Scientific research on ‘animal welfare’ began because of ethical concerns over the quality of life of animals, and the public looks to animal welfare research for guidance regarding these concerns. The conception of animal welfare used by scientists must relate closely to these ethical concerns if the orientation of the research and the interpretation of the findings is to address them successfully.

At least three overlapping ethical concerns are commonly expressed regarding the quality of life of animals: (1) that animals should lead natural lives through the development and use of their natural adaptations and capabilities, (2) that animals should feel well by being free from prolonged and intense fear, pain, and other negative states, and by experiencing normal pleasures, and (3) that animals should function well, in the sense of satisfactory health, growth and normal functioning of physiological and behavioural systems. Various scientists have proposed restricted conceptions of animal welfare that relate to only one or other of these three concerns. Some such conceptions are based on value positions about what is truly important for the quality of life of animals or about the nature of human responsibility for animals in their care. Others are operational claims: (1) that animal welfare research must focus on the functioning of animals because subjective experiences fall outside the realm of scientific enquiry, or (2) that studying the functioning of animals is sufficient because subjective experiences and functioning are closely correlated. We argue that none of these positions provides fully satisfactory guidance for animal welfare research.

We suggest instead that ethical concerns about the quality of life of animals can be better captured by recognizing three classes of problems that may arise when the adaptations possessed by an animal do not fully correspond to the challenges posed by its current environment. (I) If animals possess adaptations that no longer serve a significant function in the new environment, then unpleasant subjective experiences may arise, yet these may not be accompanied by significant disruption to biological functioning. Thus, a bucket-fed calf may experience a strong, frustrated desire to suck, even though it obtains adequate milk. (2) If the environment poses challenges for which the animal has no corresponding adaptation, then functional problems may arise, yet these may not be accompanied by significant effects on subjective feelings. Thus, a pig breathing polluted air may develop lung damage without appearing to notice or mind the problem. (3) Where animals have adaptations corresponding to the kinds of environmental challenges they face, problems may still arise if the adaptations prove inadequate. For example, an animal's thermoregulatory adaptations may be insufficient in a very cold environment such that the animal both feels poorly and functions poorly. We propose that all three types of problems are causes of ethical concern over the quality of life of animals and that they together define the subject matter of animal welfare science.

We estimate that stereotypies are currently displayed by over 85 million farm, laboratory and zoo animals worldwide. This paper investigates their reliability as welfare indicators, by surveying studies relating stereotypy to other welfare measures and by analysing the mechanisms underlying this behaviour. Where data exist, most (approximately 68%) situations that cause/increase stereotypies also decrease welfare. Stereotypy-eliciting situations are thus likely to be poor for welfare, although exceptions exist. Within such an environment, however, most (approximately 60%) accounts link individual stereotypy performance with improved welfare (cf approximately 20% linking it with reduced welfare). Thus, in a sub-optimal environment, non-stereotyping or low-stereotyping individuals could well have the poorest welfare, although again exceptions exist. Examining the mechanisms underlying stereotypy performance, we discuss four processes that could account for these complex links between stereotypy and welfare. Beneficial consequences from performing the specific source-behaviour of the stereotypy (‘do-it-yourself enrichment’), or arising from repetition per se (‘mantra effects’), may ameliorate welfare in poor environments. In addition, stereotypies that have become centrally controlled (habit-like), or that arise from autistic-like changes in the control of all behaviour (perseveration), are likely to be unreliable indicators of current state because they can be elicited by, or persist in, circumstances that improve welfare. To refine the role of stereotypy in welfare assessment, we suggest the collection of specific additional data to reveal when any of these four processes is acting. Until such research increases our understanding, stereotypies should always be taken seriously as a warning sign of potential suffering, but never used as the sole index of welfare; non-stereotyping or low-stereotyping individuals should not be overlooked or assumed to be faring well; simple measures of frequency should not be used to compare stereotypies that differ in age, form, or the biological or experiential characteristics of the performing animal; enrichments that do not immediately reduce stereotypies should not be assumed failures with respect to welfare; and finally, stereotypies should not be reduced by means other than tackling their underlying motivations.

