labelled
'aggression', was found whereas in primiparous ewes aggressive behaviours loaded on
the same dimension as 'maternal rejection'. Behavioural consistency was investigated by
correlating the behavioural data from primiparous and multiparous individuals. There was
a significant correlation between the scores received by primiparous and multiparous ewes
1) Corresponding author; address: Behavioural Sciences Department, Animal Biology

Division, SAC, Bush Estate, Penicuik, Midlothian EH26 OPH, UK; e-mail address:
c.dwyer@ed.sac.ac.uk
2) The authors wish to thank: Jack FitzSimons, Mark Ramsay, Hazel Brown and Greg
Callaghan for their assistance with animal husbandry; Kirsty McLean, Leslie Deans, Joan
Chirside, Sheena Calvert and Sheila Young for technical assistance at lambing time; and
Elizabeth Austin for statistical advice. This study was supported by the Scottish Office
Agriculture, Environment and Fisheries Department.
Koninklijke Brill NV, Leiden, 2000 Behaviour 137, 1391-1413
1392 DWYER & LAWRENCE
for both the 'care/warmth' and the 'rejection' dimensions of maternal behaviour. Our data
suggest that, in sheep, maternal behaviours are segregated along the twin axes of 'rejection'
and 'care/warmth'. Although the frequency of rejection behaviours declined with maternal
experience, individual ewes were consistent in their expression of maternal care across
parities, suggesting that the maternal behaviour of a primiparous ewe is reasonably predictive
of her behaviour in subsequent pregnancies.
Introduction
Individual differences in the relationship of a mother to her offspring, par-

ticularly in primates, have received considerable research attention in recent
years. Quantitative studies demonstrate that some mothers spend more time
grooming their young and are restrictive of the infants' attempts to break
contact, whereas other mothers are relatively inattentive and rejecting (re-
viewed by Fairbanks, 1996). These behavioural differences have been de-
scribed by the terms: protective, restrictive, laissez-faire and rejecting (Fair-
banks, 1996). Studies using Principal Components Analysis (PCA) of mater-
nal behaviours suggest that variation in maternal care can often be described
by two dimensions, labelled maternal protectiveness and maternal rejection
(Fairbanks & McGuire, 1987; Fairbanks, 1996). Some studies have also de-
scribed a third dimension, labelled maternal warmth (Maestripieri, 1998) or
infant activity (Schino et al., 1995). These PCA-based descriptions of ma-
ternal behaviour, that are based on correlation relationships between behav-
ioural variables, are often referred to as different maternal 'styles'. Maternal
styles, therefore, can be defined as consistent clusterings of maternal behav-
iours within an individual that show long-term persistence over parities.
The study of individual differences in maternal behaviour has largely
been in an attempt to identify the sources of variation in maternal care.
Individual differences in maternal styles have been shown to be consistent
with successive parities in primates (Fairbanks, 1989) and guinea pigs
(Albers et al., 1999), and within families (Fairbanks, 1989; Berman, 1990)
and to exist within many species of primate (Berman, 1990). Within the
macaques, for example, the frequency of expression of different maternal
styles within the population varies with species (Maestripieri, 1994). Several
studies are supportive of a link between the temperament or emotional
reactivity of a mother and her maternal style (Maestripieri, 1993, 1994;
Schino et al., 1995), suggesting that anxious mothers are more protective of
MATERNAL BEHAVIOUR AND EXPERIENCE 1393
their infant and confident mothers are more likely to be rejecting. Although
the twin dimensions of maternal protectiveness and maternal rejection have
been reliably observed in many primate species (Fairbanks, 1996), the
existence of these features of maternal style in species other than primates
has been subject to less investigation. Recently, however, the existence of
maternal style has also been demonstrated in the guinea pig (Albers et al.,
1999), where maternal style was found to be resistant to manipulations of the
genotype of the pups or the reproductive status (pregnant or non-pregnant
during lactation) of the sow.
In this study we looked for evidence of consistent maternal styles within
another species, the domestic sheep. Like primates, the sheep produces a
small litter size, typically one or two offspring per year, and maternal in-
vestment in the young is high. Maternal behaviour in the sheep is char-
acterised by a 'bonding' period, immediately after the birth of the young,
when the ewe displays intensive licking or grooming behaviour towards the
neonate, accompanied by frequent low-pitched vocalisation (Hersher et al.,
1963; Alexander, 1988). Successful mother-offspring bonding is accompa-
nied by the ewe accepting, or facilitating, the attempts of her lamb to find
the udder and to suck. The frequency and intensity of grooming behaviour
decline over the first few hours post partum (Dwyer & Lawrence, 1998) and
responsibility for contact between ewe and lamb gradually shifts to be mainly
that of the lamb over the subsequent few days after birth (O'Connor, 1990).
It is known that individual animals vary in their expression of bonding be-
haviours, both within and between breeds (Dwyer et al., 1998; Dwyer &
Lawrence, 1999a), and that maternal experience can affect the expression
of these behaviours (Owens et al., 1985; O'Connor et al., 1992). However,
it is not known how previous maternal experience influences the expres-
sion of maternal behaviour within individual animals, whether a consistent
clustering of maternal behaviour variables occurs within individuals, that is,
whether maternal styles exist in sheep, and whether the behaviour of individ-
ual primiparous ewes is predictive of behaviour in subsequent parities.
The following hypotheses were tested in this study, that (1) the dimensions
of the mother-offspring relationship recorded in studies of primates are also
seen in an ungulate species; and (2) individual animals show behavioural
consistency across parities, such that the behaviour of a primiparous mother
would be predictive of her maternal behaviour in subsequent parities. In this
initial study we looked only at those behaviours expressed in the immediate
postpartum period associated with the formation of an attachment between

