Spatial working memory in a disappearing object task is impaired in female but not male dogs with chronic osteoarthritis.

Chronic pain in humans is associated with impaired working memory but it is not known whether this is the case in long-lived companion animals, such as dogs, who are especially vulnerable to developing age-related chronic pain conditions. Pain-related impairment of cognitive function could have detrimental effects on an animal's ability to engage with its owners and environment or to respond to training or novel situations, which may in turn affect its quality of life. This study compared the performance of 20 dogs with chronic pain from osteoarthritis and 21 healthy control dogs in a disappearing object task of spatial working memory. Female neutered osteoarthritic dogs, but not male neutered osteoarthritic dogs, were found to have lower predicted probabilities of successfully performing the task compared to control dogs of the same sex. In addition, as memory retention interval in the task increased, osteoarthritic dogs showed a steeper decline in working memory performance than control dogs. This suggests that the effects of osteoarthritis, and potentially other pain-related conditions, on cognitive function are more clearly revealed in tasks that present a greater cognitive load. Our finding that chronic pain from osteoarthritis may be associated with impaired working memory in dogs parallels results from studies of human chronic pain disorders. That female dogs may be particularly prone to these effects warrants further investigation.

Using the conditioned place preference paradigm to assess hunger in dairy calves: Preliminary results and methodological issues.

Dairy calves are typically fed restricted amounts of milk. Although feed restrictions are predicted to result in negative affective states, the relative aversiveness of 'hunger' remains largely unexplored in this species. Here, we investigated whether the conditioned place preference paradigm can be used to explore how calves feel when experiencing different levels of satiation. This paradigm provides insight into what animals remember from past experiences, the assumption being that individuals will prefer places associated with more pleasant or less unpleasant experiences. Sixteen Holstein calves were either fed a restricted (3 L per meal totalling 6 L per day) or 'enhanced' milk allowance (ad libitum up to 6 L per meal totalling up to 12 L per day) in their home-pen. Calves were then placed in a conditioning pen for 4 h immediately after being fed their morning meal to allow them to develop an association between the pen and their state of post-prandial satiation. Calves were conditioned across four days with their satiation state alternating between days to allow them to develop an association between pen and satiation levels. On the 5th day, calves were individually allowed to roam freely between the two pens for 30 min. We expected that calves would prefer the pen where they previously experienced higher levels of satiation, but our results show no to limited effects of treatment. However, some methodological issues (colour and side bias) prevent us from drawing strong conclusions. We discuss reasons for these issues and potential solutions to avoid these in future studies.

Farming non-typical sentient species: ethical framework requires passing a high bar.

More widespread farming of species not typically used as livestock may be part of a sustainable approach for promoting human health and economic prosperity in a world with an increasing population; a current example is peccary farming in the Neotropics. Others have argued that species that are local to a region and which are usually not farmed should be considered for use as livestock. They may have a more desirable nutrient profile than species that are presently used as livestock. It may also reduce the pressure from hunting on other wild species, and cause less environmental damage than exotic species. We propose a sentiocentric utilitarian framework that could be used to decide whether species that are local, but generally not used as livestock, should be farmed. To illustrate the use of our decision-making framework, we employ two contrasting neotropical case studies: the Spotted Paca (Cuniculus paca) and the Capybara (Hydrochoerus hydrochaeris). We argue that it may be acceptable to use non-sentient species that are typically not farmed as livestock. However, research should determine whether farming them offers human, environmental or sustainability benefits. In addition, we recommend that if invertebrate species are considered for farming, research should be conducted to determine the likelihood that they are sentient. Finally, given the ethical failings of current livestock farming practices, we argue that a high bar must be met if 'new' species that are sentient are to be farmed.

A primer on the use of computational modelling to investigate affective states, affective disorders and animal welfare in non-human animals.

