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Siniscalchi, M., Sasso, R., Pepe, A. M., Dimatteo, S., Vallortigara, G., & Quaranta, A. (). Sniffing with the right nostril: lateralization of response to odour stimuli by dogs. Anim. Behav., In Press, Corrected Proof.
Abstract: Lateralization in dogs, Canis familiaris, has been reported for paw usage and response to visual and acoustic stimuli. Surprisingly, however, no investigation of possible lateralization for the most relevant sensory domain of dogs, namely olfaction, has been carried out. Here we investigated left and right nostril use in dogs freely sniffing different emotive stimuli in unrestrained conditions. When sniffing novel nonaversive stimuli (food, lemon, vaginal secretion and cotton swab odours), dogs showed initial preferential use of the right nostril and then a shift towards use of the left nostril with repeated stimulus presentation. When sniffing arousal stimuli such as adrenaline and veterinary sweat odorants, dogs showed a consistent right nostril bias all over the series of stimulus presentations. Results suggest initial involvement of the right hemisphere in processing of novel stimuli followed by the left hemisphere taking charge of control of routine behaviour. Sustained right nostril response to arousal stimuli appears to be consistent with the idea that the sympathetic hypothalamic-pituitary-adrenal axis is mainly under the control of the right hemisphere. The implications of these findings for animal welfare are discussed.
Keywords: animal welfare; Canis familiaris; dog; emotion; laterality olfaction; physiology
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Deuel, N. R., & Lawrence, L. M. (1987). Laterality in the gallop gait of horses. J Biomech, 20(6), 645–649.
Abstract: Bilateral asymmetry in gallop stride limb contact patterns of four Quarter Horse fillies was documented by high-speed cinematography. Horses were filmed with rider by two cameras simultaneously while galloping along a straightaway. Even though signaled for each gallop lead an equivalent number of times, horses frequently switched leads, selecting the left lead nearly twice as often as the right. Velocities and stride lengths were greater for the left lead than the right, but stride frequencies did not differ between leads. Velocity effects were partitioned out in limb contact data analysis to enable the determination of persistent gallop stride asymmetries. The contact duration for the trailing (right) fore limb on the left lead exceeded the contact duration for the trailing (left) fore limb on the right lead. Selecting the right fore limb as the trailing fore limb may have allowed horses to use it to withstand the greater stresses and caused them to preferentially gallop with the left fore limb leading. Laterality may have an important influence on equine gallop motion patterns and thereby influence athletic performance.
Keywords: Animals; *Functional Laterality; *Gait; Horses/*physiology; Kinesis
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Parr, L. A., Hopkins, W. D., & de Waal, F. B. (1997). Haptic discrimination in capuchin monkeys (Cebus apella): evidence of manual specialization. Neuropsychologia, 35(2), 143–152.
Abstract: Two experiments investigated the effects of haptic and visual discrimination on hand preference in 22 brown capuchin monkeys (Cebus apella). The percentage of left-handed subjects in Experiment 1 were 63.6%, 45.5%, and 18.2% for haptic, bipedal, and quadrupedal reaching, respectively. In Experiment 2, the haptic demands of the task were manipulated by using additional food types and another tactile medium. Left-hand preferences were further strengthened when reaching into water compared to pineshavings in Experiment 1. Reaching with no tactile interference resulted in equal numbers of lateralized and nonlateralized subjects. These results show that when reaching demands the use of haptic cues, as opposed to visual ones, monkeys shift towards greater left hand use. This is consistent with what is known about right hemisphere superiority for haptic discrimination in humans.
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Ruggieri, V. (1999). The running horse stops: the hypothetical role of the eyes in imagery of movement. Percept Mot Skills, 89(3 Pt 2), 1088–1092.
Abstract: To examine the hypothetical role of the eyes in visual mental imagery of movement 72 undergraduate women students in psychology were asked to imagine a running horse and then to produce the same mental image without moving the eyes and the head. In 59% of the subjects interesting modifications of the imagined movement appeared: 37% observed an inhibition of the movement and 19% an evident slowing up of the moving figure. The interpretation of this result was made by hypothesizing that the eyes are concretely involved in visual imagery processes.
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Sankey, C., Henry, S., Clouard, C., Richard-Yris, M. - A., & Hausberger, M. (2011). Asymmetry of behavioral responses to a human approach in young naive vs. trained horses. Physiology & Behavior, 104(3), 464–468.
Abstract: The aim of this study was to investigate the impact of training experience on young horses (Equus caballus)’ lateralized responses to an approaching human. The results show that the one year old untrained horses display asymmetrical responses to an approaching human, with more negative reactions (escapes, threats) when approached from the left side, while approaches towards the right shoulder elicited more positive behaviors. On the contrary, two years old trained horses reacted equally positively to approaches and contact on both sides. Our findings support those of previous studies investigating a link between emotionality and laterality and confirm the role of the left hemisphere in the processing of novel or negative stimuli. Moreover, the data underline the impact work and training can have on this laterality in horses.
