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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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Bergmüller, R., & Taborsky, M. (2010). Animal personality due to social niche specialisation. Trends in Ecology & Evolution, 25(9), 504–511.
Abstract: The existence of 'animal personality', i.e. consistent individual differences in behaviour across time and contexts, is an evolutionary puzzle that has recently generated considerable research interest. Although social factors are generally considered to be important, it is as yet unclear how they might select for personality. Drawing from ecological niche theory, we explore how social conflict and alternative social options can be key factors in the evolution and development of consistent individual differences in behaviour. We discuss how animal personality research might benefit from insights into the study of alternative tactics and illustrate how selection can favour behavioural diversification and consistency due to fitness benefits resulting from conflict reduction among social partners.
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Baragli, P., Vitale, V., Paoletti, E., Sighieri, C., & Reddon, A. R. (2011). Detour behaviour in horses (Equus caballus). J. Ethol., 29(2), 227–234.
Abstract: The objective of this study was to investigate the ability of horses (Equus caballus) to detour around symmetric and asymmetric obstacles. Ten female Italian saddle horses were each used in three detour tasks. In the first task, the ability to detour around a symmetrical obstacle was evaluated; in the second and third tasks subjects were required to perform a detour around an asymmetrical obstacle with two different degrees of asymmetry. The direction chosen to move around the obstacle and time required to make the detour were recorded. The results suggest that horses have the spatial abilities required to perform detour tasks with both symmetric and asymmetric obstacles. The strategy used to perform the task varied between subjects. For five horses, lateralized behaviour was observed when detouring the obstacle; this was consistently in one direction (three on the left and two on the right). For these horses, no evidence of spatial learning or reasoning was found. The other five horses did not solve this task in a lateralized manner, and a trend towards decreasing lateralization was observed as asymmetry, and hence task difficulty, increased. These non-lateralized horses may have higher spatial reasoning abilities.
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Cooper, J. J., & Albentosa, M. J. (2005). Behavioural adaptation in the domestic horse: potential role of apparently abnormal responses including stereotypic behaviour. Livest. Prod. Sci., 92(2), 177–182.
Abstract: Classically, biologists have considered adaptation of behavioural characteristics in terms of long-term functional benefits to the individual, such as survival or reproductive fitness. In captive species, including the domestic horse, this level of explanation is limited, as for the most part, horses are housed in conditions that differ markedly from those in which they evolved. In addition, an individual horse's reproductive fitness is largely determined by man rather than its own behavioural strategies. Perhaps for reasons of this kind, explanations of behavioural adaptation to environmental challenges by domestic animals, including the capacity to learn new responses to these challenges, tend to concentrate on the proximate causes of behaviour. However, understanding the original function of these adaptive responses can help us explain why animals perform apparently novel or functionless activities in certain housing conditions and may help us to appreciate what the animal welfare implications might be. This paper reviews the behavioural adaptation of the domestic horse to captivity and discusses how apparently abnormal behaviour may not only provide a useful practical indicator of specific environmental deficiencies but may also serve the animal as an adaptive response to these deficiencies in an “abnormal” environment.
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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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Briard, L., Deneubourg, J. - L., & Petit, O. (2017). How stallions influence the dynamic of collective movements in two groups of domestic horses, from departure to arrival. Behav. Process., 142, 56–63.
Abstract: Abstract The role of leader in polygynous species has been solely attributed to the male for some time, but recent studies shown decision making to be distributed within the group. However, the specific reproductive strategy and behavioural repertoire of males in polygynous species such as horses may mean that these individuals still have the potential to play a specific role during decision-making. To investigate this subject, we thoroughly studied the behaviour of two domestic stallions during collective movements of their group. We found that they initiated rarely and sometimes failed to recruit the entire group. When departing as followers, they did not accelerate the joining process. Both stallions preferentially occupied the rear position and exhibited numerous monitoring behaviours. Herding behaviours were performed by only one stallion and mostly occurred outside movement context. Finally, we removed this herding stallion from its group to evaluate how the group dynamic changed. As a result, half of the collective movements were five times slower and mares were more dispersed in comparison when the stallion was in the group. Overall, our results suggest that, the two stallions maintained their role of group monitors from departure to arrival. Their influence on the movement dynamic was indirect and did not play a specific role in the process of decision making.
