Parity and housing effects on the behavioural and hypothalamic-pituitary-adrenal axis responses of pregnant ewes.

It is common in many countries for sheep to be housed during winter from mid-gestation until lambing to protect ewes and lambs from adverse conditions and improve late gestation nutritional management. Keeping ewes indoors, however, has its own challenges as the animals may be mixed with unfamiliar conspecifics, have limited floor and feeding space, experience changes to their diet and increased handling by humans. Therefore, the objective of this study was to investigate the effect of variation in housing management (space allowance and social stability) on the behaviour and hypothalamic-pituitary-adrenal (HPA) axis responses of pregnant ewes from mid-to-late gestation (weeks 11-18 of pregnancy). Seventy-seven ewes (41 primiparous, 36 multiparous) were divided into two groups: 'Control' and 'Restricted space and mixed' (RS-Mix), where RS-Mix ewes were allocated half the amount of space (1.27 vs 2.5 m2 for RS-Mix and Control, respectively) and feedface (concentrate feeder space) allowance (36 vs 71 cm per ewe) given to the Control group and were also subjected to two social mixing events. Aggressive behaviour at the feedface and time spent standing, lying, walking, feeding and ruminating were recorded and faecal samples were collected for assessment of faecal glucocorticoid metabolite (FGM) concentrations. Higher aggression was observed in RS-Mix ewes during the first week of observation (P = 0.044), which gradually declined to the same level as Control ewes by the end of the study (P = 0.045). RS-Mix ewes were significantly less likely to be able to freely join the feedface compared to Controls (P = 0.022). No other significant treatment effects on aggressive behaviour or FGM during gestation were found. RS-Mix ewes displayed significantly higher ruminating behaviour at week 18 of gestation compared to Control ewes (P < 0.001), but no other effects were seen on general pen behaviour. However, the effect of indoor housing had a significant impact on primiparous ewes, who had lower weight gain (P = 0.015) and higher FGM concentrations (P = 0.014) compared to multiparous ewes regardless of treatment group. The data suggest that, although no sustained effects on behaviour or HPA axis responses were seen with the differences in space and feeder allowance or social stability at the levels used in this study, inexperienced (primiparous) ewes may find indoor housing more stressful; and are less able to adapt compared to multiparous ewes. These effects may influence the behaviour of the ewe at lambing time, and her offspring.

Epigenetics and developmental programming of welfare and production traits in farm animals.

The concept that postnatal health and development can be influenced by events that occur in utero originated from epidemiological studies in humans supported by numerous mechanistic (including epigenetic) studies in a variety of model species. Referred to as the 'developmental origins of health and disease' or 'DOHaD' hypothesis, the primary focus of large-animal studies until quite recently had been biomedical. Attention has since turned towards traits of commercial importance in farm animals. Herein we review the evidence that prenatal risk factors, including suboptimal parental nutrition, gestational stress, exposure to environmental chemicals and advanced breeding technologies, can determine traits such as postnatal growth, feed efficiency, milk yield, carcass composition, animal welfare and reproductive potential. We consider the role of epigenetic and cytoplasmic mechanisms of inheritance, and discuss implications for livestock production and future research endeavours. We conclude that although the concept is proven for several traits, issues relating to effect size, and hence commercial importance, remain. Studies have also invariably been conducted under controlled experimental conditions, frequently assessing single risk factors, thereby limiting their translational value for livestock production. We propose concerted international research efforts that consider multiple, concurrent stressors to better represent effects of contemporary animal production systems.

Invited review: Improving neonatal survival in small ruminants: science into practice.

