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Phillips, C. J. C., Oevermans, H., Syrett, K. L., Jespersen, A. Y., & Pearce, G. P. (2015). Lateralization of behavior in dairy cows in response to conspecifics and novel persons. Journal of Dairy Science, 98(4), 2389–2400.
Abstract: Abstract The right brain hemisphere, connected to the left eye, coordinates fight and flight behaviors in a wide variety of vertebrate species. We investigated whether left eye vision predominates in dairy cows’ interactions with other cows and humans, and whether dominance status affects the extent of visual lateralization. Although we found no overall lateralization of eye use to view other cows during interactions, cows that were submissive in an interaction were more likely to use their left eye to view a dominant animal. Both subordinate and older cows were more likely to use their left eye to view other cattle during interactions. Cows that predominantly used their left eye during aggressive interactions were more likely to use their left eye to view a person in unfamiliar clothing in the middle of a track by passing them on the right side. However, a person in familiar clothing was viewed predominantly with the right eye when they passed mainly on the left side. Cows predominantly using their left eyes in cow-to-cow interactions showed more overt responses to restraint in a crush compared with cows who predominantly used their right eyes during interactions (crush scores: left eye users 7.9, right eye users 6.4, standard error of the difference = 0.72). Thus, interactions between 2 cows and between cows and people were visually lateralized, with losing and subordinate cows being more likely to use their left eyes to view winning and dominant cattle and unfamiliar humans.
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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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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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Vallortigara, G., & Andrew, R. J. (1994). Differential involvement of right and left hemisphere in individual recognition in the domestic chick. Behav. Process., 33(1-2), 41–57.
Abstract: Right hemisphere advantage in individual recognition (as shown by differences between response to strangers and companions) is clear in the domestic chick. Chicks using the left eye (and so, thanks to the complete optic decussation, predominantly the right hemisphere) discriminate between stranger and companion. Chicks using the right eye discriminate less clearly or not at all. The ability of left eyed chicks to respond to differences between strangers and companions stimuli is associated with a more general ability to detect and respond to novelty: this difference between left and right eyed chicks also holds for stimuli which are not social partners. The right hemisphere also shows advantage in tasks with a spatial component (topographical learning; response to change in the spatial context of a stimulus) in the chick, as in humans. Similar specialisations of the two hemispheres are also revealed in tests which involve olfactory cues presented by social partners. The special properties of the left hemisphere are less well established in the chick. Evidence reviewed here suggests that it tends to respond to selected properties of a stimulus and to use them to assign it to a category; such assignment then allows an appropriate response. When exposed to an imprinting stimulus (visual or auditory) a chick begins by using right eye or ear (suggesting left hemisphere control), and then shifts to the left eye or ear (suggesting right hemisphere control), as exposure continues. The left hemisphere here is thus involved whilst behaviour is dominated by vigorous response to releasing stimuli presented by an object. Subsequent learning about the full detailed properties of the stimulus, which is crucial for individual recognition, may explain the shift to right hemisphere control after prolonged exposure to the social stimulus. There is a marked sex difference in choice tests: females tend to choose companions in tests where males choose strangers. It is possible that this difference is specifically caused by stronger motivation to sustain social contact in female chicks, for which there is extensive evidence. However, sex differences in response to change in familiar stimuli are also marked in tests which do not involve social partners. Finally, in both sexes there are two periods during development in which there age-dependent shifts in bias to use one or other hemisphere. These periods (days 3-5 and 8-11) coincide with two major changes in the social behaviour of chicks reared by a hen in a normal brood. It is argued that one function of these periods is to bring fully into play the hemisphere most appropriate to the type of response to, and learning about, social partners which is needed at particular points in development. Parallels are discussed between the involvement of lateralised processes in the recognition of social partners in chicks and humans.
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Tomkins, L. M., McGreevy, P. D., & Branson, N. J. (2010). Lack of standardization in reporting motor laterality in the domestic dog (Canis familiaris). Journal of Veterinary Behaviour, 5(5), 235–239.
