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Lansade, L., Colson, V., Parias, C., Trösch, M., Reigner, F., & Calandreau, L. (2020). Female horses spontaneously identify a photograph of their keeper, last seen six months previously. Scientific Reports, 10(1), 6302.
Abstract: Horses are capable of identifying individual conspecifics based on olfactory, auditory or visual cues. However, this raises the questions of their ability to recognize human beings and on the basis of what cues. This study investigated whether horses could differentiate between a familiar and unfamiliar human from photographs of faces. Eleven horses were trained on a discrimination task using a computer-controlled screen, on which two photographs were presented simultaneously (32 trials/session): touching one was rewarded (S+) and the other not (S-). In the training phase, the S+ faces were of four unfamiliar people which gradually became familiar over the trials. The S- faces were novel for each trial. After the training phase, the faces of the horses' keepers were presented opposite novel faces to test whether the horses could identify the former spontaneously. A reward was given whichever face was touched to avoid any possible learning effect. Horses touched the faces of keepers significantly more than chance, whether it was their current keeper or one they had not seen for six months (t = 3.65; p < 0.004 and t = 6.24; p < 0.0001). Overall, these results show that horses have advanced human face-recognition abilities and a long-term memory of those human faces.
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Boissy, A. (1995). Fear and Fearfulness in Animals. The Quarterly Review of Biology, 70(2), 165–191.
Abstract: Persistence of individual differences in animal behavior in reactions to various environmental challenges could reflect basic divergences in temperament, which might be used to predict details of adaptive response. Although studies have been carried out on fear and anxiety in various species, including laboratory, domestic and wild animals, no consistent definition of fearfulness as a basic trait of temperament has emerged. After a classification of the events that may produce a state of fear, this article describes the great variability in behavior and in physiological patterns generally associated with emotional reactivity. The difficulties of proposing fearfulness-the general capacity to react to a variety of potentially threatening situations-as a valid basic internal variable are then discussed. Although there are many studies showing covariation among the psychobiological responses to different environmental challenges, other studies find no such correlations and raise doubts about the interpretation of fearfulness as a basic personality trait. After a critical assessment of methodologies used in fear and anxiety studies, it is suggested that discrepancies among results are mainly due to the modulation of emotional responses in animals, which depend on numerous genetic and epigenetic factors. It is difficult to compare results obtained by different methods from animals reared under various conditions and with different genetic origins. The concept of fearfulness as an inner trait is best supported by two kinds of investigations. First, an experimental approach combining ethology and experimental psychology produces undeniable indicators of emotional reactivity. Second, genetic lines selected for psychobiological traits prove useful in establishing between behavioral and neuroendocrine aspects of emotional reactivity. It is suggested that fearfulness could be considered a basic feature of the temperament of each individual, one that predisposes it to respond similarly to a variety of potentially alarming challenges, but is nevertheless continually modulated during development by the interaction of genetic traits of reactivity with environmental factors, particularly in the juvenile period. Such interaction may explain much of the interindividual variability observed in adaptive responses.
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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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Bandini, E., Motes-Rodrigo, A., Steele, M. P., Rutz, C., & Tennie, C. (2020). Examining the mechanisms underlying the acquisition of animal tool behaviour. Biol. Lett., 16(2020122).
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Carson, K., & Wood-Gush, D. G. M. (1983). Equine behaviour: I. A review of the literature on social and dam--Foal behaviour. Applied Animal Ethology, 10(3), 165–178.
Abstract: In most cases, the social organisation of each of the seven species of Equidae existing today outside captivity is either territorial or non-territorial. The striking differences found between these two types of organisation in the social grouping and bonds, mating behaviour, leadership and dominance hierarchies of the animals are examined. It is thought that the non-territorial species show a less primitive type of organisation than the territorial animals. Infant Equidae are precocious animals and are able to follow their dams soon after birth. They stay close by their dams and travel with the herd from an early age and are therefore classified as “followers”, in contrast to the species which have a period of hiding after birth. Dams recognise their foals immediately after birth, whereas it takes 2 or 3 days for a foal to form an attachment to its dam. Being in close proximity to their dams, foals are able to nurse frequently and, unless artificially weaned, a foal will nurse until its dam foals again. Foals start to graze during their first week and as they grow older they spend more time grazing and less time nursing and resting. It is normal for foals to be corprophagic until one month old, and this provides them with bacteria essential for the digestion of fibre. Play behaviour is solitary in very young foals, but after 4 weeks of age, foals play together, with male foals playing more than females and showing more aggressive, fighting movements in play.
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Byström, A., Clayton, H. M., Hernlund, E., Rhodin, M., & Egenvall, A. (2020). Equestrian and biomechanical perspectives on laterality in the horse. Comp. Exerc. Physiol., 16(1), 35–45.