Contemporary animal welfare thinking is increasingly emphasising the promotion of positive states. There is a need for existing assessment frameworks to accommodate this shift in emphasis. This paper describes extensions to the Five Domains model, originally devised to assess welfare compromise, that facilitate consideration of positive experiences that may enhance welfare. As originally configured, the model provided a systematic method for identifying compromise in four physical/functional domains (nutrition, environment, health, behaviour) and in one mental domain that reflects the animal's overall welfare state understood in terms of its affective experiences. The specific modifications described here now facilitate additional identification in each domain of experiences animals have which may be accompanied by positive affects that would enhance welfare. It is explained why the grading scale and indices for evaluating welfare compromise necessarily differ from those for assessing welfare enhancement. Also, it is shown that the compromise and enhancement grades can be combined to provide a single informative symbol, the scaled use of which covers the range from severe welfare compromise and no enhancement to no compromise and high-level enhancement. Adapted thus, the Five Domains model facilitates systematic and structured assessment of positive as well as negative welfare-related affects, the circumstances that give rise to them and potential interactions between both types of affect, all of which extend the utility of the model. Moreover, clarification of the extended conceptual framework of the model itself contributes to the growing contextual shift in animal welfare science towards the promotion of positive states whilst continuing to minimise negative states.

The Glasgow Composite Measure Pain Scale (CMPS) for dogs suffering acute pain, developed using psychometric methodology, measures pain to a level of precision suitable for clinical trials. However, for routine clinical use, where the emphasis is on speed, ease of use, and guidance for analgesia provision, a short form (CMPS-SF) was developed. The CMPS-SF comprises six behavioural categories with associated descriptive expressions (items): vocalisation (4), attention to wound (5), mobility (5), response to touch (6), demeanour (5) and posture/activity (5). Items are placed in increasing order of pain intensity and numbered accordingly. The observer chooses that item within each category which best describes the dog's behaviour and ranked scores are summed; the maximum pain score is 24, or 20 if mobility is impossible to assess. Veterinary surgeons in Glasgow, University College Dublin and North Carolina Veterinary Schools completed the CMPS-SF for 122 dogs undergoing post-operative care and thereafter were asked “Do you think this animal requires analgesia? Yes/No”. The population difference in median pain score, for dogs considered to require analgesia (seven) compared with those that did not (three), was highly statistically significant (P < 0.001). Consideration of a clinical decision-point for analgesia gave an intervention level of 6/24, and 5/20 when section B (mobility assessment) could not be carried out. Difficulties in recognising pain contribute to the sub-optimal use of analgesics in veterinary practice. The CMPS-SF provides a practical means of assessing acute post-operative pain and provides guidance with regard to analgesic requirement, so improving pain management and welfare. The CMPS-SF can be downloaded from the Glasgow Pain and Welfare website at http://www.gla.ac.uk/vet/painandwelfare.

Milk yield per cow has more than doubled in the previous 40 years and many cows now produce more than 20,000 kg of milk per lactation. The increase in production should be viewed with concern because: i) the increase in milk yield has been accompanied by declining fertility, increasing leg and metabolic problems and declining longevity; ii) there are unfavourable genetic correlations between milk yield and fertility, mastitis and other production diseases, indicating that deterioration in fertility and health is largely a consequence of selection for increased milk yield; and iii) high disease incidence, reduced fertility, decreased longevity and modification of normal behaviour are indicative of substantial decline in cow welfare. Improving welfare is important as good welfare is regarded by the public as indicative of sustainable systems and good product quality and may also be economically beneficial. Expansion of the Profitable Lifetime Index used in the UK to include mastitis resistance and fertility could increase economic response to selection by up to 80%, compared with selection for milk production alone. In the last 10 years, several breeding organisations in Europe and North America followed the example of Nordic Countries and have included improving fertility and reducing incidence of mastitis in their breeding objectives, but these efforts are still timid. A multi-trait selection programme in which improving health, fertility and other welfare traits are included in the breeding objective, and appropriately weighted relative to production traits, should be adopted by all breeding organisations motivated in their goal of improving welfare.

This paper reviews the literature on human attitudes to animals, and postulates the existence of two primary motivational determinants of attitudes labelled ‘affect’ and ‘utility’. It also proposes that the relative strengths of these key attitude dimensions are affected by various modifying variables including the specific attributes of the animal, the individual characteristics and experience of the person evaluating the animal, and a range of cultural factors. The role of science as a cultural modifier of human attitudes to animals is also discussed.

