Humans can identify reward-related call types of chickens.

Humans can decode emotional information from vocalizations of animals. However, little is known if these interpretations relate to the ability of humans to identify if calls were made in a rewarded or non-rewarded context. We tested whether humans could identify calls made by chickens (Gallus gallus) in these contexts, and whether demographic factors or experience with chickens affected their correct identification context and the ratings of perceived positive and negative emotions (valence) and excitement (arousal) of chickens. Participants (n = 194) listened to eight calls when chickens were anticipating a reward, and eight calls in non-rewarded contexts, and indicated whether the vocalizing chicken was experiencing pleasure/displeasure, and high/low excitement, using visual analogue scales. Sixty-nine per cent of participants correctly assigned reward and non-reward calls to their respective categories. Participants performed better at categorizing reward-related calls, with 71% of reward calls classified correctly, compared with 67% of non-reward calls. Older people were less accurate in context identification. Older people's ratings of the excitement or arousal levels of reward-related calls were higher than younger people's ratings, while older people rated non-reward calls as representing higher positive emotions or pleasure (higher valence) compared to ratings made by younger people. Our study strengthens evidence that humans perceive emotions across different taxa, and that specific acoustic cues may embody a homologous signalling system among vertebrates. Importantly, humans could identify reward-related calls, and this ability could enhance the management of farmed chickens to improve their welfare.

Exploration of cultural norms and behavioural beliefs about zoo animal behaviour, welfare, ethics and husbandry practices in a sample of the international zoo community.

Beliefs influence the intentions of people to behave in certain ways towards animals. This study presents survey responses from 237 people working in zoos in China and Europe and describes their demographic characteristics. It explores their beliefs about zoo animal behaviour, welfare and ethical issues, and zoo practices, using a survey methodology. These beliefs may be influenced by individual demographic or cultural factors such as age, gender and region of employment, as well as experiential or situative 'norms' within the work environment. Beliefs were significantly influenced by the region of employment with Chinese respondents beliefs being significantly different to beliefs from respondents in the United Kingdom or the rest of Europe. Hierarchical cluster analysis of the survey generated clusters of people from both regions who indicated positive beliefs about zoo animal welfare as well as clu sters indicating a lack of understanding of some zoo animal welfare issues. In addition, a cluster suggesting cognitive dissonance between beliefs about animals welfare and zoo practices was generated from Chinese responses. Factor analysis identified that prioritisation of in-situ conservation within good animal welfare was a key feature in Chinese respondents, whereas European beliefs prioritising in-situ conservation were distinct from those on supporting good animal welfare. This paper identifies similarities and differences in beliefs about zoo animal welfare and zoo husbandry practices between Europe and China, and discusses the underlying norms and values that these beliefs may reflect.

Zookeepers - The most important animal in the zoo?

Education to improve knowledge of animal welfare is not a universal component of training for zoo staff, and little is reported about the perspectives of zoo staff on the need for such education. This paper reports results from structured telephone interviews of a diverse sample of eight Chinese and eight European zoo staff about aspects of zoological animal welfare, education and zoological practices. These qualitative data were thematically analyzed and key themes generated. Similar themes emerged across regions: Zoo staff consider professional attributes including motivation and enthusiasm to be important alongside formal training, zoo staff value learning opportunities but don't always feel supported, and contextual information including wild animal ecology was considered important content in zoo animal welfare education.

A "Good Life" for Dairy Cattle: Developing and Piloting a Framework for Assessing Positive Welfare Opportunities Based on Scientific Evidence and Farmer Expertise.

On-farm welfare assessment tends to focus on minimising negative welfare, but providing positive welfare is important in order to give animals a good life. This study developed a positive welfare framework for dairy cows based on the existing scientific literature which has focused on developing positive welfare indicators, and trialled a participatory approach with farmers; refining the framework based on their recommendations, followed by a vet pilot phase on farm. The results revealed that farmers and scientists agree on what constitutes "a good life" for dairy cattle. Farmers value positive welfare because they value their cows' quality of life, and want to be proud of their work, improve their own wellbeing as well as receive business benefits. For each good life resource, the proportion of farmers going above and beyond legislation ranged from 27 to 84%. Furthermore, barriers to achieving positive welfare opportunities, including monetary and time costs, were not apparently insurmountable if implementation costs were remunerated (by the government). However, the intrinsic value in providing such opportunities also incentivises farmers. Overall, most farmers appeared to support positive welfare assessment, with the largest proportion (50%) supporting its use within existing farm assurance schemes, or to justify national and global marketing claims. Collaborating with farmers to co-create policy is crucial to showcase and quantify the UK's high welfare standards, and to maximise engagement, relevance and uptake of animal welfare policy, to ensure continuous improvement and leadership in the quality of lives for farm animals.