mother and young. These behaviours are known to be unaffected by offspring
behaviour (Dwyer & Lawrence, 1999a), whereas subsequent measures of
the mother-offspring relationship may be more influenced by variation
in lamb behaviour (O'Connor, 1990). A number of behavioural variables
were measured and, in common with several primate studies, Principal
Components Factor Analysis was used to summarise these measures along a
smaller number of behavioural axes.
Materials and methods
Animals
Data were collected at parturition from 69 ewes of two breeds (37 Scottish Blackface and
32 Suffolk) over a period of 4 years. All ewes were recorded in their first parity as 2-year-
olds and in at least one subsequent parity during the next 3 years. In total, 33 ewes were
recorded in their second parity, 34 in their third and 24 in their fourth. Twenty-two ewes were
recorded during three parturitions. Parity 2 and 3 ewes were aged between 3 and 5 years
old, whereas parity 4 ewes were all 5-year-olds. Datasets from individual ewes were used in
analysis only if, for all parturitions, only 1 or 2 lambs were born, length of labour was not
excessively long or abnormal, and the lambs survived the recording period. In each year ewes
were mated to a ram of the same breed as themselves, and gave birth to single or twin lambs.
Ewes were housed in large straw-bedded pens (7 x 7 m) in groups of approximately 10 ewes
from mid gestation until 3 days after parturition. Ewes were given hay and fresh drinking
water ad libitum and concentrate feed at commercial farming levels over the last 6 weeks of
pregnancy, and during lactation. During the weeks prior to expected parturition dates ewes
were accustomed to the presence of observers in the walkways between pens.
All ewes were weighed prior to conception, and 3 days after delivery. In addition a
measure of body fat, a condition score (CS) was recorded at the same time by estimating
the depth of fat and muscle over the vertebrae in the lumbar region. Lamb weights and sex
were recorded at 24-h old.
Behavioural observations
A 24-h surveillance was maintained during the 2-week lambing period in each year.
Additionally, a continuous video record using 8 cameras and a Panasonic 8-channel digital
field switcher (WJ-FS20/B, Matsushita Communication Industrial, Japan) was made. Each
animal was identified by a paint-brand to facilitate recognition on the videotapes.
As far as possible, ewes were allowed to give birth to and care for their lamb unaided.
However, lambing assistance was provided if the ewes were judged to be experiencing some
difficulty with delivery as previously described (Dwyer & Lawrence, 1999a). In all cases
intervention was kept to a minimum necessary to ensure the welfare of the ewe and lamb,
and mainly involved correcting lamb presentation before the ewe continued the birth process
unaided.
TABLE 1. Definitions of ewe behaviours recorded (with codes used in Figs 3

and 4)
Behaviour Description Code
Grooming: Licking and nibbling directed towards lamb Latency: L_egrm

Proportion: P_egrm
Withdrawing Ewe moves backwards away from the lamb at her Rwdraw
head (2+ steps)
Nosing Ewe makes contact with the lamb with her nose Rnose
without licking or nibbling
Butting, Pushes Ewe knocks lamb down or away with a rapid Rbutt
downwards or sideways movement of the head
Prevention of lamb Ewe movements that occurred within 5 s of

sucking attempts: the lamb moving to the udder:
Backing Ewe steps backwards as lamb moves forwards Rback
Circling Ewe steps sideways, moving hindquarters only Rcirc
away from the lam
Forwards Ewe steps forwards over the top of the lamb Rforw
Data on maternal behaviour were collected, from videotape, by focal animal observation
in the 2 hours after the birth of a lamb using Keytime data recording software (Deag, 1993).
In each year the following behaviours were recorded: latency from birth to grooming the
lamb, proportion of time spent grooming, frequency of withdrawing from lamb, nosing lamb,
butting lamb, pushing the lamb down or away, and responses to lamb sucking attempts
(backing, circling, moving forward or standing still). The definitions of ewe behaviours are
as previously described (Dwyer & Lawrence, 1998) and are given in Table 1. Ewe responses
to lamb sucking attempts were expressed as proportions of the total number of udder-
directed movements made by the lamb to compensate for differences in lamb behaviour.
Ewe vocalisations were collected by live focal observations onto hand-held, Atari Portfolio
computers (Atari Corporation, Slough, UK). Ewe vocalisations were identified as 'low-
pitched bleats', or 'rumbles', characteristically emitted with the mouth closed, and 'high-
pitched bleats', produced with the mouth open. In addition, the change in ewe bodyweight
and CS over gestation were recorded in each year, and the overt length of labour (from the
first appearance of fluids or membranes at the vulva until the full expulsion of the lamb).
Statistical methods
The overall effect of parity on the expression of maternal behaviours was investigated by
Wilcoxon signed-rank tests as the data were not normally distributed. Individual consistency
or change in behaviour with maternal experience was examined by separate Principal
Components Factor Analysis (Minitab Statistical Software) followed by orthogonal (VMAX)
rotation of data taken from the same ewes in their first (primiparous) and one subsequent
(multiparous) parity. For the ewes with more than 2 records the record from their last parity
was used. The following variables were used in the factor analysis for both the primiparous
and multiparous parturition of each ewe: latency to groom lamb; proportion of time spent
grooming lamb; rate of withdrawing from the lamb; rate of butting or pushing the lamb; rate
of nosing (sniffing) the lamb; proportion of all sucking attempts where ewe responded by
backing; proportion of all sucking attempts where ewe responded by circling; proportion of
all sucking attempts where ewe responded by moving forward (see Table 1 for description of
behaviours and code labels used on the figures). Maternal vocalisations were not included
in the Principal Components Analysis as only a subset of ewes had a complete set of
behavioural and vocal records. The consistency of behaviour was investigated by comparing
the Factor scores of individuals derived from the primiparous and multiparous PCA analysis
using Spearman's rank correlations. For the 22 ewes where three records were available
Kendall's coefficient of concordance was used to compare behaviour across all parities.
The relationship between ewe vocal behaviour and other behaviours were determined by
correlating vocalisations and factor scores for those individual animals with vocalisation data.
The effect of ewe breed on factor scores was analysed by Kruskal-Wallis statistics.
Results
Effects of parity on parturition
There were no significant differences between parities in the length of labour