Objective measures of animal emotion-like and mood-like states are essential for preclinical studies of affective disorders and for assessing the welfare of laboratory and other animals. However, the development and validation of measures of these affective states poses a challenge partly because the relationships between affect and its behavioural, physiological and cognitive signatures are complex. Here, we suggest that the crisp characterisations offered by computational modelling of the underlying, but unobservable, processes that mediate these signatures should provide better insights. Although this computational psychiatry approach has been widely used in human research in both health and disease, translational computational psychiatry studies remain few and far between. We explain how building computational models with data from animal studies could play a pivotal role in furthering our understanding of the aetiology of affective disorders, associated affective states and the likely underlying cognitive processes involved. We end by outlining the basic steps involved in a simple computational analysis.

Do you see what I see? Testing horses' ability to recognise real-life objects from 2D computer projections.

The use of 2-dimensional representations (e.g. photographs or digital images) of real-life physical objects has been an important tool in studies of animal cognition. Horses are reported to recognise objects and individuals (conspecifics and humans) from printed photographs, but it is unclear whether image recognition is also true for digital images, e.g. computer projections. We expected that horses trained to discriminate between two real-life objects would show the same learnt response to digital images of these objects indicating that the images were perceived as objects, or representations of such. Riding-school horses (N = 27) learnt to touch one of two objects (target object counterbalanced between horses) to instantly receive a food reward. After discrimination learning (three consecutive sessions of 8/10 correct trials), horses were immediately tested with on-screen images of the objects over 10 image trials interspersed with five real object trials. At first image presentation, all but two horses spontaneously responded to the images with the learnt behaviour by contacting one of the two images, but the number of horses touching the correct image was not different from chance (14/27 horses, p > 0.05). Only one horse touched the correct image above chance level across 10 image trials (9/10 correct responses, p = 0.021). Our findings thus question whether horses recognise real-life objects from digital images. We discuss how methodological factors and individual differences (i.e. age, welfare state) might have influenced animals' response to the images, and the importance of validating the suitability of stimuli of this kind for cognitive studies in horses.

Validation of complementary non-invasive tools for stress assessment in spotted paca (Cuniculus paca).

Monitoring the concentration of glucocorticoid metabolites (GCMs) in faecal samples is a non-invasive tool for physiological stress evaluation, particularly useful when studying wild species. However, both negative and positive stimuli (distress and eustress, respectively) can lead to a rise in glucocorticoids. Thus, besides validating whether GCM concentration in faeces reflects endogenous adrenal activity, we also need to identify behavioural indicators of distress to avoid misinterpretation. Therefore, we submitted four adult male spotted pacas (Cuniculus paca) to an exogenous adrenocorticotropic hormone (ACTH) challenge-test in a Latin square design (4 × 4) to monitor changes in the GCM concentration in faeces. We also aimed to describe behaviours potentially indicative of distress. We collected excreted faeces and video-recorded the animals' behaviours for five consecutive days, one day before and four days after application of the following four treatments: 1st control (no-handling); 2nd control (intra-muscular [IM] injection of saline solution); low-dose ACTH (IM injection of 0.18 ml ACTH); and high-dose ACTH (IM injection of 0.37 ml ACTH). There was a peak in the concentration of GCM in faeces collected 24 h after the injection of the high-dose ACTH treatment. Additionally, independent of the treatments, spotted pacas spent less time on exploration and feeding states, while spending more time in the inactive but awake (IBA) state following the treatment application (challenge day). The use of GCM concentration in faecal samples together with the behavioural changes (less exploration and feeding, and more IBA) showed to be efficient as a non-invasive tool for welfare assessment of farmed spotted paca.

Dissecting the links between reward and loss, decision-making, and self-reported affect using a computational approach.

