The resting behaviour of 64 groups of sheep was observed in the lairages of two slaughterhouses after transport for about four hours in a commercial livestock transporter. The sheep were scanned at five minute intervals for the initial three hours in the lairage pen. The median percentage of time spent lying per group was 17 per cent (range 1 to 63). The percentage of time spent lying was not significantly related to the duration of transport, time of arrival or slaughterhouse. There was a significant positive relationship between space allowance (range 0.22 to 0.98 m2 per sheep) and percentage of time spent lying per group (P < 0.001). Increased space allowance also tended to be associated with a decrease in the occurrence of all interactions between the sheep (P = 0.05). Groups penned with two unfamiliar groups tended to spend less time lying than those penned on their own or with one other group (P = 0.07). There was no significant difference in the total percentage of time spent lying by groups penned on slats and those penned on straw. The results indicated that the ability of sheep to rest in the lairage may be compromised by providing low space allowances.

In a 2x2 factorial design, (n=6) sheep were either transported by road for 15h or kept in their home pens, and then either starved for 12h with access to water or offered hay ad libitum, with access to water. All groups were offered hay and water 12h after transport. Behavioural observations and measurements of dehydration and feed restriction were made before, during, and for 24h post-transport, to evaluate the implications of these procedures for the welfare of sheep.

After the journey, the immediate priority for the sheep was to eat. Consumption of hay increased water intake and reduced the time spent lying down. The plasma Cortisol concentration was greater in sheep which had been starved during the 12h post-transport period, than in those offered hay during this time; and the plasma free fatty acid concentration was greater in sheep which had been transported than in those which had not. Although transported sheep kept without hay during the first 12h post-transport drank more water than those which had not been transported, the mean time before they drank was greater than 7h. During the transport period, there was less lying behaviour in transported sheep than in non-transported sheep but transported sheep did not lie down more posttransport than non-transported ones. This work suggests that sheep should be offered both feed and water after a 15h journey. However, when feed was not available after a 15h journey, drinking and resting did not appear to be immediate priorities.

The effects of food and/or water deprivation at different ambient temperatures (7 or 35 °C) on stress hormone release in sheep (n = 8), were studied to provide background data for research into the effects of road transport. Blood samples were taken from catheterized animals at the start and, at 6h intervals, during 48h tests in an environmental chamber. Cortisol release was unaffected by temperature or deprivation state but was stimulated by introduction to the chamber. Prolactin secretion showed a similar tendency and levels of this hormone were generally higher in the first test, whichever chamber was used. Heat exposure also had a prolonged stimulatory effect on prolactin release, especially in the first test. Growth hormone concentrations were rather variable but tended to be greatest when the animals were deprived of food. Measurements of plasma osmolality indicated that sheep remained in water balance, even when water was withheld for 48h, unless they had access to food. The results suggest that under laboratory conditions, and over a wide thermal range, withholding food and water for 48h does not induce Cortisol or prolactin release in sheep. However, exposure to novel situations seems to have a stimulatory effect.

A review of the literature was undertaken to consider the possible effects of human intervention (shepherding) at around the time of parturition in extensively farmed sheep. There is little clear empirical evidence to suggest that shepherding ensures either easy births or the integrity of ewe–lamb contact — factors closely linked to the welfare of the animals at this time. There is similarly no clear support for shepherding being harmful. However, the following suggestions are made: first, human presence can inhibit or delay parturition; second, extended parturition can increase the risk of, or is associated with, dystocia; and third, disturbance at birth can compromise ewe-lamb bonding and consequently lamb survival. Furthermore, sheep populations that have undergone rigorous selection for ease of lambing and minimal shepherding in extensive environments have well-documented physical and behavioural traits underlying their predisposition for enhanced lamb survival. Although our cultural legacy may impose a duty to intensively monitor animals at lambing, it is concluded that, at least in some situations, shepherding may not be entirely beneficial. The commonly held view of the necessity for some human intervention in extensive livestock systems is perhaps overly paternalistic, and requires a more comprehensive appraisal.

