La Deans

The effect of environment on behaviour, plasma cortisol and prolactin in parturient sows

Nest-building in the pig is thought to be stimulated by a pre-parturient surge in prolactin. There is concern that sows in crates may experience psychological stress as a result of physical interference with nest-building. Thirty-three gilts were implanted with jugular catheters approximately 10–14 days before expected date of parturition (EPD). On day 5 before EPD, gilts were moved into either conventional farrowing crates without bedding (treatment C; n=16) or pens allowing freedom of movement with bedding (treatment P; n=17). The animals were blood sampled on the day before and the day after introduction to the treatments. Blood and behaviour sampling was resumed 48 h before EPD and continued until 4 h post-commencement of farrowing. As with previous studies gilts in both environments were more active (P<0.001) and performed more substrate-directed behaviour in the pre-parturient period (P<0.001). Gilts in pens spent considerable amounts of time in straw-directed behaviour, and gilts in crates increased amounts of floor- and fixture-directed behaviour (both P<0.001). Prolactin was not affected by treatment and increased in both environments over the same time period as this substrate-directed behaviour. However, the causal role of prolactin in nest-building must now be questioned given that a number of individuals showed no increase in prolactin over the 48 h before EPD, but still showed an increase in substrate-directed activity. Cortisol was strongly affected by treatment with treatment C having elevated total cortisol over much of the pre-parturient period (P<0.001). Cortisol also increased in penned animals during parturition. There were no consistent correlations among behaviour, prolactin and cortisol. The general increase in cortisol in the early post-partum period may suggest that parturition in itself has stress-inducing aspects. The additional rise in cortisol found in the crated sows may indicate that the close confinement of the crate, by interfering with the expression of maternal behaviour, induces psychological stress. There was no evidence that the elevated cortisol response in crated gilts extended beyond the end of parturition.

The responsiveness of sows to their piglets in relation to the length of parturition and the involvement of endogenous opioids

Abstract The aim of this study was to describe maternal behaviour in the pig and to investigate the effect of endogenous opioids on maternal responsiveness. The behaviour of 16 Large White×Landrace female pigs was recorded around farrowing which involved recording the pigs posture and her response when piglets were present at her nose. To determine the role of endogenous opioids, sows were injected (i.m.) with either naloxone, an opioid antagonist, (1 mg kg −1 bodyweight ( n =8)) or saline ( n =8) at 3.75 h after the birth of the first piglet. Generally the initial period following the birth of the first piglet seemed to be the most active after which the sows spent almost all of the time in lateral recumbency. The results also show that farrowing sows are generally unresponsive to their piglets during farrowing. Sows receiving naloxone became more responsive towards their piglets. The changes seen in posture and responsiveness to piglets were delayed in sows with a longer parturition suggesting some involvement of cumulative piglet births on passivity. It is proposed that opioid-mediated passivity in the pig, characterised by lateral lying and unresponsiveness to piglets, may be advantageous by maximising suckling opportunities and reducing the risk of crushing piglets and of attracting predators to the nest.

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.

The effect of environment on plasma cortisol and beta-endorphin in the parturient pig and the involvement of endogenous opioids.

Previous work has indicated that plasma cortisol increases during farrowing in the pig suggesting increasing physiological stress. The aim of this study was to determine changes in plasma cortisol and beta-endorphin over farrowing in the pig to obtain a more detailed profile of pituitary and adrenal release at this time and also to investigate the involvement of endogenous opioids in the mediation of the HPA axis. Indwelling jugular catheters were implanted, under general anaesthesia, in 31 Large White x Landrace gilts approximately 15 days before the expected parturition day (EPD). Gilts were moved into either a farrowing crate, without straw (n = 15), or a straw-bedded pen (n = 16) 5 days before the EPD. Samples were taken during the pre-farrowing period and then during farrowing itself. At 7.5 min after the birth of the first piglet (BFP), gilts either received naloxone, an opioid antagonist, (1 mg kg(-1) body weight, i.v.) or a control dose of saline. Plasma beta-endorphin increased following the BFP but remained fairly constant over the third and fourth hour of farrowing. Plasma cortisol continued to increase over the 4 h following the BFP. Changes seen in these hormones were generally insensitive to the environment and there was little evidence of opioid mediation of the HPA axis at parturition. From these results it is suggested that certain aspect(s) of parturition itself stimulate the HPA axis. However it is unknown if the rise in plasma cortisol is a result of some stress-inducing factor of the parturition process or whether it reflects a metabolic function. The study also demonstrates the lack of any inhibitory mediation of the HPA axis by endogenous opioids at parturition.

