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	Author	Ghirlanda, S.; Frasnelli, E.; Vallortigara, G.
	Title	Intraspecific competition and coordination in the evolution of lateralization			Type	Journal Article
	Year	2009	Publication	Philosophical Transactions of the Royal Society B: Biological Sciences	Abbreviated Journal	Phil. Trans. Biol. Sci.
	Volume	364	Issue	1519	Pages	861-866
	Keywords
	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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	Corporate Author				Thesis
	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
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	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5346
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	Author	Ghirlanda, S.; Vallortigara, G.
	Title	The evolution of brain lateralization: a game-theoretical analysis of population structure			Type	Journal Article
	Year	2004	Publication	Proceedings of the Royal Society of London. Series B: Biological Sciences	Abbreviated Journal
	Volume	271	Issue	1541	Pages	853-857
	Keywords
	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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	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		ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5345
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	Author	Hamilton, C.R.; Vermeire, B.A.
	Title	Complementary hemispheric specialization in monkeys			Type	Journal Article
	Year	1988	Publication	Science	Abbreviated Journal	Science
	Volume	242	Issue	4886	Pages	1691-1694
	Keywords
	Abstract	Twenty-five split-brain monkeys were taught to discriminate two types of visual stimuli that engage lateralized cerebral processing in human subjects. Differential lateralization for the two kinds of discriminations was found; the left hemisphere was better at distinguishing between tilted lines and the right hemisphere was better at discriminating faces. These results indicate that lateralization of cognitive processing appeared in primates independently of language or handedness. In addition, cerebral lateralization in monkeys may provide an appropriate model for studying the biological basis of hemispheric specialization.
	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		ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5342
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	Author	Tommasi, L.
	Title	Mechanisms and functions of brain and behavioural asymmetries			Type	Journal Article
	Year	2009	Publication	Philosophical Transactions of the Royal Society B: Biological Sciences	Abbreviated Journal	Phil. Trans. Biol. Sci.
	Volume	364	Issue	1519	Pages	855-859
	Keywords
	Abstract	For almost a century the field of brain and behavioural asymmetries has been dominated by studies on humans, resting on the evidence that the anatomical structures underlying language functions are asymmetrical, and that human handedness is lateralized at the population level. Today, there is not only evidence of population-level lateralization of brain and behaviour across a variety of vertebrate and invertebrate species, but also a growing consensus that the comparative analysis of the environmental and developmental factors that give origin to neural and behavioural laterality in animal models, together with theoretical analyses of their costs and benefits, will be crucial for understanding the evolutionary pathways that led to such a multifaceted phenomenon. The present theme issue provides a survey of theoretical, review and research work cutting across the biological and the cognitive sciences, focusing on various species of fishes, birds and primates (including humans) and emphasizing an integrative approach to the study of lateralization encompassing neural, behavioural, cognitive, developmental and environmental aspects.
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	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN		ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5340
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	Author	Weisbecker, V.; Goswami, A.
	Title	Brain size, life history, and metabolism at the marsupial/placental dichotomy			Type	Journal Article
	Year	2010	Publication	Proceedings of the National Academy of Sciences of the United States of America	Abbreviated Journal	Proc. Natl. Acad. Sci. U.S.A.
	Volume	107	Issue	37	Pages	16216-16221
	Keywords
	Abstract	The evolution of mammalian brain size is directly linked with the evolution of the brain's unique structure and performance. Both maternal life history investment traits and basal metabolic rate (BMR) correlate with relative brain size, but current hypotheses regarding the details of these relationships are based largely on placental mammals. Using encephalization quotients, partial correlation analyses, and bivariate regressions relating brain size to maternal investment times and BMR, we provide a direct quantitative comparison of brain size evolution in marsupials and placentals, whose reproduction and metabolism differ extensively. Our results show that the misconception that marsupials are systematically smaller-brained than placentals is driven by the inclusion of one large-brained placental clade, Primates. Marsupial and placental brain size partial correlations differ in that marsupials lack a partial correlation of BMR with brain size. This contradicts hypotheses stating that the maintenance of relatively larger brains requires higher BMRs. We suggest that a positive BMR–brain size correlation is a placental trait related to the intimate physiological contact between mother and offspring during gestation. Marsupials instead achieve brain sizes comparable to placentals through extended lactation. Comparison with avian brain evolution suggests that placental brain size should be constrained due to placentals’ relative precociality, as has been hypothesized for precocial bird hatchlings. We propose that placentals circumvent this constraint because of their focus on gestation, as opposed to the marsupial emphasis on lactation. Marsupials represent a less constrained condition, demonstrating that hypotheses regarding placental brain size evolution cannot be generalized to all mammals.