Is Animal Welfare an Internationally Understood Concept in the Zoo World? Thematic Analysis of Two Regional Groups of Zoo Staff.

Universal frameworks for zoo animal welfare have been suggested. However, there is little evidence of a cross-cultural understanding of zoo animal welfare. This paper reports themes emerging from a qualitative study of international (European and Chinese) zoo professionals on zoo animal welfare issues. Structured telephone interviews were conducted with eight Chinese and eight European zoo staff, covering aspects of zoological animal welfare, conservation and zoological husbandry practices. These qualitative data were thematically analysed, and key themes generated. This paper describes three themes relating to 'What is animal welfare' 'Animal welfare frameworks' and 'The human element in animal welfare'. This analysis indicates that the concept of animal welfare has cultural equivalence across Europe and between Europe and China, and that zoo staff are familiar with welfare frameworks. In China, a need for senior leadership and motivating staff to improve animal welfare emerged.

'All My Animals Are Equal, but None Can Survive without the Horse'. The Contribution of Working Equids to the Livelihoods of Women across Six Communities in the Chimaltenango Region of Guatemala.

It is widely assumed that working equid husbandry is carried out by men, and women are often not recognised as facilitating equid welfare. The aim of this study is to investigate how working equids contribute to women's livelihoods in six of the World Horse Welfare programme target communities in Guatemala and determine what roles women have in their care. Thirty-four face-to-face interviews were carried out and data were analysed using both quantitative and qualitative methods. This study found that working equids support women by reducing domestic drudgery, generating income, feeding livestock and saving time. Thirty-two women played a major role in the care of one or more equids, and overall, women did not feel that they knew enough about equid husbandry. Thirty-one women said they would attend training opportunities if the advertising was clear and they felt that women were able to join. This study recognises the contribution of working equids to women's livelihoods, describes the roles women play in equid husbandry and addresses the discrepancies between women's roles and their capacity to undertake these tasks. This emphasises the need for extension services to include and cater for women, improving equid welfare and their ability to continue supporting women's livelihoods.

A Survey of Farm Management Practices Relating to the Risk Factors, Prevalence, and Causes of Lamb Mortality in Ireland.

Lamb mortality is a key factor influencing ewe productivity and profitability. The current study investigated risk factors associated with and management practices implemented on sheep farms to reduce lamb mortality. A survey consisting of 13 multiple-part questions (57 separate questions) was administered to all sheep farmers participating in the Teagasc National Farm Survey, representative of the Irish national population of sheep farms. A total of 60% of respondents identify mating or lambing date, and this practice tended to be associated with reduced lamb mortality (1.2%, p = 0.08). Individual lambing pens were used by 88% of farmers, but 26% did not clean or disinfect them. A total of 79% and 9.5% of farmers applied iodine to all lambs' navels and administered antibiotics to all lambs to treat and/or prevent diseases, respectively. Most farmers vaccinated their ewes (86%) and lambs (79%) against clostridial diseases and/or pasteurellosis; 13% vaccinated against abortion agents. Lamb mortality tended to be lower (Kruskal-Wallis (KW) = 2.749; p = 0.09) on farms that used stomach tubing, heat box, iodine, hospital, and individual pens compared with farms that do not implement all those practices. Predators, lamb birth weight, and diseases were perceived by respondents to be the three main causes of live-born lamb mortality. The gross margin is significantly higher on lowland farms by €37 per ewe compared with hill farms (Kruskal-Wallis (KW) = 4.056; p < 0.001). The combination of full-time farming and the use of hospital and individual pens improved gross margin (€18/ewe, p = 0.028). It is concluded that on-farm management practices affect both lamb mortality and flock gross margin.