Links between affective states and risk-taking are often characterised using summary statistics from serial decision-making tasks. However, our understanding of these links, and the utility of decision-making as a marker of affect, needs to accommodate the fact that ongoing (e.g., within-task) experience of rewarding and punishing decision outcomes may alter future decisions and affective states. To date, the interplay between affect, ongoing reward and punisher experience, and decision-making has received little detailed investigation. Here, we examined the relationships between reward and loss experience, affect, and decision-making in humans using a novel judgement bias task analysed with a novel computational model. We demonstrated the influence of within-task favourability on decision-making, with more risk-averse/'pessimistic' decisions following more positive previous outcomes and a greater current average earning rate. Additionally, individuals reporting more negative affect tended to exhibit greater risk-seeking decision-making, and, based on our model, estimated time more poorly. We also found that individuals reported more positive affective valence during periods of the task when prediction errors and offered decision outcomes were more positive. Our results thus provide new evidence that (short-term) within-task rewarding and punishing experiences determine both future decision-making and subjectively experienced affective states.

Haematology, physiological parameters, morphometry and parasitological status of rescued bearded capuchin monkeys (Sapajus libidinosus).

BACKGROUND: There are few studies on the physiology and haematology of rescued bearded capuchin monkeys. These are necessary to better understand the health and welfare status of the animals, including when performing reintroductions, and to avoid zoonoses. METHODS: We aimed to obtain physiological and haematological values, morphometry and parasitological status of 26 bearded capuchins in two rescue centres in Northeast Brazil. RESULTS: We found sex- and age-related differences in respiratory rate, body weight and body mass index, but not haematology. The haematological values obtained were significantly different from published data but within the reference intervals for the species. Animals infected with Ancylostoma spp. had significantly lower values in the parameters: haemoglobin concentration, mean corpuscular volume and mean corpuscular haemoglobin concentration than non-infected individuals. CONCLUSION: Physiological and haematological values of rescued capuchins were similar to those found in previous studies. Ancylostomiasis appears to cause alterations to haematological values of this species.

Using Primary Reinforcement to Enhance Translatability of a Human Affect and Decision-Making Judgment Bias Task.

Good translatability of behavioral measures of affect (emotion) between human and nonhuman animals is core to comparative studies. The judgment bias (JB) task, which measures "optimistic" and "pessimistic" decision-making under ambiguity as indicators of positive and negative affective valence, has been used in both human and nonhuman animals. However, one key disparity between human and nonhuman studies is that the former typically use secondary reinforcers (e.g., money) whereas the latter typically use primary reinforcers (e.g., food). To address this deficiency and shed further light on JB as a measure of affect, we developed a novel version of a JB task for humans using primary reinforcers. Data on decision-making and reported affective state during the JB task were analyzed using computational modeling. Overall, participants grasped the task well, and as anticipated, their reported affective valence correlated with trial-by-trial variation in offered volume of juice. In addition, previous findings from monetary versions of the task were replicated: More positive prediction errors were associated with more positive affective valence, a higher lapse rate was associated with lower affective arousal, and affective arousal decreased as a function of number of trials completed. There was no evidence that more positive valence was associated with greater "optimism," but instead, there was evidence that affective valence influenced the participants' decision stochasticity, whereas affective arousal tended to influence their propensity for errors. This novel version of the JB task provides a useful tool for investigation of the links between primary reward and punisher experience, affect, and decision-making, especially from a comparative perspective.

Nature calls: intelligence and natural foraging style predict poor welfare in captive parrots.

Understanding why some species thrive in captivity, while others struggle to adjust, can suggest new ways to improve animal care. Approximately half of all Psittaciformes, a highly threatened order, live in zoos, breeding centres and private homes. Here, some species are prone to behavioural and reproductive problems that raise conservation and ethical concerns. To identify risk factors, we analysed data on hatching rates in breeding centres (115 species, 10 255 pairs) and stereotypic behaviour (SB) in private homes (50 species, 1378 individuals), using phylogenetic comparative methods (PCMs). Small captive population sizes predicted low hatch rates, potentially due to genetic bottlenecks, inbreeding and low availability of compatible mates. Species naturally reliant on diets requiring substantial handling were most prone to feather-damaging behaviours (e.g. self-plucking), indicating inadequacies in the composition or presentation of feed (often highly processed). Parrot species with relatively large brains were most prone to oral and whole-body SB: the first empirical evidence that intelligence can confer poor captive welfare. Together, results suggest that more naturalistic diets would improve welfare, and that intelligent psittacines need increased cognitive stimulation. These findings should help improve captive parrot care and inspire further PCM research to understand species differences in responses to captivity.