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	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN		ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5338
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	Author	Finarelli, J.A.; Flynn, J.J.
	Title	Brain-size evolution and sociality in Carnivora			Type	Journal Article
	Year	2009	Publication	Proceedings of the National Academy of Sciences of the United States of America	Abbreviated Journal	Proc. Natl. Acad. Sci. U.S.A.
	Volume	106	Issue	23	Pages	9345-9349
	Keywords
	Abstract	Increased encephalization, or larger brain volume relative to body mass, is a repeated theme in vertebrate evolution. Here we present an extensive sampling of relative brain sizes in fossil and extant taxa in the mammalian order Carnivora (cats, dogs, bears, weasels, and their relatives). By using Akaike Information Criterion model selection and endocranial volume and body mass data for 289 species (including 125 fossil taxa), we document clade-specific evolutionary transformations in encephalization allometries. These evolutionary transformations include multiple independent encephalization increases and decreases in addition to a remarkably static basal Carnivora allometry that characterizes much of the suborder Feliformia and some taxa in the suborder Caniformia across much of their evolutionary history, emphasizing that complex processes shaped the modern distribution of encephalization across Carnivora. This analysis also permits critical evaluation of the social brain hypothesis (SBH), which predicts a close association between sociality and increased encephalization. Previous analyses based on living species alone appeared to support the SBH with respect to Carnivora, but those results are entirely dependent on data from modern Canidae (dogs). Incorporation of fossil data further reveals that no association exists between sociality and encephalization across Carnivora and that support for sociality as a causal agent of encephalization increase disappears for this clade.
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	Publisher		Place of Publication		Editor
	Language		Summary Language		Original Title
	Series Editor		Series Title		Abbreviated Series Title
	Series Volume		Series Issue		Edition
	ISSN		ISBN		Medium
	Area		Expedition		Conference
	Notes				Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5337
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	Author	Leiner, L.; Fendt, M.
	Title	Behavioural fear and heart rate responses of horses after exposure to novel objects: Effects of habituation			Type	Journal Article
	Year	2011	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	131	Issue	3-4	Pages	104-109
	Keywords	Anxiety; Avoidance; Behavioural test; Emotion; Fear; Flight; Habituation; Horse; Vocalization
	Abstract	The emotion fear promotes the fitness of wild animals. In a farm environment, exaggerated fear, e.g., in horses, can cause several problems. Therefore, knowledge about fear in horses helps to prevent or to handle potential fear-inducing situations. The present study investigated which behavioural fear responses can be observed during exposure of horses to a novel stimulus, whether these behavioural responses are correlated with physiological changes, and whether and how specifically these changes are reduced after habituation training to one of the novel objects. Our data shows that behavioural and physiological fear responses in horses are correlated, are reliable to observe and to measure, and appear in a typical chronological order. Furthermore, after habituation-training to an object, the fear response to this object is specifically attenuated whereas the fear response to another object remains.
	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	5332
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	Author	Laister, S.; Stockinger, B.; Regner, A.-M.; Zenger, K.; Knierim, U.; Winckler, C.
	Title	Social licking in dairy cattle--Effects on heart rate in performers and receivers			Type	Journal Article
	Year	2011	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	130	Issue	3-4	Pages	81-90
	Keywords	Dairy cows; Allogrooming; Affiliative social behaviour; Cardiac activity; Affective states
	Abstract	Using heart rate (HR) measurements we investigated whether potential calming effects of social licking were evident for both active (performers) and passive (receivers) licking partners. A HR decline was assumed to indicate relaxation and thus the experience of positive emotions. Effects of the licking category (spontaneous, solicited), the animals' basic activity (standing, lying) and the licked body region (head, neck, rest) were also considered. Two studies (A, B) were carried out in the same loose housed Austrian Simmental dairy herd. HR was recorded in up to 20 focal animals on 16 and 18 days, respectively. Using either direct observations (A) or video recordings (B), social licking interactions were continuously observed. The cow's basic activity was recorded using scan sampling at 5 min intervals. Linear mixed effects models were applied separately for Study A and B in order to compare the mean HR of the licking bouts with the mean of the respective 5 min pre- and post-licking periods. In receivers we found a significant calming effect in terms of a HR decline during allogrooming in both studies (A: -1.3 beats per minute, B: -1.1 bpm). This effect was more pronounced when animals were standing (A/B: -2.4 bpm/-3.8 bpm). However, it was not affected by the licked body region. In dairy cows performing social licking, we did not find an overall calming effect. On the contrary, in Study B, HR significantly increased during licking in lying performers (+2.5 bpm). This reaction was even stronger, when licking was directed to the receivers' head (+3.5 bpm) or neck (+3.0 bpm) as compared to the rest of the body (+1.4 bpm). The licking category had no effect on HR changes during the licking events. Our findings suggest that relaxation effects induced by social licking differ between performers and receivers and are affected by the cows' basic activity. In receivers, there were clear indications of a calming effect implying the experience of positive affective states. In performers, such calming effects during social licking were not identified.
