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Gácsi, M., Gyoöri, B., Virányi, Z., Kubinyi, E., Range, F., Belényi, B., et al. (2009). Explaining Dog Wolf Differences in Utilizing Human Pointing Gestures: Selection for Synergistic Shifts in the Development of Some Social Skills. PLoS ONE, 4(8), e6584.
Abstract: <sec> <title>Background</title> <p>The comparison of human related communication skills of socialized canids may help to understand the evolution and the epigenesis of gesture comprehension in humans. To reconcile previously contradicting views on the origin of dogs' outstanding performance in utilizing human gestures, we suggest that dog-wolf differences should be studied in a more complex way.</p> </sec><sec> <title>Methodology/Principal Findings</title> <p>We present data both on the performance and the behaviour of dogs and wolves of different ages in a two-way object choice test. Characteristic behavioural differences showed that for wolves it took longer to establish eye contact with the pointing experimenter, they struggled more with the handler, and pups also bit her more before focusing on the human's signal. The performance of similarly hand-reared 8-week-old dogs and wolves did not differ in utilizing the simpler proximal momentary pointing. However, when tested with the distal momentary pointing, 4-month-old pet dogs outperformed the same aged hand reared wolves. Thus early and intensive socialisation does not diminish differences between young dogs and wolves in behaviour and performance. Socialised adult wolves performed similarly well as dogs in this task without pretraining. The success of adult wolves was accompanied with increased willingness to cooperate.</p> </sec><sec> <title>Conclusion/Significance</title> <p>Thus, we provide evidence for the first time that socialised adult wolves are as successful in relying on distal momentary pointing as adult pet dogs. However, the delayed emergence of utilising human distal momentary pointing in wolves shows that these wild canines react to a lesser degree to intensive socialisation in contrast to dogs, which are able to control agonistic behaviours and inhibition of actions in a food related task early in development. We suggest a “synergistic” hypothesis, claiming that positive feedback processes (both evolutionary and epigenetic) have increased the readiness of dogs to attend to humans, providing the basis for dog-human communication.</p> </sec>
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Warmuth, V., Eriksson, A., Bower, M. A., Cañon, J., Cothran, G., Distl, O., et al. (2011). European Domestic Horses Originated in Two Holocene Refugia. PLoS ONE, 6(3), e18194 EP -.
Abstract: <p>The role of European wild horses in horse domestication is poorly understood. While the fossil record for wild horses in Europe prior to horse domestication is scarce, there have been suggestions that wild populations from various European regions might have contributed to the gene pool of domestic horses. To distinguish between regions where domestic populations are mainly descended from local wild stock and those where horses were largely imported, we investigated patterns of genetic diversity in 24 European horse breeds typed at 12 microsatellite loci. The distribution of high levels of genetic diversity in Europe coincides with the distribution of predominantly open landscapes prior to domestication, as suggested by simulation-based vegetation reconstructions, with breeds from Iberia and the Caspian Sea region having significantly higher genetic diversity than breeds from central Europe and the UK, which were largely forested at the time the first domestic horses appear there. Our results suggest that not only the Eastern steppes, but also the Iberian Peninsula provided refugia for wild horses in the Holocene, and that the genetic contribution of these wild populations to local domestic stock may have been considerable. In contrast, the consistently low levels of diversity in central Europe and the UK suggest that domestic horses in these regions largely derive from horses that were imported from the Eastern refugium, the Iberian refugium, or both.</p>
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Wathan, J., Burrows, A. M., Waller, B. M., & McComb, K. (2015). EquiFACS: The Equine Facial Action Coding System. PLoS ONE, 10(8), e0131738.