(Table 2). Lambs born to primiparous ewes were significantly lighter than
lambs born to parity 2 and 3 ewes, and tended to be lighter than those born
to parity 4 ewes (Table 2). However, when considered in proportion to the
maternal bodyweight primiparous and multiparous ewes carried an equal
foetal burden through gestation (Table 2), as primiparous ewes were lighter
than multiparous ewes.
Effects of parity on ewe behaviours
The principal effect of parity was between inexperienced ewes (first parity)
and experienced ewes (all subsequent parities). All comparisons between
parities 2, 3 and 4 were not significant. Primiparous ewes took longer
to start to groom their lambs after delivery, were more likely to butt or
withdraw from their lambs and made more high pitched vocalisations than
multiparous ewes (Table 3). In addition, primiparous ewes made significantly
more backing and circling movements as the lambs attempted to suck (e.g.
Circling: parity 1 vs parity 2: T = 107, N = 33, p < 0.005; parity 1 vs
parity 3: T = 172, N = 33, p = 0.054; parity 1 vs parity 4: T = 70,
N = 24, p < 0.05; Fig. 1). Primiparous ewes also tended to make more
TABLE 2. Effects of parity on parturition
Parity 1 Parity 2 Parity 3 Parity
Length of labour (mins) 41.0 39.0 41.0 42.0 P
(26.75-77.75) (27.3-89.2) (22.0-88.0) (13.0-74
Mean lamb weight (kg)* 4.02 0.12 5.03 0.22 4.89 0.19 4.23 0.19
Lamb weight as a proportion 9.89 0.27 9.08 0.40 9.38 0.51 10.75 +
of ewe weight P1 vs P
Values are medians (with upper and lower interquartile ranges) or means SEM. Significa
rank tests (labour only) or by paired t-tests. *T-tests were carried out only on data where ew
subsequent parities - N for tests were 15 (parity 2), 21 (parity 3) and 10 (parity 4).
TABLE 3. Effects of parity on ewe behaviours
Parity 1 Parity 2 Parity 3 Parity 4
Latency to groom lamb (s) 106 39.8 32.2 10.2 P1 vs P2: T
(34-485) (12.6-95.6) (10.7-103.3) (5.9-25.5) P1 vs
Rate of low-pitched bleats 5.55 3.24 5.00 3.01 P1 vs P2: T
(2.0-7.68) (2.16-5.25) (2.98-8.55) (1.04-5.35) P1 vs
Rate of high-pitched bleats 0.38 0.067 0.038 0.030 PI vs P2: T
(0.0-1.46) (0.0-0.15) (0.0-0.07) (0.0-0.22) P1
Percent of ewes butting lamb 17.39 3.03 0 0 X2 = 14
Rate of withdrawing from lamb 0.039 0.010 0.000 0.000 P1 vs P2
(0.0-0.182) (0.0-0.03) (0.0-0.0) (0.0-0.0) P1 vs P
Rate of nosing lamb 0.45 0.658 0.592 0.817 P1 vs P2: T =
(0.25-0.62) (0.417-1.317) (0.417-1.138) (0.508-1.292) P1

Values are medians with upper and lower interquartile ranges. Significance was determined by Wilcox
and their corresponding parity record. Only the proportion of ewes butting their lamb is shown as the
0.8
A .00 L
0.6
Stood Back l Circle M Forward
Fig. 1. Effect of parity on responses of ewes to the sucking attempts of their lambs in the first
two hours after delivery. Maternal responses, expressed as a proportion of the total number
of sucking attempts received, are shown as standing and allowing sucking attempts, moving
backwards, circling or moving forward.
1.00 _
80.80 - T i1
g 0.60 - I
0.40 T-
~0.20 -
0.00 I I I
Fig. 2. The effect of parity on median (and interquartile ranges) grooming attention
(expressed as a proportion of the total observation time) for the first 30 minutes after lamb
delivery (round symbols and solid lines) and over the first 2 hours after lamb delivery (triangle
symbols and broken lines).
forward movements than parity 3 (T = 168, N = 33, p < 0.005) and parity
4 ewes (T = 86, N = 24, p = 0.07).
There were fewer influences of maternal experience on ewe affiliative
behaviours. Multiparous ewes had a higher rate of nosing their lambs than
primiparous ewes (Table 3). However, the amount of grooming carried out
over the first two hours after delivery did not differ significantly with parity
(Fig. 2). Likewise, there was no significant effect of parity on ewe low-
pitched vocalisation (Table 3), although ewes in parities 2 and 4 tended to
vocalise less than primiparous ewes.
Factor analysis of maternal behaviour
Principal Components factor Analysis (PCA) was carried out on 8 behav-

ioural variables for the analysis of the behaviour of multiparous ewes, and
primiparous ewes. The rate of ewe pushing the lamb away could not be dis-
tinguished from butting in the primiparous ewes, therefore these two vari-
ables were combined. In the analysis of primiparous ewes two factors had
eigenvalues of greater than 1.0 and accounted for 60.0% of the total varia-
tion between variables. In the analysis of multiparous animals three factors
were identified accounting for 57.8% of the variation. The loadings of behav-
iours on the factors are shown in Figs 3 and 4. For primiparous ewes factor 1
accounted for 39.8% of the total variation and had large positive loadings for
the ewe withdrawing, butting the lamb and backing away or circling as the
lamb attempted to suck, and a weakly positive loading for nosing the lamb
and moving forward (Fig. 3). Factor 1 was therefore labelled 'rejection'. Ewe
withdrawing, backing, nosing and circling also loaded together on Factor 2
(16.9% of the variation) of the multiparous ewe analysis (Fig. 4b); this fac-
tor was considered to correspond to Factor 1 of the primiparous ewes and
was also labelled 'rejection'. However ewe butting and pushing the lamb
1.00
0.80 - P_*egr
0.60
0.40
K 0.20 *Rb at
0p
0.00
> -0.20 Rnose RN draw

-0.40 * * Rcircle
-0.60
-0.40 - ~~~~~~~~Rback -
-0.80 - L_egrm * Rforw
-1.00 l_ l ll l
-1.00 -0.80 -0.60 -0.40 -0.20 0.00 0.20 0.40 0.60 0.80 1.00
Factor 1
Fig. 3. Principal Components Analysis of behavioural data from primiparous ewes showing
the loadings of the first ('rejection') and second factor('care/warmth'). The key to behaviours
is given in Table 1.
1.00
0.80
0.60 -
0.40 -
,0.20 _ P_egrm Rbutt L egrm
v-4 -0.20 - Rforw

-0.40 -
Rcircle Rnose
-0.60 - Rwdraw
0
-0.80 - Rback
-1.oo I I
-1.00 -0.80 -0.60 -0.40 -0.20 0.00 0.20 0.40 0.60 0.80 1.00
Factor I
1.00
0.80 - * Rbutt
0.60 - 0 Rnose
0.40
< 0.20 - Rback Rcircle