Although the physiological and behavioural changes that can indicate poor welfareare generally agreed upon, using these measures in practice sometimes yieldsresults that are hard to interpret. For example, different types of measure maysuggest quite different things about an animal's welfare. Suchcontradictions are often due to the differing properties of the variables beingmeasured. How each variable responds to a stressor can be affected by severalfactors - the type of unpleasant stimulus to which the animal is exposed; whenand for how long exposure occurs; the animal's psychological state, egdoes it feel that it is in control?; and the time at which the measurement ismade, relative to the stressor. Typical responses also often differ betweenspecies and between individuals, and may even change in a single individual overtime. Furthermore, some responses used to assess welfare lack specificity: theycan be elicited by neutral or even pleasant events as well as by aversive ones.Appreciating these factors is vital when designing experiments, when choosingwhat to measure along with each welfare variable, and when interpreting results.Even after taking these factors into consideration, interpreting a result canstill be difficult. One approach then is to consider the effects on welfare ofthe changes measured, eg if there is immunosuppression, does the animal succumbto disease? Another is to use the animal's behaviour to indicate itspreference for, or aversion to, particular environments. Ultimately, however,interpreting welfare measures involves subjective judgements which will beinfluenced by the nature of our concern for the animal under consideration. Byraising these problems, we hope that this review will highlight and clarify theapparent contradictions that sometimes emerge in scientific studies of animalwelfare, and help researchers improve the designs of their experiments for thebenefit of the animals concerned.

Rather than construct lists of many different welfare indicators and give each of them the same weight, I argue that the assessment of animal welfare should be directed at answering two key questions: I) Are the animals healthy? 2) Do they have what they want? Behaviour has a major role in answering both. Behaviour is currently used to help answer the first question through its use in the clinical and pre-clinical assessment of pain, injury and disease, and potentially could have an even greater role, particularly if used in conjunction with new technology. Behaviour is also of crucial importance in gauging what animals want, most obviously in the use of choice and preference tests, but also through other methods that are particularly suitable for on-farm welfare assessment. These include quantitative observations of the spatial distribution of animals and of behavioural ‘indicators’ of what animals want, such as vocalisations.

In hedonic theories of motivation, ‘motivational affective states’ (MASs) are typically seen as adaptations which motivate certain types of behaviour, especially in situations where a flexible or learned response is more adaptive than a rigid or reflexive one. MASs can be negative (eg unpleasant feelings of hunger or pain) or positive (eg pleasant feelings associated with eating and playing). Hedonic theories often portray negative and positive MASs as opposite ends of a one-dimensional scale.

We suggest that natural selection has favoured negative and positive affect as separate processes to solve two different types of motivational problems. We propose that negative MASs (eg thirst, fear) evolved in response to ‘need situations’ where the fitness benefit of an action has increased, often because the action is needed to cope with a threat to survival or reproductive success. We propose that these negative MASs develop in response to a change in the body (eg dehydration) or the environment (eg the approach of a predator) which creates the need for action, and that negative MASs can become intense and prolonged if the threat to fitness is high and persistent. We propose that positive MASs evolved in ‘opportunity situations ‘ where an action (eg playing, exploring) has become advantageous because the fitness cost of performing it has declined. We propose that these positive MASs occur during, or as a result of, the performance of types of behaviour which are beneficial for fitness at a variety of times, not only when they are required to meet immediate needs; and that the pleasure inherent in the behaviour motivates the animal to perform it when the cost of so doing is sufficiently low. Some behaviour (eg eating) can be motivated by both positive and negative affect. Other behaviour, such as playing or fleeing from a predator, may be motivated largely by positive or negative affect alone. Our hypothesis needs to be tested, but we suggest that it corresponds well to common human experience.

The hypothesis provides a basis for predicting whether an aspect of animal management is likely to cause strong and prolonged negative affect (‘suffering’), or to prevent animals from experiencing certain types of pleasure. This distinction is important for bringing animal welfare assessment into line with ethical concerns.