Determining a Welfare Prioritization for Horses Using a Delphi Method.

Equine welfare issues are receiving increasing attention in the UK, but welfare problems can arise from a wide range of causes. In order to identify the most important welfare concerns for horses, we used a Delphi method with 19 equine welfare experts. An initial list of 84 equine welfare issues was generated using an online discussion board and NVivo thematic analysis. Subsequently, experts ranked these welfare issues for perceived prevalence, severity and duration of suffering associated with each issue on a 6-point Likert scale. All issues with a mean score of 3 or above (n = 37) were included in subsequent rounds. Finally, a subset of experts attended a two-day workshop to determine the final priority list of welfare issues. The welfare issues perceived to be most prevalent were lack of biosecurity, delayed euthanasia, lack of owner knowledge of equine welfare needs, fear and stress from use, and obesity. The issues considered to cause greatest suffering for individual horses were delayed euthanasia, lack of recognition by owners of pain behaviour, large worm burdens, obesity and being fed unsuitable diets for equine feeding behaviour. These outcomes can help to focus research and education interventions on the most pressing welfare issues for horses.

Prioritization of Farm Animal Welfare Issues Using Expert Consensus.

Prioritization of animal welfare issues can help identify which areas most require research funding and raise awareness of best practices. A modified Delphi method was used to obtain expert opinion on the highest priority welfare issues for UK farmed livestock. Fifty-eight UK-based experts were recruited onto the study, with a minimum of 3 years experience of working with either cattle, pigs, poultry, or small ruminants (12-16 experts per group). Experts were chosen to represent a broad range of opinions. Two rounds of surveys were conducted online using Online Survey, and the final round was an in-person workshop with 21 experts. In the first survey, experts were provided with a comprehensive list of species-specific welfare issues derived from the literature. Participants were asked to score each welfare issue, for (i) severity, (ii) duration, and (iii) prevalence on a 6-point Likert scale. The results of the first survey were reviewed and the welfare issues which scored a neutral-to-high response (scores 3-6) were carried forward. In round 2, participants were asked whether they agreed or disagreed with the rankings that were made from the results of round 1. The final stage of the process was a workshop, which consisted of a combination of group exercises and discussions, to reach the final consensus. Welfare priority lists were divided into two categories: severity/duration, and prevalence, to identify the priority welfare issues affecting individual animals and the population, respectively. Across all farmed species common concerns were inadequate or inappropriate nutrition, inability of stockpeople to recognize or treat welfare issues (such as pain or behavioral problems), lameness, chronic or endemic health issues, euthanasia and mortality and morbidity of neonates. Specific concerns related to behavioral restriction and damaging or abnormal behavior in pigs, poultry and dairy animals, inadequate housing for pigs and poultry, consequences of breeding decisions in pigs and poultry, and lack of access to veterinary care in sheep and beef. This Delphi process resulted in consensus on the most significant welfare challenges faced by UK livestock species and can help to guide future research and education priority decisions.

Evaluation of Animal-Based Indicators to Be Used in a Welfare Assessment Protocol for Sheep.

Sheep are managed under a variety of different environments (continually outdoors, partially outdoors with seasonal or diurnal variation, continuously indoors) and for different purposes, which makes assessing welfare challenging. This diversity means that resource-based indicators are not particularly useful and, thus, a welfare assessment scheme for sheep, focusing on animal-based indicators, was developed. We focus specifically on ewes, as the most numerous group of sheep present on farm, although many of the indicators may also have relevance to adult male sheep. Using the Welfare Quality® framework of four Principles and 12 Criteria, we considered the validity, reliability, and feasibility of 46 putative animal-based indicators derived from the literature for these criteria. Where animal-based indicators were potentially unreliably or were not considered feasible, we also considered the resource-based indicators of access to water, stocking density, and floor slipperiness. With the exception of the criteria "Absence of prolonged thirst," we suggest at least one animal-based indicator for each welfare criterion. As a minimum, face validity was available for all indicators; however, for many, we found evidence of convergent validity and discriminant validity (e.g., lameness as measured by gait score, body condition score). The reliability of most of the physical and health measures has been tested in the field and found to be appropriate for use in welfare assessment. However, for the majority of the proposed behavioral indicators (lying synchrony, social withdrawal, postures associated with pain, vocalizations, stereotypy, vigilance, response to surprise, and human approach test), this still needs to be tested. In conclusion, the comprehensive assessment of sheep welfare through largely animal-based measures is supported by the literature through the use of indicators focusing on specific aspects of sheep biology. Further work is required for some indicators to ensure that measures are reliable when used in commercial settings.