	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	5331
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	Author	Lee, J.; Floyd, T.; Erb, H.; Houpt, K.
	Title	Preference and demand for exercise in stabled horses			Type	Journal Article
	Year	2011	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	130	Issue	3-4	Pages	91-100
	Keywords	Horse; Welfare; Exercise; Operant conditioning; Two choice preference; Treadmill
	Abstract	Operant conditioning and two choice preference tests were used to assess the motivation of horses to be released from straight and from box stalls. The motivations for food, a companion, and release into a paddock were compared when the horses had to work for each commodity at increasing fixed ratios of responses (panel presses) to reward in an equine operant conditioning stall. The motivation for food (mean ± SEM = 258 ± 143) responses was much greater than that for either release (38 ± 32) from a straight stall into a large paddock alone or into a small paddock with another horse (95 ± 41) (P = 0.04). When given a two choice preference test between exercise on a treadmill for 20 min or returning to their box stalls, eight of nine horses chose to return to their stalls. In a two choice preference test six of eight horses in box stalls chose to be released into a paddock alone. Horses were given a series of two choice preference tests to determine how long they preferred to be in a paddock. After 15 min in the paddock the horses were re-tested, but all chose the paddock when released into a paddock with three other horses. They were retested every 15 min until they chose to return to their stalls. They chose to stay out for 35 ± 6 min when other horses were in the paddock but for only 17 ± 2 min when they would be alone. When deprived of stall release for 48 h the horses chose to remain in the paddock with other horses for 54 ± 6 min, but showed no compensatory behavior when they were alone (duration chosen = 16 ± 4 min). These findings indicate that horses are not strongly motivated to exercise alone and will choose not to endure forced exercise on a treadmill. The social context of voluntary exercise is important; horses are willing to stay out of their stalls longer if other horses are present and will show compensatory behavior only if other horses are present. These finding have implications for optimizing turnout time for stalled horses.
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	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	5330
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	Author	Christensen, J.W.; Zharkikh, T.; Chovaux, E.
	Title	Object recognition and generalisation during habituation in horses			Type	Journal Article
	Year	2011	Publication	Applied Animal Behaviour Science	Abbreviated Journal	Appl. Anim. Behav. Sci.
	Volume	129	Issue	2-4	Pages	83-91
	Keywords	Horse; Habituation; Object recognition; Generalisation
	Abstract	The ability of horses to habituate to frightening stimuli greatly increases safety in the horse-human relationship. A recent experiment suggested, however, that habituation to frightening visual stimuli is relatively stimulus-specific in horses and that shape and colour are important factors for object generalisation (Christensen et al., 2008). In a series of experiments, we aimed to further explore the ability of horses (n = 30, 1 and 2-year-old mares) to recognise and generalise between objects during habituation. TEST horses (n = 15) were habituated to a complex object, composed of five simple objects of varying shape and colour, whereas CONTROL horses (n = 15) were habituated to the test arena, but not to the complex object. In the first experiment, we investigated whether TEST horses subsequently reacted less to i) simple objects that were previously part of the complex object (i.e. testing for object recognition) and ii) a novel object (new shape and colour, i.e. testing for object generalisation), compared to CONTROLS. In the second experiment we investigated whether TEST horses reacted to a change in object order and object location. Behavioural reactions to the object, latency to eat, total eating time and heart rate were recorded. Compared to CONTROLS, TEST horses reacted significantly less towards objects, which were previously part of the complex object (e.g. mean heart rate; P = 0.006), indicating object recognition. In contrast to our expectations, TEST horses also reacted significantly less towards the novel object (e.g. mean heart rate; P = 0.018), suggesting that they were capable of object generalisation. We also found that TEST horses showed an increase in exploratory behaviour when objects within the complex object changed order and location (both P < 0.001), whereas there was no increase in heart rate, indicating that the horses were not frightened by the changes. The results demonstrate that it is possible to increase object generalisation in horses by habituating them to a range of colours and shapes simultaneously. This knowledge greatly affects the way in which horses may be trained to react calmly towards frightening objects.
	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	5328
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