Behaviour in captivity predicts some aspects of natural behaviour,but not others,in a wild cricket population

Examining the relevance of ‘animal personality’ involves linking consistent among- and within-individual behavioural variation to fitness in the wild. Studies aiming to do this typically assay personality in captivity and rely on the assumption that measures of traits in the laboratory reflect their expression in nature. We examined this rarely tested assumption by comparing laboratory and field measurements of the behaviour of wild field crickets (Gryllus campestris) by continuously monitoring individual behaviour in nature, and repeatedly capturing the same individuals and measuring their behaviour in captivity. We focused on three traits that are frequently examined in personality studies: shyness, activity and exploration. All of them showed repeatability in the laboratory. Laboratory activity and exploration predicted the expression of their equivalent behaviours in the wild, but shyness did not. Traits in the wild were predictably influenced by environmental factors such as temperature and sunlight, but only activity showed appreciable within-individual repeatability. This suggests that some behaviours typically studied as personality traits can be accurately assayed in captivity, but the expression of others may be highly context-specific. Our results highlight the importance of validating the relevance of laboratory behavioural assays to analogous traits measured in the wild.     

Express your personality or go along with the group: what determines the behaviour of shoaling perch?

Behavioural syndromes, defined as correlated behaviours in different contexts, have been studied across species and taxa including humans as part of a personality concept. While most studies have focused on solitary individuals, less is known on how shoaling fish compromise between own personality and group behaviour. Risk-taking behaviour in 1-year-old perch (Perca fluviatilis) was observed to compare individual behaviour when in a group and when alone. An experimental design gave the fish the choice between foraging in an open area in the presence of a piscivore and hiding in the vegetation. We quantified the variation accountable by the effect of individuals being in a group, individuals alone and repeated measurements, using hierarchical mixed effects models. Within-group variances were low, but when individuals were later tested alone, individual differences explained most of the variation. Still, the individual best linear unbiased predictors (BLUPs) of time spent in the open area, extracted from the random effects of the mixed effects model, were positively correlated with the corresponding BLUPs when alone. The results indicate that individual behavioural traits are to some degree expressed also within groups. Most fish showed a shyer behaviour when alone, but bolder individuals changed less between treatments than did shyer ones, suggesting a more influential role of bold fish in the group.     

The influence of mitonuclear genetic variation on personality in seed beetles

There is a growing awareness of the influence of mitochondrial genetic variation on life-history phenotypes, particularly via epistatic interactions with nuclear genes. Owing to their direct effect on traits such as metabolic and growth rates, mitonuclear interactions may also affect variation in behavioural types or personalities (i.e. behavioural variation that is consistent within individuals, but differs among individuals). However, this possibility is largely unexplored. We used mitonuclear introgression lines, where three mitochondrial genomes were introgressed into three nuclear genetic backgrounds, to disentangle genetic effects on behavioural variation in a seed beetle. We found within-individual consistency in a suite of activity-related behaviours, providing evidence for variation in personality. Composite measures of overall activity of individuals in behavioural assays were influenced by both nuclear genetic variation and by the interaction between nuclear and mitochondrial genomes. More importantly, the degree of expression of behavioural and life-history phenotypes was correlated and mitonuclear genetic variation affected expression of these concerted phenotypes. These results show that mitonuclear genetic variation affects both behavioural and life-history traits, and they provide novel insights into the maintenance of genetic variation in behaviour and personality.     

Personality and Nest Defence in the Great Tit (Parus major)

A growing number of studies have shown that individuals differ consistently in a suite of correlated behavioural traits across various contexts and situations. Yet, most work on animal personalities has been performed under laboratory conditions and still little is known about the ecological significance of differences in personality in the wild, and the behavioural mechanisms underlying possible fitness consequences. In this study, we investigated individual differences in personality in relation to nest defence behaviour in wild great tits. Nest defence is an important aspect of parental care and involves a trade‐off between two fitness components (i.e. survival and reproduction). As a measure of personality we used exploratory behaviour in a novel environment as this has been shown to be correlated with several other behavioural traits including risk‐taking and aggression, two important behavioural components of nest defence. We found that the intensity of alarm calling towards a human intruder was positively associated with exploratory behaviour, while there was a negative association between exploration score and number of movements during nest defence. Thus, fast explorers are shown to respond more boldly towards predators in the field. More generally we show that individuals with different personalities vary in their anti‐predator and reproductive investment strategies.     