Positive and negative gestational handling influences placental traits and mother-offspring behavior in dairy goats.

Dairy animals are subjected to a number of potential stressors throughout their lives, including daily interactions with humans. The quality of these interactions may have direct consequences for the animal undergoing the experience, but if such events occur during gestation it may also affect the developing fetus. This study examined the effects of differential handling during mid-gestation in 40 twin-bearing Saanen×Toggenburg primiparous goats. Between days 80 and 115 of gestation (gestation=150 days), goats were subjected to aversive (AVS, n=13), gentle (GEN, n=13) or minimal (M, n=14) handling protocols for 10 minute periods twice daily. The control (M) group did not receive handling treatments and all goats received normal husbandry procedures outside treatment periods. Salivary cortisol measured during the treatment period was higher in AVS goats (mean cortisol (sem) in pg/µl: AVS: 176.7 (18.2), GEN: 119.6 (11.1), M: 126.5 (13.7); P=0.007). Data collection was focussed on mother-offspring behaviors 2h post-partum, placental morphology and colostrum quality. AVS goats were the only treatment group to suffer fetal loss (16% loss vs 0% in GEN and M, P=0.05). Treatment also influenced placental morphology with a tendency for fewer cotyledons evident in placentae from the aversive treatment (AVS: 87.9 (7.8), GEN: 107.1 (7.9), M: 112.1 (9.3), P=0.093), and significantly fewer medium sized cotyledons (AVS: 67.6 (7.8), GEN: 89.3 (6.4), M: 84.3 (5.4), P=0.042). GEN goats displayed more grooming and nosing behaviors towards their young during the first 2h post-partum (grooming: GEN: 89.3% (7.1), AVS: 72.6% (7.7), M: 63.4% (9.0), P=0.045; nosing frequency: GEN: 58.8 (12.5), AVS: 28.6 (11.1), M: 34.7 (6.5), P=0.021). There was an overall trend for kids from mothers experiencing the AVS treatment to take longer to stand, reach the udder and suck compared to kids from GEN and M treatment groups. Treatment significantly affected latency to perform play behavior, with kids from AVS goats taking on average 25 min longer to play for the first time than kids from GEN and M treatment groups (P<0.001). The results show that handling during gestation affects placental morphology, fetal survival and post-partum maternal behaviors, and influences kid behavioral development. Such results have important animal welfare implications, demonstrating that negative handling of pregnant females results in poorer placental quality with potential for fetal loss. It also demonstrates the beneficial effects of positive handling on enhancement of maternal behaviors.

Ewes Direct Most Maternal Attention towards Lambs that Show the Greatest Pain-Related Behavioural Responses.

Although neonatal farm animals are frequently subjected to painful management procedures, the role of maternal behaviour in pain coping, has not been much studied. We investigated whether ewes were able to distinguish between lambs in pain and those that were not, and whether their behaviour altered depending on the severity of lamb pain. Eighty male lambs were allocated to one of 4 pain treatments within 24 hours of birth. Lambs were either handled only (C), bilaterally castrated with tight rubber rings (RR), as for RR but with the application of a Burdizzo clamp immediately proximal to the ring (Combined) or subjected to short scrotum castration (SSC) where the testicles were retained within the abdomen and only the scrotum removed. The behaviour of the ewe, treated lamb and untreated sibling where present (n = 54) were recorded for 30 minutes after treatment. Castration treatment increased the expression of abnormal standing and lying postures, specific pain-related behaviours (head-turning, stamping/kicking, easing quarters, tail wagging) and composite pain scores (P<0.001 for all). The greatest expression of pain-related behaviours was shown by lambs in the RR group, which were the only group to show rolling responses indicative of severe pain, followed by the SSC group. Ewes expressed more licking/sniffing responses to the RR and SSC lambs than towards the Combined and C lambs (P<0.05), and oriented most to RR lambs and least to C lambs (P<0.001). Ewes with two lambs also directed more attention towards the treated than the untreated lamb (P<0.001). The quantity of maternal care directed towards the lamb was positively correlated with the expression of active pain behaviours. The data demonstrate that ewes are able to discriminate between lambs in pain and those that are not, and that their response is increased with a greater severity of pain.

