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	Author	Wilson, A.M.; McGuigan, M.P.; Su, A.; van Den Bogert, A.J.
	Title	Horses damp the spring in their step			Type	Journal Article
	Year	2001	Publication	Nature	Abbreviated Journal	Nature
	Volume	414	Issue	6866	Pages	895-899
	Keywords	Animals; Biomechanics; Elasticity; Forelimb; Gait; Horses/anatomy & histology/physiology; Leg Bones/physiology; Locomotion; Models, Biological; Muscle Fibers/physiology; Muscle, Skeletal/anatomy & histology/physiology; Tendons/anatomy & histology/physiology; Vibration
	Abstract	The muscular work of galloping in horses is halved by storing and returning elastic strain energy in spring-like muscle-tendon units.These make the legs act like a child's pogo stick that is tuned to stretch and recoil at 2.5 strides per second. This mechanism is optimized by unique musculoskeletal adaptations: the digital flexor muscles have extremely short fibres and significant passive properties, whereas the tendons are very long and span several joints. Length change occurs by a stretching of the spring-like digital flexor tendons rather than through energetically expensive length changes in the muscle. Despite being apparently redundant for such a mechanism, the muscle fibres in the digital flexors are well developed. Here we show that the mechanical arrangement of the elastic leg permits it to vibrate at a higher frequency of 30-40 Hz that could cause fatigue damage to tendon and bone. Furthermore, we show that the digital flexor muscles have minimal ability to contribute to or regulate significantly the 2.5-Hz cycle of movement, but are ideally arranged to damp these high-frequency oscillations in the limb.
	Address	Department of Veterinary Basic Sciences, The Royal Veterinary College, Hatfield, Herts AL9 7TA, UK. awilson@rvc.ac.uk
	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	0028-0836	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:11780059			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	2300
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	Author	Waite, T.A.
	Title	Interruptions improve choice performance in gray jays: prolonged information processing versus minimization of costly errors			Type	Journal Article
	Year	2002	Publication	Animal Cognition	Abbreviated Journal	Anim. Cogn.
	Volume	5	Issue	4	Pages	209-214
	Keywords	Animals; Behavior, Animal; Choice Behavior; Female; Learning; Male; Models, Biological; Motivation; Reinforcement Schedule; Songbirds/*physiology; Time Factors
	Abstract	Under the assumption that selection favors minimization of costly errors, erroneous choice may be common when its fitness cost is low. According to an adaptive-choice model, this cost depends on the rate at which an animal encounters the choice: the higher this rate, the smaller the cost of choosing a less valuable option. Errors should thus be more common when interruptions to foraging are shorter. A previous experiment supported this prediction: gray jays, Perisoreus canadensis, were more error prone when subjected to shorter delays to access to food rewards. This pattern, though, is also predicted by an attentional-constraints model. Because the subjects were able to inspect the rewards during delays, their improved performance when subjected to longer delays could have been a byproduct of the experimentally prolonged opportunity for information processing. To evaluate this possibility, a follow-up experiment manipulated both delay to access and whether rewards could be inspected during delays. Depriving jays of the opportunity to inspect rewards (using opaque lids) induced only a small, nonsignificant increase in error rate. This effect was independent of length of delay and so the jays' improved performance when subjected to longer delays was not simply a byproduct of prolonged information processing. More definitively, even when the jays were prevented from inspecting rewards during delays, their performance improved when subjected to longer delays. The findings are thus consistent with the adaptive-choice model.
	Address	Department of Evolution, Ecology, and Organismal Biology, Ohio State University, 1735 Neil Avenue, Columbus, OH 43210-1293, USA. waite.1@osu.edu
	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	1435-9448	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:12461598			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	2592
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	Author	Wolf, M.; van Doorn, G.S.; Leimar, O.; Weissing, F.J.