0.0
Rforw Pem
-0.20 - Rwdraw * P_egrm
-0.40 L_egnm
-0.60
-0.80
-1.00 l l l lI l l l
-1.00 -0.80 -0.60 -0.40 -0.20 0.00 0.20 0.40 0.80 0.80 1.00
Factor 2
Fig. 4. Principal components analysis of behavioural data from multiparous ewes showing
the loadings of the first ('care/warmth') and second ('rejection') factors (a) and the second
and third ('aggression') factors (b). The key to the behaviours is given in Table 1.
loaded separately on Factor 3 (13.3%; labelled 'aggression') in the multi-

parous ewe analysis. Factors are ordered by the amount of variation between
the behavioural variables that they explain. The greatest variation, between
individuals, in the primiparous ewes was, therefore, in their rejection behav-
iour. For the primiparous ewes factor 2 (20.2% of the variation) had a large
negative loading for latency to groom the lamb, and moving forward as the
lamb attempted to suck, and a large positive loading for the proportion of the
first 2 hours spent grooming the lamb (Fig. 3). For the multiparous ewes Fac-
tor 1 (20.9% of the variation) was similar to Factor 2 of the primiparous ewe
20
16 -
12 1
0
Factor scores
10
-5 -4 -3 -2 -1 0 1 2
Factor scores
Fig. 5. Distribution of scores for Blackface (solid bars) and Suffolk (hatched bars) ewes on
factors associated with rejection behaviours when (a) primiparous and (b) multiparous; and
factors associated with maternal care/warmth when (c) primiparous and (d) multiparous.
analysis, with a large negative loading for proportion of time spent grooming
the lamb and a positive loading for moving forward, latency to groom and
nosing the lamb. Factor 2 of the primiparous ewe analysis and Factor 1 of
the multiparous ewe analysis were both labelled 'care/warmth'.
Effect of breed on factor scores
The distribution of factor scores for the 'rejection' and 'care/warmth' fac-
tors are shown in Fig. 5. Although the distributions of the two breeds were
10
6)
04 .
-5 - 4 -3 -2 - 0 1
Factor scores
10
Factor scores
Fig. 5. (Continued).
overlapping the Suffolk ewe population was characterised by some individ-

uals with higher rejection scores as both primiparous (Kruskal-Wallis test:
H59 = 21.22, p < 0.001) and multiparous (H68 = 13.15, p < 0.001)
ewes than Blackface ewes. The 'aggression' Factor scores also tended to
differ between ewe breeds (H68 = 3.81, p = 0.051). The distribution of
'care/warmth' factor scores are shown in Fig. 5, c and d. Suffolk ewes hav-
ing more negative scores (indicating less grooming behaviour) than Black-
face ewes (for primiparous ewes: H59 = 5.18, p < 0.05; for multiparous
ewes: H68 = 22.24, p < 0.001).
Relationships between factor scores, ewe vocalisations and maternal traits
Maternal low-pitched vocalisations were significantly correlated with scores

on the 'care/warmth' factor (rs = 0.455; N = 47; p < 0.001) but not
with 'rejection' scores for primiparous ewes. High pitched vocalisations in
primiparous ewes were not correlated with either factor. For multiparous
ewes low-pitched vocalisation by the ewe was not correlated with any factor
scores. Ewe high pitched vocalisation, however, were correlated with scores
on the 'care/warmth' factor (r, = 0.339, N = 45, p < 0.05) and with scores
on the 'aggression' factor (r, = 0.359, N = 45, p < 0.05).
Length of labour was not significantly related to factor scores for either
primiparous or multiparous ewes. The gestational change in condition score
was significantly correlated with scores on the 'care/warmth' axis for prim-
iparous ewes (rs = 0.688, N = 59, p < 0.001), but not for multiparous
ewes (rs = 0.138, N = 68, NS). Litter size (single or twin litters) did not
affect factor scores in primiparous ewes. In the multiparous ewes single-
bearing ewes received significantly lower scores for 'care/warmth' than twin-
bearing ewes in both breeds (Blackface singles = -0.882, twins = -0.332,
H = 3.92, N = 37, p < 0.05; Suffolk singles = 0.350, twins = 0.623,
H = 4.26, N = 31, p < 0.05). There was no effect of litter size on 'rejec-
tion' scores for either breed.
Individual consistency of maternal behaviours
Correlation of the scores received by individual ewes as primiparous and

multiparous ewes for the 'care/warmth' and 'rejection' factors were in-
vestigated as an indicator of their behavioural consistency across parities.
'Care/warmth' factor scores were significantly correlated across parity (rs =
0.34, N = 58; p < 0.01). 'Rejection' factor scores were also highly corre-
lated in ewes as primiparous and multiparous animals (rs = 0.536, N = 58,
p < 0.001). The 'aggression' factor scores received by multiparous ewes
were not correlated with either 'care/warmth' or 'rejection' scores of primi-
parous ewes.
The consistency of rank order of ewes with three behavioural records is
shown in Table 4. When all animals were considered, there was good consis-
tency between individuals across all three parities for all behaviours except
backing and circling away from the lamb as it attempted to suck. It was not
possible to look at consistency of aggressive behaviour towards the lamb
TABLE 4. Individual consistency in maternal behaviour of ewes across 3

parities
Behaviour All ewes Suffolk ewes Blackface ewes
Latency to groom W = 0.585, N = 20, W = 0.661, N = 8, W = 0.382, N = 11,

lamb p < 0.05 p =0.053 NS
Proportion of first W = 0.697, N = 21, W = 0.724, N = 8, W = 0.297, N = 12,

30 minutes grooming p < 0.01 p < 0.05 NS
Total grooming (first W = 0.673, N = 19, W = 0.625, N = 7, W = 0.347, N = 12,

2 hours) p < 0.01 p < 0.05 NS
Withdrawing from W = 0.579, N = 21, W = 0.612, N = 9, N/A*

lamb p < 0.05 p = 0.066
Nosing lamb W = 0.652, N = 22, W = 0.641, N = 10, W = 0.446, N = 12,

p<0.01 p<0.05 NS
Backingatlamb W=0.412,N= 19, W=0.191,N=7, W=0.449,N= 12,

sucking attempts NS NS NS
Circling at lamb W = 0.447, N = 19, W = 0.309, N = 7, W = 0.536, N = 12,

sucking attempts NS NS p = 0.089
Moving forward at W = 0.598, N = 19, W = 0.302, N = 7, W = 0.465, N = 12,

lamb sucking p < 0.05 NS NS
attempts
Values are Kendall's coefficient of concordance, corrected for ties as necessary, for ewes
with behavioural records for their first and two subsequent parities. N values vary where
only incomplete records were available. *The analysis was not possible as no multiparous
Blackface ewes withdrew from the lamb.
as none of the ewes in this group were aggressive as multiparous animals.