Factors affecting dystocia and offspring vigour in different sheep genotypes.

Birth difficulty and poor lamb vigour are significant causes of perinatal lamb mortality. In this study we investigated whether sheep breeds differing in appearance, muscularity and selection history also had differences in dystocia and lamb vigour, and considered some of the factors that may contribute to the variation in these traits. Data were collected at birth from a total of 3252 lambs of two terminal sire breeds selected for lean growth (Suffolk [S], n=500 and Texel [T], n=1207), from a Hill breed (Scottish Blackface [B], n=610), which has been mainly selected for hardiness, and a crossbred (Mule×T [M], n=935) representing a maternal line. For each lamb the degree of assistance at delivery, lamb presentation, amount of assistance to achieve successful sucking, sex, litter size and birth weight were recorded. T lambs required the most, and B and M lambs the least assistance at birth, S lambs were intermediate (% lambs assisted: T=55.7, S=30.7, B=22.7, M=24.9, P<0.001). T and S lambs were equally likely to be malpresented at birth (29% of births) and more likely to be malpresented than B or M lambs (20%; P<0.001). In T and S breeds lambs requiring veterinary assistance at delivery were mainly heavy and singleton lambs, whereas in B and M breeds these were exclusively low birth weight lambs in multiple litters. Although heavier lambs needed greater birth assistance, T lambs were lighter than S and M lambs, but heavier than B lambs (birth weight (kg): S=4.66, M=4.56, T=4.32, B=3.67, P<0.001). S lambs were more likely to require assistance with sucking than other breeds, and T lambs also required more assistance than B or M lambs (% lambs assisted to suck: S=56.0, T=31.6, M=19.8, B=18.4, P<0.001). Heavier lambs were more likely to suck unaided than lighter lambs (P<0.001). The data suggest that the two terminal sire breeds, selected narrowly for greater productivity (muscle growth and conformation), are more likely to experience birth difficulty and poorer lamb vigour than the breed selected for hardiness, or the cross breed. Whether these effects arise as a consequence of genetic selection (e.g. for specific lamb conformation), or as a result of management practices to achieve selection goals (e.g. increased intervention at lambing) is unknown. Specific actions to improve birth difficulty and lamb vigour, such as including these traits in the selection index, would be beneficial in improving the welfare of ewes and lambs of the terminal sire breeds.

Pain in lambs castrated at 2 days using novel smaller and tighter rubber rings without and with local anaesthetic.

Without effective pain relief, rubber ring castration of lambs is acutely painful and can also produce chronic pain. The potential of novel, smaller rubber rings to reduce this pain substantially has been investigated. Three groups of eight 2-3 day old lambs, were castrated either with conventional rubber rings (cRR), or novel smaller rubber rings without (nRR) and with local anaesthetic treatment (nRR+La). Behavioural responses and chronic lesions indicative of pain were compared. No major reductions in behavioural responses were produced by castration with nRRs compared with cRRs, but injection of local anaesthetic did reduce these responses significantly. Lambs in all groups developed chronic inflammatory lesions but behavioural evidence of chronic pain was rarely seen. After 28 days, 6/8 necrotic scrotums had been shed by nRR+La; 4/8 by nRR and 2/8 by cRR lambs. The dimensions of rubber rings on necrotic scrotums, suggest that cRRs contracted to their un-stretched size before being shed, whereas nRRs did not. It is concluded that neither of these rubber rings exerted sufficient pressure on trapped nerves to produce anaesthesia rapidly and this may be explained by the physical properties of the tissues at the neck of the scrotum which support the ring.

The effect of gestational undernutrition on maternal weight change and fetal weight in lines of mice selected for different growth characteristics.

