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Among-individual variation in behaviour is a widespread phenomenon, with several frameworks developed to explain its existence. Maternal effects, which can have significant influence over evolutionary processes, are an understudied source of behavioural variation. Maternal effects are not necessarily static, however, since their importance can change over offspring ontogeny, typically declining with age relative to additive genetic effects. Here, using a quantitative genetics approach, we test the prediction that maternal effects will influence age-specific risk-taking behaviour in Trinidadian guppies, Poecilia reticulata. Individuals were subject to a single open-field trial as juveniles and up to four repeat trials as adults, with five traits indicative of risk-taking behaviour measured in each trial. We then partitioned phenotypic variance into additive genetic (V A ) and maternal identity (V M ) components, in addition to testing brood size and maternal weight as specific sources of maternal effects. We found that V M had significant influence over juvenile traits, with very low V A estimates. Whereas, in adults, all traits were significantly heritable, with little support for V M . We also found a strong influence of maternal traits on juvenile behaviours as predicted, with significant, albeit smaller, effects found in adults. Maternal weight was heritable and itself subject to maternal effects. Thus, maternal weight is a likely source of maternal genetic effects that are expected to alter response to selection on personality in this system. More generally, our study highlights that while maternal effects can be an important source of personality variation, this varies over ontogeny of offspring.

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., (© 2015 The Author(s) Published by the Royal Society. All rights reserved.)

Aggression occurs when individuals compete over limiting resources. While theoretical studies have long placed a strong emphasis on context-specificity of aggression, there is increasing recognition that consistent behavioural differences exist among individuals, and that aggressiveness may be an important component of individual personality. Though empirical studies tend to focus on one aspect or the other, we suggest there is merit in modelling both within- and among-individual variation in agonistic behaviour simultaneously. Here, we demonstrate how this can be achieved using multivariate linear mixed effect models. Using data from repeated mirror trials and dyadic interactions of male green swordtails, Xiphophorus helleri, we show repeatable components of (co)variation in a suite of agonistic behaviour that is broadly consistent with a major axis of variation in aggressiveness. We also show that observed focal behaviour is dependent on opponent effects, which can themselves be repeatable but were more generally found to be context specific. In particular, our models show that within-individual variation in agonistic behaviour is explained, at least in part, by the relative size of a live opponent as predicted by contest theory. Finally, we suggest several additional applications of the multivariate models demonstrated here. These include testing the recently queried functional equivalence of alternative experimental approaches, (e.g., mirror trials, dyadic interaction tests) for assaying individual aggressiveness.

Genetic factors underpinning phenotypic variation are required if natural selection is to result in adaptive evolution. However, evolutionary and behavioural ecologists typically focus on variation among individuals in their average trait values and seek to characterize genetic contributions to this. As a result, less attention has been paid to if and how genes could contribute towards within‐individual variance or trait 'predictability'. In fact, phenotypic 'predictability' can vary among individuals, and emerging evidence from livestock genetics suggests this can be due to genetic factors. Here, we test this empirically using repeated measures of a behavioural stress response trait in a pedigreed population of wild‐type guppies. We ask (a) whether individuals differ in behavioural predictability and (b) whether this variation is heritable and so evolvable under selection. Using statistical methodology from the field of quantitative genetics, we find support for both hypotheses and also show evidence of a genetic correlation structure between the behavioural trait mean and individual predictability. We show that investigating sources of variability in trait predictability is statistically tractable and can yield useful biological interpretation. We conclude that, if widespread, genetic variance for 'predictability' will have major implications for the evolutionary causes and consequences of phenotypic variation. [ABSTRACT FROM AUTHOR]

Latent personality traits underpinning observed behavioral variation have been studied in a great many species. However, a lack of standardized behavioral assays, coupled to a common reliance on inferring personality from a single, observed, behavioral trait makes it difficult to determine if, when, and how conclusions can be directly compared across taxa. Here, we estimate the among-individual (co)variance structure (ID) for a set of four behaviors expressed in an open field trial, putatively indicative of boldness, in seven species of small freshwater fish. We show that the ID matrices differ in terms of the total amount of variation present, and crucially the orientation, and as a consequence, biological interpretation of the first eigenvector. Specifically, loading of observed traits on the main axis of variation in ID matched a priori expectations for a shy-bold continuum in only three of the seven cases. Nonetheless, when the "shape" of the matrices was compared in higher dimensions, there was a high level of similarity among species, and weak evidence of phylogenetic signal. Our study highlights the present difficulty of trying to compare empirical inferences about specific personality traits across studies. However, it also shows how multivariate data collection and analysis allows the structure of behavioral variation to be quantitatively compared across populations or species without reliance on ambiguous verbal labels. This suggests that the field may have much to gain from greater uptake of phylogenetically informed comparative approaches when seeking to test evolutionary hypotheses about the origin and maintenance of personality variation. [ABSTRACT FROM AUTHOR]