Additionally, Blackface multiparous ewes did not show any withdrawal be-
haviours. When the data was considered by breed Suffolk ewes were, or
tended to be, consistent for all behaviours except moving as the lamb tried
to suck. By contrast Blackface ewes were not found to be consistent in any
behaviours.
Discussion
In this study the main change in maternal behaviour with experience in in-
dividual ewes was a reduction in negative maternal behaviours whereas af-
filiative maternal behaviours (primarily ewe grooming behaviour) were un-
affected. Principal Components Analysis of maternal behaviours, for prim-

iparous and multiparous ewes, demonstrated that rejection behaviours and
care or warmth behaviours varied along two separate axes each characterised
by a different clustering of maternal behaviours. For the primiparous ani-
mals there was a negative relationship between rejection behaviour and care
behaviours, but in multiparous ewes the two dimensions were independent
of one another. Evidence is presented for consistency in maternal behaviour
across parities. Firstly, correlations between factor scores of ewes across par-
ity were found, indicative of long-term consistency of the behavioural clus-
ters representing 'rejection' and 'care/warmth' styles. Secondly, some of the
specific maternal behaviours measured showed a consistency within indi-
vidual animals across parities. In general, Suffolk ewes were characterised
by a higher number of ewes with a maternally rejecting style, and a lower
number of ewes displaying affiliative behaviours towards the lamb, whereas
Blackface ewes showed a high level of maternal care and affiliation and were
infrequently rejecting mothers. Therefore, in common with studies of pri-
mates, maternal behaviour in the sheep can be characterised by independent
styles, labelled 'rejecting' and 'care/warmth' rather than variation along a
continuum of behaviour. Unlike the primate literature, however, our data is
restricted to a short period after the birth of the lamb, and whether these
styles are stable and can be seen throughout the pre-weaning relationship
between the ewe and lamb is not yet known.
The reduced maternal responsiveness of primiparous mothers is a com-
mon phenomenon across species (e.g. rat: Moltz & Robbins, 1965; sheep:
O'Connor et al., 1992; macaques: Timmermans & Vossen, 1996; horses:
Juarbe-Diaz, 1998). In the present study this could not be simply attributed
to a more arduous first birth process, suggesting that it is related to some in-
trinsic factor within primiparous mothers. Further, inexperience does not af-
fect all components of maternal behaviour equally but specifically increases
maternal rejection. The maternal behaviour of primiparous mothers may be
impaired because of their younger age, greater anxiety and neophobia at the
presence of the lamb, or because of a reduced neuroendocrine response in
the primiparous animal.
In the present study the primiparous ewes were all adult 2-year-old
animals whereas the multiparous ewes were three or older. Two-year-old
ewes are generally close to their mature size and weight although they
may still show some increase in body weight or dimensions during their
pregnancy. This probably accounted for the smaller size of the lamb in
the first parity animals but otherwise is unlikely to affect the reproductive
process. Preweaning juvenile rats exhibit maternal behaviour at a shorter
latency than older animals on exposure to rat pups (Zaias et al., 1996),
though whether juvenile or adolescent sheep are more maternally responsive
than adults has not been tested. Another study (Alexander et al., 1993)
suggested that 5-year old primiparous ewes were less maternally responsive
than similarly aged multiparous ewes, and were rather less responsive than
younger primiparous animals. Although these data suggest that the responses
of inexperienced animals are not related to their age, in our dataset age
and inexperience are confounded thus it may be that the younger age of
the primiparous animals was partly responsible for their increased negative
behaviours.
In common agricultural practice ewes are maintained in peer groups and
have little previous exposure to new-born lambs before they give birth them-
selves. The neonate lamb, therefore, represents a novel object to the primi-
parous ewe. In rats exposure to pups, and treatment with anxiolytics, facili-
tates maternal responses (Pellegrini & Dessifulgheri, 1994; Del Cerro et al.,
1995) suggesting that aversion or fear of novelty may play a role in determin-
ing the responses of the primiparae to the onset of maternal behaviour. Nul-
liparous non-pregnant ewes, treated with oestradiol and progesterone, show
a greater aversive response to exposure to a neonatal lamb than similarly
treated multiparous non-pregnant ewes (Dwyer & Lawrence, 1997). How-
ever, these results do not distinguish between a greater fear of novelty in
the nulliparous animal or a reduced response to the hormonal priming asso-
ciated with late pregnancy. Nulliparous ewes do not respond maternally to
artificial vaginocervical stimulation, unlike multiparous ewes (Kendrick &
Keverne, 1991; Dwyer & Lawrence, 1997). In addition, the onset of maternal
behaviour in primiparous ewes can be impaired by removing amniotic fluids
from the new-born lamb (Levy & Poindron, 1987) or by inducing anosmia in
the parturient ewe (Levy et al., 1995b), whereas these interventions did not
prevent multiparous ewes from displaying maternal behaviour. Furthermore,
the parturition-associated elevation of oxytocin in the ovine olfactory bulb
is greater in multiparous than primiparous ewes (Levy et al., 1995a). The
provision of exogenous oxytocin, given intracerebroventricularly, does not
facilitate maternal behaviour in nulliparous ewes, although multiparous ewes
become maternal towards a lamb after this treatment (Keverne & Kendrick,
1991). Maternal experience also alters the effects of opioids on maternal

behaviour (Kinsley & Bridges, 1988; Keverne & Kendrick, 1991; Mann &
Bridges, 1992). Thus the greater maternal rejection of ewes without pre-
vious maternal experience may be due to the compound effects of greater
fear of the novelty of the situation, and an immaturity of the neuroendocrine
processes associated with maternity.
The finding that maternal experience differentially affects some compo-
nents of maternal behaviour, whilst maternal grooming behaviour is unaf-
fected, is reflected in studies where maternal behaviour is induced in non-
pregnant animals. In nulliparous animals, treated with progesterone and oe-
strogen, either artificial stimulation of the vagina and cervix (Kendrick &
Keverne, 1991) or intracerebroventricular infusion of morphine and oxytocin
(Keverne & Kendrick, 1991) reduce maternal aggression and rejection be-
haviours without inducing affiliative behaviours towards the lamb. Similar
treatment of ewes with previous maternal experience induced the full com-
plement of maternal behaviours. Thus, the different components of maternal
behaviour appear to be differentially regulated. In the rat, where maternal be-
haviour has been most extensively studied, disruption of maternal behaviour
by knife-cut or chemical lesions of areas of the hypothalamus lead to a reduc-
tion in pup retrieval without affecting other aspects of pup-caring behaviour,
such as nest building and crouching over pups (Numan & Sheehan, 1997;
Olazabal & Ferriera, 1997). It has been suggested that, in the rat, there may
be areas of the brain that facilitate the appetitive components of maternal
behaviour, areas that potentiate the consummatory aspects and other areas
that depress aversion to pup stimuli (Numan & Sheehan, 1997). It is possible
that, in the primiparous ewe, the consummatory aspects of maternal behav-
iour, such as grooming, are unaffected whereas maternal attraction to lamb
odours may be inhibited by fear or novelty. Further, there is good evidence
that interactions with the neonate itself are reinforcing to the development
of maternal behaviour (Poindron et al., 1980; Magnusson & Fleming, 1995)
and that the presence of amniotic fluids are vital for the full expression of
maternal behaviour in the primiparous ewe (Levy et al., 1995b). Thus, the
sensory stimuli from the lamb will have served to potentiate the continuing
expression of maternal behaviour in ewes that lick the lamb.
The compartmentalisation of aspects of maternal behaviour is supported
by the division of maternal behaviours along at least two axes as shown in our
study and others (reviewed by Fairbanks, 1996). Thus maternal protective-

