Comportements maternels chez la souris: Telle mere telle fille?

Relations between mother' and daughter' maternal behaviour were surched by continuous recording of locomotor activity and time spent in the nest during the first six days postpartum in two isogenic generations of mice. The first generation was composed of either heterozygous Fl or inbred C57BL/6 (=B6) dams, both giving birth after ovarian graft to B6 pups. In each litter the maternal behaviour of a female was studied in the same conditions as for her mother. Even in developing in quite different environments, both groups of B6 daughters did not differ for the behavioural patterns under study. Links between mother' and daughter' activity during the nursing period were shown in one group by important negative correlations suggesting a feedbak regulation mechanism. This illustrate a possible way of passive hereditary transmission of acquired behavioural characteristics.     

Dopamine receptor D2 deficiency reduces mouse pup ultrasonic vocalizations and maternal responsiveness

Dopamine signalling facilitates motivated behaviours, and the D2 dopamine receptor (D2R) is important in mother–infant interactions. D2R antagonists disrupt maternal behaviour and, in isolated rat pups, reduce ultrasonic vocalizations (USVs) that promote maternal interaction. Here, we examined the effects of genetic D2R signalling deficiency on pup‐dam interaction with Drd2 knockout (D2R KO) mice. Using heterozygous (HET) cross littermates, the effect of pup genotype on isolation‐induced USVs was quantified. Independent of parental genotype, D2R‐deficient pups emitted fewer USVs than wild type (WT) littermates in a gene dose‐dependent manner. Using reciprocal D2R KO‐WT crosses, we examined how parental genotype affects pup USVs. Heterozygous pups from D2R KO dams produced fewer USVs than HET pups from WT dams. Also, exposure to USV‐emitting pups increased plasma prolactin levels in WT dams but not in D2R KO dams, and KO dams showed delayed pup retrieval and nest building. These findings indicate the importance of the interaction between pup and dam genotypes on behaviour and further support the role of D2R signalling in maternal care.     

Preference of 12-h-old kids for their mother goat is impaired by pre-partum-induced anosmia in the mother

We investigated whether kids were able to discriminate their own mother from an alien one in a two-choice test on the day of birth when they had access to acoustic, visual and olfactory cues from their mother, and whether this discrimination depended on the selective maternal behaviour of the mother (i.e. exclusive nursing of own kids). When given the choice between their own mother and an alien equivalent dam, 8-h-old kids did not show a significant preference for their dam, whereas 12- and 24-h-old kids did. When given the choice between their own and an alien mother that were both non-selective because they had been rendered peripherally anosmic by irrigation of the nostrils with zinc sulphate, 12-h-old kids did not show a significant preference for their mother. These results are similar to those reported in sheep and may suggest that the contrast of behaviour between their own and an alien mother existing in normosmic does is important for discrimination of dams by kids at this age. Finally, testing 8-h-old kids in a smaller enclosure resulted in some improvement of their performance, although they still failed to display a significant preference for their mother. On the whole, kids are able to discriminate between their own and an alien mother goat as early as previously reported in lambs. The impairment of this ability when mothers are anosmic and not selective suggests that acceptance behaviours displayed by the mother may serve as one of the cues orientating the choice of the kid when given the choice between intact mothers. Finally, the present results do not suggest the existence of fundamental differences in the establishment of a preference for the mother between lambs, which are followers, and kids, which are hiders.     

Behavioural responses of captive hippopotamus(hippopotamus amphibius)in lactation season:Maternal investment and plasticity of infant self-independence