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da Cruz, A. B., Hirata, S., dos Santos, M. E., & Mendonça, R. S. (2023). Show me your best side: Lateralization of social and resting behaviors in feral horses. Behav. Process., 206, 104839.
Abstract: Growing evidence shows a variety of sensorial and motor asymmetries in social and non-social interactions in various species, indicating a lateralized processing of information by the brain. Using digital video cameras on tripods and drones, this study investigated lateralization in frequency and duration of social behavior patterns, in affiliative, agonistic, and resting contexts, in a feral population of horses (Equus ferus caballus) in Northern Portugal, consisting of 37 individuals organized in eight harem groups. Affiliative interactions (including grooming) were more often performed, and lasted longer, when recipients were positioned to the right side. In recumbent resting (animals lying down) episodes on the left side lasted longer. Our results of an affiliative behavior having a right side tendency, provide partial support to the valence-specific hypothesis of Ahern and Schwartz (1979) – left hemisphere dominance for positive affect, affiliative behaviors. Longer recumbent resting episodes on the left side may be due to synchronization. However, in both instances it is discussed how lateralization may be context dependent. Investigating the position asymmetries of social behaviors in feral equids will contribute to a better understanding of differential lateralization and hemispheric specialization from the ecological and evolutionary perspectives.
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da Cruz, A. B., Hirata, S., dos Santos, M. E., & Mendonça, R. S. (2023). Show me your best side: Lateralization of social and resting behaviors in feral horses. Behav. Process., 206, 104839.
Abstract: Growing evidence shows a variety of sensorial and motor asymmetries in social and non-social interactions in various species, indicating a lateralized processing of information by the brain. Using digital video cameras on tripods and drones, this study investigated lateralization in frequency and duration of social behavior patterns, in affiliative, agonistic, and resting contexts, in a feral population of horses (Equus ferus caballus) in Northern Portugal, consisting of 37 individuals organized in eight harem groups. Affiliative interactions (including grooming) were more often performed, and lasted longer, when recipients were positioned to the right side. In recumbent resting (animals lying down) episodes on the left side lasted longer. Our results of an affiliative behavior having a right side tendency, provide partial support to the valence-specific hypothesis of Ahern and Schwartz (1979) – left hemisphere dominance for positive affect, affiliative behaviors. Longer recumbent resting episodes on the left side may be due to synchronization. However, in both instances it is discussed how lateralization may be context dependent. Investigating the position asymmetries of social behaviors in feral equids will contribute to a better understanding of differential lateralization and hemispheric specialization from the ecological and evolutionary perspectives.
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Meriggi, A., Dagradi, V., Dondina, O., Perversi, M., Milanesi, P., Lombardini, M., et al. (2014). Short-term responses of wolf feeding habits to changes of wild and domestic ungulate abundance in Northern Italy. Ethology Ecology & Evolution, 27(4), 389–411.
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Drevemo, S., Fredricson, I., Hjertén, G., & McMIKEN, D. (1987). Early development of gait asymmetries in trotting Standardbred colts. Equine. Vet. J., 19(3), 189–191.
Abstract: Summary Ten trotting Standardbred colts were recorded by high-speed cinematography at the ages of eight, 12 and 18 months. The horses were trotting on a treadmill operating at 4.0 m/secs. Five horses were subjected to a programme of intensified training from eight months of age, whereas the others were not trained and acted as controls. The films were analysed on a semi-automatic film-reading equipment and a number of variables used to demonstrate the gait symmetry were calculated and scaled by computer. Certain differences between left and right diagonal and contralateral pair of limbs, respectively, were noted, suggesting that laterality in horses may be inherited. The most pronounced systematic differences were found in 18-month old horses in the trained group. The results show the importance of careful gait examination and comprehensive coordination training at an early age.
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