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Goursot, C., Düpjan, S., Puppe, B., & Leliveld, L. M. C. (2021). Affective styles and emotional lateralization: A promising framework for animal welfare research. Appl. Anim. Behav. Sci., 237, 105279.
Abstract: The growing recognition of animals as individuals has broader implications for farm animal welfare research. Even under highly standardized on-farm conditions, farm animals show heterogeneous but individually consistent behavioural patterns towards various stimuli, based on how they appraise these stimuli. As a result, animal welfare is likely to be highly individual as well, and studying the proximate mechanisms underlying distinct individual behaviour patterns and appraisal will improve animal welfare research. We propose to extend the framework of affective styles to bridge the gap between existing research fields on animal personality and affective states. Affective styles refer to consistent individual differences in emotional reactivity and regulation and can be predicted by baseline cerebral lateralization. Likewise, animals with consistent left or right motor biases--a proxy measure of individual patterns in cerebral lateralization--have been shown to differ in their personality, emotional reactivity, motivational tendencies or coping styles. In this paper, we present the current knowledge of the links between laterality and stable individual traits in behaviour and affect in light of hypotheses on emotional lateralization. Within our suggested framework, we make recommendations on how to investigate affective styles in non-human animals and give practical examples. This approach has the potential to promote a science of affective styles in nonhuman animals and significantly advance research on animal welfare.
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Bandini, E., & Tennie C. (2020). Exploring the role of individual learning in animal tool-use. PeerJ, 25, 8:e9877.
Abstract: The notion that tool-use is unique to humans has long been refuted by the growing number of observations of animals using tools across various contexts. Yet, the mechanisms behind the emergence and sustenance of these tool-use repertoires are still heavily debated. We argue that the current animal behaviour literature is biased towards a social learning approach, in which animal, and in particular primate, tool-use repertoires are thought to require social learning mechanisms (copying variants of social learning are most often invoked). However, concrete evidence for a widespread dependency on social learning is still lacking. On the other hand, a growing body of observational and experimental data demonstrates that various animal species are capable of acquiring the forms of their tool-use behaviours via individual learning, with (non-copying) social learning regulating the frequencies of the behavioural forms within (and, indirectly, between) groups. As a first outline of the extent of the role of individual learning in animal tool-use, a literature review of reports of the spontaneous acquisition of animal tool-use behaviours was carried out across observational and experimental studies. The results of this review suggest that perhaps due to the pervasive focus on social learning in the literature, accounts of the individual learning of tool-use forms by naïve animals may have been largely overlooked, and their importance under-examined.
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Wallner, B., Palmieri, N., Vogl, C., Rigler, D., Bozlak, E., Druml, T., et al. (2017). Y Chromosome Uncovers the Recent Oriental Origin of Modern Stallions. Current Biology, 27(13), 2029–2035.e5.
Abstract: The Y chromosome directly reflects male genealogies, but the extremely low Y chromosome sequence diversity in horses has prevented the reconstruction of stallion genealogies [1, 2]. Here, we resolve the first Y chromosome genealogy of modern horses by screening 1.46 Mb of the male-specific region of the Y chromosome (MSY) in 52 horses from 21 breeds. Based on highly accurate pedigree data, we estimated the de novo mutation rate of the horse MSY and showed that various modern horse Y chromosome lineages split much later than the domestication of the species. Apart from few private northern European haplotypes, all modern horse breeds clustered together in a roughly 700-year-old haplogroup that was transmitted to Europe by the import of Oriental stallions. The Oriental horse group consisted of two major subclades: the Original Arabian lineage and the Turkoman horse lineage. We show that the English Thoroughbred MSY was derived from the Turkoman lineage and that English Thoroughbred sires are largely responsible for the predominance of this haplotype in modern horses.
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Dyson, S., Berger, J., Ellis, A. D., & Mullard, J. (2018). Development of an ethogram for a pain scoring system in ridden horses and its application to determine the presence of musculoskeletal pain. Journal of Veterinary Behavior, 23, 47–57.
