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The mental experiences of animals are what characterises their welfare status. The Five Domains Model for assessing welfare aligns with the understanding that physical and mental states are linked. Following measurement of indicators within each of the four physical/functional Domains (1. Nutrition; 2. Physical environment; 3. Health; and 4. Behavioural interactions), the anticipated negative or positive affective consequences (mental experiences) are cautiously inferred and assigned to Domain 5. Those inferences derive credibility from validated knowledge of the underlying systems of physiology, neurophysiology, neuroethology and affective neuroscience. Any indicators used for assessing welfare need to be scientifically validated. This requires, firstly, evidence of the links between a measurable/observable indicator and the physical/functional impact (in Domains 1 to 4), and secondly, a demonstrable relationship between the physical/functional impact and the mental experience it is inferred the indicators reflect (in Domain five). This review refers to indicators of physical/functional states in Domains 1 to 4, which have been shown to be measurable in free-roaming wild horses, and then evaluates the scientific evidence linking them to inferred mental experiences in Domain 5. This is the first time that the scientific evidence validating a comprehensive range of welfare indicators has been synthesised in this way. Inserting these indicators into the Five Domains Model enables transparently justifiable assessment and grading of welfare status in free-roaming horses.

A detailed understanding of what is usual for a species under optimal conditions is critical for identifying and interpreting different features of body function that have known impacts on animal welfare and its assessment. When applying the Five Domains Model to assess animal welfare, the key starting point is therefore to acquire extensive species-specific knowledge relevant to each of the four physical/functional Domains of the Model. These Domains, 1 to 4, address areas where objective information is evaluated and collated. They are: (1) Nutrition; (2) Physical environment; (3) Health; and (4) Behavioural interactions. It is on the basis of this detailed knowledge that cautious inferences can then be made about welfare-relevant mental experiences animals may have, aligned with Domain 5, Mental State. However, this review is focused entirely on the first four Domains in order to provide a novel holistic framework to collate the multidisciplinary knowledge of horses required for undertaking comprehensive welfare assessments. Thus, inferring the potential mental experiences aligned with Domain 5, the final step in model-based welfare assessments, is not considered here. Finally, providing extensive information on free-roaming horses enables a better understanding of the impacts of human interventions on the welfare of horses in both free-roaming and domestic situations.

A thoracic squeeze has been observed to cause low-vigour neonates of various farmed mammal species, including calves, to enter a state of reduced responsiveness. The removal of the squeeze causes rapid recovery and the expression of normal, healthy behaviours. However, the responses of healthy calves to a thoracic squeeze have not yet been characterized. The responses of 16 healthy newborn calves to a thoracic squeeze are described, along with the effect of the squeeze's application method on the response. Calves aged between 12 and 36 h were subjected to the squeeze using a rope ( n = 8) or an inflation cuff ( n = 8). In total, 13 of the 16 calves were induced into a state of reduced responsiveness, though neural reflexes persisted in nearly all of them. The squeeze was discontinued for nearly half of those induced before the end of the 10-min period, either due to spontaneous arousal or physiological instability. Both methods of application were equally effective at inducing reduced responsiveness, though responses to the cuff appeared to be more rapid than those to the rope. These findings support previous research on piglets and foals, and suggest that the behavioural responses to a thoracic squeeze are generalised across neonates of precocial farmed mammals; the findings provide a foundation for further research exploring the mechanisms underlying the response and the benefits that its application may bring for the performance of husbandry procedures.

A thoracic squeeze has been observed to cause both healthy and low vigour neonatal foals to enter a 'less-responsive state', characterised by loss of posture, eye closure and cessation of movement, from which they rapidly recover to express normal healthy behaviours when the squeeze is released. To date, there have been no systematic studies characterising the responses of healthy neonates of other mammalian species to a thoracic squeeze. We describe the responses of healthy newborn piglets ( n = 17) to a standardised application of the thoracic squeeze and evaluate the effect of the method of squeeze application on the response. Neonatal piglets were squeezed around the chest with either a soft fabric rope as has been used in foals ( n = 8) or a novel purpose-made inflation cuff ( n = 9). Both methods were effective at inducing a less-responsive behavioural state in all piglets, with neural reflexes reduced or absent in over half of them. The inflation cuff appeared to induce the less-responsive state faster than the rope, and more piglets squeezed with the cuff remained in this state for the full 10-min squeeze. These findings suggest that the behavioural response of foals to thoracic squeezing can be generalised to neonates of other precocial mammalian species. This initial study provides a foundation for further research using the inflation cuff to explore mechanisms underlying the thoracic squeeze and ways in which it may be applied whilst performing husbandry procedures.