Emotion in animal contests.

Emotions encompass cognitive and behavioural responses to reward and punishment. Using contests as a case-study, we propose that short-term emotions underpin animals' assessments, decision-making and behaviour. Equating contest assessments to emotional 'appraisals', we describe how contestants appraise more than resource value and outcome probability. These appraisals elicit the cognition, drive and neurophysiology that governs aggressive behaviour. We discuss how recent contest outcomes induce long-term moods, which impact subsequent contest behaviour. Finally, we distinguish between integral (objectively relevant) and incidental (objectively irrelevant) emotions and moods (affective states). Unlike existing ecological models, our approach predicts that incidental events influence contest dynamics, and that contests become incidental influences themselves, potentially causing maladaptive decision-making. As affective states cross contexts, a more holistic ethology (incorporating emotions and moods) would illuminate animal cognition and behaviour.

Insensitivity to reward shifts in zebrafish (Danio rerio) and implications for assessing affective states.

Theory and empirical findings predict that individuals in a negative affective state are more sensitive to unexpected reward loss and less sensitive to unexpected reward gain compared to individuals in a neutral or positive affective state. We explore the use of sensitivity to reward shifts measured during successive contrast tasks as an indicator of affect in zebrafish (Danio rerio). In line with the assumption that exposure to rewarding stimuli induces a relatively positive affective state compared to exposure to stimuli that they do not prefer, we confirmed that zebrafish prefer enriched over barren environments, suggesting that the enriched environment is associated with positive affective states. We trained individuals to swim down a channel for food rewards of differing value and then presented them with unexpected increases or decreases in reward value. Contrary to our hypothesis, individuals conditioned to a high-value reward continued swimming at the same speed when reward value was downshifted, thus showing no successive negative contrast effect and appearing insensitive to reward loss. Individuals whose rewards were upshifted gradually increased their speed, but did not display successive positive contrast effects typical of sensitivity to reward gains. In both cases, housing type did not result in differences in swim time. One potential explanation is that goal-directed control of behaviour is necessary for an animal to show a successive contrast response to unexpected reward gain or loss, and the behaviour of zebrafish in this task was under habitual control, perhaps due to over-training. If so, refinements to task design and training procedures will allow further progress with this assay.

Individual behavioral differences and health of golden-headed lion tamarins (Leontopithecus chrysomelas).

Individual behavioral differences may influence how animals cope with altered environments. Depending on their behavioral traits, individuals may thus vary in how their health is affected by environmental conditions. We investigated the relationship between individual behavior of free-living golden-headed lion tamarins (Leontopithecus chrysomelas) responding to a novel object (to assess exploration-avoidance), and their habitat use and health status (endoparasitism; clinical measures: biometric data, heart rate, respiratory frequency, and temperature; fecal glucocorticoid metabolites). As parasite transmission can be affected by individual variation in social contact and social grooming, we also evaluated whether more sociable individuals show higher endoparasite loads compared with less sociable animals. Four groups living in landscapes with different levels of human disturbance were investigated: two in degraded forest fragments in an agricultural matrix (DFAM-higher disturbance), and two in a cocoa agroforestry system (cabruca-lower disturbance) in the Atlantic forest of South Bahia, Brazil. Using a subjective ratings approach, highly correlated adjective descriptors were combined to produce z-score ratings of one derived variable ("confidence"), which was selected to characterize the tamarins' exploration/avoidance responses during a novel object test. The higher the confidence score, the longer female tamarins spent foraging for prey independent of landscape, and the greater their body mass independent of sex and landscape. Only DFAM individuals showed intestinal parasite infection. Endoparasite loads were positively correlated with the number of grooming partners, suggesting an association between social grooming and transmission (more groomers = more endoparasites). Individual behavior, including in a test situation, may thus have some predictive value for behavior in a free-living context, and for its health consequences.