Abstract: There is evidence that more than 47% of the sports horse population in normal work may be lame, but the lameness is not recognized by owners or trainers. An alternative means of detecting pain may be recognition of behavioral changes in ridden horses. It has been demonstrated that there are differences in facial expressions in nonlame and lame horses. The purpose of this study was to develop a whole horse ethogram for ridden horses and to determine whether it could be applied repeatedly by 1 observer (repeatability study, 9 horses) and if, by application of a related pain behavior score, lame horses (n = 24) and nonlame horses (n = 13) could be differentiated. It was hypothesized that there would be some overlap in pain behavior scores among nonlame and lame horses; and that overall, nonlame horses would have a lower pain behavior score than lame horses. The ethogram was developed with 117 behavioral markers, and the horses were graded twice in random order by a trained specialist using video footage. Overall, there was a good correlation between the 2 assessments (P < 0.001; R2 = 0.91). Behavioral markers that were not consistent across the 2 assessments were omitted, reducing the ethogram to 70 markers. The modified ethogram was applied to video recordings of the nonlame horses and lame horses (ethogram evaluation). There was a strong correlation between 20 behavioral markers and the presence of lameness. The ethogram was subsequently simplified to 24 behavioral markers, by the amalgamation of similar behaviors which scored similarly and by omission of markers which showed unreliable results in relation to lameness. Following this, the maximum individual occurrence score for lame horses was 14 (out of 24 possible markers), with a median and mean score of 9 (±2 standard deviation) compared with a maximum score of 6 for nonlame horses, with a median and mean score of 2 (±1.4). For lame horses, the following behaviors occurred significantly more (P < 0.05, chi-square): ears back, mouth opening, tongue out, change in eye posture and expression, going above the bit, head tossing, tilting the head, unwillingness to go, crookedness, hurrying, changing gait spontaneously, poor quality canter, resisting, and stumbling and toe dragging. Recognition of these features as potential indicators of musculoskeletal pain may enable earlier recognition of lameness and avoidance of punishment-based training. Further research is necessary to verify this new ethogram for assessment of pain in ridden horses.
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Henry, S., Fureix, C., Rowberry, R., Bateson, M., & Hausberger, M. (2017). Do horses with poor welfare show 'pessimistic' cognitive biases? Sci. Nat., 104(1), 8.
Abstract: This field study tested the hypothesis that domestic horses living under putatively challenging-to-welfare conditions (for example involving social, spatial, feeding constraints) would present signs of poor welfare and co-occurring pessimistic judgement biases. Our subjects were 34 horses who had been housed for over 3 years in either restricted riding school situations (e.g. kept in single boxes, with limited roughage, ridden by inexperienced riders; N = 25) or under more naturalistic conditions (e.g. access to free-range, kept in stable social groups, leisure riding; N = 9). The horses' welfare was assessed by recording health-related, behavioural and postural indicators. Additionally, after learning a location task to discriminate a bucket containing either edible food ('positive' location) or unpalatable food ('negative' location), the horses were presented with a bucket located near the positive position, near the negative position and halfway between the positive and negative positions to assess their judgement biases. The riding school horses displayed the highest levels of behavioural and health-related problems and a pessimistic judgment bias, whereas the horses living under more naturalistic conditions displayed indications of good welfare and an optimistic bias. Moreover, pessimistic bias data strongly correlated with poor welfare data. This suggests that a lowered mood impacts a non-human species' perception of its environment and highlights cognitive biases as an appropriate tool to assess the impact of chronic living conditions on horse welfare.
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Sighieri, C., Tedeschi, D., De Andreis, C., Petri, L., & Baragli, P. (2003). Behaviour Patterns of Horses Can be Used to Establish a Dominant-Subordinate Relationship Between Man and Horse. Animal Welfare, 12(4), 705–708.
Abstract: This paper describes how man can enter the social hierarchy of the horse by mimicking the behaviour and stance it uses to establish dominance. A herd is organised according to a dominance hierarchy established by means of ritualised conflict. Dominance relationships are formed through these confrontations: one horse gains the dominant role and others identify themselves as subordinates. This study was conducted using five females of the Haflinger breed, totally unaccustomed to human contact, from a free-range breeding farm. The study methods were based on the three elements fundamental to the equilibrium of the herd: flight, herd instinct and hierarchy. The trainer-horse relationship was established in three phases: retreat, approach and association. At the end of the training sessions, all of the horses were able to respond correctly to the trainer. These observations suggest that it is possible to manage unhandled horses without coercion by mimicking their behaviour patterns.
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Bussières, G., Jacques, C., Lainay, O., Beauchamp, G., Leblond, A., Cadoré, J. - L., et al. (2008). Development of a composite orthopaedic pain scale in horses. Research in Veterinary Science, 85(2), 294–306.
