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Summary 1.Contests may involve injurious fighting, other types of direct physical aggression and communication. They occur over ownership access to mates and other resources that may increase an individual's attractiveness and its chance of survival. Traits that enhance resource holding potential may be the result of sexual selection, natural selection or a combination of both. 2. Agonistic behaviours are expected to be demanding to perform and costly in terms of changes in physiological state. The ability to meet the physiological costs may determine contest outcomes and constrain the intensity of agonistic activities. 3. The energetic costs have been investigated in a broad range of taxa using a variety of techniques. They include the mobilization of energy reserves, but a key cost in several taxa appears to be limited anaerobic capacity and subsequent accumulation of lactic acid. Androgens, stress hormones and neurohormones have also been shown to constrain fighting behaviour. However, due to key differences in the endocrine systems of vertebrates and invertebrates, the effects of hormones are far less consistent across taxa than in the case of metabolites. 4. Physiological constraints on fighting may vary according to their importance relative to circumstantial costs, the time-scale over which they exert their effects, their effects on different roles and their causal links with behaviour. Incorporating these factors into theoretical studies of contest behaviour may give further insights of how the costs of fighting influence agonistic behaviour.

Many phenotypic traits show plasticity but behaviour is often considered the 'most plastic' aspect of phenotype as it is likely to show the quickest response to temporal changes in conditions or 'situation'. However, it has also been noted that constraints on sensory acuity, cognitive structure and physiological capacities place limits on behavioural plasticity. Such limits to plasticity may generate consistent differences in behaviour between individuals from the same population. It has recently been suggested that these consistent differences in individual behaviour may be adaptive and the term 'animal personalities' has been used to describe them. In many cases, however, a degree of both behavioural plasticity and relative consistency is probable. To understand the possible functions of animal personalities, it is necessary to determine the relative strength of each tendency and this may be achieved by comparison of statistical effect sizes for tests of difference and concordance. Here, we describe a new statistical framework for making such comparisons and investigate cross-situational plasticity and consistency in the duration of startle responses in the European hermit crab Pagurus bernhardus, in the field and the laboratory. The effect sizes of tests for behavioural consistency were greater than for tests of behavioural plasticity, indicating for the first time the presence of animal personalities in a crustacean model.

Variation in behaviour occurs at multiple levels, including between individuals (personality) and between situations ( plasticity). Behaviour also varies within individuals, and intra-individual variation (IIV) in behaviour describes within-individual residual variance in behaviour that remains after the effects of obvious external and internal influences on behaviour have been accounted for. IIV thus describes how predictable an individual's behaviour is. Differences in predictability, between individuals and between situations, might be biologically significant. For example, behaving unpredictably under predation threat might reduce the chance of capture. Here, we investigated the duration of startle responses in hermit crabs, in the presence and absence of a predator cue. Individuals differed in startle response duration ( personality) and while individuals also varied in their sensitivity to risk, mean response time was greater in the presence of a predator ( plasticity). Moreover, IIV was greater in the presence of a predator, providing some of the first evidence that the facultative injection of unpredictability into behaviour might represent a strategy for dealing with risk.

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When animals engage in fights they face a series of decisions, which are based on the value of the contested resource and either their relative or their absolute fighting ability. Certain correlates of fighting ability or 'resource holding potential' such as body size are fixed but physiological correlates are expected to vary during the encounter. We examine the role of energy reserves in determining fight outcomes and parameters during 'shell fighting' in hermit crabs. During these fights, the two contestants perform very different roles of attacker and defender. We show that the balance of the total energy pool, in the form of glucose and glycogen, determines the ability of defenders to resist eviction from their shells. Low glucose in evicted defenders is not caused by depletion of energy reserves, rather mobilization of glycogen appears to be the result of a strategic decision about whether to resist effectively, based on the perceived fighting ability of the attacker. Attackers, however, always initiate the fight so such a decision for this role appears unlikely. In addition to influencing decisions and ability during fights, physiological correlates of fighting ability can in turn be influenced by strategic decisions.

We review the evidence for a link between consistent among-individual variation in behaviour (animal personality) and the ability to win contests over limited resources. Explorative and bold behaviours often covary with contest behaviour and outcome, although there is evidence that the structure of these 'behavioural syndromes' can change across situations. Aggression itself is typically repeatable, but also subject to high within-individual variation as a consequence of plastic responses to previous fight outcomes and opponent traits. Common proximate mechanisms (gene expression, endocrine control and metabolic rates) may underpin variation in both contest behaviour and general personality traits. Given the theoretical links between the evolution of fighting and of personality, we suggest that longitudinal studies of contest behaviour, combining behavioural and physiological data, would be a useful context for the study of animal personalities.