Susan Jarvis

Investigating the behavioural and physiological indicators of neonatal survival in pigs

Survival is reduced in low birth weight piglets, which display poor thermoregulatory abilities and are slow to acquire colostrum. Our aim was to identify additional behavioural and physiological indicators of piglet survival incorporating traits reflective of both the intrauterine and extrauterine environment. Data were collected from 135 piglets from 10 Large White x Landrace sows to investigate which physiological measurements (e.g. individual placental traits), and which behavioural measurements (e.g. the quantification of piglet vigour), were the best indicators of piglet survival. Generalised linear models confirmed piglet birth weight as a critical survival factor. However, with respect to stillborn mortality, piglet shape and size, as measured by ponderal index (birth weight/(crown-rump length)(3)), body mass index (birth weight/(crown-rump length)(2)), respectively, and farrowing birth order were better indicators. With respect to live-born mortality, postnatal survival factors identified as crucial were birth weight, vigour independent of birth weight, and the latency to first suckle. These results highlight the importance of the intrauterine environment for postnatal physiological and behavioural adaptation and identify additional factors influencing piglet neonatal survival.

Programming the offspring of the pig by prenatal social stress: Neuroendocrine activity and behaviour

This study provides evidence in the pig that stress experienced during gestation has long-lasting effects on offspring daughters, including their maternal behaviour. Thirty-six primiparous sows were divided into control and two groups that were stressed (by social mixing) during either the second (Mix 2) or third (Mix 3) trimester of pregnancy. We found detrimental effects of mixing on the mothers growth, body lesions, and cortisol secretion, but did not observe any significant effects on reproductive parameters including birth weight. At 60 days of age, 48 daughters were randomly selected from the three treatments: half were challenged using a restraint and isolation test. Then, all were culled and brain tissue was collected. In situ hybridisation measurements showed increased expression of CRH mRNA in the PVN in unrestrained Mix 2 and in the amygdala of Mix 2 and Mix 3 daughters. At 67 days, 24 further daughters were mixed to measure their responses to this social stress. All showed increased salivary cortisol secretion, but Mix 2 and Mix 3 daughters showed a greater and longer response than controls. Finally, all 24 were inseminated and at parturition maternal behaviour was measured. Mix 2 and Mix 3 daughters were more restless and more responsive to piglets that approached the head of the sow, traits which previously have been shown to be a component of abnormal maternal behaviour. Indeed, Mix 2 and Mix 3 daughters also tended to bite at their piglets more than control daughters.

Opioid-mediated changes in nociceptive threshold during pregnancy and parturition in the sow

Abstract This study aimed to investigate if pregnancy‐induced hypoalgesia occurs in the sow, and to examine the role of endogenous opioids which are known to be released in response to nociception. Sixteen Large WhitexLandrace multiparous sows were tested in straw bedded pens (2.5×2.5 m) during weeks 4, 8 and 12 of pregnancy and over the farrowing period. Testing involved thermal stimulation of eight areas on the rear‐quarters of the sows with a CO2 infra‐red laser until a physical response was seen (tail flick, leg move or muscle twitch) or for a maximum of 16 s. Over the farrowing period testing was more frequent, and at 3.75 h after the birth of the first piglet, half the sows received an injection (i.m.) of an opioid antagonist naloxone (N) (1 mg kg−1 body weight) with the remainder receiving a control dose of saline (S). Responses were recorded 15 and 30 min post‐injection. There was no significant difference between response times over weeks 4, 8 and 12 of pregnancy (P=0.152), however a significant rise was seen from week 12 to 5 days before parturition (P=0.002). Response times continued to rise until the birth of the first piglet by which time the majority of sows had stopped responding within 16 s (P<0.001). Response times fell over days 1, 2 and 7 post‐partum. After administration of naloxone response times fell compared to control animals at 15 min (P<0.001) and 30 min (P<0.01) post‐injection. These results suggest that nociceptive threshold increases during late pregnancy in the sow, perhaps as an endogenous defence against labour pain, and that during parturition this change in nociceptive threshold is, at least in part, opioid‐mediated. Oxytocin is known to be inhibited by endogenous opioids at parturition, thus future research should consider the potential role of increased nociception at birth as a negative feedback to oxytocin release.