Abstract: <p>Although previous studies of horses have investigated their facial expressions in specific contexts, e.g. pain, until now there has been no methodology available that documents all the possible facial movements of the horse and provides a way to record all potential facial configurations. This is essential for an objective description of horse facial expressions across a range of contexts that reflect different emotional states. Facial Action Coding Systems (FACS) provide a systematic methodology of identifying and coding facial expressions on the basis of underlying facial musculature and muscle movement. FACS are anatomically based and document all possible facial movements rather than a configuration of movements associated with a particular situation. Consequently, FACS can be applied as a tool for a wide range of research questions. We developed FACS for the domestic horse (<italic>Equus caballus</italic>) through anatomical investigation of the underlying musculature and subsequent analysis of naturally occurring behaviour captured on high quality video. Discrete facial movements were identified and described in terms of the underlying muscle contractions, in correspondence with previous FACS systems. The reliability of others to be able to learn this system (EquiFACS) and consistently code behavioural sequences was high?and this included people with no previous experience of horses. A wide range of facial movements were identified, including many that are also seen in primates and other domestic animals (dogs and cats). EquiFACS provides a method that can now be used to document the facial movements associated with different social contexts and thus to address questions relevant to understanding social cognition and comparative psychology, as well as informing current veterinary and animal welfare practices.</p>
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Meagher, R. K., & Mason, G. J. (2012). Environmental Enrichment Reduces Signs of Boredom in Caged Mink. PLoS ONE, 7(11), e49180.
Abstract: <p>Animals housed in impoverished cages are often labelled ‘bored’. They have also been called ‘apathetic’ or ‘depressed’, particularly when profoundly inactive. However, these terms are rarely operationally defined and validated. As a negative state caused by under-stimulation, boredom should increase interest in stimuli of all kinds. Apathy (lack of interest), by contrast, should manifest as decreased interest in all stimuli, while anhedonia (loss of pleasure, a depressive symptom) should specifically decrease interest in normally rewarding stimuli. We tested the hypotheses that mink, a model carnivore, experience more boredom, depression-like apathy, or anhedonia in non-enriched (NE) cages than in complex, enriched (E) cages. We exposed 29 subjects (13 E, 16 NE) to ten stimuli categorized <italic>a priori</italic> as aversive (e.g. air puffs), rewarding (e.g. evoking chasing) or ambiguous/neutral (e.g. candles). Interest in stimuli was assessed via latencies to contact, contact durations, and durations oriented to stimuli. NE mink contacted all stimuli faster (P = 0.003) than E mink, and spent longer oriented to/in contact with them, albeit only significantly so for ambiguous ones (treatment*type P<0.013). With stimulus category removed from statistical models, interest in all stimuli was consistently higher among NE mink (P<0.0001 for all measures). NE mink also consumed more food rewards (P = 0.037). Finally, we investigated whether lying down while awake and stereotypic behaviour (both increased by NE housing) predicted these responses. Lying awake positively co-varied with certain measures of increased exploration. In contrast, stereotypic ‘scrabbling’ or locomotion (e.g. pacing) did not. Overall, NE mink showed no evidence of apathy or depression, but instead a heightened investigation of diverse stimuli consistent with boredom. This state was potentially indicated by spending much time lying still but awake (although this result requires replication). Boredom can thus be operationalized and assessed empirically in non-human animals. It can also be reduced by environmental enrichment.</p>
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Peugnet, P., Wimel, L., Duchamp, G., Sandersen, C., Camous, S., Guillaume, D., et al. (2014). Enhanced or Reduced Fetal Growth Induced by Embryo Transfer into Smaller or Larger Breeds Alters Post-Natal Growth and Metabolism in Pre-Weaning Horses. PLoS ONE, 9(7), e102044 EP -.
Abstract: <p>In equids, placentation is diffuse and nutrient supply to the fetus is determined by uterine size. This correlates with maternal size and affects intra-uterine development and subsequent post-natal growth, as well as insulin sensitivity in the newborn. Long-term effects remain to be described. In this study, fetal growth was enhanced or restricted through ET using pony (P), saddlebred (S) and draft (D) horses. Control P-P (n = 21) and S-S (n = 28) pregnancies were obtained by AI. Enhanced and restricted pregnancies were obtained by transferring P or S embryos into D mares (P-D, n = 6 and S-D, n = 8) or S embryos into P mares (S-P, n = 6), respectively. Control and experimental foals were raised by their dams and recipient mothers, respectively. Weight gain, growth hormones and glucose homeostasis were investigated in the foals from birth to weaning. Fetal growth was enhanced in P-D and these foals remained consistently heavier, with reduced T 3 concentrations until weaning compared to P-P. P-D had lower fasting glucose from days 30 to 200 and higher insulin secretion than P-P after IVGTT on day 3. Euglycemic clamps in the immediate post-weaning period revealed no difference in insulin sensitivity between P-D and P-P. Fetal growth was restricted in S-P and these foals remained consistently lighter until weaning compared to S-D, with elevated T 3 concentrations in the newborn compared to S-S. S-P exhibited higher fasting glycemia than S-S and S-D from days 30 to 200. They had higher maximum increment in plasma glucose than S-D after IVGTT on day 3 and clamps on day 200 demonstrated higher insulin sensitivity compared to S-D. Neither the restricted nor the enhanced fetal environment affected IGF-1 concentrations. Thus, enhanced and restricted fetal and post-natal environments had combined effects that persisted until weaning. They induced different adaptive responses in post-natal glucose metabolism: an early insulin-resistance was induced in enhanced P-D, while S-P developed increased insulin sensitivity.</p>
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Freidin, E., Putrino, N., D’Orazio, M., & Bentosela, M. (2013). Dogs’ Eavesdropping from People’s Reactions in Third Party Interactions. PLoS ONE, 8(11), e79198 EP -.
