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Leliveld, L. M. C., Düpjan, S., Tuchscherer, A., & Puppe, B. (2020). Hemispheric Specialization for Processing the Communicative and Emotional Content of Vocal Communication in a Social Mammal, the Domestic Pig. Front. Behav. Neurosci., 14, 596758.
Abstract: In humans, speech perception is lateralized, with the left hemisphere of the brain dominant in processing the communicative content and the right hemisphere dominant in processing the emotional content. However, still little is known about such a division of tasks in other species. We therefore investigated lateralized processing of communicative and emotionally relevant calls in a social mammal, the pig (Sus scrofa). Based on the contralateral connection between ears and hemispheres, we compared the behavioural and cardiac responses of 36 young male pigs during binaural and monaural (left or right) playback to the same sounds. The playback stimuli were calls of social isolation and physical restraint, whose communicative and emotional relevance, respectively, were validated prior to the test by acoustic analyses and during binaural playbacks. There were indications of lateralized processing mainly in the initial detection (left head-turn bias, indicating right hemispheric dominance) of the more emotionally relevant restraint calls. Conversely, there were indications of lateralized processing only in the appraisal (increased attention during playback to the right ear) of the more communicative relevant isolation calls. This implies differential involvement of the hemispheres in the auditory processing of vocalizations in pigs and thereby hints at similarities in the auditory processing of vocal communication in non-human animals and speech in humans. Therefore, these findings provide interesting new insight in the evolution of human language and auditory lateralization.
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Janczarek, I., Stachurska, A., Kedzierski, W., Wisniewska, A., Ryzak, M., & Koziol, A. (2020). The intensity of physiological and behavioral responses of horses to predator vocalizations. BMC Veterinary Research, 16(1), 431.
Abstract: Predatory attacks on horses can become a problem in some parts of the world, particularly when considering the recovering gray wolf populations. The issue studied was whether horses transformed by humans and placed in stable-pasture environments had retained their natural abilities to respond to predation risk. The objective of the study was to determine the changes in cardiac activity, cortisol concentrations, and behavior of horses in response to the vocalizations of two predators: the gray wolf (Canis lupus), which the horses of the breed studied had coevolved with but not been exposed to recently, and Arabian leopard (Panthera pardus nimr), from which the horses had been mostly isolated. In addition, we hypothesized that a higher proportion of Thoroughbred (TB) horse ancestry in the pedigree would result in higher emotional excitability in response to predator vocalizations. Nineteen horses were divided into groups of 75%, 50% and 25% TB ancestry. The auditory test conducted in a paddock comprised a 10-min prestimulus period, a 5-min stimulus period when one of the predators was heard, and a 10-min poststimulus period without any experimental stimuli.
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Chaplin, S. J., & Gretgrix, L. (2010). Effect of housing conditions on activity and lying behaviour of horses. animal, 4(5), 792–795.
Abstract: Housing conditions for horses impose various levels of confinement, which may compromise welfare. Lying behaviour and activity can be used as welfare indicators for domestic animals and rebound behaviour suggests a build-up of motivation resulting from deprivation. The objective of this study was to determine if activity and lying behaviour of horses are affected by housing conditions and to investigate the occurrence of rebound behaviour after release from confinement. Eight horses were subjected, in pairs, to each of four experimental treatments; paddock (P), fully stabled (FS), partly stabled (PS) and yard (Y). Each horse received 6 days acclimatisation prior to the 24 h recording period. Time spent in lying and activity were electronically recorded using a tilt switch and motion sensor connected to a data logger worn on the horse's left foreleg. Time spent active during the first 5 min of release from stable to paddock in the PS treatment (days 1 and 5) and at the same time of day in the P treatment was used as a measure of rebound behaviour. Effect of housing conditions on total time spent active was highly significant (FS = 123 s, PS = 158 s, Y = 377 s, P = 779 s, P < 0.001). Housing conditions did not significantly affect total time spent lying (P = 0.646). Horses were significantly more active, compared with baseline paddock behaviour, on release from stabling on both days 1 (P = 0.006) and 5 (P = 0.025) of PS treatment. These results suggest that activity patterns of horses, but not lying behaviour, are affected by the housing conditions tested and that rebound activity occurs in horses after a period of confinement.
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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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López-Bao, J. V., Sazatornil, V., Llaneza, L., & Rodríguez, A. (2013). Indirect Effects on Heathland Conservation and Wolf Persistence of Contradictory Policies that Threaten Traditional Free-Ranging Horse Husbandry. Conservation Letters, 6(6), 448–455.
Abstract: Abstract Conservation agencies within the European Union promote the restoration of traditional land uses as a cost-effective way to preserve biodiversity outside reserves. Although the European Union pursues the integration of the environment into strategic decision-making, it also dictates sectoral policies that may damage farmland biodiversity. We illustrate this point by outlining the socioeconomic factors that allow the persistence of traditional free-ranging horse husbandry in Galicia, northwestern Spain. Free-ranging Galician mountain ponies provide ecological and socioeconomic services including the prevention of forest fires, the maintenance of heathlands and wolves, and the attenuation of wolf-human conflicts. This traditional livestock system may have persisted because it entails negligible costs for farmers. Wolf predation upon Galician mountain ponies does not threaten farmer's economies and seems to be tolerated better than attacks to more valuable stock. Recently, European Union's regulations on animal welfare, carcass management, or meat production put new economic and administrative burdens on farmers, make free-ranging horse rearing economically unsustainable, and incentivize its abandonment. The aim of the European Union to integrate environmental policies may be successful to preserve farmland biodiversity only through careful anticipation of the side effects of apparently unrelated regulations on the fragile equilibrium that sustain traditional land uses.