Chronic stress occurs when animals are unable to deal with a persistent stressor with species-typical responses, or when several stressors are present concurrently. Chronic stress is most frequently considered in intensive systems, but it may also be a welfare concern for extensively managed species, such as the sheep. Here we review behavioural and physiological responses of sheep to experimentally induced chronic stressors to determine relevant indicators of chronic stress. Neuroendocrine responses to chronic stress are difficult to interpret because initial responses are followed by an apparent normalisation. Thus, cortisol or catecholamines may be at or below pre-stress levels during chronic stress, but this varies with different stressors. Chronic stress can also affect reproductive function, impair body and wool growth and meat quality, reduce immune function, and is associated with greater parasite burdens in sheep. Chronic stress induces alterations in behaviour patterns, particularly activity and feeding, and circadian rhythms of behaviour. Stereotypic behaviours, however, are infrequent in sheep and may occur only in experimental conditions of social isolation. Behavioural and physiological data suggest that rough handling and sheepdogs may be sources of chronic stress for sheep. Social subordination and weaning also act as chronic stressors, leading to higher parasitism in these animals and a greater response to additional stressors. Lameness and parasitism are associated with physiological and behavioural responses indicating that these are severe forms of chronic stress in sheep. It is unclear whether environmental stressors, such as weather and food availability, induce chronic stress in sheep. Under-nutrition may, however, be a welfare concern through its impact on lamb survival. The existence of many sources of chronic stress in the management of sheep suggests that the welfare of this species requires more attention than it has currently received.

When sheep are transported they are potentially exposed to a number of factors, including water and feed deprivation, low space allowance and elevated environmental temperature, that are not related to vehicle motion but could result in animal welfare problems, either on their own or in combination. In a 2x2 factorial experiment, groups of sheep (n = 6) were moved from individual pens where they had access to hay and water to environmental chambers kept at either 14°C or 21°C. Within each chamber, half the sheep had access to water but they were all kept at a space allowance of 041m2 sheep−1 without feed. After 24h they were returned to their individual pens and offered hay and water. Behaviour and a range of biochemical measurements of dehydration and feed restriction were recorded before, during and after the treatment period. During the treatment period there was no evidence of dehydration, and sheep with access to water drank less than they did before the treatment. The plasma concentration of free fatty acids increased during fasting and, post-treatment, the intake of hay was greater than before treatment. The rapid post-treatment intake of dry feed was associated with some evidence of dehydration, as indicated by increased plasma osmolality and plasma vasopressin concentration. This suggests that following provision and consumption of large quantities of feed after a period without access to feed and water during transportation, sheep must be allowed sufficient time to drink before a subsequent journey is undertaken.

The return to consciousness of 24 electrically stunned lambs was assessed by measurement of physiological reflexes and electroencephalography (EEG) recordings. The physical activity of the lambs after head-only electrical stunning included one tonic phase and two clonic phases. The tonic phase began immediately after the stun and ended 10 s after the stun; the first clonic phase started immediately after the tonic phase and ended 36 s after the stun, and the second clonic phase started immediately after the first clonic phase and ended 70 s after the stun. During the tonic phase and the first clonic phase, electrical activity recordings of the brain showed a dramatic increase in the relative power spectra of alpha and beta frequencies. Both frequencies returned to pre-stun levels with the end of the first clonic phase. During the second clonic phase, the relative power spectrum of theta frequency was higher than that before stunning. These results suggest that during the tonic phase and the first clonic phase, the animal was unconscious, whereas during the second clonic phase the return of some conscious function began. Spontaneous breathing returned at about 29 s post-stun, whereas the corneal reflex returned at about 38 s. It is therefore suggested that the return of spontaneous breathing is the safest indicator that the animal is close to recovering consciousness.

The traditional concerns about farm animal welfare have centred around the impact of intensive environments and management practices on the animal. This emphasis on the physical environment is changing, however, with greater consideration being given to animal factors and in particular to the selective breeding of farm animals. In this paper we use examples from our own research on dairy cattle and sheep breeding that have made positive and practical contributions towards reducing welfare problems by creating more balanced breeding programmes. In both examples, inclusion of health and fitness traits into breeding indexes can be shown to be more profitable than selecting on production traits alone. In addition, we propose that in principle animal breeding combined with economics research can make a more general contribution towards resolving animal welfare issues, by providing a framework for the quantitative evaluation of the costs and benefits of an animal production system. The advantage of the approaches currently used in multi-trait selection is that they transform all traits (production-based or welfare-based) to a common currency allowing direct comparisons of costs and benefits. Currently the weights applied to traits reflect their economic value to the producer. This approach is likely to underestimate non-economic welfare aspects such as the pain or discomfort associated with lameness, and new approaches are needed to more fully account for these non-economic welfare costs. We therefore propose that consideration is given to the use of approaches such as contingent valuation, which has been widely used in economics to derive values for non-economic activities. The question of who would pay for the addition of these welfare costs to a breeding index remains open, but it would seem most reasonable to treat these as a public good and pay for them as such through appropriate mechanisms.

