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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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Ghirlanda, S., Frasnelli, E., & Vallortigara, G. (2009). Intraspecific competition and coordination in the evolution of lateralization. Phil. Trans. Biol. Sci., 364(1519), 861–866.
Abstract: Recent studies have revealed a variety of left–right asymmetries among vertebrates and invertebrates. In many species, left- and right-lateralized individuals coexist, but in unequal numbers (‘population-level’ lateralization). It has been argued that brain lateralization increases individual efficiency (e.g. avoiding unnecessary duplication of neural circuitry and reducing interference between functions), thus counteracting the ecological disadvantages of lateral biases in behaviour (making individual behaviour more predictable to other organisms). However, individual efficiency does not require a definite proportion of left- and right-lateralized individuals. Thus, such arguments do not explain population-level lateralization. We have previously shown that, in the context of prey–predator interactions, population-level lateralization can arise as an evolutionarily stable strategy when individually asymmetrical organisms must coordinate their behaviour with that of other asymmetrical organisms. Here, we extend our model showing that populations consisting of left- and right-lateralized individuals in unequal numbers can be evolutionarily stable, based solely on strategic factors arising from the balance between antagonistic (competitive) and synergistic (cooperative) interactions.
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Ghirlanda, S., & Vallortigara, G. (2004). The evolution of brain lateralization: a game-theoretical analysis of population structure. Proceedings of the Royal Society of London. Series B: Biological Sciences, 271(1541), 853–857.
Abstract: In recent years, it has become apparent that behavioural and brain lateralization at the population level is the rule rather than the exception among vertebrates. The study of these phenomena has so far been the province of neurology and neuropsychology. Here, we show how such research can be integrated with evolutionary biology to understand lateralization more fully. In particular, we address the fact that, within a species, left– and right–type individuals often occur in proportions different from one–half (e.g. hand use in humans). The traditional explanations offered for lateralization of brain function (that it may avoid unnecessary duplication of neural circuitry and reduce interference between functions) cannot account for this fact, because increased individual efficiency is unrelated to the alignment of lateralization at the population level. A further puzzle is that such an alignment may even be disadvantageous, as it makes individual behaviour more predictable to other organisms. Here, we show that alignment of the direction of behavioural asymmetries in a population can arise as an evolutionarily stable strategy when individual asymmetrical organisms must coordinate their behaviour with that of other asymmetrical organisms. Brain and behavioural lateralization, as we know it in humans and other vertebrates, may have evolved under basically ‘social’ selection pressures.
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Siniscalchi, M., Sasso, R., Pepe, A. M., Vallortigara, G., & Quaranta, A. (2010). Dogs turn left to emotional stimuli. Behav. Brain. Res., 208(2), 516–521.
Abstract: During feeding behaviour, dogs were suddenly presented with 2D stimuli depicting the silhouette of a dog, a cat or a snake simultaneously into the left and right visual hemifields. A bias to turn the head towards the left rather than the right side was observed with the cat and snake stimulus but not with the dog stimulus. Latencies to react following stimulus presentation were lower for left than for right head turning, whereas times needed to resume feeding behaviour were higher after left rather than after right head turning. When stimuli were presented only to the left or right visual hemifields, dogs proved to be more responsive to left side presentation, irrespective of the type of stimulus. However, cat and snake stimuli produced shorter latencies to react and longer times to resume feeding following left rather than right monocular visual hemifield presentation. Results demonstrate striking lateralization in dogs, with the right side of the brain more responsive to threatening and alarming stimuli. Possible implications for animal welfare are discussed.
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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.
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Quaranta, A., Siniscalchi, M., Frate, A., & Vallortigara, G. (2004). Paw preference in dogs: relations between lateralised behaviour and immunity. Behavioural Brain Research, 153(2), 521–525.
Abstract: Paw use in a task consisting of the removal of a piece of adhesive paper from the snout was investigated in 80 mongrel and pure-bred domestic dogs (Canis familiaris). Population lateralisation was observed, but in opposite directions in the two sexes (animals were not desexed): males preferentially used their left paw, females their right paw. The relationship between immune function and paw preference was then investigated. Some immune parameters (total number of white blood cells including lymphocytes, granulocytes and monocytes; leukocyte formula; total proteins; γ-globulins) were investigated in a sample of left-pawed (n=6), right-pawed (n=6) and ambidextrous (n=6) dogs. The results showed that the percentage of lymphocytes was higher in left-pawed than in right-pawed and ambidextrous dogs, whereas granulocytes percentage was lower in left-pawed than in right-pawed and ambidextrous dogs. Moreover, total number of lymphocytes cells was higher in left-pawed than in right-pawed and ambidextrous dogs, whereas the number of γ-globulins was lower in left-pawed than in right-pawed and ambidextrous dogs. These findings represent the first evidence that brain asymmetry modulates immune responses in dogs.
