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Reddon, A. R., & Hurd, P. L. (2009). Acting unilaterally: Why do animals with strongly lateralized brains behave differently than those with weakly lateralized brains? Bioscience Hypotheses, 2(6), 383–387.
Abstract: Cerebral lateralization was once thought to be unique to humans, but is now known to be widespread among the vertebrates. Lateralization appears to confer cognitive advantages upon those that possess it. Despite the taxonomic ubiquity and described advantages of lateralization, substantial individual variation exists in all species. Individual variation in cerebral lateralization may be tied to individual variation in behaviour and the selective forces that act to maintain variation in behaviour may also act to maintain variation in lateralization. The mechanisms linking individual variation in the strength of cerebral lateralization to individual variation in behaviour remain obscure. We propose here a general hypothesis which may help to explain this link. We suggest that individuals with strong and weak lateralizations behave differently because of differences in the ability of one hemisphere to inhibit the functions of the other in each type of brain organization. We also suggest a specific mechanism involving the asymmetric epithalamic nucleus, the habenula. We conclude by discussing some predictions and potential tests of our hypothesis.
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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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Siniscalchi, M., Sasso, R., Pepe, A. M., Dimatteo, S., Vallortigara, G., & Quaranta, A. (2010). Catecholamine plasma levels following immune stimulation with rabies vaccine in dogs selected for their paw preferences. Neuroscience Letters, 476(3), 142–145.
Abstract: Epinephrine and norepinephrine plasma levels were assessed in dogs in relation to paw preference following an immune challenge with rabies vaccine. The results showed that both catecholamines increased after the vaccine administration, confirming the main role of the sympathetic nervous system in the modulation activity between the brain and the immune system. Moreover, ambidextrous dogs showed a significantly higher increase of epinephrine levels 8 days after immunization with respect to right- and left-pawed dogs, suggesting that the biological activity of this molecule could be key for a different immune response with regard to laterality.
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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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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.
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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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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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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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Marinsek, N. L., Gazzaniga, M. S., & Miller, M. B. (2016). Chapter 17 – Split-Brain, Split-Mind. In S. Laureys, O. Gosseries, & G. Tononi (Eds.), The Neurology of Conciousness (Second Edition) (pp. 271–279). San Diego: Academic Press.
Abstract: The corpus callosum anatomically and functionally connects the two cerebral hemispheres. Despite its important role in interhemispheric communication however, severing the corpus callosum produces few--if any--noticeable cognitive or behavioral abnormalities. Incredibly, split-brain patients do not report any drastic changes in their conscious experience even though nearly all interhemispheric communication ceases after surgery. Extensive research has shown that both hemispheres remain conscious following disconnection and the conscious experience of each hemisphere is private and independent of the other. Additionally, the conscious experiences of the hemispheres appear to be qualitatively different, such that the consciousness of the left hemisphere is more enriched than the right. In this chapter, we offer explanations as to why split-brain patients feel unified despite possessing dual conscious experiences and discuss how the divided consciousness of split-brain patients can inform current theories of consciousness.
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