ness is not on the opposite end of a linear scale to maternal rejection and ewes
can show high levels of rejecting behaviours without showing a reduction in
grooming behaviour. Although a maternal style where a high level of rejec-
tion behaviour was expressed was more prevalent amongst the primiparous
ewes in this study this pattern of maternal behaviour was also observed in
multiparous ewes, particularly in ewes of the Suffolk breed. In primates an
increase in the frequency of maternal rejection behaviours was seen in dif-
ferent species of macaques (Maestripieri, 1994), in situations where social or
other risks were low (Fairbanks & McGuire, 1993; Fairbanks, 1996), and in
older mothers (Schino et al., 1995). Individual differences in mothering style
in primates may have an underlying emotional basis (reviewed by Fairbanks,
1996), with a high level of social anxiety resulting in a more protective style.
Of the two breeds used in our study the Blackface ewes were more reactive,
particularly to the presence of humans, had a longer flight distance in the
field (C. Dwyer, unpubl. obs.), and maintained a larger spatial distance be-
tween themselves and other ewes (Dwyer & Lawrence, 1999b) than Suffolk
ewes. Blackface ewes also maintain a shorter spatial distance between them-
selves and their lamb than Suffolk ewes (Dwyer & Lawrence, 1999b). The
greater maternal care of Blackface ewes, and maternal rejection of Suffolks
may reflect differences in the underlying emotionality of the two breeds.
Suffolk ewes were fairly consistent in all aspects of their maternal
behaviour across parities, whereas Blackface ewes were less consistent,
particularly in their rejection behaviours. Studies of temperament suggest
that it is children or animals with the extremes of reaction that remain the
most consistent over time (reviewed by Clarke & Boinski, 1995), whereas
the intermediate majority are less predictable. As Suffolk ewes showed
considerably more variation in maternal behaviours, particularly in their
rejection behaviours, it is perhaps not surprising that they were the most
consistent group over time. Studies in rhesus monkeys also show significant
consistency within individual mothers over parities (Berman, 1990). She
suggests that consistency in maternal responses may be influenced by a stable
social environment, as well as an underlying consistency in temperament
rooted in both genetics and early life experience (Fairbanks, 1989). It is
likely that the Suffolk ewes also had more variation in early life experiences
than the Blackface ewes, as their own experience of maternal care would
have been variable. When more than two parities were compared there was
little evidence that a similar rank order was maintained between individuals
in Blackface ewes. This does not, however, necessarily suggest that ewes
were not consistent in their responses to the lamb over parities, but may have
been due to a lack of sufficient differences in behaviour between individual
ewes. In primates a rejecting maternal style is associated with a greater
infant mortality (Fairbanks, 1996), and this is also likely to be the case in
sheep, particularly in hill sheep that inhabit a relatively hostile environment.
It is possible that environmental pressure has led to the Blackface breed
expressing an almost exclusively caring or protective maternal style. Suffolk
ewes, by contrast, perhaps due to the greater intervention by humans, have
maintained individual differences in maternal behaviour over several parities.
It seems that, in this breed, a greater range of maternal styles is displayed.
In the analysis of primiparous ewes maternal rejection was also accom-
panied by aggressive behaviours towards that lamb, whereas in the analysis
of multiparous ewes behaviours indicative of maternal aggression (butting
and pushing) were on a separate dimension to behaviours associated with
rejection (withdrawal, moving away from the lamb as it attempts to suck).
In group-living, seasonally breeding animals, like the sheep, there is a re-
quirement for ewes to restrict their maternal care to their own offspring. In
the ewe this is achieved by the formation of a selective olfactory memory of
the odours of their lamb in a short time-window after the birth of the lamb
(Poindron & Le Neindre, 1980). In the immediate post partum period ewes
are receptive to any lamb but the exclusivity of maternal behaviour develops
rapidly and with a shorter latency in multiparous compared to primiparous
ewes (Poindron & Le Neindre, 1980; Levy et al., 1991). In the primiparous
ewes maternal aggression was generally associated with elevated maternal
rejection behaviours, sometimes leading to the total rejection and abandon-
ment of the lamb. However, in multiparous ewes aggressive behaviours were
associated with high maternal care, including grooming attention, but deficits
in maternal selectivity in that some ewes did not reliably recognise their own
lamb and were intermittently aggressive towards it.
In conclusion, this study demonstrated that maternal experience causes a
reduction in ewe withdrawal, aggression and non-cooperation with sucking
attempts, without affecting maternal affiliative behaviours such as grooming.
Although maternal rejection behaviours were more prevalent amongst the
inexperienced ewes, individuals with a greater propensity to display caring or
rejecting maternal styles were seen among both primiparous and multiparous
ewes. In addition, there are both breed and individual consistency in m