Hormone levels predict individual differences in reproductive success in a passerine bird

Hormones mediate major physiological and behavioural components of the reproductive phenotype of individuals. To understand basic evolutionary processes in the hormonal regulation of reproductive traits, we need to know whether, and during which reproductive phases, individual variation in hormone concentrations relates to fitness in natural populations. We related circulating concentrations of prolactin and corticosterone to parental behaviour and reproductive success during both the pre-breeding and the chick-rearing stages in both individuals of pairs of free-living house sparrows, Passer domesticus. Prolactin and baseline corticosterone concentrations in pre-breeding females, and prolactin concentrations in pre-breeding males, predicted total number of fledglings. When the strong effect of lay date on total fledgling number was corrected for, only pre-breeding baseline corticosterone, but not prolactin, was negatively correlated with the reproductive success of females. During the breeding season, nestling provisioning rates of both sexes were negatively correlated with stress-induced corticosterone levels. Lastly, individuals of both sexes with low baseline corticosterone before and high baseline corticosterone during breeding raised the most offspring, suggesting that either the plasticity of this trait contributes to reproductive success or that high parental effort leads to increased hormone concentrations. Thus hormone concentrations both before and during breeding, as well as their seasonal dynamics, predict reproductive success, suggesting that individual variation in absolute concentrations and in plasticity is functionally significant, and, if heritable, may be a target of selection.     

A strong genetic correlation underlying a behavioural syndrome disappears during development because of genotype–age interactions

In animal populations, as in humans, behavioural differences between individuals that are consistent over time and across contexts are considered to reflect personality, and suites of correlated behaviours expressed by individuals are known as behavioural syndromes. Lifelong stability of behavioural syndromes is often assumed, either implicitly or explicitly. Here, we use a quantitative genetic approach to study the developmental stability of a behavioural syndrome in a wild population of blue tits. We find that a behavioural syndrome formed by a strong genetic correlation of two personality traits in nestlings disappears in adults, and we demonstrate that genotype–age interaction is the likely mechanism underlying this change during development. A behavioural syndrome may hence change during organismal development, even when personality traits seem to be strongly physiologically or functionally linked in one age group. We outline how such developmental plasticity has important ramifications for understanding the mechanistic basis as well as the evolutionary consequences of behavioural syndromes.     

Singing Activity Reveals Personality Traits in Great Tits

In animal communication, sexually selected signals have been shown to often signal individual attributes such as motivation or quality. Birdsong is among the best studied signalling systems, and song traits vary substantially among individuals. The question remains if variation in signalling also reflects more general and consistent individual characteristics. Such consistent individual differences in behaviour that are relatively stable over time and contexts are referred to as personality or behavioural syndromes. Here, we studied the relation between singing and explorative behaviour, a well‐studied personality trait, using great tits (Parus major) under standardized aviary conditions. The results show that singing activity measured as the number of songs sung in spring prior to breeding correlated with male but not with female explorative behaviour. In contrast, song repertoire was not related to explorative behaviour but varied over the day. The link between explorative and singing behaviour suggests that sexually selected signals are more than signals of quality but can also reflect other intrinsic behavioural characteristics such as personality traits.     

Urine marking in male common voles: Does behavioural activity matter?

Rodent urine provides animals with a large amount of information, from the identity of the animal through its physical condition to social status. Many studies therefore focus on rodent urine-marking behaviour and use marking frequency as an indicator of social status or competitive ability. However, marking, like many other aspects of rodent behaviour, may be affected by individual behavioural activity, a factor that has not been examined so far. We therefore studied a relationship between male urine-marking in reaction to another male's marks (standard opponent) and individual personality profile, characterised by behavioural activity in an open field test (OFT). The marking appeared to be consistent and specific for particular individuals as there was a significant positive relationship between individual markings in two different phases of the experiment. The linkage between behavioural activity in the OFT and urine-marking frequency was non-linear (quadratic), which suggested that males with intermediate activity marked more intensively than males from the extremes of the behavioural spectra. The relationship between the opponent's and the tested males' markings was positive, however, we found no statistically significant evidence that the voles would attempt to overmark the opponent. Marking thus seems to have more of a self-advertising than a competitive function in the common vole. Further, as high marking activity is under strong intra- or intersexual selection, the result might suggest a stabilising selection of the personality trait described as behavioural activity in our study.     