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Having recognized that variation around the population-level "Golden Mean" of labile traits contains biologically meaningful information, behavioural ecologists have focused increasingly on exploring the causes and consequences of individual variation in behaviour. These are exciting new directions for the field, assisted in no small part by the adoption of mixed-effects modelling techniques that enable the partitioning of among- and within-individual behavioural variation. It has become commonplace to extract predictions of individual random effects from such models for use in subsequent analyses (for example, between a personality trait and other individual traits such as cognition, physiology, or fitness-related measures). However, these predictions are made with large amounts of error that is not carried forward, rendering further tests susceptible to spurious P values from these individual-level point estimates. We briefly summarize the problems with such statistical methods that are used regularly by behavioural ecologists, and highlight the robust solutions that exist within the mixed model framework, providing tutorials to aid in their implementation. [ABSTRACT FROM AUTHOR]

Animal ‘personality’, the phenomenon of consistent individual differences in behaviour within populations, has been documented widely, yet its functional significance and the reasons for its persistence remain unclear. One possibility is that among-individual behavioural variation is linked to fitness-determining traits via effects on resource acquisition. In this study, we tested this idea, using rockpool prawns, Palaemon elegans , to test for a correlation between ‘high-risk exploration’ and the ability to monopolize a limited resource. Modified open field trials (OFTs) confirmed that consistent among-individual (co)variation in high-risk exploratory behaviours does exist in this species, and multivariate analysis showed trait variation is consistent with a major axis of personality variation. Subsequent feeding trials in size-matched groups where competition was possible revealed a high repeatability of feeding duration, used here as a proxy for RHP (resource-holding potential). We found significant negative correlations between feeding duration and two ‘risky’ behaviours, such that individuals that took fewer risks fed more. Our results are not consistent with the widely hypothesized idea of a ‘proactive syndrome’ in which bolder, risk-taking personalities are positively associated with RHP. Rather they suggest the possibility of a trade-off, with some individuals successful at monopolizing limited, high-value resources, while others are more willing to engage in potentially risky exploration (which may increase the likelihood of encountering novel resource patches). We speculate that alternative strategies for acquiring limited resources might thereby contribute to the maintenance of personality variation observed in wild populations. [ABSTRACT FROM AUTHOR]

Cognition is vital for carrying out behaviours required for survival and reproduction. Cognitive performance varies between species, but also between individuals within populations. While variation is a prerequisite for natural selection, selection does not act on traits in isolation. The extent to which cognitive traits covary with other aspects of phenotype (e.g. personality traits) may be important in shaping evolutionary dynamics. Here we adopted a multivariate approach to test spatial learning in male Poecilia reticulata and asked whether differences in cognitive performance are associated with (repeatable) differences in stress response behaviour. Functional links between cognitive traits and 'stress coping style' have been hypothesized. Furthermore, individual level studies of cognitive performance typically rely on multiple testing paradigms that may themselves be a stressor. There is a risk that variation in stress responsiveness is itself a cause of apparent, but artefactual variance in cognitive ability. Fish repeatedly experienced two spatial learning tasks (maze layouts) and an acute stress response test (open field trial). We found repeatable differences between individuals in performance within and across maze layouts. On average, performance improved with experience in the first maze, consistent with spatial learning, but not in the second. Individuals varied in the trajectory of mean performance with trial number in both mazes, suggesting they differ in 'learning rate'. Acute stress response behaviour was repeatable but predicted neither average time to solve the maze nor learning rate. We found no support for between-individual correlation between acute stress response and cognitive performance. However, we highlight the possibility that cumulative, chronic stress effects may nevertheless cause declines in performance for some individuals (leading to lack of improvement in mean time to solve the second maze). If so, this may represent a pervasive but difficult challenge for our ability to robustly estimate learning rates in studies of animal cognition. • Multivariate approach to test spatial learning and acute stress response in guppies. • Spatial learning performance is repeatable among individuals across contexts. • Acute stress response behaviour is also repeatable among individuals. • No support for correlation between acute stress response and cognitive performance. • Cumulative chronic stress may cause declines in performance for some individuals. [ABSTRACT FROM AUTHOR]