Neonatal mortality in small ruminant livestock has remained stubbornly unchanging over the past 40 years, and represents a significant loss of farm income, contributes to wastage and affects animal welfare. Scientific knowledge about the biology of neonatal adaptation after birth has been accumulating but does not appear to have had an impact in improving survival. In this paper, we ask what might be the reasons for the lack of impact of the scientific studies of lamb and kid mortality, and suggest strategies to move forward. Biologically, it is clear that achieving a good intake of colostrum, as soon as possible after birth, is crucial for neonatal survival. This provides fuel for thermoregulation, passive immunological protection and is involved in the development of attachment between the ewe and lamb. The behaviour of the lamb in finding the udder and sucking rapidly after birth is a key component in ensuring sufficient colostrum is ingested. In experimental studies, the main risk factors for lamb mortality are low birthweight, particularly owing to poor maternal nutrition during gestation, birth difficulty, litter size and genetics, which can all be partly attributed to their effect on the speed with which the lamb reaches the udder and sucks. Similarly, on commercial farms, low birthweight and issues with sucking were identified as important contributors to mortality. In epidemiological studies, management factors such as providing assistance with difficult births, were found to be more important than risk factors associated with housing. Social science studies suggest that farmers generally have a positive attitude to improving neonatal mortality but may differ in beliefs about how this can be achieved, with some farmers believing they had no control over early lamb mortality. Facilitative approaches, where farmers and advisors work together to develop neonatal survival strategies, have been shown to be effective in achieving management goals, such as optimising ewe nutrition, that lead to reductions in lamb mortality. We conclude that scientific research is providing useful information on the biology underpinning neonatal survival, such as optimal birthweights, lamb vigour and understanding the importance of sufficient colostrum intake, but the transfer of that knowledge would benefit from an improved understanding of the psychology of management change on farm. Developing tailored solutions, on the basis of adequate farm records, that make use of the now substantial body of scientific literature on neonatal mortality will help to achieve lower neonatal mortality.

Pen size and parity effects on maternal behaviour of Small-Tail Han sheep.

The aim of this experiment was to study the effects of pen size and parity on maternal behaviour of twin-bearing Small-Tail Han ewes. A total of 24 ewes were allocated to a 2×2 design (six per pen), with parity (primiparous or multiparous) and pen size (large: 6.0×3.0 m; small: 6.0×1.5 m) as main effects at Linyi University, Shandong Province, China. Behaviour was observed from after parturition until weaning. All ewes were observed for 6 h every 5 days from 0700 to1000 h and from 1400 to 1700 h. Continuous focal animal sampling was used to quantify the duration of maternal behaviours: sucking, grooming and following as well as the frequency of udder accepting, udder refusing and low-pitched bleating. Oestradiol and cortisol concentrations in the faeces (collected in the morning every 5 days) were detected using EIA kits. All lambs were weighed 24 h after parturition and again at weaning at 35 days of age. The small pen size significantly reduced following (P<0.005), grooming (P<0.001) and suckling durations (P<0.05), as well as the frequency of udder refusals (P<0.001). However, there was a significant interaction with ewe parity, with decreased grooming and suckling in the small pen largely seen in the multiparous ewes (P<0.001). Independent of pen size, multiparous ewes accepted more sucking attempts by their lambs (P<0.05) and made more low-pitched bleats than primiparous ewes (P<0.001). Multiparous ewes had higher faecal oestradiol concentrations than primiparous ewes (P<0.001), and ewes in small pens had higher faecal cortisol levels compared with ewes in larger pens (P<0.001). As lambs increased in age, the duration of maternal grooming, following and suckling as well as frequency of udder acceptance and low-pitched bleating all declined, and the frequency of udder refusing increased (P<0.001 for all). Ewe parity, but not pen size, affected lamb weight gain during the period of observation (P<0.001). This is the first study to show that pen size, interacting with parity, can affect the expression of maternal behaviour in sheep during lactation. The study is also the first to report on the maternal behaviour of Chinese native sheep breeds (Small-Tail Han sheep), with implications for the production of sheep in China.

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.

Stillbirth in dairy calves is influenced independently by dystocia and body shape.

The objectives of this study were to determine (1) if stillborn calves born following dystocia present with specific injuries/pathological changes compared to stillborns delivered without difficulty, and (2) whether such stillborns differ in conformation from dystocic calves that survive. Post-mortem examinations were carried out on 20 stillborns that were either unassisted (N) or were 'farm-staff'-assisted/normally presented (FN) at birth. Evidence of greater trauma and bruising was observed in the FN calves and parameters such as body length, birth-weight and thyroid:body weight were similar. In a second part of the study birth-weight, body length and height, girth length, body mass (BMI), and ponderal (PI) indices were assessed in 490 calves. Regardless of the severity of dystocia, stillborns had greater body lengths and lower BMIs and PIs than calves born alive (P<0.05), suggesting prenatal factors contribute to their post-natal survival. FN calves were heavier than N calves (P<0.05), and both FN and farm-staff-assisted/malpresented calves had lower PIs than N calves (P<0.05). The study found that criteria such as grossly visible carcass haemorrhage, bruising, and brain congestion were not reliable in terms of identifying calves that had experienced dystocia. Half of the stillborns had breathed indicating they were alive and possibly had experienced pain/distress at time of delivery. Body conformation was related to stillbirth independently of dystocia, a finding likely reflecting inadequate prenatal development.

