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Zucca, P., Cerri, F., Carluccio, A., & Baciadonna, L. (). Space availability influence laterality in donkeys (Equus asinus). Behav. Process., In Press, Uncorrected Proof.
Abstract: Cerebral lateralization is the portioning of the cognitive functions between the two cerebral hemispheres. Several factors, like embryological manipulations, light exposure, health conditions, sex and age can influence the left-right brain asymmetries and contribute to increasing the variability in the strength and direction of laterality within most species. We investigated the influence of an environmental constraint, namely space availability, as a new source of variation on laterality in an adult vertebrate model, the donkey. In a baseline condition we tested whether donkeys show a motor lateralization bias at population level, while in an experimental condition we manipulated space availability to verify if a reduction in this parameter could represent a new source of variation in laterality. Results show that donkeys are lateralized at population level with a strong bias to standing with the right forelimb advanced over the left and that a reduction of space availability is an important source of variation in the laterality strength and direction within this species. The comparative analysis of the environmental and developmental factors that give origin to neural and behavioural laterality in animal models will be very important for a better understanding of the evolutionary origin of such multifaceted phenomenon.
Keywords: Cerebral lateralization; Donkey; Footedness; Welfare; Equus asinus
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Rogers, L. J. (2017). A Matter of Degree: Strength of Brain Asymmetry and Behaviour. Symmetry, 9(4).
Abstract: Research on a growing number of vertebrate species has shown that the left and right sides of the brain process information in different ways and that lateralized brain function is expressed in both specific and broad aspects of behaviour. This paper reviews the available evidence relating strength of lateralization to behavioural/cognitive performance. It begins by considering the relationship between limb preference and behaviour in humans and primates from the perspectives of direction and strength of lateralization. In birds, eye preference is used as a reflection of brain asymmetry and the strength of this asymmetry is associated with behaviour important for survival (e.g., visual discrimination of food from non-food and performance of two tasks in parallel). The same applies to studies on aquatic species, mainly fish but also tadpoles, in which strength of lateralization has been assessed as eye preferences or turning biases. Overall, the empirical evidence across vertebrate species points to the conclusion that stronger lateralization is advantageous in a wide range of contexts. Brief discussion of interhemispheric communication follows together with discussion of experiments that examined the effects of sectioning pathways connecting the left and right sides of the brain, or of preventing the development of these left-right connections. The conclusion reached is that degree of functional lateralization affects behaviour in quite similar ways across vertebrate species. Although the direction of lateralization is also important, in many situations strength of lateralization matters more. Finally, possible interactions between asymmetry in different sensory modalities is considered.
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Shen, Y. - Q., Hebert, G., Lin, L. - Y., Luo, Y. - L., Moze, E., Li, K. - S., et al. (2005). Interleukine-1β and interleukine-6 levels in striatum and other brain structures after MPTP treatment: influence of behavioral lateralization. Journal of Neuroimmunology, 158(1–2), 14–25.
Abstract: MPTP (N-methyl-4-phenyl-1,2,3,6-tetrahydropyridine) induces diminution of the dopamine in nigrostriatal pathway and cognitive deficits in mice. MPTP treatment also increases pro-inflammatory cytokine production in substantia nigra and striatum. Since, pro-inflammatory cytokines influence striatal dopamine content and provoke cognitive impairments, the cognitive defects induced by MPTP may be partly due to brain cytokine induction in other structures than nigrostriatal pathway. Furthermore, behavioral lateralization, as assessed by paw preference, influences cytokine production at the periphery and in the central nervous system. Behavioral lateralization may thus influence brain cytokine levels after MPTP. In order to address these issues, mice selected for paw preference were injected with 25 mg/kg MPTP i.p. for five consecutive days after which striatal dopamine and DOPAC contents were measured by HPLC and IL-1β and IL-6 quantified by ELISA in the striatum, cerebral cortex, hippocampus and hypothalamus. The results showed that MPTP treatment induced dramatic loss of DA in striatum, simultaneously, IL-6 levels decreased in the striatum and increased in hippocampus and hypothalamus, while IL-1β levels decreased in the striatum, cerebral cortex and hippocampus. Interestingly, striatal dopamine turnover under basal conditions as well as striatal IL-1β and IL-6 levels under basal conditions and after MPTP depended on behavioral lateralization. Left pawed mice showed a higher decrease in dopamine turnover and lower cytokine levels as compared to right pawed animals. Behavioral lateralization also influenced IL-6 hippocampal levels under basal conditions and IL-1β cortical levels after MPTP. From these results, it can be concluded that MPTP-induced cognitive defects are accompanied by an alteration of pro-inflammatory cytokine levels in brain structures other than those involved in the nigrostriatal pathway. In addition, MPTP-induced dopamine decrease is influenced by behavioral lateralization, possibly through an effect on brain cytokine levels.
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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.
Keywords: Right hemisphere; Left hemisphere; Domestic fowl; Lateralization; Chick
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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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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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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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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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Branson, N. J., & Rogers, L. J. (2006). Relationship between paw preference strength and noise phobia in Canis familiaris. J. Comp. Psychol., 120(3), 176–183.
Abstract: The authors investigated the relationship between degree of lateralization and noise phobia in 48 domestic dogs (Canis familiaris) by scoring paw preference to hold a food object and relating it to reactivity to the sounds of thunderstorms and fireworks, measured by playback and a questionnaire. The dogs without a significant paw preference were significantly more reactive to the sounds than the dogs with either a left-paw or right-paw preference. Intense reactivity, therefore, is associated with a weaker strength of cerebral lateralization. The authors note the similarity between their finding and the weaker hand preferences shown in humans suffering extreme levels of anxiety and suggest neural mechanisms that may be involved. (PsycINFO Database Record (c) 2010 APA, all rights reserved)
Keywords: noise phobia; lateralization; paw preference; dog; fear
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