The present study investigated whether the genetic growth characteristics (fast or slow growing, lean or fat) of a mother influences her ability to partition nutrients to developing offspring. A total of sixty-one pregnant mice of three selected lines were used: fast-growing, relatively fat (FF, n 19); fast-growing, relatively lean (FL, n 23); and normal growth, relatively lean (NL, n 19). On day 1 of pregnancy, mice were given either ad libitum access to food (control (C): n 32) or pair-fed at 80 % of C intake (restricted (R): n 29). Feed intake and dam weight were measured daily. The weight of the mouse, organs, mammary tissue and the weight of fetuses and placentas were determined at day 18 of gestation. Overall, R dams gained less than half the weight of C dams during gestation. NL dams gained the most weight, and FF dams gained the least weight (P < 0·001). R dams in the fast-growing lines mobilised significantly more body fat during gestation than the NL line (P < 0·001) and had a greater reduction in mammary tissue growth. The relative weight of the litter increased in R dams of the FF line but was reduced in both the lean lines. Undernutrition reduced fetal and placental weight, and reduced placental efficiency in all the lines. The reduction was least in the FF line and greatest in the FL line. The data suggest that selection of animals for different growth characteristics alters their response to undernutrition during pregnancy, the relatively fat line was better able to buffer its offspring from the effects of undernutrition than the lean lines, regardless of their underlying rate of growth.

Parity effects on maternal behaviour are not related to circulating oestradiol concentrations in two breeds of sheep.

Primiparous females of many species, including the sheep, do not show as competent maternal behaviour as multiparous mothers. In this study we investigated whether circulating concentrations of oestradiol might be related to this response, since breed differences in oestradiol concentrations have been shown to correlate with maternal behaviours. Oestradiol concentrations were measured in plasma collected at 2-weekly intervals from primiparous and multiparous ewes of two breeds (Scottish Blackface and Suffolk) throughout gestation, and maternal behaviour was recorded at parturition. Circulating oestradiol concentrations, and the ratio of oestradiol to progesterone concentration (O:P ratio) were higher in late pregnancy in Blackface ewes, but there were no significant effects of ewe parity. However, there was a significant interaction between breed and parity as O:P ratio was lower in primiparous Suffolk ewes compared to multiparous. This relationship was not seen in Blackface ewes. Blackface ewes of both parities groomed their lambs more frequently and made more low-pitched vocalisations than Suffolk ewes. Primiparous ewes were more likely to move as the lamb attempted to suck, and to show withdrawal or aggression towards the lamb. Primiparous Suffolk ewes also made more high pitched bleats than other classes of ewe. The data support previous studies which show an effect of breed on circulating oestradiol and O:P ratio but suggest that parity effects on maternal behaviour may be mediated by sensitivity to circulating concentration of oestradiol rather than variations in the plasma concentrations of this hormone.

Effects of cobalt/vitamin B12 status in ewes on ovum development and lamb viability at birth.

Scottish Blackface ewes from cobalt-deficient farmland were fed a diet containing 0.06 mg cobalt per kg dry matter from approximately 30 days before embryo recovery/transfer until lambing. Ewes remained untreated (-Co; n = 82) or were given an intraruminal cobalt-containing bolus to compensate for the dietary deficit (+Co; n = 82). Ewes used as embryo donors (-Co, n = 17; +Co, n = 16) were artificially inseminated with semen from a single Suffolk sire. Day 6 embryos obtained from -Co and +Co donors were transferred in singleton to -Co and +Co recipients in a 2 x 2 factorial-designed experiment to determine the effects of cobalt/vitamin B12 status during the periconception period (factor 1) and pregnancy (factor 2) on lamb viability at birth. Mean (+/- s.e.m.) circulating concentrations of vitamin B12 in -Co and +Co donors at ovum recovery were 182 +/- 10 and 1288 +/- 64 pmol L(-1), respectively (P < 0.001), and the number of corpora lutea per ewe ovulating was 9.9 +/- 1.6 and 14.4 +/- 1.3, respectively (P < 0.05). Treatment did not affect the proportion of recovered ova that contained >32 cells (viable) or the median stage of development (late morula), but viable ova recovered from -Co v. +Co ewes had a better morphological grade (2.0 +/- 0.1 v. 2.20 +/- 0.04, respectively; P < 0.01). There was no effect of treatment on the proportion of recipient ewes that became pregnant. Circulating concentrations of vitamin B12 were lower in -Co than +Co ewes during pregnancy (P < 0.001) and at birth in lambs born to -Co ewes compared with those born to +Co ewes (P < 0.001). There was no effect of donor or recipient cobalt/vitamin B12 status on lamb birthweight, neonatal vigour or neonatal rectal temperatures, but lambs derived from +Co v. -Co embryo donors were more active in the first 3 days after birth (P < 0.05). Results show that sub-clinical cobalt/vitamin B12 deficiency reduces ovulatory response in superovulated ewes and that periconception nutrition can affect neonatal lamb behaviour.