The impact of dystocia on dairy calf health, welfare, performance and survival.

Up to one-third of dairy calves are born after dystocia and this is a major cause of calf mortality. This study investigated the neonatal physiology, survival, health and subsequent growth of dairy calves following dystocia and is the first longitudinal study to analyse multiple effects and to look beyond the perinatal period. A total of 455 live born Holstein calves (N: No assistance, n=360; FN: Farmer assistance but normally presented calf, n=82; FM: Farmer assistance of malpresented calf, n=13) were followed from birth to first service (heifers) or until leaving the farm (bulls). Compared to N calves, FN and FM animals had higher salivary cortisol concentrations at day 1 (P<0.001) and FN calves had lower passive immune transfer (P=0.03). Dystocia had no biologically significant impact on rectal temperature throughout the first 4 days (P>0.05). During the first 60 days, FM calves had a higher proportion of days with non-routine health treatments (P<0.05) and, by the time of weaning, mortality in FN and FM heifers was higher than in N calves (2.8×; P<0.01). However, in surviving calves, growth to first service was not affected by dystocia category (P>0.05). Calves which survive dystocia experience lower passive immunity transfer, higher mortality and higher indicators of physiological stress. Such calves have poorer welfare in the neonatal period and possibly beyond. Strategies need to be implemented to improve the subsequent health and welfare of such calves and to lower the incidence of dystocia.

Genetic parameters for fitness and neonatal behavior traits in sheep.

Poor neonatal survival constrains productivity and good welfare. The heritability of survival in sheep is very low, suggesting that genetic progress will be slow. Previously we have shown that a difficult birth and low neonatal lamb vigor are important predictors of future survival. In this study we investigated the heritability of these traits, and their relationship to production traits, as an alternative indirect route to improve lamb survival. Neonatal lamb data from 11,092 animals were collected over 2 years from 290 commercial sheep flocks, using previously developed methods to rapidly assess three traits (birth assistance, lamb vigor, sucking ability) on farm. Heritabilities for neonatal traits were moderate: birth assistance (mean ± standard error; 0.26 ± 0.03), lamb vigor (0.40 ± 0.04) and sucking ability (0.32 ± 0.03). Genetic correlations between neonatal traits were moderate to high, and positive. Heritabilities for production traits were also moderate: 8-week weight (0.27 ± 0.06), 20-week weight (0.39 ± 0.07), ultrasound muscle depth (0.37 ± 0.06). Genetic and phenotypic correlations between the neonatal traits and production traits were not significantly different from zero. However, lambs that were scored as of poor vigor at birth were less likely to be recorded at 8 or 20 weeks, indicating that they may have died. The data demonstrate that the neonatal survival traits of birth assistance, lamb vigor and sucking assistance are moderately heritable when treated as a lamb trait, indicating that selection to target these lamb traits would successfully, and efficiently, improve survival without influencing productivity.

Short communication: survival, growth to weaning, and subsequent fertility of live-born dairy heifers after a difficult birth.

The experience of a difficult birth (dystocia) is traumatic and has adverse effects on the newborn in various species. Despite affecting up to 1 in 3 births in dairy cattle, studies on calves have been mostly limited to the first day of life. The objective of this study was to investigate the effects of dystocia on the survival to calving, growth to weaning, and subsequent fertility as nulliparous animals. Historical data from live-born Holstein heifer calves born from cows with various birth difficulty scores (no assistance; moderate; high difficulty) were obtained from 2 herds (Edinburgh herd: n=1,237; Crichton Royal Farm herd: n=721). Each herd was analyzed separately for birth weights, weaning weights, growth rate to weaning, number of services to conception, and age at first calving using REML and generalized linear mixed model analyses. Survival analysis (Cox proportional hazards model) was used in the Edinburgh herd to analyze the subsequent survival of live-born heifers, whereas descriptive data are presented for the Crichton Royal Farm herd. A higher mortality risk to weaning and to first service was observed in the live-born heifers that experienced moderate difficulty at birth compared with heifers born naturally. Surviving dystocial heifers had similar growth-to-weaning and fertility performance as heifers born naturally in both herds. It could be that the performance of dystocial heifers that survived to weaning was not affected or that it was compensated for by farm management. This study highlights long-term effects of the early experience of a difficult birth and thereby stresses the importance of preventing dystocia not just from the point of view of the adult cow, but also from the perspective of the calf. This would also improve farm efficiency and calf welfare.