Individual-level personality influences social foraging and collective behaviour in wild birds

There is increasing evidence that animal groups can maintain coordinated behaviour and make collective decisions based on simple interaction rules. Effective collective action may be further facilitated by individual variation within groups, particularly through leader–follower polymorphisms. Recent studies have suggested that individual-level personality traits influence the degree to which individuals use social information, are attracted to conspecifics, or act as leaders/followers. However, evidence is equivocal and largely limited to laboratory studies. We use an automated data-collection system to conduct an experiment testing the relationship between personality and collective decision-making in the wild. First, we report that foraging flocks of great tits (Parus major) show strikingly synchronous behaviour. A predictive model of collective decision-making replicates patterns well, suggesting simple interaction rules are sufficient to explain the observed social behaviour. Second, within groups, individuals with more reactive personalities behave more collectively, moving to within-flock areas of higher density. By contrast, proactive individuals tend to move to and feed at spatial periphery of flocks. Finally, comparing alternative simulations of flocking with empirical data, we demonstrate that variation in personality promotes within-patch movement while maintaining group cohesion. Our results illustrate the importance of incorporating individual variability in models of social behaviour.     

Personality-dependent dispersal in the invasive mosquitofish: group composition matters

Understanding/predicting ecological invasions is an important challenge in modern ecology because of their immense economical and ecological costs. Recent studies have revealed that within-species variation in behaviour (i.e. animal personality) can shed light on the invasion process. The general hypothesis is that individuals' personality type may affect their colonization success, suggesting that some individuals might be better invaders than others. We have recently shown that, in the invasive mosquitofish (Gambusia affinis), social personality trait was an important indicator of dispersal distance, with more asocial individuals dispersing further. Here, we tested how mean personality within a population, in addition to individual personality type, affect dispersal and settlement decisions in the mosquitofish. We found that individual dispersal tendencies were influenced by the population's mean boldness and sociability score. For example, individuals from populations with more asocial individuals or with more bold individuals are more likely to disperse regardless of their own personality type. We suggest that identifying behavioural traits facilitating invasions, even at the group level, can thus have direct applications in pest management.     

Among‐individual heterogeneity in maternal behaviour and physiology affects reproductive allocation and offspring life‐history traits in the garter snake Thamnophis elegans

Accumulating evidence suggests that within‐individual plasticity of behavioural and physiological traits is limited, resulting in stable among‐individual differences in these aspects of the phenotype. Furthermore, these traits often covary within individuals, resulting in a continuum of correlated phenotypic variation among individuals within populations and species. This heterogeneity, in turn, affects individual fitness and can have cross‐generational effects. Patterns of trait covariation, among‐individual differences, and subsequent fitness consequences have long been recognized in reptiles. Here, we provide a test of patterns of among‐individual heterogeneity in behaviour and physiology and subsequent effects on reproduction and offspring fitness in the garter snake Thamnophis elegans. We find that measures of activity levels vary among individuals and are consistent within individuals in reproductive female snakes, indicating stable behavioural phenotypes. Blood hormone and glucose concentrations are not as stable within individuals, indicating that these traits do not describe consistent physiological phenotypes. Nonetheless, the major axes of variation in maternal traits describe behavioural and physiological phenotypes that interact to predict offspring body condition and mass at birth. This differential allocation of energy to offspring, in turn, strongly influences subsequent offspring growth and survival. This pattern suggests the potential for strong selection on phenotypes defined by behaviour–physiology interactions.     