	Title	Life-history trade-offs favour the evolution of animal personalities			Type	Journal Article
	Year	2007	Publication	Nature	Abbreviated Journal	Nature
	Volume	447	Issue	7144	Pages	581-584
	Keywords	Aggression/physiology/psychology; Animals; Behavior, Animal/physiology; Evolution; Exploratory Behavior/physiology; Models, Biological; Personality/*physiology; Predatory Behavior/physiology; Reproduction/physiology; Risk-Taking; Selection (Genetics)
	Abstract	In recent years evidence has been accumulating that personalities are not only found in humans but also in a wide range of other animal species. Individuals differ consistently in their behavioural tendencies and the behaviour in one context is correlated with the behaviour in multiple other contexts. From an adaptive perspective, the evolution of animal personalities is still a mystery, because a more flexible structure of behaviour should provide a selective advantage. Accordingly, many researchers view personalities as resulting from constraints imposed by the architecture of behaviour (but see ref. 12). In contrast, we show here that animal personalities can be given an adaptive explanation. Our argument is based on the insight that the trade-off between current and future reproduction often results in polymorphic populations in which some individuals put more emphasis on future fitness returns than others. Life-history theory predicts that such differences in fitness expectations should result in systematic differences in risk-taking behaviour. Individuals with high future expectations (who have much to lose) should be more risk-averse than individuals with low expectations. This applies to all kinds of risky situations, so individuals should consistently differ in their behaviour. By means of an evolutionary model we demonstrate that this basic principle results in the evolution of animal personalities. It simultaneously explains the coexistence of behavioural types, the consistency of behaviour through time and the structure of behavioural correlations across contexts. Moreover, it explains the common finding that explorative behaviour and risk-related traits like boldness and aggressiveness are common characteristics of animal personalities.
	Address	Theoretical Biology Group, Centre for Ecological and Evolutionary Studies, University of Groningen, Kerklaan 30, 9751 NN Haren, The Netherlands
	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	1476-4687	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:17538618			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	4098
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	Author	Bell, A.M.
	Title	Evolutionary biology: animal personalities			Type
	Year	2007	Publication	Nature	Abbreviated Journal	Nature
	Volume	447	Issue	7144	Pages	539-540
	Keywords	Aggression/physiology/psychology; Animals; Behavior, Animal/physiology; Evolution; Humans; Models, Biological; Personality/genetics/physiology; Reproduction/genetics/physiology; Risk-Taking; Selection (Genetics)
	Abstract
	Address
	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	1476-4687	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:17538607			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	4099
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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	Harcourt, J.L.; Ang, T.Z.; Sweetman, G.; Johnstone, R.A.; Manica, A.
	Title	Social feedback and the emergence of leaders and followers			Type	Journal Article
	Year	2009	Publication	Current Biology : CB	Abbreviated Journal	Curr Biol
	Volume	19	Issue	3	Pages	248-252
	Keywords	Analysis of Variance; Animals; Appetitive Behavior/physiology; Feedback; Great Britain; Leadership; Markov Chains; Models, Biological; Monte Carlo Method; Smegmamorpha/physiology; Social Behavior; Video Recording
	Abstract	In many animal groups, certain individuals consistently appear at the forefront of coordinated movements [1-4]. How such leaders emerge is poorly understood [5, 6]. Here, we show that in pairs of sticklebacks, Gasterosteus aculeatus, leadership arises from individual differences in the way that fish respond to their partner's movements. Having first established that individuals differed in their propensity to leave cover in order to look for food, we randomly paired fish of varying boldness, and we used a Markov Chain model to infer the individual rules underlying their joint behavior. Both fish in a pair responded to each other's movements-each was more likely to leave cover if the other was already out and to return if the other had already returned. However, we found that bolder individuals displayed greater initiative and were less responsive to their partners, whereas shyer individuals displayed less initiative but followed their partners more faithfully; they also, as followers, elicited greater leadership tendencies in their bold partners. We conclude that leadership in this case is reinforced by positive social feedback.
	Address	Department of Zoology, University of Cambridge, Cambridge, UK
	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	0960-9822	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:19185497			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5123
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	Author	Rands, S.A.; Cowlishaw, G.; Pettifor, R.A.; Rowcliffe, J.M.; Johnstone, R.A.
	Title	The emergence of leaders and followers in foraging pairs when the qualities of individuals differ			Type	Journal Article
	Year	2008	Publication	BMC Evolutionary Biology	Abbreviated Journal	BMC Evol Biol
	Volume	8	Issue		Pages	51
	Keywords	Animals; Feeding Behavior; Food Chain; Models, Biological; Social Dominance
	Abstract	BACKGROUND: Foraging in groups offers animals a number of advantages, such as increasing their likelihood of finding food or detecting and avoiding predators. In order for a group to remain together, there has to be some degree of coordination of behaviour and movement between its members (which may in some cases be initiated by a decision-making leader, and in other cases may emerge as an underlying property of the group). For example, behavioural synchronisation is a phenomenon where animals within a group initiate and then continue to conduct identical behaviours, and has been characterised for a wide range of species. We examine how a pair of animals should behave using a state-dependent approach, and ask what conditions are likely to lead to behavioural synchronisation occurring, and whether one of the individuals is more likely to act as a leader. RESULTS: The model we describe considers how the energetic gain, metabolic requirements and predation risks faced by the individuals affect measures of their energetic state and behaviour (such as the degree of behavioural synchronisation seen within the pair, and the value to an individual of knowing the energetic state of its colleague). We explore how predictable changes in these measures are in response to changes in physiological requirements and predation risk. We also consider how these measures should change when the members of the pair are not identical in their metabolic requirements or their susceptibility to predation. We find that many of the changes seen in these measures are complex, especially when asymmetries exist between the members of the pair. CONCLUSION: Analyses are presented that demonstrate that, although these general patterns are robust, care needs to be taken when considering the effects of individual differences, as the relationship between individual differences and the resulting qualitative changes in behaviour may be complex. We discuss how these results are related to experimental observations, and how the model and its predictions could be extended.