behaviour across parities. The consistency in maternal behaviours
by individuals may be related to the underlying emotionality of the an
primates, the temperament of the infant can be predicted from the m
style of the mother (Fairbanks, 1996), and in rats variations in mat
alters the stress responses of the adult (Liu et al., 1997; Francis &
1999). Longitudinal studies in primates suggest that individual diff
maternal style are related to the previous mother-infant contact ex
by the mother herself, thus a continuity of maternal style persis
generations (Fairbanks, 1989). Whether a similar maternal style exis
families in sheep requires further research, but it is possible that the
behaviour of a ewe is a consequence not only of her genes and the i
environment but is also acquired via her own previous experience of m
care. Thus the greater prevalence of a rejecting maternal style in
ewes could be attributable to a greater likelihood of having experie
pattern of maternal care as a lamb.
References
Albers, P.C.H., Timmermans, P.J.A. & Vossen, J.M.H. (1999). Evidence for the existence of
mothering styles in guinea pigs (Cavia aperea F porcellus). - Behaviour 136, p. 469-
479.
Alexander, G. (1988). What makes a good mother? Components and comparative aspects of
maternal behaviour in ungulates. - Proc. Austr. Soc. Anim. Prod. 17, p. 25-41.
- -, Bradley, L.R. & Stevens, D. (1993). Effect of age and parity on maternal behaviour in
single-bearing Merino ewes. - Austr. J. Exp. Agric. 33, p. 721-728.
Berman, C.M. (1990). Consistency in maternal behavior within families of free-ranging
rhesus monkeys: An extension of the concept of maternal style. - Am. J. Primatol.
22, p. 159-169.
Clarke, A.S. & Boinski, S. (1995). Temperament in nonhuman primates. - Am. J. Primatol.
37, p. 103-125.
Deag, J.M. (1993). Keytime: a program system for recording and analysing behavioural data.
- Copyright John M. Deag. 1987-1993.
Del Cerro, M.C.R., Izquierdo, M.A.P., Perez-Laso, C., Rodriguez-Zafra, M., Guillamon, A.
& Segovia, S. (1995). Early postnatal diazepam exposure facilitates maternal behavior
in virgin female rats. - Brain Res. Bull. 38, p. 143-148.
Dwyer, C.M. & Lawrence, A.B. (1997). Induction of maternal behaviour in non-pregnant,
hormone-primed ewes. - Anim. Sci. 65, p. 403-408.
- & - - (1998). Variability in the expression of maternal behaviour in primiparous
sheep: Effects of genotype and litter size. - Appl. Anim. Behav. Sci. 58, p. 311-330.
- & - - (1999a). Does the behaviour of the neonate influence the expression of
maternal behaviour in sheep? - Behaviour 136, p. 367-389.
- - & - (1999b). Ewe-ewe and ewe-lamb behaviour in a hill and lowland breed of
sheep: a study using embryo transfer. - Appl. Anim. Behav. Sci. 61, p. 319-334.
- , McLean, K.A., Deans, L.A., Chirnside, J., Calvert, S.K. & Lawrence, A.B. (1998).
Vocalisations between mother and young in the sheep: Effects of breed and maternal
experience. - Appl. Anim. Behav. Sci. 58, p. 105-119.
Fairbanks, L.A. (1989). Early experience and cross-generational continuity of mother-infant
contact in Vervet monkeys. - Dev. Psychobiol. 22, p. 669-681.
- (1996). Individual differences in maternal style: Causes and consequences for mothers
and offspring. - Adv. Study Behav. 25, p. 579-611.
- & McGuire, M.T. (1987). Mother-infant relationships in Vervet monkeys: response to
new adult males. Int. J. Primatol. 8, p. 351-366.
& - (1993). Maternal protectiveness and response to the unfamiliar in Vervet
monkeys. Am. J. Primatol. 30, p. 119-129.
Francis, D.D. & Meaney, M.J. (1999). Maternal care and the development of stress responses.
- Current Opinion Neurobiol. 9, p. 128-134.
Herscher, L., Richmond, J.B. & Moore, A.U. (1963). Maternal behaviour in sheep and goats.
In: Maternal behaviour in mammals (H. Rheingold ed.). John Wiley and Sons,
London, p. 203-232.
Juarbe-Diaz, S.V., Houpt, K.A. & Kusunose, R. (1998). Prevalence and characteristics of foal
rejection in Arabian mares. - Equine Vet. J. 30, p. 424-428.
Kendrick, K.M. & Keverne, E.B. (1991). Importance of progesterone and estrogen priming
for the induction of maternal behaviour by vaginocervical stimulation in sheep: Effects
of maternal experience. - Physiol. Behav. 49, p. 745-750.
Keverne, E.B. & Kendrick, K.M. (1991). Morphine and corticotrophin-releasing factor
potentiate maternal acceptance in multiparous ewes after vaginocervical stimulation.
- Brain Res. 540, p. 55-62.
Kinsley, C.H. & Bridges, R.S. (1988). Parity-associated reductions in behavioral sensitivity
to opiates. Biol. Reprod. 39, p. 270-278.
Levy, F., Gervais, R., Kindermann, U., Litterio, M., Poindron, P. & Porter, R. (1991).
Effects of early post-partum separation on maintenance of maternal responsiveness and
selectivity in parturient ewes. - Appl. Anim. Behav. Sci. 31, p. 101-110.
- , Kendrick, K.M., Goode, J.A., Guevara-Guzman, R. & Keverne, E.B. (1995a).
Oxytocin and vasopressin release in the olfactory bulb of parturient ewes - changes
with maternal experience and effects of acetylcholine, K-butyric acid, glutamate, and
noradrenaline release. - Brain Res. 669, p. 197-206.
- , Locatelli, A., Piketty, V., Tillet, Y. & Poindron, P. (1995b). Involvement of the
main but not the accessory olfactory system in maternal behaviour of primiparous and
multiparous ewes. - Physiol. Behav. 57, p. 97-104.
- & Poindron, P. (1987). The importance of amniotic fluids for the establishment
of maternal behaviour in experienced and inexperienced ewes. Anim. Behav. 35,
p. 1188-1192.
Liu, D., Diorio, J., Tannenbaum, B., Caldji, C., Francis, D., Freedman, A., Sharma, S., Pear-
son, D., Plotsky, P.M. & Meaney, M.J. (1997). Maternal care, hippocampal glucocorti-
coid receptors, and hypothalamic-pituitary-adrenal responses to stress. - Science 277,
p. 1659-1662.
Maestripieri, D. (1993). Maternal anxiety in rhesus macaques (Macaca mulatta). II. Emo-
tional bases of individual differences in mothering style. - Ethology 95, p. 32-42.
(1994). Mother-infant relationships in three species of macaques (Macaca mulatta,