In order to promote hippopotamus management in the captive and ex-situ environment, especially the control of behavioural and physiological status during breeding and lactation seasons, we conducted a preliminary study on behavioural responses of a pair of hippos including both mother and infant in Hangzhou Wildlife Park, China. The study of the captive hippos for about 1-month in the lactation season was carried out during August and September, 2009. The behavioural patterns were identified by all occurrence sampling and instantaneous scanning sampling methods with 5-10 min intervals. As a result, mother-offspring conflicts and interactions did occur throughout the whole study period. Early maternal investment showed a positive trend in activity rhythms (slope = 0.0014, Z = 0.3027, P ＜0.001) and a negative trend (slope = -0.0066, Z = 0.8807, P＜0.001) in territorial occupation of water, all of which supported our hypotheses that the mother hippo might exert less care for the infant and cut down on her own obligations in nursing. For infant self-independence, during the whole lactation season, the primary trends of activities and territorial occupation dynamics of the infant hippo were slightly different from before, judging from linear models (slope = -0.0017, Z = 0. 3309, P＜0.001). However, the frequencies of activities were not stable, especially at around 12 days of age. The trends of territorial occupation (slope = -0.0071, Z= 0. 904, P＜0.001) also showed negative dynamics in water body occupation by the time the infant hippo grew up. The general trend (slope = -0.005, Z = 0.06, P＜0.001) of suckling dynamics was demonstrably negative, with an upwards fluctuation at period 3 (10-15th day). This also illustrated that as the infant developed, the dependency on the mother was reduced at the end of the lactation season. In addition, a sharp decline between P3 and P4 also supported the mother-offspring conflict theory. In general, time budgets of hippos in active behaviour were (31.8±2.1)% for the mother and (32.1 ±2.6)% for the infant. Spatial distributions in water within temporal limitations were (80.1 ±2.7)% for the mother and (81.8±2.7)% for the infant. Behavioural dynamics showed strong synchronous relations between maternal investment and infant independence. Our current short-term investigation proves to be a key in management and conservation of hippopotami during the lactation season.     

Low-frequency sound communication by mouse pups (Mus musculus): wriggling calls release maternal behaviour

Low-frequency calls (wriggling calls), emitted mainly between days 2 and 15 after birth by mouse pups while pushing for the teats during suckling by the mother, are studied with regard to their significance in intraspecific communication. Wriggling calls release maternal behaviour and ‘attention’ responses in the mother. Licking of the pups is the most prominent maternal response, followed by changes of suckling position and by nest building. One function of the wriggling calls is to keep these maternal behaviours at a high level: otherwise their spontaneous rates of occurrence would decrease rapidly after the birth of the pups. Maternal and attention responses can be released in females lying on paralysed pups by playbacks of wriggling calls. The behavioural contexts of the production of wriggling calls and pure ultrasounds are mutually exclusive and the perception and recognition of each type of pup call cannot be confused by the mother.     

Impact of intra- and interstrain cross-fostering on mouse maternal care

The importance of maternal care in shaping an individual's phenotype in health and disease is becoming more and more apparent in both human and animal studies. However, in mouse studies using inbred strains or knockout mice to analyze the genetic influences on the development of normal and aberrant behavioral phenotypes, maternal behavior is very poorly characterized and often ignored. This study provides an extensive analysis of spontaneous maternal behavior of inbred mice in three conditions: (1) comparing two commonly used strains, (2) analyzing the impact of adopting pups from the same strain (intrastrain cross-fostering) and (3) analyzing the impact of adopting pups from a different strain (interstrain cross-fostering). For each condition, maternal behavior was analyzed continuously over 23-h periods on postnatal days 2, 4, 6 and 9. We report that (1) the maternal behavior of C57BL/6J and DBA/2J dams toward their biological offspring is highly similar, (2) intrastrain cross-fostering has minimal impact on maternal behavior of C57BL/6J and DBA/2J dams, (3) interstrain cross-fostering does not modify the strain differences in maternal care observed between AKR and C3H/He mothers and (4) the pup strain does influence the amount of maternal behavior shown by both mothers in interstrain cross-fostering. These latter findings demonstrate that both mother strain and pup strain are key determinants of maternal behavior.     

Effects of maternal strain on ethanol responses in reciprocal F1 C57BL/6J and DBA/2J hybrid mice

Variations in maternal behavior, either occurring naturally or in response to experimental manipulations, have been shown to exert long-lasting consequences on offspring behavior and physiology. Despite previous research examining the effects of developmental manipulations on drug-related phenotypes, few studies have specifically investigated the influence of strain-based differences in maternal behavior on drug responses in mice. The current experiments used reciprocal F1 hybrids of two inbred mouse strains (i.e. DBA/2J and C57BL/6J) that differ in both ethanol (EtOH) responses and maternal behavior to assess the effects of maternal environment on EtOH-related phenotypes. Male and female DBA/2J and C57BL/6J mice and their reciprocal F1 hybrids reared by either DBA/2J or C57BL/6J dams were tested in adulthood for EtOH intake (choice, forced), EtOH-induced hypothermia, EtOH-induced activity and EtOH-induced conditioned place preference (CPP). C57BL/6J and DBA/2J mice showed differences on all EtOH responses. Consistent with previous reports that maternal strain can influence EtOH intake, F1 hybrids reared by C57BL/6J dams consumed more EtOH during forced exposure than did F1 hybrids reared by DBA/2J dams. Maternal strain also influenced EtOH-induced hypothermic responses in F1 hybrids, producing differences in hybrid mice that paralleled those of the inbred strains. In contrast, maternal strain did not influence EtOH-induced activity or CPP in hybrid mice. The current findings indicate that maternal environment may contribute to variance in EtOH-induced hypothermia and EtOH intake, although effects on EtOH intake appear to be dependent upon the type of EtOH exposure.     