We previously developed a Ten-Stage Protocol for scientifically assessing the welfare of individual free-roaming wild animals using the Five Domains Model. The protocol includes developing methods for measuring or observing welfare indices. In this study, we assessed the use of remote camera traps to evaluate an extensive range of welfare indicators in individual free-roaming wild horses. Still images and videos were collected and analysed to assess whether horses could be detected and identified individually, which welfare indicators could be reliably evaluated, and whether behaviour could be quantitatively assessed. Remote camera trapping was successful in detecting and identifying horses (75% on still images and 72% on video observation events), across a range of habitats including woodlands where horses could not be directly observed. Twelve indicators of welfare across the Five Domains were assessed with equal frequency on both still images and video, with those most frequently assessable being body condition score (73% and 79% of observation events, respectively), body posture (76% for both), coat condition (42% and 52%, respectively), and whether or not the horse was sweating excessively (42% and 45%, respectively). An additional five indicators could only be assessed on video; those most frequently observable being presence or absence of weakness (66%), qualitative behavioural assessment (60%), presence or absence of shivering (51%), and gait at walk (50%). Specific behaviours were identified in 93% of still images and 84% of video events, and proportions of time different behaviours were captured could be calculated. Most social behaviours were rarely observed, but close spatial proximity to other horses, as an indicator of social bonds, was recorded in 36% of still images, and 29% of video observation events. This is the first study that describes detailed methodology for these purposes. The results of this study can also form the basis of application to other species, which could contribute significantly to advancing the field of wild animal welfare.

Throughout its 25-year history, the Five Domains Model for animal welfare assessment has been regularly updated to include at each stage the latest authenticated developments in animal welfare science thinking. The domains of the most up-to-date Model described here are: 1 Nutrition, 2 Physical Environment, 3 Health, 4 Behavioural Interactions and 5 Mental State. The first four domains focus attention on factors that give rise to specific negative or positive subjective experiences (affects), which contribute to the animal's mental state, as evaluated in Domain 5. More specifically, the first three domains focus mainly on factors that disturb or disrupt particular features of the body's internal stability. Each disturbed or disrupted feature generates sensory inputs which are processed by the brain to form specific negative affects, and these affects are associated with behaviours that act to restore the body's internal stability. As each such behaviour is essential for the survival of the animal, the affects associated with them are collectively referred to as "survival-critical affects". In contrast, Domain 4, now named Behavioural Interactions, focusses on evidence of animals consciously seeking specific goals when interacting behaviourally with (1) the environment, (2) other non-human animals and (3) as a new feature of the Model outlined here, humans. The associated affects, evaluated via Domain 5, are mainly generated by brain processing of sensory inputs elicited by external stimuli. The success of the animals' behavioural attempts to achieve their chosen goals is reflected in whether the associated affects are negative or positive. Collectively referred to as "situation-related affects", these outcomes are understood to contribute to animals' perceptions of their external circumstances. These observations reveal a key distinction between the way survival-critical and situation-related affects influence animals' aligned behaviours. The former mainly reflect compelling motivations to engage in genetically embedded behavioural responses, whereas the latter mainly involve conscious behavioural choices which are the hallmarks of agency. Finally, numerous examples of human-animal interactions and their attendant affects are described, and the qualitative grading of interactions that generate negative or positive affect is also illustrated.