Historically, pet dogs were trained using mainly negative reinforcement or punishment, but positive reinforcement using rewards has recently become more popular. The methods used may have different impacts on the dogs’ welfare. We distributed a questionnaire to 364 dog owners in order to examine the relative effectiveness of different training methods and their effects upon a pet dog's behaviour. When asked how they trained their dog on seven basic tasks, 66% reported using vocal punishment, 12% used physical punishment, 60% praise (social reward), 51% food rewards and II% play. The owner's ratings for their dog's obedience during eight tasks correlated positively with the number of tasks which they trained using rewards (P< 0.01), but not using punishment (P = 0.5). When asked whether their dog exhibited any of 16 common problematic behaviours, the number of problems reported by the owners correlated with the number of tasks for which their dog was trained using punishment (P< 0.001), but not using rewards (P = 0.17). Exhibition of problematic behaviours may be indicative of compromised welfare, because such behaviours can be caused by— or result in — a state of anxiety and may lead to a dog being relinquished or abandoned. Because punishment was associated with an increased incidence of problematic behaviours, we conclude that it may represent a welfare concern without concurrent benefits in obedience. We suggest that positive training methods may be more useful to the pet-owning community.

Welfare is multidimensional, comprising good health, comfort, expression of behaviour, and so on. Its overall assessment therefore requires a multicriteria evaluation. The set of criteria shall be exhaustive (no missing item), minimal (only necessary items), agreed by stakeholders, and legible (a limited number of criteria). Furthermore, the interpretation from one criterion shall not depend on that from another. We propose a set of 12 subcriteria grouped into four criteria: feeding, housing, health and optimised emotional states. This work will assist in developing measures to be used on-farm to form a European standard for overall assessment of animal welfare.

Increasing litter size has long been a goal of pig breeders and producers, and may have implications for pig (Sus scrofa domesticus) welfare. This paper reviews the scientific evidence on biological factors affecting sow and piglet welfare in relation to large litter size. It is concluded that, in a number of ways, large litter size is a risk factor for decreased animal welfare in pig production. Increased litter size is associated with increased piglet mortality, which is likely to be associated with significant negative animal welfare impacts. In surviving piglets, many of the causes of mortality can also occur in non-lethal forms that cause suffering. Intense teat competition may increase the likelihood that some piglets do not gain adequate access to milk, causing starvation in the short term and possibly long-term detriments to health. Also, increased litter size leads to more piglets with low birth weight which is associated with a variety of negative long-term effects. Finally, increased production pressure placed on sows bearing large litters may produce health and welfare concerns for the sow. However, possible biological approaches to mitigating health and welfare issues associated with large litters are being implemented. An important mitigation strategy is genetic selection encompassing traits that promote piglet survival, vitality and growth. Sow nutrition and the minimisation of stress during gestation could also contribute to improving outcomes in terms of piglet welfare. Awareness of the possible negative welfare consequences of large litter size in pigs should lead to further active measures being taken to mitigate the mentioned effects.

The notion ‘quality of life’ (QoL) suggests that welfare in animals encompasses more than just an absence of suffering; it concerns the quality of an animal's entire relationship with its environment, of how it lives its life. Judgements of such quality are based on the integration of perceived details of how animals behave over time in different contexts. The scientific status of such judgements has long been ambiguous, but in recent decades has begun to be addressed by animal scientists. This paper starts with a brief review of qualitative approaches to the study of animal behaviour, which tend to address characteristics such as individuality, personality, and emotionality. The question then arises whether such characteristics involve a subjective, experiential aspect, and identify animals as sentient beings. The second half of this paper argues that taking the integrative nature of qualitative judgements seriously enables a ‘whole animal’ perspective, through which it becomes possible to view behaviour as a dynamic, expressive body language that provides a basis for assessing the quality of an animal's experience (eg contented, anxious). Judging this quality is a skill that requires knowledge of species-specific behaviour, experience in observing and interacting with animals in different contexts, and a willingness to communicate with animals as sentient beings. A substantial body of research indicates that this skill can function reliably in a scientific context, and can be applied usefully as a practical welfare assessment tool. Thus qualitative approaches to the study of animal behaviour should make an important contribution to the growing interest in animal QoL.

