Breed, litter and parity effects on placental weight and placentome number, and consequences for the neonatal behaviour of the lamb.

Lamb survival is impaired in low birth weight lambs, and those that are slow to stand and suck. Many of the factors that influence lamb vigour, such as parity, litter size, and breed, may exert their effects, at least partially, before birth by influencing placenta development. Our hypothesis was that retarded lamb behavioural development was due to differences in placentation in these animals. Data were collected from Blackface and Suffolk lambs in the first 2 h after birth and placentas were collected when delivered. Suffolk lambs, which were behaviourally slower and had lower rectal temperatures than Blackface lambs, were associated with larger but less efficient placentas (placental efficiency defined as foetal weight supported per g placenta) with fewer foetal cotyledons than Blackface placentas. Triplet lambs were significantly slower than twin or single lambs to suck and had lower rectal temperatures. Although placenta efficiency increased with litter size, placenta and cotyledon weight, and cotyledon number increased with twinning but not thereafter. It seemed likely that triplet lambs suffered some placental insufficiency in comparison to other litter sizes. Lambs born to first parity mothers were slower to stand and reach the udder than lambs of more experienced ewes, and first parity ewes also had smaller and less efficient placentas although cotyledon number was not affected. Male lambs tended to be slower than female lambs for most behaviours, although rectal temperatures were not affected. The sire of the lamb also influenced lamb behaviour and rectal temperature. Both lamb sex and lamb sire influenced the average weight of placental cotyledons, thus some of the sire effect on the behaviour and birth weight of his progeny might be mediated through placental development. Lamb neonatal vigour was correlated with placental efficiency suggesting that lamb behaviour immediately after birth is related to placental development and function.

The effect of parity and environmental restriction on behavioural and physiological responses of pre-parturient pigs.

There is increasing evidence that restriction of pre-parturient behaviour in pigs is stress-inducing, characterised by an elevation in hypothalamic-pituitary-adrenal (HPA) activity in gilts. To determine whether pigs adapt to behavioural restriction, through modification of nest-building behaviour, we studied pre-parturient pigs in either farrowing crates (no bedding, n=7) or straw-bedded pens (n=7) in their first (gilts) and second (sows) parity, with physiological measurements being taken in the second parity. Observations and blood sampling were carried out during the pre-parturient phase. Crated pigs changed posture more often than penned pigs (F(1,12)=7.06, P<0.05), with the number of posture changes reducing across parities in both environments. The reduction in posture changing was more apparent in the crated sows which may indicate that attempted nest-building behaviour of sows with prior experience of farrowing crates is less fragmented. The crated pigs spent a greater proportion of time sitting across both parities (F(1,12)=9.4, P<0.01), and spent less time manipulating available substrates (F(1,12)=10.67, P<0.05). There was a tendency for penned pigs to spend a greater proportion of time standing (F(1,12)=3.77, P=0.076) with peak nesting behaviour occurring earlier in relation to parturition than in crated pigs. In addition penned sows performed more floor-directed behaviour than penned gilts, and at an earlier stage in relation to parturition. However, crated sows also performed peak nest-building earlier than crated gilts. Plasma cortisol profiles indicated elevated HPA activity in crated sows during the pre-parturient period (F(42,303)=1.43, P<0.05) suggesting increased physiological stress, however, the difference between crated and penned sows was less than that previously seen in gilts. The increased range of pre-parturient behaviours seen in the penned sows suggests that experience may result in an 'improvement' in their nest-building behaviour: earlier preparation of the nest site and then subsequent manipulation of substrates. The crated sows appeared to show some behavioural adaptation to the crate environment; earlier peak in floor directed behaviour and total substrate directed behaviour, reduced posture changing. In conclusion the nest-building behaviour of pigs is modified over parities with adaptation to the behavioural restrictions imposed by the farrowing crate. However, this adaptation, through prior experience, does not completely reduce the elevation in HPA activity previously reported in pre-parturient crated gilts.

The effect of opioid antagonism and environmental restriction on plasma oxytocin and vasopressin concentrations in parturient gilts.

Oxytocin plays an important role at parturition due to its involvement in uterine contractions, foetal expulsion and the onset of maternal behaviour. The role of the related neurohypophysial hormone, vasopressin, is less clear; however, there is some evidence that it is also involved in maternal behaviour and its role in osmotic regulation is well established. The aim of this study was to investigate the inhibitory effects of endogenous opioids on these hormones during the expulsive phase of parturition in the pig, and to examine how opioid restraint interacts with environmental restriction. The subjects of this study were 31 Large Whitex Landrace primiparous sows (gilts). An indwelling jugular catheter was implanted under general anaesthesia at 12 days before the expected parturition day (EPD). From 5 days before the EPD 15 of the gilts were individually housed in a restrictive parturition crate without straw and 16 were individually housed in a straw-bedded pen. Blood samples were taken with increasing frequency towards and during parturition through a catheter extension to reduce disturbance. At 7.5 min after the birth of the first piglet half of the gilts in each environment received a dose of the opioid receptor antagonist naloxone (1 mg/kg, i.v.) with the remaining gilts receiving saline as a control. Overall, there was no effect of environment on either circulating oxytocin or vasopressin. However, both oxytocin and vasopressin were inhibited by endogenous opioids during the expulsive phase. The inhibitory effects of opioids on these hormones did not appear to have any adverse effects on the progress of parturition as judged by cumulative piglet birth intervals. The regulation of the opioid inhibition of oxytocin and vasopressin during parturition is discussed in relation to other neurotransmitters and whether opioid inhibition of these neurohypophysial hormones is part of the 'normal' physiological response to parturition or whether it is stress-induced.

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.

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

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 White x Landrace multiparous sows were tested in straw bedded pens (2.5 x 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.