A proposition addressed here is that, although bitted horses are viewed by many equestrians as being largely free of bit-related mouth pain, it seems likely that most behavioural signs of such pain are simply not recognised. Background information is provided on the following: the major features of pain generation and experience; cerebrocortical involvement in the conscious experience of pain by mammals; the numerous other subjective experiences mammals can have; adjunct physiological responses to pain; some general feature of behavioural responses to pain; and the neural bases of sensations generated within the mouth. Mouth pain in horses is then discussed. The areas considered exclude dental disease, but they include the stimulation of pain receptors by bits in the interdental space, the tongue, the commissures of the mouth, and the buccal mucosa. Compression, laceration, inflammation, impeded tissue blood flow, and tissue stretching are evaluated as noxious stimuli. The high pain sensitivity of the interdental space is described, as are likely increases in pain sensitivity due to repeated bit contact with bruises, cuts, tears, and/or ulcers wherever they are located in the mouth. Behavioural indices of mouth pain are then identified by contrasting the behaviours of horses when wearing bitted bridles, when changed from bitted to bit-free bridles, and when free-roaming unbitted in the wild. Observed indicative behaviours involve mouth movements, head-neck position, and facial expression ("pain face"), as well as characteristic body movements and gait. The welfare impacts of bit-related pain include the noxiousness of the pain itself as well as likely anxiety when anticipating the pain and fear whilst experiencing it, especially if the pain is severe. In addition, particular mouth behaviours impede airflow within the air passages of the upper respiratory system, effects that, in their turn, adversely affect the air passages in the lungs. Here, they increase airflow resistance and decrease alveolar gas exchange, giving rise to suffocating experiences of breathlessness. In addition, breathlessness is a likely consequence of the low jowl angles commonly maintained during dressage. If severe, as with pain, the prospect of breathlessness is likely to give rise to anxiety and the direct experience of breathlessness to fear. The related components of welfare compromise therefore likely involve pain, breathlessness, anxiety, and fear. Finally, a 12-point strategy is proposed to give greater impetus to a wider adoption of bit-free bridles in order to avoid bit-induced mouth pain.

Knowledge of the welfare status of wild animals is vital for informing debates about the ways in which we interact with wild animals and their habitats. Currently, there is no published information about how to scientifically assess the welfare of free-roaming wild animals during their normal day-to-day lives. Using free-roaming horses as an example, we describe a ten-stage protocol for systematically and scientifically assessing the welfare of individual non-captive wild animals. The protocol starts by emphasising the importance of readers having an understanding of animal welfare in a conservation context and also of the Five Domains Model for assessing welfare. It goes on to detail what species-specific information is required to assess welfare, how to identify measurable and observable indicators of animals' physical states and how to identify which individuals are being assessed. Further, it addresses how to select appropriate methods for measuring/observing physical indicators of welfare, the scientific validation of these indicators and then the grading of animals' welfare states, along with assigning a confidence score. Finally, grading future welfare risks and how these can guide management decisions is discussed. Applying this ten-stage protocol will enable biologists to scientifically assess the welfare of wild animals and should lead to significant advances in the field of wild animal welfare., Competing Interests: The authors declare no conflict of interest.

Presented is an updated understanding of the development of sensory systems in the offspring of a wide range of terrestrial mammals, the prenatal exposure of those systems to salient stimuli, and the mechanisms by which that exposure can embed particular sensory capabilities that prepare newborns to respond appropriately to similar stimuli they may encounter after birth. Taken together, these are the constituents of the phenomenon of "trans-natal sensory continuity" where the embedded sensory capabilities are considered to have been "learnt" and, when accessed subsequently, they are said to have been "remembered". An alternative explanation of trans-natal sensory continuity is provided here in order to focus on the mechanisms of "embedding" and "accessing" instead of the potentially more subjectively conceived outcomes of "learning" and "memory". Thus, the mechanistic concept of "intrauterine sensory entrainment" has been introduced, its foundation being the well-established neuroplastic capability of nervous systems to respond to sensory inputs by reorganising their neural structures, functions, and connections. Five conditions need to be met before "trans-natal sensory continuity" can occur. They are (1) sufficient neurological maturity to support minimal functional activity in specific sensory receptor systems in utero; (2) the presence of sensory stimuli that activate their aligned receptors before birth; (3) the neurological capability for entrained functions within specific sensory modalities to be retained beyond birth; (4) specific sensory stimuli that are effective both before and after birth; and (5) a capability to detect those stimuli when or if they are presented after birth in ways that differ (e.g., in air) from their presentation via fluid media before birth. Numerous beneficial outcomes of this process have been reported for mammalian newborns, but the range of benefits depends on how many of the full set of sensory modalities are functional at the time of birth. Thus, the breadth of sensory capabilities may be extensive, somewhat restricted, or minimal in offspring that are, respectively, neurologically mature, moderately immature, or exceptionally immature at birth. It is noted that birth marks a transition from intrauterine sensory entrainment to extrauterine sensory entrainment in all mammalian young. Depending on their neurological maturity, extrauterine entrainment contributes to the continuing maturation of the different sensory systems that are operational at birth, the later development and maturation of the systems that are absent at birth, and the combined impact of those factors on the behaviour of newborn and young mammals. Intrauterine sensory entrainment helps to prepare mammalian young for life immediately after birth, and extrauterine sensory entrainment continues this process until all sensory modalities develop full functionality. It is apparent that, overall, extrauterine sensory entrainment and its aligned neuroplastic responses underlie numerous postnatal learning and memory events which contribute to the maturation of all sensory capabilities that eventually enable mammalian young to live autonomously.