Environmental and genetic determinants of innovativeness in a natural population of birds

Much of the evidence for the idea that individuals differ in their propensity to innovate and solve new problems has come from studies on captive primates. Increasingly, behavioural ecologists are studying innovativeness in wild populations, and uncovering links with functional behaviour and fitness-related traits. The relative importance of genetic and environmental factors in driving this variation, however, remains unknown. Here, we present the results of the first large-scale study to examine a range of causal factors underlying innovative problem-solving performance (PSP) among 831 great tits (Parus major) temporarily taken into captivity. Analyses show that PSP in this population: (i) was linked to a variety of individual factors, including age, personality and natal origin (immigrant or local-born); (ii) was influenced by natal environment, because individuals had a lower PSP when born in poor-quality habitat, or where local population density was high, leading to cohort effects. Links with many of the individual and environmental factors were present only in some years. In addition, PSP (iii) had little or no measurable heritability, as estimated by a Bayesian animal model; and (iv) was not influenced by maternal effects. Despite previous reports of links between PSP and a range of functional traits in this population, the analyses here suggest that innovativeness had weak if any evolutionary potential. Instead most individual variation was caused by phenotypic plasticity driven by links with other behavioural traits and by environmentally mediated developmental stress. Heritability estimates are population, time and context specific, however, and more studies are needed to determine the generality of these effects. Our results shed light on the causes of innovativeness within populations, and add to the debate on the relative importance of genetic and environmental factors in driving phenotypic variation within populations.     

Frequency-dependent payoffs and sequential decision-making favour consistent tactic use

Although natural selection should have favoured individuals capable of adjusting the weight they give to personal and social information according to circumstances, individuals generally differ consistently in their individual weighting of both types of information. Such individual differences are correlated with personality traits, suggesting that personality could directly affect individuals' ability to collect personal or social information. Alternatively, the link between personality and information use could simply emerge as a by-product of the sequential decision-making process in a frequency-dependent context. Indeed, when the gains associated with behavioural options depend on the choices of others, an individual's sequence of arrival could constrain its choice of options leading to the emergence of correlated behaviours. Any factor such as personality that affects decision order could thus be correlated with information use. To test this new explanation, we developed an individual-based model that simulates a group of animals engaged in a game of sequential frequency-dependent decision: a producer-scrounger game. Our results confirm that the sequence of decision, in this case enforced by the order in which animals enter a foraging area, consistently influences their mean tactic use and their individual plasticity, an outcome reminiscent of the correlation reported between personality and social information use.     

Delayed song maturation and territorial aggression in a songbird

Asymmetries in competitive ability can determine the outcome of social interactions in animals and are often expressed through differences in sexual traits. Competitive ability (resource holding potential, RHP), trait expression and ultimately reproductive success may vary with an individual's age or experience. In some species, reproductively mature males delay acquisition of some adult traits and thereby signal their young age. Theory on animal contests predicts that individuals assess the RHP of an opponent relative to their own, such that escalation is more common between evenly matched opponents. Here, we test predictions from this hypothesis that males respond to a territorial intruder based on their RHP relative to the intruder's RHP. We simulated white-crowned sparrows (Zonotrichia leucophrys) intruding into the territory of a recruit or return. Playback of a song repertoire simulating a young male (recruit) elicited a weaker response from established territory holders (return), but a stronger response from recruits. Playback of a single song type simulating an older male elicited the opposite responses. This indicates that males distinguished between simulated young and old intruders based on song, and responded differently depending on their own experience. Our study highlights the possibility that receiver as well as sender traits should be considered when interpreting animal interactions.     

Variable individual consistency in timing and destination of winter migrating fish

Migration is an important event in the life history of many animals, but there is considerable variation within populations in the timing and final destination. Such differential migration at the population level can be strongly determined by individuals showing different consistencies in migratory traits. By tagging individual cyprinid fish with uniquely coded electronic tags, and recording their winter migrations from lakes to streams for 6 consecutive years, we obtained highly detailed long-term information on the differential migration patterns of individuals. We found that individual migrants showed consistent site fidelities for over-wintering streams over multiple migratory seasons and that they were also consistent in their seasonal timing of migration. Our data also suggest that consistency itself can be considered as an individual trait, with migrants that exhibit consistent site fidelity also showing consistency in migratory timing. The finding of a mixture of both consistent and inconsistent individuals within a population furthers our understanding of intrapopulation variability in migration strategies, and we hypothesize that environmental variation can maintain such different strategies.     