Effect of a difficult calving on the vigour of the calf, the onset of maternal behaviour, and some behavioural indicators of pain in the dam.

The neonate's development and survival is dependent upon being vigorous at birth and receiving appropriate maternal care. However, difficulty at delivery can result in less vigorous offspring and maternal care can be altered, probably as a consequence of exhaustion, pain and human intervention. The first 3h after expulsion of the calf were observed continuously from videos following twelve natural calvings and sixteen calvings assisted by farm staff (including four malpresentations) from Holstein cows. Calvings were balanced within groups for parity of the dam, genetic group, sex and birth weight of the calf, calving pen and calving season. Assisted calves were less vigorous with higher latencies to attempt to stand, achieve standing, walk and reach the udder than unassisted calves (P<0.05). Furthermore, assisted calves also tended to be less likely to stand and walk within the first 3h after birth (P<0.1), spent more time lying on their flank (P=0.019) and had more frequent bouts of this behaviour (P=0.033). Assisted dams did not take longer to lick the calf and performed as much licking as unassisted dams (P>0.05), indicating no delayed onset or impaired expression of maternal behaviour in dams given assistance at delivery. Study of potential pain-related behaviours revealed that assisted dams spent less time self-grooming (P=0.033) than dams delivering naturally, which could suggest greater pain. However, there were no significant differences in any of the other pain-related behaviours. Our results suggest that, although maternal behaviour was unaffected by a difficult delivery, dairy calves born following difficult calvings have lower vigour in the first 3h after birth than unassisted calves. This might have longer-term effects on the health and survival of the calves.

Breed differences in fetal and placental development and feto-maternal amino acid status following nutrient restriction during early and mid pregnancy in Scottish Blackface and Suffolk sheep.

This study assessed the effect of feeding 0.75 energy requirements between Days 1 and 90 of pregnancy on placental development and feto-placental amino acid status on Day 125 of pregnancy in Scottish Blackface and Suffolk ewes carrying a single fetus. Such moderate nutrient restriction did not affect placental size, placentome number or the distribution of placentome types. Although fetal weight was unaffected by maternal nutrition, fetuses carried by nutrient restricted mothers had relatively lighter brains and gastrocnemius muscles. Suffolk fetuses were heavier and longer with a greater abdominal circumference, relatively lighter brains, hearts and kidneys, but heavier spleens, livers and gastrocnemius muscles than Blackface fetuses. Total placentome weight was greater in Suffolk than Blackface ewes. Ewe breed had a greater effect on amino acid concentrations than nutrition. Ratios of maternal to fetal amino acid concentrations were greater in Suffolk ewes than Blackface ewes, particularly for some essential amino acids. The heavier liver and muscles in Suffolk fetuses may suggest increased amino acid transport across the Suffolk placenta in the absence of breed differences in gross placental efficiency. These data provide evidence of differences in nutrient handling and partitioning between the maternal body and the fetus in the two breeds studied.

Development and validation of on-farm behavioural scoring systems to assess birth assistance and lamb vigour.

Lamb mortality remains a significant welfare and economic issue for sheep production. Lamb survival is to a degree dependent upon an easy delivery and the expression of appropriate behaviours from both mother and offspring, such as rapid standing, udder seeking and sucking by the lamb. Genetic solutions have the potential to improve birth assistance and lamb behaviour but large amounts of data are needed. Therefore, to achieve this objective, simple, proxy methods (scoring systems) were developed to quantify the level of birth difficulties and lamb vigour on farm. In the first study, detailed historical behavioural data from 1156 lambs (Scottish Blackface and Suffolk (S)) were analysed to develop criteria for 3 scores: birth assistance, lamb vigour and sucking assistance. The birth assistance score was developed by analysing the relationships between birth presentation and intervention levels, and intervention level and labour length. Lambs with abnormal birth positions required more assistance than normally presented lambs and lambs with long labours required more and greater assistance than those with short labours. Lamb vigour score was developed by analysing the latencies for the lamb to first perform specific behaviours; more vigorous lambs reach landmark behaviours faster than low vigour lambs. The sucking assistance score was developed from the relationship between the latency to suck successfully and assistance level, where lambs that were slow to suck required more assistance than lambs that were quick to suck. In the second study, the behaviour scoring systems (5-point categorical scales) were validated using a commercial flock of 80 twin-bearing crossbred ewes mated with either Texel (T) or S sires by simultaneously recording scores and the latency to perform specific landmark behaviours (i.e. to stand, seek the udder and suck). The vigour scores (recorded at 5 min of age) were compared with the latency from birth to standing and showed that lambs with lower (better) vigour scores were faster to stand after birth than those with higher scores. The sucking assistance scores were compared with the latency from birth to sucking, and showed that lambs with lower sucking assistance scores are quicker to suck than those with high scores. These results showed that the scoring systems could provide a practical and reliable assessment of birth assistance and lamb behaviour on farm and were sufficiently sensitive to discriminate vigour levels between lambs sired by either S or T rams.