The presence of human visitors has been shown to affect the behaviour of several different mammalian species in a number of different zoos, but the behavioural changes observed are not always consistent with a simple ‘stressful influence’ explanation. Data for non-primate species are too sparse to draw meaningful conclusions; but for primates, the evidence reviewed in this paper allows several hypotheses to be tested. Neither a social facilitation nor an audience attraction hypothesis can be generally supported by the available studies. However, these studies are consistent with a general stressful influence hypothesis, although the extent of this influence is itself affected by other variables, notably species and housing differences. There is some evidence that chronic exposure to human audiences may lessen this stressful influence in some species; and in certain circumstances (notably where some members of the public throw food) the effect of the audience is almost an enriching one.

Growing public concern regarding animal welfare and consumer demand for humanely produced products have placed pressure on the meat, wool and dairy industries to improve and confirm the welfare status of their animals. This has increased the need for reliable methods of assessing animal welfare during commercial farm practices. The measurement of the stress caused by commercial farm practices is a major component of animal welfare assessment. However, a major issue for animal welfare science is that many of the techniques used to measure stress involve invasive procedures, such as blood sampling, which may themselves cause a stress response and therefore affect the measurement of interest. To reduce this problem, a number of non-invasive or minimally invasive methods and devices have been developed to measure stress. These include the measurement of cortisol concentrations in saliva and faeces, and remote devices for recording body temperature, heart rate and the collection of blood samples. This review describes the benefits and limitations of some of these methods for measuring stress. In particular, the review focuses on recent advances and current research in the use of infrared thermography (IRT) for measuring stress. Specific applications for IRT in the dairy and beef industries are also described including an automated, non-invasive system for early diagnosis of infection in cattle. It is essential that non-invasive measures of acute and chronic stress are developed for reliable assessment of animal welfare during standard farm management practices and IRT may be a useful tool for this purpose. IRT may offer advantages over many other non-invasive systems as it appears to be capable of measuring different components of the stress axis, including acute sympathetic and hypothalamic-pituitary-adrenocortical responses.

Four groups of dogs, which had been subjected to housing conditions of varying quality for years, were assumed to experience different levels of stress. The groups were compared for behavioural and hormonal parameters in order to identify measures that indicate chronic stress in the dog and which may help to identify poor welfare in this species. As a standard for comparison, one of the four groups was composed of privately owned dogs; we assumed that chronic stress levels were relatively low in this group (GI). The three remaining groups of dogs (GII, Gill and GIV) were kept under conditions of low to relatively high austerity, and had basal urinary ratios of Cortisol to creatinine, adrenaline to creatinine and, to a lesser extent, noradrenaline to creatinine, that varied from low to high, respectively. Significant differences (P < 0.05) were found in Cortisol to creatinine ratios when comparing GI to GII, GUI and GIV and when GII was compared to GIV. Statistical analyses indicated that the mean adrenaline to creatinine ratio in GI differed from that in the remaining groups and that the ratio in GII differed from that in GUI. Noradrenaline to creatinine ratios differed significantly only between GI and GUI. Dopamine to creatinine ratios and noradrenaline to adrenaline ratios did not differ significantly between groups. When dogs were not disturbed, those that were kept under the most austere conditions typically had high levels of locomotor activity, nosing, urinating and paw lifting. After mild disturbance by a slamming door or in the presence of a researcher these animals reacted actively, with increased locomotor activity, circling and nosing, and they showed high levels of behaviours that have previously been associated with acute stress: body shaking, yawning, ambivalent postures and displacement behaviours. Chronic stress in dogs may be identified by increased paw lifting when animals are not disturbed and by ample behavioural expressions of arousal when they are mildly stimulated. Since some behaviours may occur in contexts not related to stress, behavioural data are easily misinterpreted with regard to chronic stress. Interpretation will only be meaningful when physiological measures such as urinary adrenaline to creatinine ratios and, especially, urinary Cortisol to creatinine ratios are also determined.

For cats, appropriate housing conditions and a quick adjustment to new surroundings should be promoted during temporary stays in animal shelters and boarding catteries. In this study the development of stress in 140 boarding cats during a two-week stay under single-, pairand group-housing conditions in a boarding cattery was investigated and compared with the stress levels of 45 control cats which had been at the animal shelter for several weeks. Signs of stress were recorded by a non-invasive Cat-Stress-Score.