Comparison of personality between juveniles and adults in clonal gecko species

The developmental perspectives of animal personality enhance our understanding of how personality structure changes in relation to life stage. Clonal animals are ideal models for developmental studies because personality differences can be solely attributed to environmental factors. Here, I investigated the presence of personality within a species of clonal gecko, Lepidodactylus lugubris, at different developmental stages. For juveniles and adult geckos, I measured exploration (reaction to a novel situation) and boldness (risk-prone tendency) and evaluated repeatability and correlation of these behavioural traits. Each gecko exhibited different exploration and boldness with significant repeatability through time but no correlation between these behavioural traits. Small juveniles were composed of only bold and low explorative individuals but large juveniles and adults were composed of various personality type individuals. These results demonstrate that subject geckos have a similar personality structure across life stages and that exploration and boldness are independent personality without forming behavioural syndrome structure. Biased composition of personality type between life stages suggests that appearance of different personality type individuals during an early ontogenetic stage generates personality variation within the clonal population. This study provides developmental insight about personality structure and its composition in clonal animals living in the wild.     

动物个性研究进展

"个性"是指不同时空条件下动物种群个体间行为的稳定差异。大量的理论和实验性研究表明,个性差异在动物界普遍存在,其是种群多度和分布、物种共存及群落构建的重要驱动因子。介绍了动物个性的概念、分类及衡量指标,将前人测量个性类型的方法加以总结;随后介绍了动物个性的生态学意义,尤其是个性对动物生活史策略、种群分布与多度、群落结构和动态、生态系统功能和过程以及疾病与信息传播等的影响。在此基础上,进一步分析了在人类活动增加等全球变化背景下,动物个性如何调控动物个体行为、种群和群落动态对这些环境变化的响应。阐述了动物个性的形成与维持机制,并对未来的研究方向进行了展望。     

Genetic variance for behavioural ‘predictability’ of stress response

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.     

Parasitism and the evolutionary ecology of animal personality

The ecological factors responsible for the evolution of individual differences in animal personality (consistent individual differences in the same behaviour across time and contexts) are currently the subject of intense debate. A limited number of ecological factors have been investigated to date, with most attention focusing on the roles of resource competition and predation. We suggest here that parasitism may play a potentially important, but largely overlooked, role in the evolution of animal personalities. We identify two major routes by which parasites might influence the evolution of animal personality. First, because the risk of acquiring parasites can be influenced by an individual's behavioural type, local parasite regimes may impose selection on personality traits and behavioural syndromes (correlations between personality traits). Second, because parasite infections have consequences for aspects of host 'state', parasites might induce the evolution of individual differences in certain types of host behaviour in populations with endemic infections. Also, because infection often leads to specific changes in axes of personality, parasite infections have the potential to decouple behavioural syndromes. Host-parasite systems therefore provide researchers with valuable tools to study personality variation and behavioural syndromes from a proximate and ultimate perspective.     

Adaptive strategies for managing uncertainty may explain personality‐related differences in behavioural plasticity

There is growing evidence that individuals within populations show consistent differences in their behaviour across contexts (personality), and that personality is associated with the extent to which individuals adjust their behaviour as function of changing conditions (behavioural plasticity). We propose an evolutionary explanation for a link between personality and plasticity based upon how individuals manage uncertainty. Individuals can employ three categories of tactics to manage uncertainty. They can 1) gather information (sample) to reduce uncertainty, 2) show strategic (state‐dependent) preferences for options that differ in their associated variances in rewards (i.e. variance‐sensitivity), or 3) invest in insurance to mitigate the consequences of uncertainty. We explicitly outline how individual differences in the use of any of these tactics can generate personality‐related differences in behavioural plasticity. For example, sampling effort is likely to co‐vary with individual activity and exploration behaviours, while simultaneously creating population variation in reactions to changes in environmental conditions. Individual differences in the use of insurance may be associated with differences in risk‐taking behaviours, such as boldness in the face of predation, thereby influencing the degree of adaptive plasticity across individuals. Population variation in responsiveness to environmental changes may also reflect individual differences in variance‐sensitivity, because stochastic change in the environment increases variances in rewards, which may both attract and benefit variance‐prone individuals, but not variance‐averse individuals. We review the existing evidence that individual variation in strategies for managing uncertainty exist, and describe how positive‐feedbacks between sampling, variance‐sensitivity and insurance can maintain and exaggerate even small initial differences between individuals in the relative use of these tactics. Given the pervasiveness of the problem of uncertainty, alternative strategies for managing uncertainty may provide a powerful explanation for consistent differences in behaviour and behavioural plasticity for a wide range of traits.