Prepartum plasma estradiol and postpartum cortisol, but not oxytocin, are associated with interindividual and breed differences in the expression of maternal behaviour in sheep.

Consistent, individual differences in the expression of maternal behaviour have been described in several species including the sheep. The neuroendocrine mechanisms underlying the onset of maternal behaviour in the sheep have been described, although the relationship between hormonal events and individual differences in behaviour has yet to be determined. In this study, we examined whether the individual differences in plasma estradiol, progesterone, oxytocin and cortisol concentrations were related to observed individual and breed differences in maternal behaviours in two breeds of sheep (Scottish Blackface and Suffolk) known to differ in maternal behavioural expression. Maternal estradiol concentration increased rapidly before parturition and was higher in Blackface ewes than Suffolk ewes. Plasma progesterone declined before parturition and was higher in Suffolk than Blackface ewes. Prepartum estradiol, but not progesterone, was related to individual differences in maternal grooming of the lamb. Plasma oxytocin did not differ between breeds in late gestation. There was a tendency for oxytocin to be higher in Blackface than Suffolk ewes immediately after birth. However, there were no significant relationships between prepartum or postpartum oxytocin and any maternal behaviours. Plasma cortisol was higher in Blackface than Suffolk ewes in the last days of pregnancy but rose in both breeds over the last 24 h before parturition and did not differ at delivery. Cortisol peripartum was negatively related to individual differences in maternal affiliative behaviours. These data suggest that estradiol, and potentially cortisol, may mediate individual differences in maternal behaviour in sheep.

Ewe-lamb bonding behaviours at birth are affected by maternal undernutrition in pregnancy.

Maternal undernutrition in pregnancy results in low birth-weights and impaired postnatal survival in sheep. Largely anecdotal evidence suggests that the expression of appropriate maternal and neonate behaviours may also be disrupted by undernutrition. In the present study, we investigated the effect of a moderate (35 %) reduction in ewe nutritional intake in pregnancy on the expression of ewe-lamb bonding behaviours in primiparous Scottish Blackface ewes. Low-intake (L) ewes had significantly higher plasma progesterone than high-intake (H) ewes from mid-gestation onwards (e.g. plasma progesterone at 20 weeks (ng/ml): H 15.72, L 22.38, sed 1.80, P<0.001), and a lower oestradiol: progesterone value than H ewes at delivery (H 0.46, L 0.35, sed 0.05, P<0.05). Lamb birth-weight was reduced in the L lambs compared with H lambs (mean body weight (kg): H 3.31, L 3.00, sed 0.14, P<0.05), but the incidence of malpresentation at delivery was greater in L lambs. L ewes spent significantly less time licking their lambs than H ewes after delivery (time grooming in 2 h after birth (%): H 56.12, L 48.17, sed 2.639, P<0.01) and were more aggressive towards the lambs. Lamb behaviours were not directly affected by maternal nutritional treatment, but lamb birth-weight had a significant effect on neonatal developmental progress. Low-birth-weight lambs were slower than heavier lambs to stand and sucked less frequently. In tests of maternal attachment to the lamb, H ewes received higher scores than L ewes at both 24 h after birth (ewes receiving high scores (%): H 41.3, L 21.4, P<0.05) and at 3 d postnatal. We conclude that even a moderate level of undernutrition impairs the attachment between ewes and lambs by affecting maternal behaviours expressed at birth. In addition, the results suggest that levels of nutrition resulting in a decrease in birth weight will affect neonatal lamb behavioural progress.