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Reinhardt, I., Kluth, G., Nowak, C., Szentiks, C. A., Krone, O., Ansorge, H., et al. (2019). Military training areas facilitate the recolonization of wolves in Germany. Conservation Letters, 12(3), e12635.
Abstract: Abstract Wolves (Canis lupus) are currently showing a remarkable comeback in the highly fragmented cultural landscapes of Germany. We here show that wolf numbers increased exponentially between 2000 and 2015 with an annual increase of about 36%. We demonstrate that the first territories in each newly colonized region were established over long distances from the nearest known reproducing pack on active military training areas (MTAs). We show that MTAs, rather than protected areas, served as stepping-stones for the recolonization of Germany facilitating subsequent spreading of wolf territories in the surrounding landscape. We did not find any significant difference between MTAs and protected areas with regard to habitat. One possible reason for the importance of MTAs may be their lower anthropogenic mortality rates compared to protected and other areas. To our knowledge, this is the first documented case where MTAs facilitate the recolonization of an endangered species across large areas.
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López-Bao, J. V., Sazatornil, V., Llaneza, L., & Rodríguez, A. (2013). Indirect Effects on Heathland Conservation and Wolf Persistence of Contradictory Policies that Threaten Traditional Free-Ranging Horse Husbandry. Conservation Letters, 6(6), 448–455.
Abstract: Abstract Conservation agencies within the European Union promote the restoration of traditional land uses as a cost-effective way to preserve biodiversity outside reserves. Although the European Union pursues the integration of the environment into strategic decision-making, it also dictates sectoral policies that may damage farmland biodiversity. We illustrate this point by outlining the socioeconomic factors that allow the persistence of traditional free-ranging horse husbandry in Galicia, northwestern Spain. Free-ranging Galician mountain ponies provide ecological and socioeconomic services including the prevention of forest fires, the maintenance of heathlands and wolves, and the attenuation of wolf-human conflicts. This traditional livestock system may have persisted because it entails negligible costs for farmers. Wolf predation upon Galician mountain ponies does not threaten farmer's economies and seems to be tolerated better than attacks to more valuable stock. Recently, European Union's regulations on animal welfare, carcass management, or meat production put new economic and administrative burdens on farmers, make free-ranging horse rearing economically unsustainable, and incentivize its abandonment. The aim of the European Union to integrate environmental policies may be successful to preserve farmland biodiversity only through careful anticipation of the side effects of apparently unrelated regulations on the fragile equilibrium that sustain traditional land uses.
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d'Ingeo, S., Quaranta, A., Siniscalchi, M., Stomp, M., Coste, C., Bagnard, C., et al. (2019). Horses associate individual human voices with the valence of past interactions: a behavioural and electrophysiological study. Scientific Reports, 9(1), 11568.
Abstract: Brain lateralization is a phenomenon widely reported in the animal kingdom and sensory laterality has been shown to be an indicator of the appraisal of the stimulus valence by an individual. This can prove a useful tool to investigate how animals perceive intra- or hetero-specific signals. The human-animal relationship provides an interesting framework for testing the impact of the valence of interactions on emotional memories. In the present study, we tested whether horses could associate individual human voices with past positive or negative experiences. Both behavioural and electroencephalographic measures allowed examining laterality patterns in addition to the behavioural reactions. The results show that horses reacted to voices associated with past positive experiences with increased attention/arousal (gamma oscillations in the right hemisphere) and indicators of a positive emotional state (left hemisphere activation and ears held forward), and to those associated with past negative experiences with negative affective states (right hemisphere activation and ears held backwards). The responses were further influenced by the animals' management conditions (e.g. box or pasture). Overall, these results, associating brain and behaviour analysis, clearly demonstrate that horses' representation of human voices is modulated by the valence of prior horse-human interactions.
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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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Camerlink, I., Menneson, S., Turner, S. P., Farish, M., & Arnott, G. (2018). Lateralization influences contest behaviour in domestic pigs. Scientific Reports, 8(1), 12116.
Abstract: Cerebral lateralization, i.e. hemispheric asymmetries in structure and function, relates in many species to a preference to attack from their left. Lateralization increases cognitive capacity, enabling the simultaneous processing of multiple sources of information. Therefore, lateralization may constitute a component of fighting ability (Resource Holding Potential), and/or influence the efficiency of information-gathering during a contest. We hypothesized that lateralization will affect contest outcome and duration, with an advantage for more strongly lateralized individuals. In 52 dyadic contests between weight-matched pigs (Sus scrofa; n = 104; 10 wk age), the direction of orientation towards the opponent was scan sampled every 10 s. Laterality indexes (LI) were calculated for the direction and strength of lateralization. Up to 12.5% of the individuals showed significant lateralization towards either the right or left but lateralization was absent at the population level. In line with our hypothesis, animals showing strong lateralization (irrespective of direction) had a shorter contest duration than animals showing weak lateralization. Winners did not differ from losers in their strength or direction of lateralization. Overall the results suggest that cerebral lateralization may aid in conflict resolution, but does not directly contribute to fighting ability, and will be of value in the study of animal contests.
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