State-dependent judgement bias in Drosophila: evidence for evolutionarily primitive affective processes.

Affective states influence decision-making under ambiguity in humans and other animals. Individuals in a negative state tend to interpret ambiguous cues more negatively than individuals in a positive state. We demonstrate that the fruit fly, Drosophila melanogaster, also exhibits state-dependent changes in cue interpretation. Drosophila were trained on a Go/Go task to approach a positive (P) odour associated with a sugar reward and actively avoid a negative (N) odour associated with shock. Trained flies were then either shaken to induce a purported negative state or left undisturbed (control), and given a choice between: air or P; air or N; air or ambiguous odour (1 : 1 blend of P : N). Shaken flies were significantly less likely to approach the ambiguous odour than control flies. This 'judgement bias' may be mediated by changes in neural activity that reflect evolutionarily primitive affective states. We cannot say whether such states are consciously experienced, but use of this model organism's versatile experimental tool kit may facilitate elucidation of their neural and genetic basis.

Animal emotion: Descriptive and prescriptive definitions and their implications for a comparative perspective.

In recent years there has been a growing research interest in the field of animal emotion. But there is still little agreement about whether and how the word "emotion" should be defined for use in the context of non-human species. Here, we make a distinction between descriptive and prescriptive definitions. Descriptive definitions delineate the ways in which the word emotion is used in everyday life. Prescriptive definitions are used to pick out the set of events that scientific theories of emotion purport to explain. Picking out three prescriptive definitions, we show that the different ways in which emotions are defined correspond to processes that are distributed differentially across the animal kingdom. We propose that these definitions provide a useful starting point for investigating the varying emotional capacities of a wide range of animals, providing a basis for a new, comparative science of emotion.

Long-term and trans-generational effects of neonatal experience on sheep behaviour.

Early life experiences can have profound long-term, and sometimes transgenerational, effects on individual phenotypes. However, there is a relative paucity of knowledge about effects on pain sensitivity, even though these may impact on an individual's health and welfare, particularly in farm animals exposed to painful husbandry procedures. Here, we tested in sheep whether neonatal painful and non-painful challenges can alter pain sensitivity in adult life, and also in the next generation. Ewes exposed to tail-docking or a simulated mild infection (lipopolysaccharide (LPS)) on days 3­4 of life showed higher levels of pain-related behaviour when giving birth as adults compared with control animals. LPS-treated ewes also gave birth to lambs who showed decreased pain sensitivity in standardized tests during days 2­3 of life. Our results demonstrate long-term and trans-generational effects of neonatal experience on pain responses in a commercially important species and suggest that variations in early life management can have important implications for animal health and welfare.

Can male Eurasian jays disengage from their own current desire to feed the female what she wants?

Humans' predictions of another person's behaviour are regularly influenced by what they themselves might know or want. In a previous study, we found that male Eurasian jays (Garrulus glandarius) could cater for their female partner's current desire when sharing food with her. Here, we tested the extent to which the males' decisions are influenced by their own current desire. When the males' and female's desires matched, males correctly shared the food that was desired by both. When the female's desire differed from their own, the males' decisions were not entirely driven by their own desires, suggesting that males also took the female's desire into account. Thus, the male jays' decisions about their mates' desires are partially biased by their own desire and might be based upon similar processes as those found in humans.

Exploring the similarities between risk factors triggering depression in humans and elevated in-cage "inactive but awake" behavior in laboratory mice.