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Quaresmini, C., Forrester, G. S., Spiezio, C., & Vallortigara, G. (2014). Social environment elicits lateralized behaviors in gorillas (Gorilla gorilla gorilla) and chimpanzees (Pan troglodytes). Journal of Comparative Psychology, 128(3), 276–284.
Abstract: The influence of the social environment on lateralized behaviors has now been investigated across a wide variety of animal species. New evidence suggests that the social environment can modulate behavior. Currently, there is a paucity of data relating to how primates navigate their environmental space, and investigations that consider the naturalistic context of the individual are few and fragmented. Moreover, there are competing theories about whether only the right or rather both cerebral hemispheres are involved in the processing of social stimuli, especially in emotion processing. Here we provide the first report of lateralized social behaviors elicited by great apes. We employed a continuous focal animal sampling method to record the spontaneous interactions of a captive zoo-living colony of chimpanzees (Pan troglodytes) and a biological family group of peer-reared western lowland gorillas (Gorilla gorilla gorilla). We specifically focused on which side of the body (i.e., front, rear, left, right) the focal individual preferred to keep conspecifics. Utilizing a newly developed quantitative corpus-coding scheme, analysis revealed both chimpanzees and gorillas demonstrated a significant group-level preference for focal individuals to keep conspecifics positioned to the front of them compared with behind them. More interestingly, both groups also manifested a population-level bias to keep conspecifics on their left side compared with their right side. Our findings suggest a social processing dominance of the right hemisphere for context-specific social environments. Results are discussed in light of the evolutionary adaptive value of social stimulus as a triggering factor for the manifestation of group-level lateralized behaviors. (PsycINFO Database Record (c) 2016 APA, all rights reserved)
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Sovrano, V. A., Rainoldi, C., Bisazza, A., & Vallortigara, G. (1999). Roots of brain specializations: preferential left-eye use during mirror-image inspection in six species of teleost fish. Behav. Brain. Res., 106(1-2), 175–180.
Abstract: It has recently been reported that predator inspection is more likely to occur when a companion (i.e. the mirror image of the test animal) is visible on the left rather than on the right side of mosquitofish Gambusia holbrooki. This very unexpected outcome could be consistent with the hypothesis of a preferential use of the right eye during sustained fixation of a predator as well as of a preferential use of the left eye during fixation of conspecifics. We measured the time spent in monocular viewing during inspection of their own mirror images in females of six species of fish, belonging to different families--G. holbrooki, Xenotoca eiseni, Phoxinus phoxinus, Pterophyllum scalare, Xenopoecilus sarasinorum, and Trichogaster trichopterus. Results revealed a consistent left-eye preference during sustained fixation in all of the five species. Males of G. holbrooki, which do not normally show any social behaviour, did not exhibit any eye preferences during mirror-image inspection. We found, however, that they could be induced to manifest a left-eye preference, likewise females, if tested soon after capture, when some affiliative tendencies can be observed. These findings add to current evidence in a variety of vertebrate species for preferential involvement of structures located in the right side of the brain in response to the viewing of conspecifics.
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Ventolini, N., Ferrero, E. A., Sponza, S., Della Chiesa, A., Zucca, P., & Vallortigara, G. (2005). Laterality in the wild: preferential hemifield use during predatory and sexual behaviour in the black-winged stilt. Anim. Behav., 69(5), 1077–1084.
Abstract: We recorded preferential use of the left and right monocular visual field in black-winged stilts, Himantopus himantopus, during predatory pecking and during courtship and mating behaviour in a naturalistic setting. The stilts had a population-level preference for using their right monocular visual field before predatory pecking; pecks that followed right-hemifield detection were more likely to be successful than pecks that followed left-hemifield detection, as evinced by the occurrence of swallowing and shaking head movements after pecking. In contrast, shaking behaviour, a component of courtship displays, and copulatory attempts by males were more likely to occur when females were seen with the left monocular visual field. Asymmetric hemifield use observed in natural conditions raises interesting issues as to the costs and benefits of population-level behavioural lateralization in wild animals.
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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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