Abstract: This study addresses development and validation of a composite multifactorial pain scale (CPS) in an experimental equine model of acute orthopaedic pain. Eighteen horses were allocated to control (sedation with/without epidural analgesia – mixture of morphine, ropivacaine, detomidine and ketamine) and experimental groups: amphotericin-B injection in the tarsocrural joint induced pain and analgesia was either i.v. phenylbutazone administered post-induction of synovitis, or pre-emptive epidural mixture, or a pre-emptive combination of the 2. Inter- and intra-observer reproducibility was good (0.8<K<1). The key specific and sensitive behavioural indices were response to palpation of the painful area, posture, and, of lesser value, pawing on the floor, kicking at abdomen and head movement. Of particular interest was the statistical correlation observed between the CPS and both non-invasive blood pressure (P<0.0001) and blood cortisol (P<0.002). This study established the value of some behavioural and physiological criteria in determining equine orthopaedic pain intensity and clearly demonstrated that pre-emptive, multimodal analgesia provided better management than the two other protocols tested.
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Versace, E., Morgante, M., Pulina, G., & Vallortigara, G. (2007). Behavioural lateralization in sheep (Ovis aries). Behav. Brain. Res., 184(1), 72–80.
Abstract: This study investigates behavioural lateralization in sheep and lambs of different ages. A flock was tested in a task in which the animals were facing an obstacle and should avoid it on either the right or left side to rejoin flock-mates (adult sheep) or their mothers (lambs). A bias for avoiding the obstacle on the right side was observed, with lambs apparently being more lateralized than sheep. This right bias was tentatively associated with the left-hemifield laterality in familiar faces recognition which has been documented in this species. Differences between adult sheep and lambs were likely to be due to differences in social reinstatement motivation elicited by different stimuli (flock-mates or mothers) at different ages. Preferential use of the forelegs to step on a wood-board and direction of jaw movement during rumination was also tested in adult animals. No population bias nor individual-level lateralization was observed for use of the forelegs. At the same time, however, there was a large number of animals showing individual-level lateralization for the direction of jaw movement during rumination even though there was no population bias. These findings highlight that within the same species individual- and population-level lateralization can be observed in different tasks. Moreover, the results fit the general hypothesis that population-level asymmetries are more likely to occur in tasks that require social coordination among behaviourally asymmetric individuals.
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Krueger, K., Trager, L., Farmer, K., & Byrne, R. (2022). Tool Use in Horses. Animals, 12(15), 1876.
Abstract: Tool use has not yet been confirmed in horses, mules or donkeys. As this subject is difficult to research with conventional methods, we used a crowdsourcing approach to gather data. We contacted equid owners and carers and asked them to report and video examples of �unusual� behaviour via a dedicated website. We also searched YouTube and Facebook for videos of equids showing tool use. From 635 reports, including 1014 behaviours, we found 20 cases of tool use, 13 of which were unambiguous in that it was clear that the behaviour was not trained, caused by reduced welfare, incidental or accidental. We then assessed (a) the effect of management conditions on tool use and (b) whether the animals used tools alone, or socially, involving other equids or humans. We found that management restrictions were associated with corresponding tool use in 12 of the 13 cases (p = 0.01), e.g., equids using sticks to scrape hay within reach when feed was restricted. Furthermore, 8 of the 13 cases involved other equids or humans, such as horses using brushes to groom others. The most frequent tool use was for foraging, with seven examples, tool use for social purposes was seen in four cases, and there was just one case of tool use for escape. There was just one case of tool use for comfort, and in this instance, there were no management restrictions. Equids therefore can develop tool use, especially when management conditions are restricted, but it is a rare occurrence.
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Janczarek, I., Wisniewska, A., Chruszczewski, M. H., Tkaczyk, E., & Górecka-Bruzda, A. (2020). Social Behaviour of Horses in Response to Vocalisations of Predators. Animals, 10(2331).
Abstract: We tested the hypothesis that social defensive responses to the vocalisation of a predator still exist in horses. The recordings of a grey wolf, an Arabian leopard and a golden jackal were played to 20 Konik polski and Arabian mares. Durations of grazing, standing still, standing alert and the number of steps in walk and trot/canter were measured. In one-minute scans, the distances of the focal horse from the reference horse (DIST-RH) and from the nearest loudspeaker (DIST-LS) were approximated. The vocalisation of a leopard aroused the Arabians more than the Koniks (less grazing, stand-still and walk, more stand-alert and trotting/cantering). Koniks showed more relaxed behaviours to the leopard vocalisation (more grazing, stand-still and walk), but high alertness to the wolf playback (stand-alert, trotting/cantering). Spatial formation of the herd of Koniks showed tight grouping (lower DIST-RH) and maintaining distance from the potential threat (DIST-LS) in response to the wolf howling, while the Arabians approached the loudspeakers in linear herd formation when the leopard growls were played. Adult horses responded to potential predation by changing spatial group formations. This ability to apply a social strategy may be one of the explanations for the least number of horses among all hunted farm animal species.
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