Abstract: <p>Eavesdropping involves the acquisition of information from third-party interactions, and can serve to indirectly attribute reputation to individuals. There is evidence on eavesdropping in dogs, indicating that they can develop a preference for people based on their cooperativeness towards others. In this study, we tested dogs’ eavesdropping abilities one step further. In a first experiment, dogs could choose between cooperative demonstrators (the donors) who always gave food to an approaching third person (the beggar); here, the only difference between donors was whether they received positive or negative reactions from the beggar (through verbal and gestural means). Results showed that dogs preferentially approached the donor who had received positive reactions from the beggar. By contrast, two different conditions showed that neither the beggar’s body gestures nor the verbal component of the interaction on their own were sufficient to affect the dogs’ preferences. We also ran two further experiments to test for the possibility of dogs’ choices being driven by local enhancement. When the donors switched places before the choice, dogs chose at random. Similarly, in a nonsocial condition in which donors were replaced by platforms, subjects chose at chance levels. We conclude that dogs’ nonrandom choices in the present protocol relied on the simultaneous presence of multiple cues, such as the place where donors stood and several features of the beggar’s behavior (gestural and verbal reactions, and eating behavior). Nonetheless, we did not find conclusive evidence that dogs discriminated the donors by their physical features, which is a prerequisite of reputation attribution.</p>
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Fraser, O. N., & Bugnyar, T. (2010). Do Ravens Show Consolation? Responses to Distressed Others. PLoS ONE, 5(5), e10605.
Abstract: <sec> <title>Background</title> <p>Bystander affiliation (post-conflict affiliation from an uninvolved bystander to the conflict victim) may represent an expression of empathy in which the bystander consoles the victim to alleviate the victim's distress (“consolation”). However, alternative hypotheses for the function of bystander affiliation also exist. Determining whether ravens spontaneously offer consolation to distressed partners may not only help us to understand how animals deal with the costs of aggressive conflict, but may also play an important role in the empathy debate.</p> </sec><sec> <title>Methodology/Principal findings</title> <p>This study investigates the post-conflict behavior of ravens, applying the predictive framework for the function of bystander affiliation for the first time in a non-ape species. We found weak evidence for reconciliation (post-conflict affiliation between former opponents), but strong evidence for both bystander affiliation and solicited bystander affiliation (post-conflict affiliation from the victim to a bystander). Bystanders involved in both interactions were likely to share a valuable relationship with the victim. Bystander affiliation offered to the victim was more likely to occur after intense conflicts. Renewed aggression was less likely to occur after the victim solicited affiliation from a bystander.</p> </sec><sec> <title>Conclusions/Significance</title> <p>Our findings suggest that in ravens, bystanders may console victims with whom they share a valuable relationship, thus alleviating the victims' post-conflict distress. Conversely victims may affiliate with bystanders after a conflict in order to reduce the likelihood of renewed aggression. These results stress the importance of relationship quality in determining the occurrence and function of post-conflict interactions, and show that ravens may be sensitive to the emotions of others.</p> </sec>
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Sankey, C., Henry, S., André, N., Richard-Yris, M. - A., & Hausberger, M. (2011). Do Horses Have a Concept of Person? PLoS ONE, 6(3), e18331 EP -.