A questionnaire survey of farm experience, undertaken during lambing time in the year 2000 by 95 second-year veterinary undergraduate students, highlighted numerous areas of concern. On those farms attended, more than one third of shepherds (32; 34%) neither washed their hands in an approved scrub nor used arm-length disposable plastic gloves before attempted correction of a lambing problem. Sheep received a prophylactic antibiotic injection after an assisted lambing on just 33 farms (35%), while the majority of farmers (62 farms; 65%) treated only those ewes that became sick some days after assisted lambing. Veterinary assistance was requested to only 22 of 359 (6.1%) dystocia cases from a sample population of 79 610 lowground ewes. When lambs could not be delivered by farm staff the ewes were either humanely destroyed (65) or injected with antibiotics but subsequently died because of ensuing toxaemia from the emphysematous lambs in utero (272).

The effect of transport on core and peripheral body temperatures and heart rate was assessed in ten 18-month-old Coopworth ewes (Ovis aries) Manual recordings of core (rectal) temperatures were obtained, and automated logging of peripheral (external auditory canal and pinna) temperatures and heart rate was carried out on the day prior to (day 1) and during (day 2) a standardised transport procedure. Transport produced a significant increase in the rectal temperature, which declined following unloading. Peripheral measures of body temperature also exhibited changes with transport. However, both ear-canal and pinna temperatures declined during actual transport, reflecting to some extent the decline in ambient temperatures recorded externally by sensors on the ear tags of the animals. Peripheral measurement of temperature, particularly at the readily accessible ear canal, may offer potential as a technique for the long-term monitoring of thermal responses to stress. However, further research is required into the potentially confounding effects of ambient temperature and wind chill factors.

In the test described here, sheep are exposed to a situation of conflict between the motivation to approach other sheep and the motivation to avoid a human handler. The distance that the test sheep keep from the handler is a reflection of the relative aversiveness of this handler to the sheep. The test itself requires only a minimum amount of handling and gives the test animals the opportunity to choose their distance from the aversive stimulus, thereby reducing stress during the test itself. The two aversive stimuli chosen for comparison were a human handler facing toward the test arena (more aversive) or the same handler turning his back to the arena (less aversive). Ten Scottish Blackface sheep were tested individually a total of ten times, five times with each of the two stimuli in alternate tests. During the first two tests, nine of the sheep stayed further away when the human was facing toward the arena, compared to when he was facing away; this shows that the test is able to discriminate differences in aversiveness between two stimuli as perceived by individual sheep. This difference was not apparent in the following eight tests, probably because of the fact that the stimuli were not reinforced during the tests. Because the test is concerned with sheep's reaction to a stimulus (eg handler), the procedure associated with the stimulus itself (eg shearing, castration) does not have to be repeated in the test, which means that this method is ideal for studying procedures which cause distress to the animals or which are difficult to repeat.

Deep body temperature was measured in four wethers and four ewes surgically implanted with biotelemetry devices. Records were taken over several days in the home pen (baseline data) and also in response to three potentially stressful procedures: transport, exposure to a sheepdog, and forced exercise. Loading the animals into a vehicle and transporting them for 2.5h produced a rise in core temperature that, in males, persisted for several hours. Moving the sheep into an outside pen and subsequent exposure to the dog appeared to produce transient increases in body temperature, although these changes were not statistically significant. By contrast, exercise for 30min resulted in a rapid and pronounced (approximately 2°C) temperature rise that was followed by an equally abrupt return to baseline. Sustained increases in deep body temperature or changes in circadian temperature rhythms in healthy sheep may be a response to psychological distress and, therefore, indicative of poor welfare.

The question of whether witnessing slaughter of conspecifics is distressing was investigated in sheep. Previously catheterized sheep were allowed to see the stunning and sticking (exsanguination) of other sheep. Heart rate was monitored and serial blood samples were taken to assess stress responses. Although the measurable parameter levels were generally high due to human contact and handling, there were no specific increases in response to witnessing stunning and slaughter. This work failed to produce any evidence to suggest that sheep are distressed by witnessing the slaughter act.