The effects of chronic environmental stress on parturition and on oxytocin and vasopressin secretion in the pig

Previous work has suggested that an acute behavioural confinement in mid-partum can inhibit oxytocin secretion and prolong delivery in the pig, an effect that is opioid mediated. The present experiment investigated the effect of longer-term (chronic) behavioural confinement, that has previously been shown to elevate total plasma cortisol, on speed of delivery and on plasma oxytocin and lysine vasopressin concentrations during the peri-parturient period in primiparous pigs (gilts). Five days before their expected parturition (farrowing) date, gilts with preplaced jugular catheters were either confined to farrowing crates that severely restricted maternal behaviour, or housed in pens that permitted free movement and maternal behaviour (e.g. nest building). Blood samples were taken continuously from 24 h before the birth of the first piglet (BFP) to 6 h post-BFP, and for oxytocin on Days 1, 2, and 7 following parturition (Days P1, P2, P7). Both oxytocin and vasopressin were strongly influenced by parturition (P<0.001). There was no overall effect of chronic crating on either hormone, but crated and penned gilts did show significant differences with respect to the pattern of both oxytocin and vasopressin concentrations over time (P<0.05 in both cases). Oxytocin and vasopressin first increased in crated and penned gilts from 3 h pre-BFP (P<0.05). Crated gilts subsequently showed greater increases in both oxytocin and vasopressin over the first hour of delivery than penned gilts (mean oxytocin (pmol 1−1): 53.3±8.5 vs. 39.7±5.0 for crated vs. penned gilts; mean vasopressin (pmol l−1):4.4±0.7 vs. 2.0±04 for crated vs. penned gilts; both P<0.05). For oxytocin, crated gilts then showed subsequent declining concentrations relative to penned gilts (P<0.05). For vasopressin, penned gilts reached similar concentrations as crated gilts in the third hour post-BFP before vasopressin concentrations in both groups declined. Crated gilts also gave birth to piglets faster in the early stages of delivery (e.g. mean interval between Piglets 2 and 3 (min): 9.6±2.5 vs. 25.6±8.54 for crated and penned gilts, respectively: P<0.02). We conclude that confinement of gilts to a farrowing crate for 5 days neither adversely affects the progress of delivery in the primiparous pig nor the secretion of posterior pituitary hormones involved in parturition.

Investigation of the relationship between farrowing environment, sex steroid concentrations and maternal aggression in gilts

Maternal oestrogen and progesterone have been shown to be important in the initiation of maternal behaviour. Thirty-three Large White x Landrace gilts, housed in groups during pregnancy, were observed and aggressive interactions recorded. Individuals had jugular catheters implanted 14.5 (s.e. 0.34) days before their expected parturition date (EPD). Five days before EPD gilts were randomly allocated and moved to either a conventional farrowing crate (C; without straw, 16 gilts) or a pen (P; 2.1 x 3.1 m2; with straw bedding, 17 gilts). Blood samples were taken at frequencies determined by the proximity to farrowing onset. Piglets were removed at birth and returned 2 h after placental expulsion. The reaction of each gilt to her piglets was monitored. Gilts savaging piglets were sedated with azaperone (n = 8). There was no overall effect of farrowing environment on oestradiol and progesterone concentrations. The pre-farrowing ratio of progesterone to oestradiol was higher for (P) gilts (0.45 vs. 0.25, (P) vs. (C); S.E.D. 0.085, P < 0.05) as was their overall maximum oestradiol level (3.39 vs. 2.29 ng/ml, (P) vs. (C); S.E.D. 0.39, P < 0.01). In contrast to progesterone, oestradiol patterns varied considerably between individuals. Dominance rank value during pregnancy, but not levels of aggression, correlated positively to pre-farrowing oestradiol concentrations. Treatment with azaperone was not related to farrowing environment, piglet weight or litter size. Azaperone treated gilts showed a higher pre-farrowing oestradiol to progesterone ratio (0.55 vs. 0.29, +/- azaperone; S.E.D. 0.10, P < 0.05), significantly higher levels of oestradiol post-partum (0.7 vs. 0.19 ng/ml, +/- azaperone; S.E.D. 0.20, P < 0.001) and significantly lower levels of aggression during pregnancy (1.68 vs. 2.23 aggressive interactions/h, +/- azaperone; S.E.D. 0.15, P < 0.001). The results indicate that there are no major effects of farrowing environment on sex steroid concentrations. Maternal aggression under these conditions appears to be negatively related to aggression during pregnancy, but this is not reflected in plasma concentrations of sex steroids around parturition.