Abstract: Over the past 2 decades, numerous studies have been undertaken to assess motor laterality in the domestic dog. In anticipation of growth in this area of enquiry, we decided to review the literature on canine motor biases to identify any shortcomings, reflect on the lessons to be learned from and offer ways forward for future research into canine laterality. The aim of this review is to (i) summarize motor laterality findings in the dog, (ii) highlight areas lacking in standardization, and (iii) propose necessary criteria for future tests and global reporting protocols. Our review of the literature highlighted the lack of standardization between studies in task selection, sample size, number of behavior scores recorded, and the methods by which motor laterality were classified and reported. This review illustrates the benefits of standardizing methods of motor laterality assessment so that comparisons can be made between the populations sampled. By adopting such an approach, researchers should mutually benefit as motor laterality data could then be compared and subjected to meta-analysis.
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Tomkins, L. M., Williams, K. A., Thomson, P. C., & McGreevy, P. D. (2010). Sensory Jump Test as a measure of sensory (visual) lateralization in dogs (Canis familiaris). Journal of Veterinary Behavior, 5(5), 256–267.
Abstract: Sensory lateralization in dogs (n = 74) was investigated in this study using our innovation, the Sensory Jump Test. This required the modification of head halters to create three different ocular treatments (binocular, right, and left monocular vision) for eye preference assessment in a jumping task. Ten jumps were recorded as a jump set for each treatment. Measurements recorded included (i) launch and landing paws, (ii) type of jump, (iii) approach distance, (iv) clearance height of the forepaw, hindpaw, and the lowest part of the body to clear the jump, and (v) whether the jump was successful. Factors significantly associated with these jump outcomes included ocular treatment, jump set number, and replication number. Most notably, in the first jump set, findings indicated a left hemispheric dominance for the initial navigation of the Sensory Jump Test, as left monocular vision (LMV) compromised of jumping more than right monocular (RMV) and binocular vision, with a significantly reduced approach distance and forepaw clearance observed in dogs with LMV. However, by the third jump set, dogs undergoing LMV launched from a greater approach distance and with a higher clearance height, corresponding to an increase in success rate of the jump, in comparison with RMV and binocular vision dogs. A marginally non-significant RMV bias was observed for eye preference based on the laterality indices for approach distance (P = 0.060) and lowest body part clearance height (P = 0.067). A comparison between eye preference and launching or landing paws showed no association between these measures of sensory and motor laterality. To our knowledge, this is the first study to report on sensory lateralization in the dog, and furthermore, to compare both motor and sensory laterality in dogs.
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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.
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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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Perez-Cruz, C., Simon, M., Czéh, B., Flügge, G., & Fuchs, E. (2009). Hemispheric differences in basilar dendrites and spines of pyramidal neurons in the rat prelimbic cortex: activity- and stress-induced changes. Eur. J. Neurosci., 29(4), 738–747.
Abstract: Abstract Pyramidal neurons of the rat medial prefrontal cortex have been shown to react to chronic stress by retracting their apical dendrites and by spine loss. We extended these findings by focusing on the basilar dendritic tree of layer III pyramidal neurons in both hemispheres of the rat prelimbic cortex. Animals were subjected to daily restraint stress for 1 week (6 h/day), during either the resting or the activity period. The morphology of basilar dendrites and spines of Golgi–Cox-stained neurons in the left and right hemispheres was digitally reconstructed and analyzed. We observed the following: (i) there was an inherent hemispheric asymmetry in control rats during the resting period: the number of spines on proximal dendrites was higher in the left than in the right hemisphere; (ii) basal dendrites in controls displayed a diurnal variation: there was more dendritic material during the resting period than in the activity period; (iii) chronic stress reduced the length of basal dendrites in only the right prelimbic cortex; (iv) chronic stress reduced spine density on proximal basal dendrites; (v) restraint stress during the activity period had more pronounced effects on the physiological stress parameters than restraint stress during the resting period. Our results show dynamic hemisphere-dependent structural changes in pyramidal neurons of the rat prelimbic cortex that are tightly linked to periods of resting and activity. These morphological alterations reflect the capacity of the neurons to react to external stimuli and mirror presumptive changes in neuronal communication.
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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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