Savaging gilts are more restless and more responsive to piglets during the expulsive phase of parturition

Abstract Piglet-directed aggression by farrowing pigs seriously threatens the welfare of the piglets and also results in economic waste within the pig industry. Pre-weaning piglet mortality in the UK is around 11% with savaging accounting for up to 25% of deaths. The aim of this study was to investigate whether gilts that savage piglets behave differently to non-savaging gilts during the expulsive phase of parturition. The behaviour of the gilts during the 8xa0h following the birth of the first piglet was recorded. The gilts were categorised as savagers (n=4) or non-savagers (n=15) depending on their response to piglets during farrowing, and then behaviour during farrowing was compared between the two groups. The maternal behaviour during the first 8xa0h of farrowing differed significantly between the savagers and non-savagers. Savaging gilts were significantly more restless and more responsive towards their piglets during farrowing as characterised by increased posture changing, higher frequency of walking and time spent investigating piglets that approached their head. Overall the results of this study suggest that aberrant maternal behaviour during farrowing involves increased restlessness and responsiveness to piglets, particularly in the early stages of the expulsive phase, which are opposite to the inactivity and passivity suggested to reflect good maternal care in the pig. It is suggested that this restlessness and responsiveness of savaging gilts may not necessarily reflect poor maternal ability, but may be related to the individual’s inability to cope with restrictive environments around farrowing.

The effect of confinement during lactation on the hypothalamic–pituitary–adrenal axis and behaviour of primiparous sows

Commercial sows are typically confined in crates before and during parturition and remain there throughout lactation. In various animal species including non-lactating pigs, confinement over similar periods leads to adaptive changes in the HPA axis, consistent with chronic stress. To investigate evidence for chronic stress in lactating sows, primiparous sows (gilts) were kept in behaviourally confining crates with straw bedding (CS, n = 8) or without bedding (C, n = 8) or in larger strawed pens (PS, n = 16) between 5 days before parturition until 29 days postpartum (piglets were weaned on day 28). Behavioural and physiological recordings (Plasma ACTH and cortisol) were taken at intervals (baseline), and CRH injections were given on five occasions (days 2, 8, 15, 22 and 29 postpartum). The PS gilts spent more time in substrate-directed behaviour and lying ventrally, and less time lying laterally and sitting than the two crated treatments (C and CS) throughout lactation. Baseline plasma ACTH and cortisol levels showed no treatment differences, although we confirmed that a diurnal pattern exists, with morning (1000 h) cortisol being higher than later in the day. CRH challenge tests suggested changes in the HPA axis, consistent with chronic stress, by the end of the lactation period. Cortisol response to CRH tended to be higher in CS than PS across all days, and by day 29 cortisol response to CRH was significantly higher in CS compared to PS and tended to be higher in C than PS. Cortisol/ACTH ratio following CRH challenge also tended to be higher in the crate treatments (C and CS) by day 29. These data suggest that prolonged confinement in farrowing crates may have a negative impact on sow welfare.

Genetic parameters of piglet survival and birth weight from a two-generation crossbreeding experiment under outdoor conditions designed to disentangle direct and maternal effects.

Multivariate Bayesian linear-threshold models were used to estimate genetic parameters of peri- and postnatal piglet survival and individual birth weight of piglets reared under outdoor conditions. Data of 21,835 individual piglet observations were available from a 2-generation crossbreeding experiment selected for direct and maternal genetic effects of postnatal piglet survival on piglet and dam levels, respectively. In the first generation, approximately one-half of the Landrace sires used were selected for large or average breeding values of maternal genetic effects on postnatal piglet survival, whereas in the second generation the Large White sires used were selected for direct genetic effects of the same trait. Estimates of direct and maternal heritability were 0.21 and 0.15, 0.24 and 0.14, and 0.36 and 0.28 for piglet survival at birth and during the nursing period, and individual birth weight, respectively. In particular, direct heritabilities are substantially larger than those from the literature estimated for indoor-reared piglets, suggesting that genetic effects of these traits are substantially greater under outdoor conditions. Direct or maternal genetic correlations between survival traits or with birth weight were small (ranging from 0.06 to 0.17), indicating that peri- and postnatal survival are genetically under rather different control, and survival was only slightly positively influenced by birth weight. There were significant (P < 0.05) negative genetic correlations between direct and maternal genetic effects within each of the analyzed traits ranging from -0.36 to -0.45, which have to be considered when selecting for piglet survival. Adjustment of traits for litter size or inclusion of genetic groups showed insignificant effects on the magnitude of the estimated genetic parameters. The magnitude of genetic parameters suggested that there is substantial potential for genetic improvement of survival traits and birth weight in direct and maternal genetic effects, especially when piglets are kept under outdoor conditions.