Maternal buffering of stress response in free‐ranging Pacific harbor seal pups in Alaska

We examined the effects of maternal buffering in free‐ranging Pacific harbor seal pups during capture and handling research procedures. We predicted that pups held with their mother would benefit from social buffering and exhibit lower cortisol concentrations resulting from capture and handling than dependent pups caught without their mothers and weaned pups. We expected that pups captured with their mother that experienced a short separation would exhibit increased stress‐induced vocal behavior and activity level compared to dependent weaned pups caught alone. The results showed that the presence of the mother significantly buffered the stress response, as measured by reduced serum cortisol concentrations, in pups captured with their mothers as compared to dependent pups captured alone. Cortisol concentrations of mothers with pups initially were higher than nonlactating females, then diminished. Pups showed a significantly higher rate of vocalization soon after maternal separation compared to single pups separated for a longer period of time. Newly separated pups, especially males, showed a high level of activity compared to the other pups. The results provide unique quantitative evidence of the physiology underlying the maternal‐pup bond in a marine mammal, and the role that maternal buffering may play on the stress response of the offspring.     

Differential effects of undernutrition during pregnancy on the behaviour of does and their kids at parturition and on the establishment of mutual recognition

We investigated whether undernutrition during the second half of pregnancy impaired the behaviour of does and their kids at parturition and early mutual recognition. Twenty-two control and 22 underfed mixed-breed, multiparous dairy goats were used, together with their respective kids (control, n = 31: nine singles, 16 twins and six triplets; underfed, n = 32: 11 singles, 18 twins and three triplets). Undernutrition involved limiting protein and energy intake at 70% of the nutritional requirements for maintenance and foetal growth from day 70 of pregnancy until birth. The behaviour of mothers and their two first-born kids was observed for 90 min from the birth of each kid. Maternal olfactory recognition of the kid was assessed at 4 h post partum by testing selective nursing behaviour. Non-olfactory recognition was assessed at 8 h in a two-choice test excluding olfactory cues. In kids, preference for the mother was assessed in a two-choice test at either 12 or 24 h post partum. Bodyweight of does and kids were lower in the underfed group up to 2 weeks post partum. At parturition, licking, maternal bleating frequency and latency to nursing did not differ between nutritional groups. Control kids were faster than underfed kids to stand, search for and reach the udder, but underfed kids bleated more and tended to spend more time at the udder. Both control and underfed does accepted their own kid and rejected the alien in the selectivity test at 4 h. In contrast, at 8 h post partum, only control goats showed a significant preference for their own kid in the non-olfactory recognition test. Both control and underfed kids showed a preference for their own mother at 12 and 24 h and undernutrition during pregnancy had little influence on the performance of kids. However, 12 h-old underfed kids tended to be less active than control kids and visited their own mothers less than control kids. There were no significant correlations between the behaviour of the mother or of the kid at parturition and their performance in the discrimination tests. Overall, undernutrition in the second half of pregnancy appears to be more detrimental for the behaviour of the mother than for the kid. Furthermore, it has more impact on the establishment of maternal non-olfactory recognition than on maternal care at parturition or the establishment of maternal selectivity.     

The influence of the sex of litter-mates on subsequent maternal behaviour in Rattus norvegicus.

Female rats reared in unisexual (U) or bisexual (B) cross-fostered litters were mated at 75 days of age and their maternal behaviour compared by periodic observation over 20 days following birth of their litters. In two separate experiments, U-females showed a lower incidence of pregnancy and, at birth, of litters with ten or more pups than B-females (P less than 0-05). Litters culled to ten animals on the day after birth were significantly heavier in B- than in U-females, although this difference was not evident at weaning as litters reared by U-females showed a relatively greater (P less than 0-01) increase in weight during the investigation than the corresponding litters reared by B-females. Using a scoring method, no difference was evident in the opportunities that U- and B-females offered for their pups to suck, and, when not nursing pups in the nest, both groups of females engaged equally in activities such as grooming, sleeping, eating, and rearing on their hind legs. Throughout the 20-day observation period, B-females maintained significantly more elaborate nests (as scored on an arbitrary scale) than U-females. This difference was evident in females rearing litters of both five and ten animals, and was observed in two separate experiments. In addition, whereas only 15 per cent of U-females were observed nest-building, over 60 per cent of B-females were noted to be engaged in this activity on one or more occasions (P less than 0-001). It is suggested that, in the light of previous findings, these results may reflect differences in oestrogen and/or progesterone metabolism in unisexually-reared and bisexually-reared female rats.     