Differential effects of maternal undernutrition between days 1 and 90 of pregnancy on ewe and lamb performance and lamb parasitism in hill or lowland breeds.

This study examined the hypothesis that responses of Scottish Blackface (BF, a hill breed) and Suffolk (SUF, a lowland breed) ewes to undernutrition between d 1 and 90 of pregnancy would differ. Over 2 consecutive breeding seasons, ewes (4 pens/treatment; 15 to 20 ewes/pen) were artificially inseminated and from d 1 to 90 after AI allocated to 0.75 (restricted; RES) or 1.0 (control; CON) energy requirements for ewe maintenance and fetal growth. From d 90 to term, all ewes were allocated feed to meet requirements for stage of pregnancy and expected litter size. On d 90, RES ewes had lighter BW (P = 0.001) and smaller BCS (P = 0.019) than CON ewes. Plasma NEFA concentrations were greater in RES than CON ewes (P = 0.048) at d 60 of pregnancy. Pregnancy length was longer for RES than CON ewes (P = 0.003). Lambs from SUF-RES ewes had lighter birth weights than SUF-CON lambs, but BF-RES lamb birth weights were not different from BF-CON lambs (interaction, P = 0.066). However, maternal undernutrition did not affect BW at weaning (P > 0.45). Between birth and 3 d of age, BF lambs maintained greater body temperatures (P < 0.001) and plasma concentrations of triiodothyronine (T3, P < 0.001) and thyroxine (P < 0.001) than SUF lambs. Lambs from RES ewes had greater concentrations of T3 (P = 0.026) than CON lambs, whereas these differences were greater between BF-RES and BF-CON lambs than between SUF-RES and SUF-CON lambs (interaction, P = 0.028). Maternal undernutrition did not affect lamb rectal temperature (P > 0.27). In yr 1 only, fewer lambs (P = 0.022) were reared to weaning by RES than CON ewes. Similarly, in yr 1 only, other strongyle fecal egg counts at weaning were greater in SUF-RES lambs than SUF-CON, BF-CON, or BF-RES lambs (interaction, P = 0.099). This experiment showed that lambs of a breed selected for lean tissue growth and normally maintained in a lowland environment were more affected by maternal undernutrition between d 1 and 90 of pregnancy than lambs of a hill breed managed in a more adverse environment.

Assessing the welfare challenges to out-wintered pregnant suckler cows.

Out-wintering beef cows reduces annual housing costs and bedding requirements and there is less exposure to diseases associated with housing. However, to counter these benefits cows may be exposed to conditions that pose a significant challenge to welfare, and ways of assessing this are required. Two feeding treatments were applied to four groups of 10 cows (two groups/treatment), one to maintain condition score (H) and the other to allow a modest loss of condition score (L), which is commonly applied in farm practice. Cow groups were rotated around four paddocks in a Latin Square design of four periods each of 3 weeks, and they were weighed and condition was scored at the end of each period. Their behaviour and location was recorded at 30-min intervals with six 3-h sessions in each period. Ambient temperature, wind speed, rainfall and solar radiation were recorded every 30 min to enable calculation of cow lower critical temperature (LCT). The climatic conditions were wet at the start of the experiment with moderate wind speeds throughout (5 m/s) and relatively mild ambient temperature (5°C). Feeding treatment had no significant effect on any of the variables measured. Cows spent most of the observation sessions standing, particularly at the beginning of the experiment when the soil conditions were wettest. They sought sheltered locations when wind speeds were high and thus their calculated LCT was near or below ambient temperature. Nutritional models predicted periods of cold stress but the cows adapted their behaviour to counteract this, emphasising the need for a combined physical and behavioural approach to assessing welfare challenges.