Abstract: <sec> <title>Background</title> <p>Animals' ability for cross-modal recognition has recently received much interest. Captive or domestic animals seem able to perceive cues of human attention and appear to have a multisensory perception of humans.</p> </sec> <sec> <title>Methodology/Principal Findings</title> <p>Here, we used a task where horses have to remain immobile under a vocal order to test whether they are sensitive to the attentional state of the experimenter, but also whether they behave and respond differently to the familiar order when tested by a familiar or an unknown person. Horses' response varied according to the person's attentional state when the order was given by an unknown person: obedience levels were higher when the person giving the order was looking at the horse than when he was not attentive. More interesting is the finding that whatever the condition, horses monitored much more and for longer times the unknown person, as if they were surprised to hear the familiar order given by an unknown voice.</p> </sec> <sec> <title>Conclusion/Significance</title> <p>These results suggest that recognition of humans may lie in a global, integrated, multisensory representation of specific individuals, that includes visual and vocal identity, but also expectations on the individual's behaviour in a familiar situation.</p> </sec>
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Dalla Costa, E., Minero, M., Lebelt, D., Stucke, D., Canali, E., & Leach, M. C. (2014). Development of the Horse Grimace Scale (HGS) as a Pain Assessment Tool in Horses Undergoing Routine Castration. PLoS ONE, 9(3), e92281 EP -.
Abstract: <sec><title>Background</title><p>The assessment of pain is critical for the welfare of horses, in particular when pain is induced by common management procedures such as castration. Existing pain assessment methods have several limitations, which reduce the applicability in everyday life. Assessment of facial expression changes, as a novel means of pain scoring, may offer numerous advantages and overcome some of these limitations. The objective of this study was to develop and validate a standardised pain scale based on facial expressions in horses (Horse Grimace Scale [HGS]).</p></sec><sec><title>Methodology/Principal Findings</title><p>Forty stallions were assigned to one of two treatments and all animals underwent routine surgical castration under general anaesthesia. Group A (n = 19) received a single injection of Flunixin immediately before anaesthesia. Group B (n = 21) received Flunixin immediately before anaesthesia and then again, as an oral administration, six hours after the surgery. In addition, six horses were used as anaesthesia controls (C). These animals underwent non-invasive, indolent procedures, received the same treatment as group A, but did not undergo surgical procedures that could be accompanied with surgical pain. Changes in behaviour, composite pain scale (CPS) scores and horse grimace scale (HGS) scores were assessed before and 8-hours post-procedure. Only horses undergoing castration (Groups A and B) showed significantly greater HGS and CPS scores at 8-hours post compared to pre operatively. Further, maintenance behaviours such as explorative behaviour and alertness were also reduced. No difference was observed between the two analgesic treatment groups.</p></sec><sec><title>Conclusions</title><p>The Horse Grimace Scale potentially offers an effective and reliable method of assessing pain following routine castration in horses. However, auxiliary studies are required to evaluate different painful conditions and analgesic schedules.</p></sec>
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Hausberger, M., Gautier, E., Biquand, V., Lunel, C., & Jégo, P. (2009). Could Work Be a Source of Behavioural Disorders? A Study in Horses. PLoS ONE, 4(10), e7625 EP -.
Abstract: <p>Stress at work, as shown by a number of human studies, may lead to a variety of negative and durable effects, such as impaired psychological functioning (anxiety, depression…). Horses share with humans this characteristic of working on a daily basis and are submitted then to work stressors related to physical constraints and/or more “psychological” conflicts, such as potential controversial orders from the riders or the requirement to suppress emotions. On another hand, horses may perform abnormal repetitive behaviour (“stereotypies”) in response to adverse life conditions. In the present study, we investigated whether the type of work the horses are used for may have an impact on their tendency to show stereotypic behaviour (and its type) outside work. Observations in their box of 76 horses all living in the same conditions, belonging to one breed and one sex, revealed that the prevalence and types of stereotypies performed strongly depended upon the type of work they were used for. The stereotypies observed involved mostly mouth movements and head tossing/nodding. Work constraints probably added to unfavourable living conditions, favouring the emergence of chronic abnormal behaviours. This is especially remarkable as the 23 hours spent in the box were influenced by the one hour work performed every day. To our knowledge, this is the first evidence of potential effects of work stressors on the emergence of abnormal behaviours in an animal species. It raises an important line of thought on the chronic impact of the work situation on the daily life of individuals.</p>
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