Maternally derived hormones,neurosteroids and the development of behaviour

In a wide range of taxa, there is evidence that mothers adaptively shape the development of offspring behaviour by exposing them to steroids. These maternal effects have major implications for fitness because, by shaping early development, they can permanently alter how offspring interact with their environment. However, theory on parent–offspring conflict and recent physiological studies showing that embryos rapidly metabolize maternal steroids have placed doubt on the adaptive significance of these hormone-mediated maternal effects. Reconciling these disparate perspectives requires a mechanistic understanding of the pathways by which maternal steroids can influence neural development. Here, we highlight recent advances in developmental neurobiology and psychiatric pharmacology to show that maternal steroid metabolites can have direct neuro-modulatory effects potentially shaping the development of neural circuitry underlying ecologically relevant behavioural traits. The recognition that maternal steroids can act through a neurosteroid pathway has critical implications for our understanding of the ecology and evolution of steroid-based maternal effects. Overall, compared to the classic view, a neurosteroid mechanism may reduce the evolutionary lability of hormone-mediated maternal effects owing to increased pleiotropic constraints and frequently influence long-term behavioural phenotypes in offspring.     

The behaviour of twin rhesus monkeys and comparisons with the behaviour of single infants

Twin rhesus monkeys, born in a laboratory cage, were reared alone with their mother for a year. Observations were made of mother-infant interactions. Differences between the infants which concerned the amount of time spent off the mother seem to have been due to differences in the behaviour of the mother towards the two twins, whereas differences between measures of behaviour while off the mother seem to have been due to differences between the twins. The twins also differed in activity and in responses to mildly frightening or strange situations. The behaviour of the twins was compared with that of two single infants reared under the same cage conditions and also alone with their mothers. Some differences were found between the twins and the singles in mother-infant interaction, and there were also some differences in activity and in responses to mildly frightening or strange situations.     

Maternal behaviour and peripartum levels of oestradiol and progesterone show little difference in Merino ewes selected for calm or nervous temperament under indoor housing conditions

Temperament influences the expression of maternal behaviour in sheep under experimental field conditions. We investigated whether maternal behaviour between ewes selected for a calm or nervous temperament is independent from environmental conditions. In addition, the level of maternal behaviour expressed by mothers is correlated with the concentration of hormones during the peripartum period. Therefore, we investigated whether the selection for temperament had resulted in hormonal differences between the two lines with regard to the hormones that could be involved in the onset of maternal behaviour. Oestradiol, progesterone and cortisol concentrations from 4 days before parturition to 24 h after parturition were determined from blood samples collected from 10 calm and 12 nervous ewes. Behavioural interactions between ewe and lamb were also recorded for 2 h starting at parturition. Mothers of both temperament lines showed adequate maternal behaviour under the controlled conditions of the study. Therefore, the results of the study do not suggest that selection for a calm or nervous temperament has profoundly affected the intrinsic ability of mothers to display adequate maternal behaviour. Hormonal differences between the two temperament lines were generally small and their possible influence on the display of maternal behaviour in the two temperament lines would have to be demonstrated.     

Within-litter variance in rat maternal behaviour

As a first step in determining the influence of maternal behaviour on sibling behavioural variance, we tested whether rat mothers differentially interact with neonates within the same litter. We also tested whether fading of an ink-mark on individual pups could provide an index of within-litter variance in maternal licking in laboratory rats. In Study 1, during the first postnatal week we distinguished individual Sprague-Dawley rat pups across 4 litters by placing an ink-mark on the skin and quantified variance in maternal licking frequency toward each pup and compared fading of individual pup marks to the frequency of maternal licks received and to four pup characteristics that could influence mark-fading. In Study 2, neonate mark-fading (a proxy for maternal licking) was compared to adolescent and adult offspring behaviour across 8 litters. Results indicated that: (1) there are substantial and consistent differences in how much rat mothers lick same-sex siblings within a litter, (2) differential licking rates can be documented with a non-observational method (ink-mark-fading), and (3) within-litter variance in maternal behaviour may relate to sibling behavioural variance. The findings indicate a viable research model for future experimental studies on causes and consequences of differential maternal investment within families.     