The effect of late pregnancy supplementation of ewes with vitamin E on lamb vigour.

The experiment measured lamb responses to supplementation of the pregnant ewe diet with vitamin E above requirement. Crossbred ewes were mated with either Suffolk or Texel rams. Twin-bearing ewes were randomly allocated (approximately 21 months of age at allocation) to one of four treatment groups (20 ewes per group, 10 mated with Suffolk and 10 with Texel rams). Treatments imposed were 50, 100, 150 or 250 IU supplementary vitamin E per ewe per day to give a four treatment by two sire-type factorial experimental design. Ewes were fed concentrates to meet energy requirements for stage of pregnancy and hay ad libitum. Diets were introduced approximately 6 weeks before lambing. Blood samples were obtained prior to introduction of diets, 17 days after introduction of diets and within 24 h of lambing from a subset of eight ewes per treatment (32 total). Colostrum samples were obtained from 10 ewes per treatment, 12 h after birth of the first lamb. All births were observed and a lamb vigour score was assigned to each lamb 5 min after birth. At 1 and 12 h after birth, rectal temperature, and at 12 h after birth, sex, crown-rump length and BW of each lamb were recorded. Mean ewe plasma α-tocopherol concentration prior to introduction of the diets was 1.5 µg/ml (s.e.m. 0.09) and did not differ between groups. There were positive linear (P < 0.001) effects of dietary vitamin E on plasma (17 days after introduction of diets) and colostrum (12 h after birth) α-tocopherol concentrations. Lamb vigour scores were superior (P < 0.001) for lambs sired by Texel rather than Suffolk rams but there were no differences as a result of vitamin E supplementation. Lamb mortality was low and unrelated to either sire or supplementary vitamin E. Lamb birth and weaning weights were also unaffected by vitamin E supplementation. Supplementing the ewe with vitamin E therefore had no effect on any lamb measurements.

Individual variation in the expression of maternal behaviour: a review of the neuroendocrine mechanisms in the sheep.

Individual variation in the expression of maternal behaviour can affect offspring development and survival. Ewes showing a poorer quality of maternal behaviour (low levels of licking and grooming, infrequent low-pitched bleating, inconsistent udder acceptance) at parturition and throughout lactation (large ewe-lamb distance, low vigilance) form a weaker bond with their lambs, and have higher lamb mortality, than ewes which show high levels of grooming and low-pitched bleating, close ewe-lamb distances and high levels of vigilance. Using two models of variation in maternal behaviour in the sheep (differences between ewes of two breeds, and differences between primiparous and multiparous ewes), the neuroendocrine mechanisms that underpin these differences are explored. In both cases, significant variation in oestrogen priming is observed, with breed differences in circulating concentrations of oestrogen in late gestation, and parity effects on receptor density but not circulating concentrations. The consequences of these differences for central oxytocin release and binding are discussed. In addition, preliminary data suggest that opioid modulation, in both breed and parity models, may also play a role in mediating individual variation in maternal behaviour in the sheep.

The potential for improving physiological, behavioural and immunological responses in the neonatal lamb by trace element and vitamin supplementation of the ewe.

Neonatal lamb mortality represents both a welfare issue (due to the considerable suffering and distress) and an important production inefficiency. In lambs, approximately 80% of mortality can be attributed to the starvation-mismothering-exposure complex and occurs in the first 3 days after birth. It was the object of this review to determine the micronutrient(s) most likely to have a positive effect on neonatal lamb survival when included above the requirement for that micronutrient. Micronutrients discussed were Co, Cu, I, Fe, Mn, Se, Zn, vitamins A and E and n-3 fatty acids. For Co, Fe, Mn and Zn, there was no evidence of positive responses to supplementation. Cu and I had toxicity thresholds that were sufficiently close to requirement that supplementing above requirement presented a risk of inducing toxicity. In the case of vitamin A, while serum concentrations indicated that sub-optimal status did exist, long-term buffering from liver stores (from grazing) makes experimentation difficult and practical benefits to supplementation unlikely. Therefore, the most likely candidates for supplementation were Se, vitamin E and fatty acids. Fatty acid supplementation with fish oils or docosahexaenoic acid-containing algal biomass consistently improved lamb vigour but it is unlikely that supplementation will be economic. Positive responses to Se supplementation throughout gestation were recorded. However, in many studies the Se status of control ewes was marginal and there is a need for more studies where control ewes are clearly adequate in Se. Positive responses to vitamin E supplementation above requirement in the last-third of gestation were observed but the optimum dietary inclusion of vitamin E and period of feeding during pregnancy still require clarification.