This study was designed to investigate the physiological responses induced in sheep (n = 18) by two different loading techniques followed by a short road journey. All animals were prepared with venous catheters, to minimize the disturbing effects of blood sampling, and nine sheep were fitted with heart rate monitors. The animals were loaded onto a transport vehicle in groups of three, alternately using a conventional tailgate ramp or a crate raised with a hydraulic lift. When all of the sheep were loaded, they were taken on a journey lasting 195min. Blood samples were collected in the home pen, directly after loading, and at 15min intervals during the journey. Measurements were made of plasma concentrations of Cortisol, prolactin and catecholamines (adrenaline and noradrenaline). The results indicated that heart rate increased during loading, regardless of the method used. No changes in concentrations of Cortisol or the catecholamines were detected, although a small increase in prolactin was noted when animals were loaded using the ramp. During transport, all sheep exhibited increases in plasma Cortisol concentrations which were greatest during the first 2h of the journey. The results suggest that, under the conditions employed in this experiment, the effects of the two loading procedures were similar and that transport appeared to be more stressful than loading.

Appropriate space allowances for animals are yet to be specifically determined for lairage. Space allowances that may be suitable for animals in lairage are suggested, based on reviewed studies of animals in transport, lairage and on farm. The longer animals are in lairage the more space they require, in order to be able to get up and lie down and lie undisturbed by congeners. Little work has been done on air quality and air flow characteristics in lairages. The range of ventilation must be sufficient to control levels of toxic or irritant gases such as carbon dioxide and ammonia and to remove excess heat and humidity; the latter being particularly relevant for pig lairages in hot weather. Intensities of sound measured in lairages often exceed 85 dB and there is evidence to suggest that such levels can be stressful especially for pigs; and human shouting appears particularly aversive to animals. Cattle vocalise in response to painful stimuli and to convey information to conspecifics that may be related to fear and distress. There is limited evidence that sheep adapt to continuous sound, provided it is not too loud, but respond to intermittent sounds such as gates banging and human shouting. Vocal communication between sheep may be less important than that between cattle and pigs. Levels of vocalisation are potential indices of animal welfare. Animals' prior experiences and factors such as sex, group size and constitution, pen design, and climatic or environmental conditions affect their welfare and responses to conditions in lairage.

Determining objective measures for proof of consciousness in non-human animals has been helped by improved understanding of neural correlates of human consciousness. Functional imaging and neuropsychological studies have shown remarkable overlap between structures involved in actual perception of social and non-social objects and those involved in forming mental images of them. One area of particular interest is individual face recognition. This involves regions of the temporal lobe that are mainly only activated by actual perception or mental imagery of faces. Using behavioural, neuroanatomical and neurophysiological approaches in sheep, we have found that they have similar specialized abilities for recognizing many individuals from their faces. They have developed the same specialized neural processing regions in the temporal lobe for aiding such recognition. Furthermore, parallel activation of other brain regions controlling behavioural and emotional responses only occurs when they are overtly interested in the individuals whose faces they perceive. Such interest might therefore equate to their becoming consciously aware of them. Preliminary experiments have indicated that sheep may form and use mental images and that the regions of the temporal lobe that respond to faces can also do so under conditions where faces are suggested but do not actually appear. Such similarities between humans and sheep in this form of social recognition make it difficult to claim that humans can form mental images of faces whereas sheep cannot. While the ability to form and use mental imagery is not in itself definitive proof of consciousness, it is an important component part.

Survival and mortality of extensive hill ewes are important commercial factors and valuable indicators of welfare, but difficult to study. Body condition scoring is a tried-and-tested management and monitoring tool which has been defined as a good predictor of fitness in ewes and is easily measured under hill farm conditions. This paper presents the results of a study on ewe survival rates in hill conditions in Scotland. Ewe performance and survival rates were measured in two contrasting hill flocks over a five-year period. Statistical analysis showed that mid-pregnancy body condition score was the most reliable indicator of subsequent ewe survival, followed closely by age of the ewe and environmental and management conditions. This study confirms that there are considerable welfare issues related to hill flocks and that improved winter nutritional management is a key component to good welfare and productivity. It also reinforces the view that body condition score is a good quantitative predictor of animal welfare and that poor mid-winter score indicates high risk of mortality, both at the flock and individual ewe level.