	Address	Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK. sean.rands@bristol.ac.uk
	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	1471-2148	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:18282297			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5126
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	Author	Conradt, L.; Roper, T.J.
	Title	Group decision-making in animals			Type	Journal Article
	Year	2003	Publication	Nature	Abbreviated Journal	Nature
	Volume	421	Issue	6919	Pages	155-158
	Keywords	Animals; Behavior, Animal/physiology; Decision Making; Democracy; Group Processes; *Models, Biological; Population Density; Social Behavior
	Abstract	Groups of animals often need to make communal decisions, for example about which activities to perform, when to perform them and which direction to travel in; however, little is known about how they do so. Here, we model the fitness consequences of two possible decision-making mechanisms: 'despotism' and 'democracy'. We show that under most conditions, the costs to subordinate group members, and to the group as a whole, are considerably higher for despotic than for democratic decisions. Even when the despot is the most experienced group member, it only pays other members to accept its decision when group size is small and the difference in information is large. Democratic decisions are more beneficial primarily because they tend to produce less extreme decisions, rather than because each individual has an influence on the decision per se. Our model suggests that democracy should be widespread and makes quantitative, testable predictions about group decision-making in non-humans.
	Address	School of Biological Sciences, University of Sussex, Brighton BN1 9QG, UK. l.conradt@sussex.ac.uk
	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	0028-0836	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:12520299			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5136
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	Author	Rands, S.A.; Cowlishaw, G.; Pettifor, R.A.; Rowcliffe, J.M.; Johnstone, R.A.
	Title	Spontaneous emergence of leaders and followers in foraging pairs			Type	Journal Article
	Year	2003	Publication	Nature	Abbreviated Journal	Nature
	Volume	423	Issue	6938	Pages	432-434
	Keywords	Animals; Energy Metabolism; Food; Food Chain; Models, Biological; Motor Activity; Social Behavior; Time Factors
	Abstract	Animals that forage socially often stand to gain from coordination of their behaviour. Yet it is not known how group members reach a consensus on the timing of foraging bouts. Here we demonstrate a simple process by which this may occur. We develop a state-dependent, dynamic game model of foraging by a pair of animals, in which each individual chooses between resting or foraging during a series of consecutive periods, so as to maximize its own individual chances of survival. We find that, if there is an advantage to foraging together, the equilibrium behaviour of both individuals becomes highly synchronized. As a result of this synchronization, differences in the energetic reserves of the two players spontaneously develop, leading them to adopt different behavioural roles. The individual with lower reserves emerges as the 'pace-maker' who determines when the pair should forage, providing a straightforward resolution to the problem of group coordination. Moreover, the strategy that gives rise to this behaviour can be implemented by a simple 'rule of thumb' that requires no detailed knowledge of the state of other individuals.
	Address	Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK. s.rands@zoo.cam.ac.uk
	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	0028-0836	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:12761547			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5138
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	Author	Seyfarth, R.M.
	Title	A model of social grooming among adult female monkeys			Type	Journal Article
	Year	1977	Publication	Journal of Theoretical Biology	Abbreviated Journal	J. Theor. Biol.
	Volume	65	Issue	4	Pages	671-698
	Keywords	Animals; Behavior, Animal; Female; Grooming; Haplorhini/physiology; *Models, Biological; Reproduction; Social Dominance; Time Factors
	Abstract	Grooming networks among adult female monkeys exhibit two similar features across a number of different species. High-ranking animals receive more grooming than others, and the majority of grooming occurs between females of adjacent rank. A theoretical model which duplicates these features is presented, and the properties of the model are used to explain the possible causation and function of female grooming behaviour. The model illustrates how relatively simple principles governing the behaviour of individuals may be used to explain more complex aspects of the social structure of non-human primate groups.
	Address
	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	0022-5193	ISBN		Medium
	Area		Expedition		Conference
	Notes	PMID:406485			Approved	no
	Call Number	Equine Behaviour @ team @			Serial	5259
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