Abstract: It has been suggested that one of the underlying causes of asymmetrical performance and left/right bias in sound riding horses is laterality originating in the cerebral cortices described in many species. The aim of this paper is to review the published evidence for inherent biomechanical laterality in horses deemed to be clinically sound and relate these findings to descriptions of sidedness in equestrian texts. There are no established criteria to determine if a horse is left or right dominant but the preferred limb has been defined as the forelimb that is more frequently protracted during stance and when grazing. Findings on left-right differences in forelimb hoof shape and front hoof angles have been linked to asymmetric forelimb ground reaction forces. Asymmetries interpreted as motor laterality have been found among foals and unhandled youngsters, and the consistency or extent of asymmetries seems to increase with age. Expressions of laterality also vary with breed, sex, training and handling, stress, and body shape but there are no studies of the possible link between laterality and lameness. In a recent study of a group of seven dressage horses, a movement pattern in many ways similar to descriptions of sidedness in the equestrian literature, e.g. one hind limb being more protracted and placed more laterally than the other, has been documented. The role of innate laterality versus painful conditions, training, human handedness and simply habit remains to be determined. Understanding the biomechanical manifestations of laterality in healthy horses, including individual variation, would yield a potential basis for how laterality should be taken into account in relation to training/riding and rehabilitation of lameness.
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Suagee-Bedore, J. K., Linden, D. R., & Bennett-Wimbush, K. (2021). Effect of Pen Size on Stress Responses of Stall-Housed Horses Receiving One Hour of Daily Turnout. J. Equine Vet. Sci., 98, 103366.
Abstract: Group turnout provides important socializing opportunities for horses, particularly those that are primarily stalled. A high percentage of equine injuries occur during group turnout, which could partly be due to the physical constraints of fencing. To investigate appropriate paddock sizes for group turnouts, horses (n = 12) from a single herd were divided into groups of 4, stalled for 24 hours, and then turned out for 1 hour into one of three differently sized pens: 342, 263, and 184 m2 per horse. Groups rotated through pens across 3 days, receiving one treatment per day. Blood was sampled for cortisol concentrations at 08:00 hours each morning, and then at 15 and 60 minutes into the turn out sessions, and 60 minutes after return to individual stalls. Groups rotated through three turnout times: 09:00, 12:00, 14:00 hours. Counts of agonistic behaviors (chasing, contact biting, and kicking) and low-level threats (pinned ears, tail swishing, bite and kick threats) were recorded. When turned out in pens that provided 342 m2 per horse, horses exhibited reduced plasma cortisol concentrations by 15 minutes after turnout and at 1 hour after return to their stalls (P < .05). Horses in pens providing 184 m2 per horse exhibited greater agonistic (P < .001) and low-level threat (P < .01) behaviors than horses in larger pens. These data provide insight into appropriate pen sizes for horses from established herds. Providing at least 342 m2 per horse may reduce the chance of injury in horses accustomed to group turnout.
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Brinkmann, L., Gerken, M., & Riek, A. (2013). Effect of long-term feed restriction on the health status and welfare of a robust horse breed, the Shetland pony (Equus ferus caballus). Res. Vet. Sci., 94(3), 826–831.
Abstract: Outdoor group housing is increasingly recognized as an appropriate housing system for domesticated horses. The objective of this study was therefore to investigate the effect of potential feed shortage in semi-natural horse keeping systems in winter on animal health and welfare. In 10 female Shetland ponies blood concentrations (NEFA, total protein (TP), total bilirubin (TB), beta-hydroxybutyrate (BHB) and thyroxine (T4)), body mass and the body condition score (BCS) were monitored for 7months including a 4months period of feed restriction in five of the 10 ponies. Restrictively fed animals lost 18.4±2.99% of their body mass and the BCS decreased by 2.2±0.8 points (BCS scale: 0=emaciated, 5=obese). Feed restriction led to a continuous increase in TB (P<0.001) and NEFA (P<0.01) concentrations compared to control ponies. The TP and BHB values only differed at the end of the trial with lower concentrations in restricted fed mares (P<0.05). Feed restriction had no effect on thyroxine concentrations. TB concentrations in the feed restricted group were out of the reference range during the entire feeding trial. The increased NEFA concentrations in feed restricted compared to control ponies suggest that fat was mobilized. The BCS, as well as plasma NEFA and TB concentrations were good indicators for a rapid detection of possible health problems caused by undernourishment in horses when kept under semi-natural conditions. In contrast, blood parameters of the control animals were within the reference ranges, suggesting that a year round outdoor housing with additional feed supply is an adequate housing system for a robust horse breed like the Shetland pony.