Activation of the sympathetic nervous system, with associated increases in heart rate and the redistribution of blood in preparation for ‘fight or flight’, is an integral part of the ‘defence reaction’. In sheep, the defence reaction involves vasoconstriction in the ear-pinna. If decreases in ear-pinna temperature (Tp) can be used to indicate vasoconstriction, then it may be possible to use changes in Tp as a measure of the defence reaction. Ewe lambs were exposed to stressors including mustering into pens, moving between pens, isolation from conspecifics, and prolonged periods of exercise. Measurements of heart rate (HR), Tp, vaginal temperature (Tv), and salivary cortisol and urinary catecholamine concentrations were used to assess stress responses. A repeatable pattern of changes in HR, Tp and Tv was observed in response to stressors. Short-term disturbances resulted in increased HR, reduced Tp, and increased Tv. More sustained disturbances — for example, prolonged periods of exercise — resulted in a sustained elevation in HR, a sustained decrease in Tp, and a sustained elevation in Tv. The highest levels of cortisol and catecholamines were associated with the treatments that resulted in the longest periods of decreased Tp. We infer that changes in Tp occur largely in response to changes in sympathetic nervous activity, and that the potential exists to measure elements of stress responses by monitoring Tp in freely behaving animals. This is a minimally invasive measure that allows the monitoring of modest numbers of animals over prolonged periods with minimal handling.

An adaptive conjoint analysis was used to evaluate stakeholders’ opinion of welfare indicators for ship-transported sheep and cattle, both onboard and in pre-export depots. In consultations with two nominees of each identified stakeholder group (government officials, animal welfare representatives, animal scientists, stockpersons, producers/pre-export depot operators, exporters/ship owners and veterinarians), 18 potential indicators were identified. Three levels were assigned to each using industry statistics and expert opinion, representing those observed on the best and worst 5% of voyages and an intermediate value. A computer-based questionnaire was completed by 135 stakeholders (48% of those invited). All indicators were ranked by respondents in the assigned order, except fodder intake, in which case providing the amount necessary to maintain bodyweight was rated better than over or underfeeding, and time in the pre-export assembly depot, in which case 5 days was rated better than 0 or 10 days. The respective Importance Values (a relative rating given by the respondent) for each indicator were, in order of declining importance: mortality (8.6%), clinical disease incidence (8.2%), respiration rate (6.8%), space allowance (6.2%), ammonia levels (6.1%), weight change (6.0%), wet bulb temperature (6.0%), time in assembly depot (5.4%), percentage of animals in hospital pen (5.4%), fodder intake (5.2%), stress-related metabolites (5.0%), percentage of feeding trough utilised (5.0%), injuries (4.8%), percentage of animals able to access food troughs at any one time (4.8%), percentage of animals lying down (4.7%), cortisol concentration (4.5%), noise (3.9%), and photoperiod (3.4%). The different stakeholder groups were relatively consistent in their ranking of the indicators, with all groups nominating the same top two and at least five of the top seven indicators. Some of the top indicators, in particular mortality, disease incidence and temperature, are already recorded in the Australian industry, but the study identified potential new welfare indicators for exported livestock, such as space allowance and ammonia concentration, which could be used to improve welfare standards if validated by scientific data. The top indicators would also be useful worldwide for countries engaging in long distance sea transport of livestock.

The aim of this paper is to discuss the relationship between qualitative categories of sheep welfare and associated quantitative behavioural observations. Most scientific studies rely on quantitative measures, however to interpret those measures in terms of an animal's experience of welfare, the use of qualitative terminologies denoting various emotional states (eg fear, pain, distress) seems hard to avoid. This is especially so in cases where the same behavioural or physiological indicator could have different meanings for welfare; high levels of locomotion, for example, could indicate fear, aggression, or both. To resolve such dilemmas, scientists often resort to judging an animal's most likely state through direct observation; indeed qualitative characterisations are often intertwined with reported quantitative results. This paper reviews frequently used qualitative categories of sheep welfare and the behavioural context in which these categories are applied. Qualitative judgements of welfare tend to habitually be regarded as ‘subjective’, ie as anthropomorphic and unreliable; however from this review it appears that such judgements reflect the overall demeanour of an animal's behaviour, and as such may provide a useful empirical basis for the investigation of emotional expression in sheep. Further development of a qualitative behavioural approach may provide a more complete picture of the expressive repertoire of sheep, and enhance our understanding of ways in which these animals experience well-being or distress.