Low heritabilities, but genetic and maternal correlations between red squirrel behaviours

Consistent individual differences in behaviour, and behavioural correlations within and across contexts, are referred to as animal personalities. These patterns of variation have been identified in many animal taxa and are likely to have important ecological and evolutionary consequences. Despite their importance, genetic and environmental sources of variation in personalities have rarely been characterized in wild populations. We used a Bayesian animal model approach to estimate genetic parameters for aggression, activity and docility in North American red squirrels (Tamiasciurus hudsonicus). We found support for low heritabilities (0.08-0.12), and cohort effects (0.07-0.09), as well as low to moderate maternal effects (0.07-0.15) and permanent environmental effects (0.08-0.16). Finally, we found evidence of a substantial positive genetic correlation (0.68) and maternal effects correlation (0.58) between activity and aggression providing evidence of genetically based behavioural correlations in red squirrels. These results provide evidence for the presence of heritable variation in red squirrel behaviour, but also emphasize the role of other sources of variation, including maternal effects, in shaping patterns of variation and covariation in behavioural traits.     

INHERITANCE OF ENVIRONMENTAL VARIATION IN BODY SIZE: SUPERPARASITISM OF SEEDS AFFECTS PROGENY AND GRANDPROGENY BODY SIZE VIA A NONGENETIC MATERNAL EFFECT

Maternal effects provide the most common mechanism by which environmental variation in one generation affects the phenotype of individuals in subsequent generations. In egg-laying animals, however, we typically observe that maternal effects can have large influences on early growth (egg size and early development), but these effects gradually disappear and become undetectable by the time progeny mature due to developmental plasticity in progeny. We describe a system in which an environmentally induced reduction in body size is inherited by progeny via a nongenetic maternal effect. The seed beetle, Callosobruchus maculatus, completes development inside a discrete resource package (a seed) selected by its mother. Due to superparasitism in response to low host availability, progeny frequently develop at high densities, resulting in intense larval competition and pupation at a smaller body size. Females reared at higher density (and thus emerging smaller) lay smaller eggs than females reared at lower density. Progeny from these smaller eggs mature at a smaller size than progeny reared from the larger eggs laid by females reared at lower density. Crosses between high and low density lines demonstrated that treatment differences in body size are maternally inherited, confirming that the inheritance of body size variation in part involves an environmentally based maternal effect.     

Endogenous 24-hour cycle of core temperature and oxygen consumption in week-old Zucker rat pups

Summary Experiments were designed to test whether or not the 24-h core temperature fluctuations in week-old rat pups are of endogenous origin. Lean (Fa/-) Zucker rat pups born on the same day to mothers maintained in two different colonies with light/dark cycles 12 h out of phase with each other were mother-reared through the first 3–4 days of life and then artificially reared simultaneously in constant dim light. Continuous, automatic measurement of core temperature and oxygen consumption during artificial rearing showed clear 24-h rhythms in 5- to 8-day-old pups. Each rhythm reached a daily minimum at a time corresponding to the beginning of the light period in the colony of origin. The amplitude of these rhythms did not diminish during artificial rearing, nor did the phase difference between the rhythms of pups originating in the two colonies systematically change. The persistent 12-h phase differences between these two groups of pups prove that the observed rhythms are not caused by exogenous stimuli. We conclude that the rat pup possesses an endogenous time-keeping mechanism that permits the expression of overt rhythmicity at the age of 1 week.Abbreviations Tc core temperature - LD-pups born to a mother entrained to a 7:00 to 19:00 light cycle, then artificially reared in constant dim light - DL-pups born to a mother entrained to a 19:00 to 7:00 light cycle, then artificially reared in constant dim light - SCN suprachiasmatic nucleus     

Quantitative genetics of growth and development time in the burying beetle Nicrophorus pustulatus in the presence and absence of post-hatching parental care