The focus of this opinion is on the key features of sentience in animals which can experience different states of welfare, encapsulated by the new term 'welfare-aligned sentience'. This term is intended to exclude potential forms of sentience that do not enable animals in some taxa to have the subjective experiences which underlie different welfare states. As the scientific understanding of key features of sentience has increased markedly during the last 10 to 15 years, a major purpose here is to provide up-to-date information regarding those features. Eleven interconnected statements about sentience-associated body functions and behaviour are therefore presented and explained briefly. These statements are sequenced to provide progressively more information about key scientifically-supported attributes of welfare-aligned sentience, leading, in their entirety, to a more comprehensive understanding of those attributes. They are as follows: (1) Internal structure-function interactions and integration are the foundations of sentience; (2) animals posess a capacity to respond behaviourally to a range of sensory inputs; (3) the more sophisticated nervous systems can generate subjective experiences, that is, affects; (4) sentience means that animals perceive or experience different affects consciously; (5) within a species, the stage of neurobiological development is significant; (6) during development the onset of cortically-based consciousness is accompanied by cognitively-enhanced capacities to respond behaviourally to unpredictable postnatal environments; (7) sentience includes capacities to communicate with others and to interact with the environment; (8) sentience incorporates experiences of negative and positive affects; (9) negative and positive affective experiences 'matter' to animals for various reasons; (10) acknowledged obstacles inherent in anthropomorphism are largely circumvented by new scientific knowledge, but caution is still required; and (11) there is increasing evidence for sentience among a wider range of invertebrates. The science-based explanations of these statements provide the foundation for a brief definition of 'welfare-aligned sentience', which is offered for consideration. Finally, it is recommended that when assessing key features of sentience the same emphasis should be given to positive and negative affective experiences in the context of their roles in, or potential impacts on, animal welfare.

There is a growing interest and need for zoos to develop and implement welfare assessment tools that are practical to use and provide meaningful results that can inform management decisions. This paper presents a process that was developed to support this type of evidence-based management in zoo animal welfare. The process is configured to facilitate institutional risk assessment, using an adapted version of the Five Domains Model for animal welfare assessment. It is designed to systematically analyse information gathered from zoo personnel in order to highlight areas of welfare risk, as well as areas that are performing well and areas requiring further investigation. A trial was conducted on three zoos over three years. Results of the trial suggest the process developed is practical and effective in identifying areas of welfare risk in a wide range of species in a zoo setting. It represents a further step towards achieving high-level animal welfare in zoos by integrating animal welfare as an institutional priority. The more zoos that employ such strategies, the greater the ability of the sector to advance the welfare of the animals in their care.

