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Austin, N. P., & Rogers, L. J. (2014). Lateralization of agonistic and vigilance responses in Przewalski horses (Equus przewalskii). Applied Animal Behaviour Science, 151, 43–50.
Abstract: tEye and limb preferences were scored in the closest undomesticated relative of Equuscaballus using the same methods as used previously to study laterality in feral horses.Observations were made of 33 Przewalski horses (Equus ferus przewalskii) (male N = 20,female N = 13) living under natural social conditions on a large reserve in France. Signifi-cant left-eye/side biases were found in agonistic interactions within harem bands (M ± SEbias to left 58% ± 0.01 for threats, P < 0.001; 68% ± 0.05 for attacks; P < 0.001) and in stallionfights (threats, 52% ± 0.01 left, P < 0.001; attacks, 63% ± 0.02 left, P < 0.001): as many as 80%of the horses were significantly lateralized in attack responses within harem bands. Lat-erality of vigilance was measured as lifting up the head from grazing and turning it to theleft or right side: a directional bias to the left was found (M ± SE 53% ± 0.02 left, P < 0.001).Side bias in reactivity was calculated as the percent of head lifts above the level of thewithers on the left or right side and this was also left side biased (M ± SE 73% ± 0.03 left,P < 0.001). These results indicate right-hemisphere specialization for control of aggressionand responses to novelty. The left bias in attack scores within harem bands was strongerin males than females (P = 0.024) and in immature than adult horses (P = 0.032). Immaturehorses were also more strongly lateralized than adults in vigilance scores (P = 0.022), whichmay suggest that experience reduces these side biases. Our results show that Przewalskihorses exhibit left eye preferences, as do feral horses, and do so even more strongly thanferal horses. Considering feral and Przewalski horses together, we deduce that ancestralhorses had similar lateral biases. Also similar to feral horses, the Przewalski horses showedno significant forelimb preference at the group level or in the majority of horses at theindividual level, confirming the hypothesis that previously reported limb preferences indomestic breeds are entrained or generated by breed-specific selection.
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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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Esch, L., Wöhr, C., Erhard, M., & Krueger, K. (2019). Horses� (Equus Caballus) Laterality, Stress Hormones, and Task Related Behavior in Innovative Problem-Solving. Animals, 9(5), 265.
Abstract: Domesticated horses are constantly confronted with novel tasks. A recent study on anecdotal data indicates that some are innovative in dealing with such tasks. However, innovative behavior in horses has not previously been investigated under experimental conditions. In this study, we investigated whether 16 horses found an innovative solution when confronted with a novel feeder. Moreover, we investigated whether innovative behavior in horses may be affected by individual aspects such as: age, sex, size, motor and sensory laterality, fecal stress hormone concentrations (GCMs), and task-related behavior. Our study revealed evidence for 25% of the horses being capable of innovative problem solving for operating a novel feeder. Innovative horses of the present study were active, tenacious, and may be considered to have a higher inhibitory control, which was revealed by their task related behavior. Furthermore, they appeared to be emotional, reflected by high baseline GCM concentrations and a left sensory and motor laterality. These findings may contribute to the understanding of horses� cognitive capacities to deal with their environment and calls for enriched environments in sports and leisure horse management.
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Reddon, A. R., & Hurd, P. L. (2009). Individual differences in cerebral lateralization are associated with shy-bold variation in the convict cichlid. Anim. Behav., 77(1), 189–193.
Abstract: Cerebral lateralization, the preferential use of one hemisphere of the brain to perform certain cognitive functions, is a widespread and evolutionarily ancient adaptation. Lateralization appears to enhance cognitive capacity, yet substantial individual variation in the strength cerebral lateralization is apparent in all species studied so far. It is puzzling that cerebral lateralization, a seemingly advantageous trait, has not been driven to fixation. It has been suggested that variation in lateralization may be linked to individual variation in behaviour, which itself may be subject to disruptive selection. We examined the relation between cerebral lateralization and individual variation in boldness in the convict cichlid, Archocentrus nigrofasciatus. We show that convict cichlids that are more strongly lateralized when exploring a familiar environment, but not a novel one, are quicker to emerge from a shelter in a test for boldness. The possibility that cerebral lateralization is linked to life history strategy is discussed.
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Jennings, D. J. (2014). Limited evidence that visual lateralization is associated with fitness in rutting male fallow deer. Anim. Behav., 92, 85–91.
Abstract: Under certain models of animal competition, individuals are expected to gather information about opponent quality in order to determine whether they should fight or withdraw. However, the ability to process complex information differs between individuals and across brain hemispheres: a feature of vertebrate cognition known as lateralization that is not anticipated by contest models. I investigated the relationship between aggressive behaviour and mating success during the fallow deer, Dama dama, rut and a measure of lateralization derived from eye preference during parallel walking. Results show that there was no relationship between the tendency to escalate to fighting or predictability in the tendency to engage in fighting and lateralization. Conversely, there was a quadratic relationship between third-party intervention behaviour and lateralization: the greater the tendency to intervene in ongoing fights the lower the degree of lateralization. However, individuals that showed lateralization for right-eye use were least likely to be targeted by the intervening male; thus lateralization is beneficial in this context because targeted males are highly likely to lose this subsequent encounter. The relationship between lateralization and mating success was also nonlinear: males that showed little evidence for an eye bias during lateral displays had the greatest mating success. Taken together, individuals that showed lateralization benefited from avoiding being targeted after third-party intervention; conversely, individuals that showed little evidence for lateralization actively intervened during ongoing fights and had higher mating success. These results suggest that, although lateralization does appear to confer a fitness advantage on individuals, this is not as extensive as anticipated.
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Komárková, M., & Bartošová, J. (2013). Lateralized suckling in domestic horses (Equus caballus). Animal Cognition, 16(3), 343–349.
Abstract: Brain lateralization enables preferential processing of certain stimuli and more effective utilization of these stimuli in either the left or the right cerebral hemisphere. Horses show both motor and sensory lateralization patterns. Our aim was to determine whether a lateralized response could be detected in foals during the naturally side-biased behaviour, suckling. The foals’ preferred suckling side could be the effect of either visual or motor lateralization. In the case of a visual lateralized response, foals are expected to suck more often from the mother’s right side, so potential danger can be detected by the better adapted right hemisphere (i.e. left eye). Motor lateralization can be identified when a foal will suck predominantly from one side, either left or right. We found no population trend in the preferred suckling side, but we detected significant differences amongst individual foals. One-third (35.4 %) of 79 foals showed a strong, either right or left side preference which increased with age. The mothers did not influence the foals’ suckling side preferences either by side-biased rejection or termination of suckling. According to our findings, a general pattern of sucking with the left eye open for better danger detection and recognition is unlikely in foals up to 7 months old. Foals of this age are probably young or fully focused on suckling and rely on their mothers’ vigilance. Individual side preferences amongst foals are suggested to be based on motor lateralization.
Keywords: Domestic horse; Foal; Suckling; Lateralization
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