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Batt, L. S., Batt, M. S., Baguley, J. A., & McGreevy, P. D. (2009). The relationships between motor lateralization, salivary cortisol concentrations and behavior in dogs. Journal of Veterinary Behaviour, 4(6), 216–222.
Abstract: The degree of lateralization (LI) indicates both the direction and strength of a paw preference. Here, a positive value is indicative of a right paw bias, and a negative value of a left paw bias. Higher numbers on the positive side of the scale and lower numbers on the negative side of the scale indicate a greater strength of that lateralization. The strength of motor lateralization (|LI|) is the absolute value of the LI. The use of absolute value removes directionality (i.e., does not indicate left or right paw bias) and instead indicates only the strength of the paw preference. Both LI and |LI| have been associated with behavioral differences in a range of species. The assessment of motor lateralization in the dog can be conducted by observing the paw used to perform motor tasks. Elevated cortisol concentrations have been associated with fearfulness in many species. Additionally, fearfulness and boldness can be assessed in response to so-called temperament tests. Consequently, in this study we examine the relationship between lateralization, temperament test results, and cortisol concentrations in 43 potential guide dogs, of which 38 were Labrador retrievers and 5 were golden retrievers. Over a 14-month period, the current study assessed motor lateralization and salivary cortisol concentrations 3 times (approximately 6 months of age, 14 months of age, and after the dogs' performance in the guide dog program had been determined) and behavior twice (approximately 6 and 14 months of age). This study is the first to examine the relationship between behavior, lateralization, and cortisol concentrations in dogs. It implemented an objective and quantifiable assessment of behavior that may be of use to a variety of dog-focused stakeholders. Findings show that during the Juvenile testing period (6 months of age), dogs with higher cortisol concentrations were typically less able to rest when exposed to the unfamiliar testing room. Results from both Juvenile and Adult Test (14 months of age) periods showed that a greater |LI| and LI were associated with more confident and relaxed behavior when dogs were exposed to novel stimuli and unfamiliar environments. Significant elevations of cortisol concentrations were found at the completion of guide dog training when compared with results from the 2 prior test periods. This finding may reflect maturation or the effect of the prolonged kenneling which occurred during this period.
Keywords: Dog; temperament; motor lateralization; cortisol; behavior; cortisol EIA
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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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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.
Keywords: Detour behaviour; Equus caballus; Horses; Lateralization; Spatial reasoning
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Neveu, P. J. (1993). Brain Lateralization and Immunomodulation. Int J Neurosci, 70(1-2), 135–143.
Abstract: The two sides of the brain may be differently involved in the modulation of immune responses as demonstrated by lesional and behavioral approaches in rodents. Lesions of right or left neocortex induced opposite effects on various immune parameters including mitogen-induced lymphoproliferation, interleukin-2 production, macrophage activation or natural killer cell activity. This animal model, useful to elucidate whereby the brain and the immune system can communicate, appears to be suitable for studying the immune perturbations observed during stroke in humans. Brain asymmetry in modulation of immune reactivity may also be demonstrated in intact animal using a behavioral paradigm. The direction of a lateralized motor behavior ie paw preference in a food reaching task, correlated with an asymmetrical brain organization, was shown to be associated with lymphocyte reactivity, natural killer cell activity and auto-antibody production. The association between paw preference and immune reactivity in mice varies according to the immune parameters tested and is a sex-dependent phenomenon in which genetic background may be involved. The experimental models for investigating asymmetrical brain modulation of the immune system should be useful for studying several physiological, pathological and genetic aspects of neuroimmunomodulation.
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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.
Keywords: Lateralization; Laterality; Brain asymmetry; Hemisphere; Sheep; Lamb; Strength of lateralization
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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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