Overall, the levels of stress in boarding cats declined during the two weeks of boarding, with a pronounced decline in the first days, but did not reach the stress levels of the control group by the end of the second week of housing. In the second week, the average stress level of about one third of all boarding cats was rated higher than ‘weakly tense’ with 4 per cent of cats rated even higher than ‘very tense’. Neither housing style (single, paired or grouped) nor age had an influence on stress levels.

It was concluded that about two thirds of the boarding cats adjusted well to the boarding cattery during a two-week stay, while for the other third, temporary boarding was more stressful. For 4 per cent of the animals the two-week stay in a boarding cattery was classified as inappropriate because no reduction of their high stress levels occurred.

Improving the quality of life of captive animals is dependent on developing valid measures of how animals feel about their lives. It has recently been suggested that biases in information processing may offer a novel means of understanding animal emotions. Anxious and depressed people tend to interpret ambiguous information negatively. We explored the proposal that such cognitive biases also exist in non-human animals and could therefore be used as novel measures of animal welfare. We used a novel cognitive bias task based on a learnt taste aversion to determine whether birds deprived of environmental enrichment show biases in their classification of ambiguous signals. We hypothesised that starlings in enriched cages should be more likely to classify ambiguous signals as being associated with a positive outcome than starlings housed in standard, unenriched cages. Starlings were trained on a go/no-go procedure to discriminate between two visual stimuli (cardboard lids of white and dark grey) associated with outcomes of a different value (palatable and unpalatable mealworms hidden underneath). Individual birds' responses to unreinforced, intermediate stimuli (various shades of grey between white and dark grey) were subsequently examined while each bird was housed sequentially in both standard and enriched cages. The probability of a bird classifying an ambiguous pale grey lid as hiding a palatable mealworm was lower in standard cages than enriched cages, but this difference was found only in birds that received enriched cages first. Our results can be interpreted as showing a pessimistic bias in birds that have recently experienced a decline in environmental quality. These findings support the use of cognitive bias-based tasks as a novel, non-invasive technique for assessing welfare in non-human animals.

A multitude of endocrine mechanisms are involved in coping with challenges. Glucocorticoids, secreted by the adrenal glands, are in the front line of the battle to overcome stressful situations. They are usually measured in plasma samples as parameters of adrenal activity and thus of disturbance. Unfortunately, collecting blood samples itself can disturb an animal. Thus, non-invasive methods for the determination of glucocorticoids or their metabolites have become increasingly popular. The pros and cons of various non-invasive sample materials (saliva, excreta, milk, hair/feathers and eggs) for glucocorticoid determination are given. Above all, faecal samples offer the advantage that they can be collected easily. In faecal samples, circulating hormone levels are integrated over a certain period of time and represent the cumulative secretion of hormones. Thus, the levels are less affected by short fluctuations or the pulse-like nature of hormone secretion. However, using this technique to assess an animal's adrenocortical activity is not especially simple. Whether frequent sampling is necessary or single samples will suffice depends upon the study's aim (whether one is examining the impact of acute or chronic stressors). Background knowledge of the metabolism and excretion of cortisol/corticosterone metabolites is required and a careful validation for each species and sex investigated is obligatory. The present review also addresses analytical issues regarding sample storage, extraction procedures and immunoassays and includes a comprehensive list of published studies (up to 2011) describing the use of such methods in farmed animals. Applied properly, non-invasive techniques to monitor glucocorticoid metabolites in faecal samples of various species are a useful tool for welfare assessment, especially as they are easily applied at farm or group level.

Pain experienced by lame cattle is often masked by their instinctive stoicism, leading to delayed detection and treatment of lameness. This paper investigates the usefulness of posture scoring during locomotion and the evaluation of daily activity levels as indicators of pain and discomfort resulting from lameness in dairy cattle. In this study, 345 lactating cattle were observed over a seven-month winter housing period. The posture of each cow was scored twice weekly using a subjective five-point numerical rating scale. Daily activity levels were measured objectively using pedometers. The effect of lameness on behaviour was addressed. Increased posture scores were associated with the presence of foot lesions (P < 0.001) and with reduced daily activity levels (P < 0.001). In comparison to sound cows, lame cows showed lower daily activity levels (P < 0.001). These results improve understanding of pain-related behaviours in cattle, and such an approach may assist future development of welfare assessment systems.