Despite a growing interest in the evolutionary aspects of maternal effects, few studies have examined the genetic consequences of maternal effects associated with parental care. To begin to provide data on nonlaboratory or nondomestic animals, we compared the effect of presence and absence of parental care on phenotype expression of larval mass and development time at different life-history stages in the burying beetle Nicrophorus pustulatus. This beetle has facultative care; parents can feed their larvae through regurgitation of digested carrion or offspring can feed by themselves from previously prepared carrion. To investigate larval responses to these two levels of care, including estimates of additive genetic effects, maternal effects, and genotype-by-environment interactions, we used a half-sibling split-family breeding experiment-raising half of the offspring of a family in the presence of their mother and the other half without their mother present. Larvae reared with their mother present were on average heavier and developed faster, although some of the differences in development decreased or were eliminated by the adult stage. These results suggest that presence or absence of post-hatching maternal care plays an important role in phenotype expression early in life, whereas later the phenotype of the offspring is determined mainly by the genotype and/or unshared environmental effects. Our study also permitted us to examine the differences in genetic effects between the two care environments. Heritabilities, maternal/common environment effect, and most genetic correlations did not differ between the care treatments. Genetic analyses revealed substantial additive genetic effects for development time but small effects for measures of body mass. Maternal plus common environment effects were high for measures of mass but low for development time, suggesting that indirect genetic effects of maternal and/or common environment are less important for the evolution of development time than for mass. Estimates of genetic correlations revealed a trade-off between the duration of the two development stages after the offspring left the carrion. There was also a negative genetic correlation between the time spent on carrion and the mass at 72 h, when mothers usually stop feeding. The analysis of genotype-by-environment interactions indicates substantial variation among maternal families in response to care. Presence or absence of parental care may therefore contribute to the additive genetic variance through its interaction with the maternal component of the additive genetic variance. The presence of this interaction further suggests that parents may vary in care strategies, with some parents dispersing after preparation of the carrion and some parents staying with the larvae. This interaction may help maintain genetic variation in growth, development time, and parental care behavior. Additional work is needed, however, to quantify indirect genetic effects and genetic variation in parental care behavior itself.     

Maternal Epigenetic Transmission of Social Motivation in Birds

Behavioural development is a complex phenomenon involving interactions between genetic constraints and environmental influences. One of the most potent environmental influences during the ontogeny of the behavioural characteristics of young is played by mothers. In particular, mammalian mothers modulate the social competences of their young, influencing all their future social life. Here, we investigated the influence of the social characteristics of adoptive mother birds ( Coturnix coturnix japonica ) on the social motivation (origin of every social relationship) of the young they reared. We characterized the social behaviour of standard stock chicks reared by female quail, genetically selected either for low (LSR) or high (HSR) levels of social reinstatement (index of treadmill behaviour which combined the tendency to run towards conspecifics with the tendency to move away from them), to investigate epigenetic transmission of social motivation. Our results show that HSR and LSR adoptive mothers partially transmitted their social characteristics to their young: chicks reared by HSR females presented higher levels of social motivation than chicks reared by LSR females. This maternal influence appeared much clearer in young males than in young females. Our study reveals that, as in mammals, bird mothers influence epigenetically the development of the social behaviour of their young.     

Maternal behaviour and lamb survival: from neuroendocrinology to practical application

Parental care promotes offspring survival and, for livestock species, this care is provided solely by the mother. Maternal behaviour in the sheep has been exceptionally well-studied compared with other species and many of the underpinning biological processes leading to the expression of maternal care are known. In this review the current state of play with regard to the biology of maternal care will be reviewed, and its application to provide practical solutions to reduce lamb mortality considered. For maternal care to be elicited at birth the ewe requires elevated circulating oestradiol in late gestation, which stimulates the expression of oxytocin receptors in both peripheral and central areas (particularly the hypothalamic and limbic areas of the brain). At birth stretching of the vaginocervical canal elicits a spinal reflex which triggers the release of oxytocin primarily from neurones within the paraventricular nucleus of the hypothalamus. Oxytocin release causes an increase in the neurotransmitters noradrenaline, acetylcholine, glutamate and γ-aminobutyric acid (GABA) in the olfactory bulb, and other brain regions important for maternal behaviour. Finally, sensory cues provided by the lamb, in particular the amniotic fluids surrounding it, lead to the expression of maternal behaviours (licking, low-pitched bleats, acceptance of the lamb at the udder and suckling). This allows the expression of the two facets of maternal behaviour in the ewe: nurturance of the young and maternal selectivity, whereby a specific olfactory memory for the ewes own lamb is formed and the expression of maternal care is restricted to this lamb. Variation in the expression of maternal care has been demonstrated in primiparous ewes compared with multiparous, in different sheep genotypes, with undernutrition, stress in pregnancy, following a difficult delivery, and may occur with variation in ewe temperament. An understanding of the importance of the timing of various events in late pregnancy and during parturition, as well as the factors that can disrupt these events, can help to design management activities to minimise risks to the successful onset of maternal behaviour. Management practices that work with the biology of the ewe will be the most successful in ensuring that maternal care is expressed, so improving the welfare of the ewe and lamb, and the profitability of the farm.