Keywords: Laterality; Emotionality; Human approach; Horse
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Lucidi, P., Bacco, G., Sticco, M., Mazzoleni, G., Benvenuti, M., Bernabò, N., et al. (2013). Assessment of motor laterality in foals and young horses (Equus caballus) through an analysis of derailment at trot. Physiol. Behav., 109, 8–13.
Abstract: The conflicting results regarding the study of motor laterality in horses may indicate that there does not exist a proper method to assess the degree and the direction of motor bias in these animals. Unfortunately, even less is known about the development of laterality in horses, and to what extent early manipulations can still exert their effects in adulthood. We propose a new method that can be easily applied at a very early age thus avoiding testing adult horses eventually biased by human handling and/or training. Forty-six horses (29 nine-month-old foals and 17 two-year old horses) were handled since birth bilaterally and housed in groups in wide areas. At the time of the analysis, in order to minimize environmental and sensorial disturbances, each horse was tested in a round pen individually or as dyad mother-foal. The ability/inability to properly execute a circle at trot was then recorded, assuming the direction of derailment, i.e. the cutting of the circle, as an indicator of motor bias. From the results of the study it is arguable that motor laterality in horses is acquired over time: in fact foals tested while their mothers were being subjected to longeing showed a higher percentage of ambidextrous animals, while two-year-old horses appeared biased toward the right (p<0.05). Results are discussed in the light of the scientific knowledge about equine biomechanics, taking into account horses' locomotion that leads to the advancement of the body mass through the activation of a kinetic chain that originates from the hindquarters.
Keywords: Horse; Laterality; Longeing; Sidedness
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Rogers, L. J. (2000). Evolution of hemispheric specialization: advantages and disadvantages. Brain Lang, 73(2), 236–253.
Abstract: Lateralization of the brain appeared early in evolution and many of its features appear to have been retained, possibly even in humans. We now have a considerable amount of information on the different forms of lateralization in a number of species, and the commonalities of these are discussed, but there has been relatively little investigation of the advantages of being lateralized. This article reports new findings on the differences between lateralized and nonlateralized chicks. The lateralized chicks were exposed to light for 24 h on day 19 of incubation, a treatment known to lead to lateralization of a number of visually guided responses, and the nonlateralized chicks were incubated in the dark. When they were feeding, the lateralized chicks were found to detect a stimulus resembling a raptor with shorter latency than nonlateralized chicks. This difference was not a nonspecific effect caused by the light-exposed chicks being more distressed by the stimulus. Instead, it appears to be a genuine advantage conferred by having a lateralized brain. It is suggested that having a lateralized brain allows dual attention to the tasks of feeding (right eye and left hemisphere) and vigilance for predators (left eye and right hemisphere). Nonlateralized chicks appear to perform these dual tasks less efficiently than lateralized ones. Reference is made to other species in discussing these results.
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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.
Keywords: Dog; Laterality; Vision; Behaviour; Physiology; Cognition; Emotion; Animal welfare
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Sakai, M., Hishii, T., Takeda, S., & Kohshima, S. (2006). Laterality of flipper rubbing behaviour in wild bottlenose dolphins (Tursiops aduncus): Caused by asymmetry of eye use? Behav. Brain. Res., 170(2), 204–210.
Abstract: To determine whether wild Indo-Pacific bottlenose dolphins (Tursiops aduncus) at Mikura Island, Japan, show asymmetry of eye or flipper use during a social behaviour, we investigated the laterality of flipper-to-body (F-B) rubbing, in which one dolphin (“rubber”) rubs the body of another (“rubbee”) with its flipper. We analysed 382 episodes of video-recorded F-B rubbings performed by identified individuals (N = 111 rubbers). F-B rubbing was conducted significantly more frequently with the left flipper than with the right flipper. The duration of F-B rubbings was also significantly longer with the left flipper than with the right flipper. Of 20 dolphins, nine individuals showed significant left-side bias as the rubber in this behaviour, whereas no dolphins showed significant right-side bias. The results indicate a population-level left-side bias of the rubber in F-B rubbing. An analysis of the swimming configurations during this behaviour suggests that the asymmetry in F-B rubbing was caused not only by the laterality of the rubber, but by a preference for use of the left eye in both dolphins during this behaviour. Dolphins used the left eye significantly more frequently than the right eye during the inquisitive behaviour, while they showed no significant bias in flipper use during the object-carrying behaviour. These facts also suggest that the asymmetry of F-B rubbing is caused by the preference for using the left eye. Significant left-side bias was observed only in F-B rubbings initiated by the rubbee, in which the rubbee determined its position during this behaviour. This suggests that this behavioural asymmetry was enhanced by the rubbees choosing the left side of the rubber to ensure better and longer rubs.
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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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