M. nemestrina, M. arctoides). I. Development of the mother-infant relationship in the
first three months. - Behaviour 131, p. 75-96.
- (1998). Social and demographic influences on mothering style in pigtail macaques.
Ethololy 104, p. 379-385.
Magnusson, J.E. & Fleming, A.S. (1995). Rat pups are reinforcing to the matemal rat - role
of sensory cues. - Psychobiol. 23, p. 69-75.
Mann, P.E. & Bridges, R.S. (1992). Neural and endocrine sensitivities to opioids decline as a
function of multiparity in the rat. - Brain Res. 580, p. 241-248.
Moltz, H. & Robbins, D. (1965). The matemal behaviour of primiparous and multiparous
rats. - J. Comp. Physiol. Psychol. 61, p. 455-460.
Numan, M. & Sheehan, T.P. (1997). Neuroanatomical circuitry for mammalian matemal
behavior. - Ann. N.Y. Acad. Sci. 807, p. 101-125.
O'Connor, C.E. (1990). Mother-offspring relationships in Scottish Blackface sheep. - PhD
thesis, University of Edinburgh.
- -, Lawrence, A.B. & Wood-Gush, D.G.M. (1992). Influence of litter size and parity on
matemal behaviour at parturition in Scottish Blackface sheep. - Appl. Anim. Behav.
Sci. 33, p. 345-355.
Olazabel, D.E. & Ferreira, A. (1997). Matemal behavior in rats with kainic acid-induced
lesions of the hypothalamic paraventricular nucleus. - Physiol. Behav. 61, p. 779-784.
Owens, J.L., Bindon, B.M., Edey, T.N. & Piper, L.R. (1985). Behaviour at parturition and
lamb survival of Boorola Merino sheep. - Livest. Prod. Sci. 13, p. 359-372.
Pelligrini, V. & Dessifulgheri, F. (1994). Sex differences in matemal behaviors of immature
rats - the role of emotionality. Aggr. Behav. 20, p. 257-265.
Poindron, P. & Le Neindre, P. (1980). Endocrine and sensory regulation of matemal behaviour
in the ewe. - Adv. Study Behav. 11, p. 75-119.
- -, - -, Raksanyi, I., Trillat, G. & Orgeur, P. (1980). Importance of the characteristics
of the young in the manifestation and establishment of matemal behaviour in sheep. -
Reprod. Nutr. Dev. 20, p. 817-826.
Schino, G., D'Amato, F.R. & Troisi, A. (1995). Mother-infant relationships in Japanese
macaques: sources of inter-individual variation. - Anim. Behav. 49, p. 151-158.
Timmermans, P.J.A. & Vossen, J.M.H. (1996). The influence of repeated motherhood on
periparturitional behaviour in cynomolgus macaques (Macaca fascicularis). - Int.
J. Primatol. 17, p. 277-296.
Zaias, J., Okimoto, L., Trivedi, A., Mann, P.E. & Bridges, R.S. (1996). Inhibitory effects
of naltrexone on the induction of parental behavior in juvenile rats. - Pharmacol.
Biochem. Behav. 53, p. 987-993.

Brill Behaviour: This Content Downloaded From 175.45.191.251 On Thu, 04 May 2017 08:57:32 UTC

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Brill Behaviour: This Content Downloaded From 175.45.191.251 On Thu, 04 May 2017 08:57:32 UTC

Uploaded by

Copyright:

Available Formats

Maternal Behaviour in Domestic Sheep (Ovis aries): Constancy and Change with Maternal

CAXIHERINE M. DWYERI) and ALISTAIR B. LAWRENCE2)

1) Corresponding author; address: Behavioural Sciences Department, Animal Biology

Koninklijke Brill NV, Leiden, 2000 Behaviour 137, 1391-1413

Individual differences in the relationship of a mother to her offspring, par-

postpartum period associated with the formation of an attachment between

Materials and methods

TABLE 1. Definitions of ewe behaviours recorded (with codes used in Figs 3

Behaviour Description Code

Grooming: Licking and nibbling directed towards lamb Latency: L_egrm

Prevention of lamb Ewe movements that occurred within 5 s of

Effects of parity on parturition

There were no significant differences between parities in the length of labour

Effects of parity on ewe behaviours

(26.75-77.75) (27.3-89.2) (22.0-88.0) (13.0-74

(34-485) (12.6-95.6) (10.7-103.3) (5.9-25.5) P1 vs

Rate of low-pitched bleats 5.55 3.24 5.00 3.01 P1 vs P2: T

(2.0-7.68) (2.16-5.25) (2.98-8.55) (1.04-5.35) P1 vs

Rate of high-pitched bleats 0.38 0.067 0.038 0.030 PI vs P2: T

(0.0-1.46) (0.0-0.15) (0.0-0.07) (0.0-0.22) P1

Percent of ewes butting lamb 17.39 3.03 0 0 X2 = 14

Rate of withdrawing from lamb 0.039 0.010 0.000 0.000 P1 vs P2

(0.0-0.182) (0.0-0.03) (0.0-0.0) (0.0-0.0) P1 vs P

Rate of nosing lamb 0.45 0.658 0.592 0.817 P1 vs P2: T =

(0.25-0.62) (0.417-1.317) (0.417-1.138) (0.508-1.292) P1

Parity 1 Parity 2 Parity 3 Parity 4

Stood Back l Circle M Forward

Factor analysis of maternal behaviour

Principal Components factor Analysis (PCA) was carried out on 8 behav-

> -0.20 Rnose RN draw

-0.80 - L_egrm * Rforw

,0.20 _ P_egrm Rbutt L egrm

v-4 -0.20 - Rforw

< 0.20 - Rback Rcircle

loaded separately on Factor 3 (13.3%; labelled 'aggression') in the multi-

Effect of breed on factor scores

overlapping the Suffolk ewe population was characterised by some individ-

Relationships between factor scores, ewe vocalisations and maternal traits

Maternal low-pitched vocalisations were significantly correlated with scores

Individual consistency of maternal behaviours

Correlation of the scores received by individual ewes as primiparous and

TABLE 4. Individual consistency in maternal behaviour of ewes across 3

Behaviour All ewes Suffolk ewes Blackface ewes

Latency to groom W = 0.585, N = 20, W = 0.661, N = 8, W = 0.382, N = 11,

Proportion of first W = 0.697, N = 21, W = 0.724, N = 8, W = 0.297, N = 12,

Total grooming (first W = 0.673, N = 19, W = 0.625, N = 7, W = 0.347, N = 12,

Withdrawing from W = 0.579, N = 21, W = 0.612, N = 9, N/A*

Nosing lamb W = 0.652, N = 22, W = 0.641, N = 10, W = 0.446, N = 12,

Backingatlamb W=0.412,N= 19, W=0.191,N=7, W=0.449,N= 12,

Circling at lamb W = 0.447, N = 19, W = 0.309, N = 7, W = 0.536, N = 12,

Moving forward at W = 0.598, N = 19, W = 0.302, N = 7, W = 0.465, N = 12,

as none of the ewes in this group were aggressive as multiparous animals.

affected. Principal Components Analysis of maternal behaviours, for prim-

1991). Maternal experience also alters the effects of opioids on maternal

study and others (reviewed by Fairbanks, 1996). Thus maternal protective-

ewes. In addition, there are both breed and individual consistency in m

(1994). Mother-infant relationships in three species of macaques (Macaca mulatta,

You might also like