Conceptual frameworks for understanding animal welfare scientifically are widely influential. An early “biological functioning” framework still influences expert opinions prepared for Courts hearing animal cruelty cases, despite deficiencies in it being revealed by the later emergence and wide scientific adoption of an “affective state” framework. According to “biological functioning” precepts, indices of negative welfare states should predominantly be physical and/or clinical and any that refer to animals’ supposed subjective experiences, i.e., their “affective states”, should be excluded. However, “affective state” precepts, which have secure affective neuroscience and aligned animal behaviour science foundations, show that behavioural indices may be utilised to credibly identify negative welfare outcomes in terms of negative subjective experiences, or affects. It is noted that the now very wide scientific acceptance of the “affective state” framework is entirely consistent with the current extensive international recognition that animals of welfare significance are “sentient” beings. A long list of negative affects is discussed and each one is described as a prelude to updating the concept of “suffering” or “distress”, often referred to in animal welfare legislation and prosecutions for alleged ill-treatment of animals. The Five Domains Model for assessing and grading animal welfare compromise is then discussed, highlighting that it incorporates a coherent amalgamation of “biological functioning” and “affective state” precepts into its operational features. That is followed by examples of severe-to-very-severe ill-treatment of dogs. These include inescapable psychological and/or physical abuse or mistreatment, excessively restrictive or otherwise detrimental housing or holding conditions, and/or seriously inadequate provision of the necessities of life, in each case drawing attention to specific affects that such ill-treatment generates. It is concluded that experts should frame their opinions in ways that include negative affective outcomes. Moreover, the cogency of such analyses should be drawn to the attention of the Judiciary when they are deliberating on suffering in animals, thereby providing a basis for them to move from a current heavy reliance on physical and/or clinical indices of cruelty or neglect towards including in their decisions careful evaluations of animals’ negative affective experiences.

Laws, regulations and professional standards increasingly aim to ban or restrict non-therapeutic tail docking in canine puppies. These constraints have usually been justified by reference to loss of tail participation in communication between dogs, the acute pain presumed to be caused during docking itself, subsequent experiences of chronic pain and heightened pain sensitivity, and the occurrence of other complications. These areas are reconsidered here. First, a scientifically robust examination of the dynamic functional foundations, sensory components and key features of body language that are integral to canine communication shows that the role of the tail has been greatly underestimated. More specifically, it shows that tail behaviour is so embedded in canine communication that docking can markedly impede unambiguous interactions between different dogs and between dogs and people. These interactions include the expression of wide ranges of both negative and positive emotions, moods and intentions that are of daily significance for dog welfare. Moreover, all docked dogs may experience these impediments throughout their lives, which challenges assertions by opponents to such bans or restrictions that the tail is a dispensable appendage. Second, and in contrast, a re-examination of the sensory capacities of canine puppies reveals that they cannot consciously experience acute or chronic pain during at least the first week after birth, which is when they are usually docked. The contrary view is based on questionable between-species extrapolation of information about pain from neurologically mature newborns such as calves, lambs, piglets and human infants, which certainly can consciously experience pain in response to injury, to neurologically immature puppies which remain unconscious and therefore unable to experience pain until about two weeks after birth. Third, underpinned by the incorrect conclusion that puppies are conscious at the usual docking age, it is argued here that the well-validated human emotional drive or desire to care for and protect vulnerable young, leads observers to misread striking docking-induced behaviour as indicating that the puppies consciously experience significant acute pain and distress. Fourth, updated information reaffirms the conclusion that a significant proportion of dogs docked as puppies will subsequently experience persistent and significant chronic pain and heightened pain sensitivity. And fifth, other reported negative consequences of docking should also be considered because, although their prevalence is unclear, when they do occur they would have significant negative welfare impacts. It is argued that the present analysis strengthens the rationale for such bans or restrictions on docking of puppies by clarifying which of several justifications previously used are and are not scientifically supportable. In particular, it highlights the major roles the tail plays in canine communication, as well as the lifetime handicaps to communication caused by docking. Thus, it is concluded that non-therapeutic tail docking of puppies represents an unnecessary removal of a necessary appendage and should therefore be banned or restricted.

In accord with contemporary animal welfare science understanding, the Five Domains Model has a significant focus on subjective experiences, known as affects, which collectively contribute to an animal's overall welfare state. Operationally, the focus of the Model is on the presence or absence of various internal physical/functional states and external circumstances that give rise to welfare-relevant negative and/or positive mental experiences, i.e., affects. The internal states and external circumstances of animals are evaluated systematically by referring to each of the first four domains of the Model, designated "Nutrition", "Environment", "Health" and "Behaviour". Then affects, considered carefully and cautiously to be generated by factors in these domains, are accumulated into the fifth domain, designated "Mental State". The scientific foundations of this operational procedure, published in detail elsewhere, are described briefly here, and then seven key ways the Model may be applied to the assessment and management of animal welfare are considered. These applications have the following beneficial objectives-they (1) specify key general foci for animal welfare management; (2) highlight the foundations of specific welfare management objectives; (3) identify previously unrecognised features of poor and good welfare; (4) enable monitoring of responses to specific welfare-focused remedial interventions and/or maintenance activities; (5) facilitate qualitative grading of particular features of welfare compromise and/or enhancement; (6) enable both prospective and retrospective animal welfare assessments to be conducted; and, (7) provide adjunct information to support consideration of quality of life evaluations in the context of end-of-life decisions. However, also noted is the importance of not overstating what utilisation of the Model can achieve., Competing Interests: The author declares no conflict of interest.

