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	Author	Austin, N.P.; Rogers, L.J.
	Title	Lateralization of agonistic and vigilance responses in Przewalski horses (Equus przewalskii)			Type	Journal Article
	Year	2014	Publication	Applied Animal Behaviour Science	Abbreviated Journal
	Volume	151	Issue		Pages	43-50
	Keywords	Behavioural lateralization: Eye preference; Limb preference; Aggression; Vigilance; Reactivity; Przewalski horses
	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.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0168-1591	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5768
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	Author	Siniscalchi, M.; McFarlane, J.R.; Kauter, K.G.; Quaranta, A.; Rogers, L.J.
	Title	Cortisol levels in hair reflect behavioural reactivity of dogs to acoustic stimuli			Type	Journal Article
	Year	2013	Publication	Research in Veterinary Science	Abbreviated Journal
	Volume	94	Issue	1	Pages	49-54
	Keywords	Dogs; Behaviour; Cortisol; Hair; Acoustic stimuli
	Abstract	Cortisol levels in hair samples were examined in fourteen domestic dogs and related to the dogs’ responses to different acoustic stimuli. Stimuli were playbacks of species-typical vocalizations recorded during three different situations (“disturbance”, “isolation” and “play” barks) and the sounds of a thunderstorm. Hair samples were collected at 9:00 h and 17:00 h two weeks after the behavioural tests. Results showed that behavioural reactivity to playback of the various stimuli correlates with cortisol levels in hair samples collected at 9:00 h, and the same was the case for the separate measures of behaviour (i.e. hiding, running away, seeking attention from the tester, panting and lowering of the body posture). Hence, levels of cortisol in hair appear to reflect the dog’s chronic state of emotional reactivity, or temperament.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0034-5288	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5833
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	Author	Austin, N.P.; Rogers, L.J.
	Title	Limb preferences and lateralization of aggression, reactivity and vigilance in feral horses, Equus caballus			Type	Journal Article
	Year	2012	Publication	Animal Behaviour	Abbreviated Journal	Anim. Behav.
	Volume	83	Issue	1	Pages	239-247
	Keywords	aggression; behavioural asymmetry; Equus caballus; eye preference; feral horse; limb preference; reactivity; side bias
	Abstract	Observational field studies were conducted on two remote populations of feral horses in Australia to determine whether lateralization is a characteristic of Equus caballus as a species or results from handling by humans. Group 1 had been feral for two to five generations and Group 2 for 10–20 generations. In both groups, left-side biases were present during agonistic interactions and in reactivity and vigilance. Therefore, as in other vertebrates, the right hemisphere appears to be specialized to control agonistic behaviour and responses to potential threats. The leftwards bias was stronger in measures of behaviour involving more aggression and reactivity. Preferences to place one forelimb in front of the other during grazing were also determined. No population bias of forelimb preference was found, suggesting that such limb preferences present in domestic horses may be entrained. Since stronger individual limb preferences were found in immature than in adult feral horses, limb preference may be modified by maturation or experience in the natural habitat. Stronger limb preference was associated significantly with elevated attention to the environment but only in younger feral horses. No sex differences in lateralization were found. The findings are evidence that horses show visual lateralization, as in other vertebrates, not dependent on handling by humans. Limb preference during grazing, by contrast, does appear to depend on experience.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0003-3472	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5651
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	Author	Robins, A.; Rogers, L.J.
	Title	Lateralized prey-catching responses in the cane toad, Bufo marinus: analysis of complex visual stimuli			Type	Journal Article
	Year	2004	Publication	Animal Behaviour.	Abbreviated Journal	Anim. Behav.
	Volume	68	Issue	4	Pages	767-775
	Keywords
	Abstract	We tested the responses of Bufo marinus to prey stimuli of varying visual complexity that were moved around the toads in either a clockwise or anticlockwise direction at 1.7 revolutions/min. Predatory responses directed at prey resembling an insect were frequent when the model insect moved clockwise across the visual midline into the right visual hemifield. In contrast, the toads tended to ignore such stimuli when they moved anticlockwise across the midline into the left hemifield. No such lateralization was found when a rectangular strip moved along its longest axis was presented in a similar way. The toads also directed more responses towards the latter stimulus than towards the insect prey. Hence, the results suggest that lateralized predatory responses occur for considered decisions on whether or not to respond to complex insect-like stimuli, but not for decisions on comparatively simple stimuli. We discuss similarities between the lateralized feeding responses of B. marinus and those of avian species, as support for the hypothesis that lateralized brain function in tetrapods may have arisen from a common lateralized ancestor.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0003-3472	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5365
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	Author	Rogers, L.J.