The Weaker Sex? The Propensity for Male-Biased Piglet Mortality

For the most part solutions to farm animal welfare issues, such as piglet mortality, are likely to lie within the scientific disciplines of environmental design and genetic selection, however understanding the ecological basis of some of the complex dynamics observed between parent and offspring could make a valuable contribution. One interesting, and often discussed, aspect of mortality is the propensity for it to be sex-biased. This study investigated whether known physiological and behavioural indicators of piglet survival differed between the sexes and whether life history strategies (often reported in wild or feral populations) relating to parental investment were being displayed in a domestic population of pigs. Sex ratio (proportion of males (males/males+females)) at birth was 0.54 and sex allocation (maternal investment measured as piglet birth weight/litter weight) was statistically significantly male-biased at 0.55 (t35u200a=u200a2.51 Pu200a=u200a0.017), suggesting that sows invested more in sons than daughters during gestation. Despite this investment in birth weight, a known survival indicator, total pre-weaning male mortality was statistically significantly higher than female mortality (12% vs. 7% respectively zu200a=u200a2.06 Pu200a=u200a0.040). Males tended to suffer from crushing by the sow more than females and statistically significantly more males died from disease-related causes. Although males were born on average heavier, with higher body mass index and ponderal index, these differences were not sustained. In addition male piglets showed impaired thermoregulation compared to females. These results suggest male-biased mortality exists despite greater initial maternal investment, and therefore reflects the greater susceptibility of this sex to causal mortality factors. Life history strategies are being displayed by a domestic population of pigs with sows in this study displaying a form of parental optimism by allocating greater resources at birth to males and providing an over-supply of this more vulnerable sex in expectation of sex-biased mortality.

Early weaning results in less active behaviour, accompanied by lower 5-HT1A and higher 5-HT2A receptor mRNA expression in specific brain regions of female pigs

In rodents and humans stressful events in early life e.g. maternal deprivation, can increase sensitivity to stress in later life. Humans may become more susceptible to mood disorders, e.g. depression. In livestock species, such as pigs, early weaning is a form of maternal deprivation. We investigated behavioural consequences in 99 female pigs weaned at three different ages (12, 21 and 42 days; d12, d21, d42). Pigs were habituated to an open field arena over 6 days before being given 5-min open-field tests over three subsequent days (days 77-79). Early-weaned pigs (d12) showed behavioural inhibition (reduced vocalisations and lower activity) compared with later-weaned pigs, although in all groups these measures declined over the three tests, so this treatment difference might reflect more rapid habituation to the test in d12 pigs. Long-term effects on mood-related 5-HT receptor subtypes were measured in the brain at 90 days in a random sample of the d12 (n=8) and d42 pigs (n=8), using (3)H-ligand-binding and autoradiography and in situ hybridisation histochemistry. There were no differences between weaning ages in binding of (3)H-8-OH-DPAT (5-HT(1A) receptor agonist) or of (3)H-ketanserin (5-HT(2A) receptor antagonist) to any brain region studied. In d12 pigs, 5-HT(1A) receptor mRNA expression per unit area was 29%, 63%, 52% and 64% lower than in d42 pigs in the parvocellular PVN, amygdala and hippocampal dentate gyrus and pyramidal cell layer, respectively. The ratio of expression per cell to expressing cells per unit area was also lower, by 31%, in the pars horizontalis of the PVN in d12 pigs. Conversely, 5-HT(2A) receptor mRNA was expressed at a 25% and 28% higher density per unit area in the amygdala and pyramidal cell layer of the hippocampus, respectively, in these d12 pigs. In individual pigs, across brain regions, 5-HT(1A) receptor mRNA data were 70-79% correlated with binding data but no correlation was found for 5-HT(2A) data, suggesting different regulatory mechanisms. The behavioural and neurobiological responses to early weaning might represent either dysfunction or adaptation. Further investigation is required.