Genetic and physiological determinants of maternal behavior and lamb survival: implications for low-input sheep management.

The relatively intensive supervision afforded many ewes at lambing time is a barrier to the development of low-input sheep management systems. However, in some flocks, reduction in this level of supervision may initially affect lamb mortality and animal welfare. In this review, possibilities for optimizing behavioral interaction between the ewe and lamb are considered, with the goal of improving lamb survival without the need for high levels of human supervision. At birth, ewes show specific behavioral patterns (e.g., licking or grooming, low-pitched bleats, udder acceptance) that facilitate the transition of the lamb from pre- to postnatal life and that accompany the formation of an exclusive olfactory memory for the lamb. The lamb also performs a specific sequence of behaviors directed toward standing, finding the udder, and sucking. The successful accomplishment of these behavior patterns is vital for the formation of a strong attachment between both partners, and for lamb survival. The expression of maternal behavior in the ewe is affected by her previous maternal experience, by nutrition in pregnancy, by breed, by temperament, and, to some extent, by the behavior of her lamb. The maternal care expressed by a ewe at parturition is indicative of her behavior throughout that lactation and in successive pregnancies, suggesting an underlying basis to maternal care intrinsic to that ewe. Studies with Scottish Blackface and Suffolk ewes show that ewes expressing high levels of maternal care have elevated plasma estradiol in late gestation compared with ewes with poorer maternal care, and that circulating estradiol concentration is correlated with maternal behaviors. Although the genetic basis of maternal behaviors has still to be fully determined, there are possibilities of improving maternal behavior by selection, and a better understanding of the neuroendocrine processes underlying individual differences in maternal behavior may help in developing selection strategies. In addition, selection on lamb behaviors, which show some genetic basis, may also be a route to improve lamb survival. Because behavior of both the ewe and lamb is affected by environmental factors, appropriate management, through pregnancy and at parturition, will enhance the expression of maternal behavior and lamb vigor, and so contribute to improving lamb survival.

Maintenance of body temperature in the neonatal lamb: effects of breed, birth weight, and litter size.

To survive, the newborn lamb must be able to maintain body temperature and to stand and move to the udder to suck colostrum to fuel heat production. The objective of this study was to investigate whether neonatal lambs showing slow behavioral progress to standing and sucking also have an impaired ability to maintain body temperature. The time taken to stand and suck after birth, rectal temperatures, and blood samples were collected from 115 newborn single, twin, and triplet lambs of 2 breeds, Scottish Blackface and Suffolk, which are known to show variations in their neonatal behavioral progress. Blood samples were assayed for thyroid hormones, known to be involved in heat production, and cortisol, which plays a role in tissue maturation before birth. In addition, colostrum samples were collected from the 56 ewes that gave birth to the lambs, and assayed for protein, fat, and vitamin contents. Heavy lambs (more than 1 SD above the breed mean), Blackface lambs, and singleton or twin lambs were quicker to stand and suck from their mothers than lightweight (more than 1 SD below the breed mean), Suffolk, or triplet lambs. Low birth weight lambs also had lower rectal temperatures than heavier lambs (P < 0.01), as did Suffolk compared with Blackface lambs (P < 0.001). Lambs that were slow to suck after birth had lower rectal temperatures than quick lambs, and this difference persisted for at least 3 d after birth. Within breed, heavy lambs had greater plasma triiodothyronine and thyroxine immediately after birth than light lambs. Blackface lambs had greater plasma triiodothyronine and thyroxine than Suffolk lambs but tended to have less cortisol. Colostrum produced by Blackface ewes had a greater fat content than that of Suffolk ewes (P < 0.001). Thus, lambs that are behaviorally slow at birth are also less able to maintain their body temperature after birth. Although part of their lower body temperature might be attributable to behavioral influences on thermoregulation, the data also suggest that physiological differences exist between these animals. These differences may be related to different degrees of maturity at birth.