	Title	Relevance of brain and behavioural lateralization to animal welfare			Type	Journal Article
	Year	2010	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	127	Issue	1-2	Pages	1-11
	Keywords	Hemispheric specialization; Cognitive bias; Stress; Limb preference; Experience; Development
	Abstract	The left and right sides of the brain are specialised to process information in different ways and to control different categories of behaviour. Research on a range of species has shown that the left hemisphere controls well-established patterns of behaviour performed in non-stressful situations, whereas the right hemisphere responds to unexpected stimuli and controls escape and other emergency responses. The known functions of each hemisphere are summarised in this paper. Then it is hypothesised that stressed animals rely on predominant use of the right hemisphere, and that a bias to use the right or left hemisphere, respectively, may explain the behavioural differences between animals with a negative cognitive bias and those with a positive cognitive bias. In some species of primates it has been shown that the preferred limb used to pick up food when the animal is in a relaxed state reflects the dominant hemisphere and may be an accessible measure indicating susceptibility to stress and tendency towards positive versus negative cognitive bias. Hence, limb preference might be a useful measure of such tendencies in domesticated species. Some difficulties in determining a relevant measure of limb preference in non-primate species are mentioned, followed by the suggestion that eye preferences for viewing certain stimuli may be a useful measure in species with laterally placed eyes. Finally, effects of experience on the development of hemispheric dominance are discussed, leading to a suggestion that the welfare of domestic animals may be enhanced by ensuring development of left hemisphere dominance (e.g. by exposing chick embryos to light) and by shifting right to left hemisphere dominance in animals with negative cognitive bias.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0168-1591	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5296
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	Author	Vallortigara, G.; Rogers, L.J.
	Title	Survival with an asymmetrical brain: advantages and disadvantages of cerebral lateralization			Type	Journal Article
	Year	2005	Publication	The Behavioral and Brain Sciences	Abbreviated Journal	Behav Brain Sci
	Volume	28	Issue	4	Pages	575-89; discussion 589-633
	Keywords	Animals; Attention/physiology; Behavior/physiology; Behavior, Animal/physiology; Dominance, Cerebral/physiology; *Evolution; Humans; Models, Biological; Visual Perception/physiology
	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.
	Address	Department of Psychology and B.R.A.I.N. Centre for Neuroscience, University of Trieste, 34123 Trieste, Italy. vallorti@univ.trieste.it
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0140-525X	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:16209828			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	4622
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	Author	Rogers, L.J.
	Title	Evolution of hemispheric specialization: advantages and disadvantages			Type	Journal Article
	Year	2000	Publication	Brain and Language	Abbreviated Journal	Brain Lang
	Volume	73	Issue	2	Pages	236-253
	Keywords	Aggression/psychology; Animals; Behavior, Animal/physiology; Brain/physiology; Chickens/physiology; Evolution; Feeding Behavior/physiology; Functional Laterality/*physiology; Visual Fields/physiology; Visual Perception/physiology
	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.
	Address	Division of Zoology, University of New England, Armidale, New South Wales, Australia. lrogers@metz.une.edu.au
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language	English	Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	0093-934X	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:10856176			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	4621
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	Author	Rogers, L.J.
	Title	Advantages and disadvantages of lateralization			Type	Book Chapter
	Year	2002	Publication		Abbreviated Journal
	Volume		Issue		Pages	126-153
	Keywords
	Abstract
	Address
	Corporate Author				Thesis
	Publisher	Cambridge University Press	Place of Publication	New York	Editor	L. J. Rogers,; R. Andrew,
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN		ISBN	9780521781619	Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @ L.J.Rogers+R.Andrew(eds)2002			Serial	4624
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	Author	Rogers, L.J.
	Title	Early Experiential Effects on Laterality: Research on Chicks has Relevance to Other Species			Type	Journal Article
	Year	1997	Publication	Laterality	Abbreviated Journal	Laterality
	Volume	2	Issue	3-4	Pages	199-219
	Keywords
	Abstract	The influence of early experience on the development of lateralisation of hemispheric function was further investigated, using the chick as a model. A range of functions are lateralised in the chick and these correlate with asymmetry in the organisation of the visual projections. Chicks using the right eye and, therefore, primarily the left hemisphere are able to switch from pecking randomly at grain and pebbles to pecking mainly at grain, whereas those using the left eye and primarily the right hemisphere continue to peck at random. Exposure to light during the last days of incubation establishes this lateralisation in males, as a consequence of the embryo being oriented in the egg so that the left eye only is occluded. Males incubated in the dark peck at random when using either the right or left eye. Irrespective of light experience, females perform the same as darkincubated males: they are not influenced by light exposure. Monocular performance of the pebble-grain task is compared to binocular performance, and the sensitive period for the influence of light is delineated. The interactive effects of sex hormone levels on the differentiation of lateralisation are discussed and also the relevance of the results to other species, including humans.
	Address
	Corporate Author				Thesis
	Publisher	Psychology Press	Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN	1357-650x	ISBN		Medium
	Area		Expedition		Conference
	Notes	doi: 10.1080/713754277			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5587
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	Author	Rogers, L.J.
	Title	A Matter of Degree: Strength of Brain Asymmetry and Behaviour			Type
	Year	2017	Publication	Symmetry	Abbreviated Journal	Symmetry
	Volume		Issue		Pages
	Keywords	functional asymmetry; strength of lateralization; direction of lateralization; advantages; disadvantages; vertebrate species; limb preference; eye bias
	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.
	Address
	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title	Symmetry
	Series Volume	9	Series Issue	4	Edition
	ISSN	2073-8994	ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	6167
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