Horses engaged in strenuous exercise display physiological responses that approach the upper functional limits of key organ systems, in particular their cardiorespiratory systems. Maximum athletic performance is therefore vulnerable to factors that diminish these functional capacities, and such impairment might also lead to horses experiencing unpleasant respiratory sensations, i.e., breathlessness. The aim of this review is to use existing literature on equine cardiorespiratory physiology and athletic performance to evaluate the potential for various types of breathlessness to occur in exercising horses. In addition, we investigate the influence of management factors such as rein and bit use and of respiratory pathology on the likelihood and intensity of equine breathlessness occurring during exercise. In ridden horses, rein use that reduces the jowl angle, sometimes markedly, and conditions that partially obstruct the nasopharynx and/or larynx, impair airflow in the upper respiratory tract and lead to increased flow resistance. The associated upper airway pressure changes, transmitted to the lower airways, may have pathophysiological sequelae in the alveolae, which, in their turn, may increase airflow resistance in the lower airways and impede respiratory gas exchange. Other sequelae include decreases in respiratory minute volume and worsening of the hypoxaemia, hypercapnia and acidaemia commonly observed in healthy horses during strenuous exercise. These and other factors are implicated in the potential for ridden horses to experience three forms of breathlessness-"unpleasant respiratory effort", "air hunger" and "chest tightness"-which arise when there is a mismatch between a heightened ventilatory drive and the adequacy of the respiratory response. It is not known to what extent, if at all, such mismatches would occur in strenuously exercising horses unhampered by low jowl angles or by pathophysiological changes at any level of the respiratory tract. However, different combinations of the three types of breathlessness seem much more likely to occur when pathophysiological conditions significantly reduce maximal athletic performance. Finally, most horses exhibit clear behavioural evidence of aversion to a bit in their mouths, varying from the bit being a mild irritant to very painful. This in itself is a significant animal welfare issue that should be addressed. A further major point is the potential for bits to disrupt the maintenance of negative pressure in the oropharynx, which apparently acts to prevent the soft palate from rising and obstructing the nasopharynx. The untoward respiratory outcomes and poor athletic performance due to this and other obstructions are well established, and suggest the potential for affected animals to experience significant intensities of breathlessness. Bitless bridle use may reduce or eliminate such effects. However, direct comparisons of the cardiorespiratory dynamics and the extent of any respiratory pathophysiology in horses wearing bitted and bitless bridles have not been conducted. Such studies would be helpful in confirming, or otherwise, the claimed potential benefits of bitless bridle use., Competing Interests: The authors declare no conflict of interest.

Although the Five Freedoms paradigm has been very influential in shaping animal welfare thinking for the last two decades, it has two key disadvantages. First, the focus on "freedom" from a range of negative experiences and states has been misunderstood in a number of quarters to mean that complete freedom from these experiences and states is possible, when in fact the best that can be achieved is for them to be minimised. Second, the major focus of the Freedoms on negative experiences and states is now seen to be a disadvantage in view of current understanding that animal welfare management should also include the promotion of positive experiences and states. The challenge therefore was to formulate a paradigm that overcame these two main problems and yet was straightforward enough to be accessible to non-specialists, including members of the lay public who are interested in animal welfare. This was achieved by highlighting the Five Provisions, originally aligned with the Five Freedoms, but now updated to direct welfare management towards activities that both minimise negative experiences or states and promote positive experiences or states as specified by particular Animal Welfare Aims assigned to each Provision. Aspects of the four welfare principles from the European Welfare Quality assessment system (WQ (®) ) and elements of all domains of the Five Domains Model for animal welfare assessment have been incorporated into the new Five Provisions/Welfare Aims paradigm. Thus, the paradigm is easily understood and provides clear guidance on beneficial objectives for animal welfare management. It is anticipated that the paradigm will have application to many species found in a wide range of circumstances., Competing Interests: The author declares no conflict of interest.