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Weishaupt, M. A., Wiestner, T., von Peinen, K., Waldern, N., Roepstorff, L., van Weeren, R., et al. (2006). Effect of head and neck position on vertical ground reaction forces and interlimb coordination in the dressage horse ridden at walk and trot on a treadmill. Equine Vet J Suppl, (36), 387–392.
Abstract: REASONS FOR PERFORMING STUDY: Little is known in quantitative terms about the influence of different head-neck positions (HNPs) on the loading pattern of the locomotor apparatus. Therefore it is difficult to predict whether a specific riding technique is beneficial for the horse or if it may increase the risk for injury. OBJECTIVE: To improve the understanding of forelimb-hindlimb balance and its underlying temporal changes in relation to different head and neck positions. METHODS: Vertical ground reaction force and time parameters of each limb were measured in 7 high level dressage horses while being ridden at walk and trot on an instrumented treadmill in 6 predetermined HNPs: HNP1 – free, unrestrained with loose reins; HNP2 – neck raised, bridge of the nose in front of the vertical; HNP3 – neck raised, bridge of the nose behind the vertical; HNP4 – neck lowered and flexed, bridge of the nose considerably behind the vertical; HNP5 – neck extremely elevated and bridge of the nose considerably in front of the vertical; HNP6 – neck and head extended forward and downward. Positions were judged by a qualified dressage judge. HNPs were assessed by comparing the data to a velocity-matched reference HNP (HNP2). Differences were tested using paired t test or Wilcoxon signed rank test (P<0.05). RESULTS: At the walk, stride duration and overreach distance increased in HNP1, but decreased in HNP3 and HNP5. Stride impulse was shifted to the forehand in HNP1 and HNP6, but shifted to the hindquarters in HNP5. At the trot, stride duration increased in HNP4 and HNP5. Overreach distance was shorter in HNP4. Stride impulse shifted to the hindquarters in HNP5. In HNP1 peak forces decreased in the forelimbs; in HNP5 peak forces increased in fore- and hindlimbs. CONCLUSIONS: HNP5 had the biggest impact on limb timing and load distribution and behaved inversely to HNP1 and HNP6. Shortening of forelimb stance duration in HNP5 increased peak forces although the percentage of stride impulse carried by the forelimbs decreased. POTENTIAL RELEVANCE: An extremely high HNP affects functionality much more than an extremely low neck.
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Boyce, P. N., & McLoughlin, P. D. (2021). Ecological Interactions Involving Feral Horses and Predators: Review with Implications for Biodiversity Conservation. Jour. Wild. Mgmt., n/a(n/a).
Abstract: ABSTRACT For many ecosystems, feral horses are increasingly becoming an important if not dominant component of ungulate biomass and hence influence on community dynamics. Yet we still know little of how horses contribute to key ecological interactions including predator-prey and indirect competitive relationships at a community level. Notably, feral species like horses can exhibit life-history traits that differ from that of native (mainly artiodactyl) herbivore competitors. Artificial selection for traits like increased, early, or extended reproduction that have yet to be reversed by natural selection, coupled with naturally selected differences in anatomy and behavior, in addition to unique management objectives for horses compared to other species, means that the dynamics of feral horse populations are not likely to align with what might be expected of other large herbivores. Unexpected population dynamics and inherent biological asymmetries between native ungulates and feral horses may therefore influence the former via direct competition for shared resources and through enemy-mediated interactions like apparent competition. In several localities feral horses now co-exist with multiple native prey species, some of which are in decline or are species at risk. Compounding risks to native species from direct or indirect competitive exclusion by horses is the unique nature and socio-political context of feral horse management, which tends towards allowing horse populations to be limited largely by natural, density-dependent factors. We summarize the inherent asymmetries between feral horse biology and that of other ungulate prey species with consequences for conservation, focusing on predator-prey and emerging indirect interactions in multi-prey systems, and highlight future directions to address key knowledge gaps in our understanding of how feral horses may now be contributing to the (re)structuring of food webs. Observations of patterns of rapid growth and decline, and associated skews in sex ratios of feral horse populations, indicate a heightened potential for indirect interactions among large ungulate prey species, where there is a prevalence of feral horses as preferred prey, particularly where native prey are declining. In places like western North America, we expect predator-prey interactions involving feral horses to become an increasingly important factor in the conservation of wildlife. This applies not only to economically or culturally important game species but also at-risk species, both predators (e.g., wolves [Canis lupus], grizzly bears [Ursus arctos]) and prey (e.g., woodland caribou [Rangifer tarandus caribou]), necessitating an ecological understanding of the role of horses in natural environments that goes beyond that of population control. ? 2021 The Wildlife Society.
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