Introduction: Depression is a human mental disorder that can also be inferred in non-human animals. This study explored whether time spent inactive but awake ("IBA") in the home-cage in mice was further triggered by risk factors similar to those increasing vulnerability to depression in humans (early life stress, genetic predispositions, adulthood stress). Methods: Eighteen DBA/2 J and 18 C57BL/6 J females were tested, of which half underwent as pups a daily maternal separation on post-natal days 2-14 (early-life stress "ELS") (other half left undisturbed). To assess the effect of the procedure, the time the dams from which the 18 subjects were born spent active in the nest (proxy for maternal behavior) was recorded on post-natal days 2, 6, 10 and 14 for 1 h before separation and following reunion (matched times for controls), using live instantaneous scan sampling (total: 96 scans/dam). For each ELS condition, about half of the pups were housed post-weaning (i.e., from 27 days old on average) in either barren (triggering IBA and depression-like symptoms) or larger, highly enriched cages (n = 4-5 per group). Time mice spent IBA post-weaning was observed blind to ELS treatment using live instantaneous scan sampling in two daily 90-min blocks, two days/week, for 6 weeks (total: 192 scans/mouse). Data were analyzed in R using generalized linear mixed models. Results: The dams were significantly more active in the nest over time (p = 0.016), however with no significant difference between strains (p = 0.18), ELS conditions (p = 0.20) and before/after separation (p = 0.83). As predicted, post-weaning barren cages triggered significantly more time spent IBA in mice than enriched cages (p < 0.0001). However, neither ELS (p = 0.4) nor strain (p = 0.84) significantly influenced time mice spent IBA, with no significant interaction with environmental condition (ELS × environment: p = 0.2861; strain × environment: p = 0.5713). Discussion: Our results therefore only partly support the hypothesis that greater time spent IBA in mice is triggered by risk factors for human depression. We discuss possible explanations for this and further research directions.

Developing guidelines for pet rat housing through expert consultation.

BACKGROUND: Pet care guidelines play an important role in ensuring that owners are well informed about good husbandry practices, allowing them to provide the best care for their animals. However, the development of such guidelines is difficult when there is little appropriate empirical evidence on which to base guidelines, as in the case of pet rats. The consultation of multiple experts can help to surmount this challenge. METHODS: We developed a set of guidelines for pet rat housing by consulting with a group of experts, including veterinarians, veterinary nurses, animal welfare scientists and experienced pet rat owners. The consultation involved two rounds of online surveys (n = 13) and one online discussion (n = 8). RESULTS: The resulting guidelines cover a broad range of features within pet rat housing, including injury prevention, details of suitable refuges and substrates, and suitable cage sizing. The guidelines may evolve as more information about pet rats comes to light but may nonetheless provide a useful starting point for any future guidelines. CONCLUSIONS: At present, these guidelines may not only be useful for pet rat owners and those advising pet rat owners, such as veterinarians, but may also be useful in the design of housing, including for laboratory rodents.

A mapping review of refinements to laboratory rat housing and husbandry.

Refining the housing and husbandry of laboratory rats is an important goal, both for ethical reasons and to allow better quality research. We conducted a mapping review of 1,017 studies investigating potential refinements of housing and husbandry of the laboratory rat to assess what refinements have, and have not, been studied, and to briefly assess whether there is evidence to support any impact on rat welfare. Among the many refinements studied, the majority involve changes to the cage, but some also involve alterations to the wider environment. The effects of these refinements were assessed using a range of readouts, many of which are difficult to interpret from a welfare perspective. Preference studies, which are easier to interpret, provide evidence that rats prefer complex environments, including shelters and multiple objects, which offer different areas/resources allowing the rat to engage in diverse behaviors. The reporting of methodology in papers was often poor, indicating that studies were potentially subject to biases. Given that many refinements co-occurred, it was often difficult to tease apart which ones were most beneficial for rat welfare. Effects of refinements were also moderated by a number of factors including age, sex, strain and photoperiod. Altogether our findings show that a one-size-fits-all approach to refinements is not appropriate, because different refinements will impact different rats in different ways. Our review has also produced a database of >1,000 articles that can be used for further and more detailed analyses. Our findings have also highlighted areas where future research is likely to be valuable, including refinements to rat transport, handling and the use of training.