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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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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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Vallortigara, G., & Rogers, L. J. (2005). Survival with an asymmetrical brain: advantages and disadvantages of cerebral lateralization. Behav Brain Sci, 28(4), 575–89; discussion 589–633.
Abstract: Recent evidence in natural and semi-natural settings has revealed a variety of left-right perceptual asymmetries among vertebrates. These include preferential use of the left or right visual hemifield during activities such as searching for food, agonistic responses, or escape from predators in animals as different as fish, amphibians, reptiles, birds, and mammals. There are obvious disadvantages in showing such directional asymmetries because relevant stimuli may be located to the animal's left or right at random; there is no a priori association between the meaning of a stimulus (e.g., its being a predator or a food item) and its being located to the animal's left or right. Moreover, other organisms (e.g., predators) could exploit the predictability of behavior that arises from population-level lateral biases. It might be argued that lateralization of function enhances cognitive capacity and efficiency of the brain, thus counteracting the ecological disadvantages of lateral biases in behavior. However, such an increase in brain efficiency could be obtained by each individual being lateralized without any need to align the direction of the asymmetry in the majority of the individuals of the population. Here we argue that the alignment of the direction of behavioral asymmetries at the population level arises as an “evolutionarily stable strategy” under “social” pressures occurring when individually asymmetrical organisms must coordinate their behavior with the behavior of other asymmetrical organisms of the same or different species.
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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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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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Tommasi, L., & Vallortigara, G. (2000). Searching for the center: spatial cognition in the domestic chick (Gallus gallus). J Exp Psychol Anim Behav Process, 26(4), 477–486.
Abstract: Chicks learned to find food hidden under sawdust by ground-scratching in the central position of the floor of a closed arena. When tested inan arena of identical shape but a larger area, chicks searched at 2 different locations, one corresponding to the correct distance (i.e., center) in the smaller (training) arena and the other to the actual center of the test arena. When tested in an arena of the same shape but a smaller area, chicks searched in the center of it. These results suggest that chicks are able to encode information on the absolute and relative distance of the food from the walls of the arena. After training in the presence of a landmark located at the center of the arena, animals searched at the center even after the removal of the landmark. Marked changes in the height of the walls of the arena produced some displacement in searching behavior, suggesting that chicks used the angular size of the walls to estimate distances.
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Chiesa, A. D., Pecchia, T., Tommasi, L., & Vallortigara, G. (2006). Multiple landmarks, the encoding of environmental geometry and the spatial logics of a dual brain. Anim. Cogn., 9(4), 281–293.
Abstract: A series of place learning experiments was carried out in young chicks (Gallus gallus) in order to investigate how the geometry of a landmark array and that of a walled enclosure compete when disoriented animals could rely on both of them to re-orient towards the centre of the enclosure. A square-shaped array (four wooden sticks) was placed in the middle of a square-shaped enclosure, the two structures being concentric. Chicks were trained to ground-scratch to search for food hidden in the centre of the enclosure (and the array). To check for effects of array degradation, one, two, three or all landmarks were removed during test trials. Chicks concentrated their searching activity in the central area of the enclosure, but their accuracy was inversely contingent on the number of landmarks removed; moreover, the landmarks still present within the enclosure appeared to influence the shape of the searching patterns. The reduction in the number of landmarks affected the searching strategy of chicks, suggesting that they had focussed mainly on local cues when landmarks were present within the enclosure. When all the landmarks were removed, chicks searched over a larger area, suggesting an absolute encoding of distances from the local cues and less reliance on the relationships provided by the geometry of the enclosure. Under conditions of monocular vision, chicks tended to rely on different strategies to localize the centre on the basis of the eye (and thus the hemisphere) in use, the left hemisphere attending to details of the environment and the right hemisphere attending to the global shape.
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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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Zucca, P., Milos, N., & Vallortigara, G. (2007). Piagetian object permanence and its development in Eurasian jays (Garrulus glandarius). Anim. Cogn., 10(2), 243–258.
Abstract: Object permanence in Eurasian jays (Garrulus glandarius) was investigated using a complete version of the Uzgiris and Hunt scale 1. Nine hand-raised jays were studied, divided into two groups according to their different developmental stages (experiment 1, older jays: 2-3 months old, n = 4; experiment 2, younger jays: 15 days old, n = 5). In the first experiment, we investigated whether older jays could achieve piagetian stage 6 of object permanence. Tasks were administered in a fixed sequence (1-15) according to the protocols used in other avian species. The aim of the second experiment was to check whether testing very young jays before their development of “neophobia” could influence the achievement times of piagetian stages. Furthermore, in this experiment tasks were administered randomly to investigate whether the jays' achievement of stage 6 follows a fixed sequence related to the development of specific cognitive abilities. All jays tested in experiments 1 and 2 fully achieved piagetian stage 6 and no “A not B” errors were observed. Performance on visible displacement tasks was better than performance on invisible ones. The results of experiment 2 show that “neophobia” affected the response of jays in terms of achievement times; the older jays in experiment 1 took longer to pass all the tasks when compared with the younger, less neophobic, jays in experiment 2. With regard to the achieving order, jays followed a fixed sequence of acquisition in experiment 2, even if tasks were administered randomly, with the exception of one subject. The results of these experiments support the idea that piagetian stages of cognitive development exist in avian species and that they progress through relatively fixed sequences.
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