The present structured, systematic and comprehensive welfare evaluation of an injured working farm dog using the Five Domains Model is of interest in its own right. It is also an example for others wanting to apply the Model to welfare evaluations in different species and contexts. Six stages of a fictitious scenario involving the dog are considered: (1) its on-farm circumstances before one hind leg is injured; (2) its entanglement in barbed wire, cutting it free and transporting it to a veterinary clinic; (3) the initial veterinary examination and overnight stay; (4) amputation of the limb and immediate post-operative recovery; (5) its first four weeks after rehoming to a lifestyle block; and (6) its subsequent life as an amputee and pet. Not all features of the scenario represent average-to-good practice; indeed, some have been selected to indicate poor practice. It is shown how the Model can draw attention to areas of animal welfare concern and, importantly, to how welfare enhancement may be impeded or facilitated. Also illustrated is how the welfare implications of a sequence of events can be traced and evaluated, and, in relation to specific situations, how the degrees of welfare compromise and enhancement may be graded. In addition, the choice of a companion animal, contrasting its welfare status as a working dog and pet, and considering its treatment in a veterinary clinical setting, help to highlight various welfare impacts of some practices. By focussing attention on welfare problems, the Model can guide the implementation of remedies, including ways of promoting positive welfare states. Finally, wider applications of the Five Domains Model are noted: by enabling both negative and positive welfare-relevant experiences to be graded, the Model can be applied to quality of life assessments and end-of-life decisions and, with particular regard to negative experiences, the Model can also help to strengthen expert witness testimony during prosecutions for serious ill treatment of animals., Competing Interests: The authors declare no conflicts of interest.

The Five Freedoms have had major impact on animal welfare thinking internationally. However, despite clear initial statements that the words 'freedom from' should indicate 'as free as possible from', the Freedoms have come to be represented as absolute or fundamental freedoms, even rights, by some animal advocate and other groups. Moreover, a marked increase in scientific understanding over the last two decades shows that the Freedoms do not capture the more nuanced knowledge of the biological processes that is germane to understanding animal welfare and which is now available to guide its management. For example, the named negative experiences of thirst, hunger, discomfort and pain, and others identified subsequently, including breathlessness, nausea, dizziness, debility, weakness and sickness, can never be eliminated, merely temporarily neutralised. Each one is a genetically embedded element that motivates animals to behave in particular ways to obtain specific life-sustaining resources, avoid or reduce physical harm or facilitate recovery from infection or injury. Their undoubted negativity creates a necessary sense of urgency to respond, without which animals would not survive. Also, the temporary neutralisation of these survival-critical affects does not in and of itself generate positive experience. This questions the commonly held assumption that good animal welfare will result when these internally generated negative affects are minimised. Animals may also experience other negative affects that include anxiety, fear, panic, frustration, anger, helplessness, loneliness, boredom and depression. These situation-related affects reflect animals' perceptions of their external circumstances. Although they are elicited by threatening, cramped, barren and/or isolated conditions, they can often be replaced by positive affects when animals are kept with congenial others in spacious, stimulus-rich and safe environments which provide opportunities for them to engage in behaviours they find rewarding. These behaviours may include environment-focused exploration and food acquisition activities as well as animal-to-animal interactive activities, all of which can generate various forms of comfort, pleasure, interest, confidence and a sense of control. Animal welfare management should aim to reduce the intensity of survival-critical negative affects to tolerable levels that nevertheless still elicit the required behaviours, and should also provide opportunities for animals to behave in ways they find rewarding, noting that poor management of survival-critical affects reduces animals' motivation to utilize such rewarding opportunities. This biologically more accurate understanding provides support for reviewing the adequacy of provisions in current codes of welfare or practice in order to ensure that animals are given greater opportunities to experience positive welfare states. The purpose is to help animals to have lives worth living, which is not possible when the predominant focus of such codes is on survival-critical measures. Finally, an updated characterisation of animal welfare that incorporates this more accurate understanding is presented.