Peter A. Biro

Are animal personality traits linked to life-history productivity?

Animal personality traits such as boldness, activity and aggressiveness have been described for many animal species. However, why some individuals are consistently bolder or more active than others, for example, is currently obscure. Given that life-history tradeoffs are common and known to promote inter-individual differences in behavior, we suggest that consistent individual differences in animal personality traits can be favored when those traits contribute to consistent individual differences in productivity (growth and/or fecundity). A survey of empirical studies indicates that boldness, activity and/or aggressiveness are positively related to food intake rates, productivity and other life-history traits in a wide range of taxa. Our conceptual framework sets the stage for a closer look at relationships between personality traits and life-history traits in animals.

Do consistent individual differences in metabolic rate promote consistent individual differences in behavior

Consistent individual differences (CIDs) in behavior are a widespread phenomenon in animals, but the proximate reasons for them are unresolved. We discuss evidence for the hypothesis that CIDs in energy metabolism, as reflected by resting metabolic rate (RMR), promote CIDs in behavior patterns that either provide net energy (e.g. foraging activity), and/or consume energy (e.g. courtship activity). In doing so, we provide a framework for linking together RMR, behavior, and life-history productivity. Empirical studies suggest that RMR is (a) related to the capacity to generate energy, (b) repeatable, and (c) correlated with behavioral output (e.g. aggressiveness) and productivity (e.g. growth). We conclude by discussing future research directions to clarify linkages between behavior and energy metabolism in this emerging research area.

Rapid depletion of genotypes with fast growth and bold personality traits from harvested fish populations

The possibility for fishery-induced evolution of life history traits is an important but unresolved issue for exploited fish populations. Because fisheries tend to select and remove the largest individuals, there is the evolutionary potential for lasting effects on fish production and productivity. Size selection represents an indirect mechanism of selection against rapid growth rate, because individual fish may be large because of rapid growth or because of slow growth but old age. The possibility for direct selection on growth rate, whereby fast-growing genotypes are more vulnerable to fishing irrespective of their size, is unexplored. In this scenario, faster-growing genotypes may be more vulnerable to fishing because of greater appetite and correspondingly greater feeding-related activity rates and boldness that could increase encounter with fishing gear and vulnerability to it. In a realistic whole-lake experiment, we show that fast-growing fish genotypes are harvested at three times the rate of the slow-growing genotypes within two replicate lake populations. Overall, 50% of fast-growing individuals were harvested compared with 30% of slow-growing individuals, independent of body size. Greater harvest of fast-growing genotypes was attributable to their greater behavioral vulnerability, being more active and bold. Given that growth is heritable in fishes, we speculate that evolution of slower growth rates attributable to behavioral vulnerability may be widespread in harvested fish populations. Our results indicate that commonly used minimum size-limits will not prevent overexploitation of fast-growing genotypes and individuals because of size-independent growth-rate selection by fishing.

Predators select against high growth rates and risk-taking behaviour in domestic trout populations

Domesticated (farm) salmonid fishes display an increased willingness to accept risk while foraging, and achieve high growth rates not observed in nature. Theory predicts that elevated growth rates in domestic salmonids will result in greater risk–taking to access abundant food, but low survival in the presence of predators. In replicated whole–lake experiments, we observed that domestic trout (selected for high growth rates) took greater risks while foraging and grew faster than a wild strain. However, survival consequences for greater growth rates depended upon the predation environment. Domestic trout experienced greater survival when risk was low, but lower survival when risk was high. This suggests that animals with high intrinsic growth rates are selected against in populations with abundant predators, explaining the absence of such phenotypes in nature. This is, to our knowledge, the first large–scale field experiment to directly test this theory and simultaneously quantify the initial invasibility of domestic salmonid strains that escape into the wild from aquaculture operations, and the ecological conditions affecting their survival.

Small within-day increases in temperature affects boldness and alters personality in coral reef fish

Consistent individual differences in behaviour, termed personality, are common in animal populations and can constrain their responses to ecological and environmental variation, such as temperature. Here, we show for the first time that normal within-daytime fluctuations in temperature of less than 3°C have large effects on personality for two species of juvenile coral reef fish in both observational and manipulative experiments. On average, individual scores on three personality traits (PTs), activity, boldness and aggressiveness, increased from 2.5- to sixfold as a function of temperature. However, whereas most individuals became more active, aggressive and bold across temperature contexts (were plastic), others did not; this changed the individual rank order across temperatures and thus altered personality. In addition, correlations between PTs were consistent across temperature contexts, e.g. fish that were active at a given temperature also tended to be both bold and aggressive. These results (i) highlight the importance of very carefully controlling for temperature when studying behavioural variation among and within individuals and (ii) suggest that individual differences in energy metabolism may contribute to animal personality, given that temperature has large direct effects on metabolic rates in ectotherms.

Sampling bias resulting from animal personality

Sampling is something most ecologists do, whether to estimate population size, determine trait distributions or collect samples for laboratory studies. Ecologists are well aware that most sampling techniques possess inherent sampling inefficiency and bias, and techniques have therefore been developed to minimise bias [1]. One form of sampling bias of great significance, gone largely unnoticed, is due to consistent individual differences in behaviour, termed personality [2]. Personality exists in diverse animal taxa, and is assumed to result from underlying behavioural tendencies that affect behaviour in different contexts, that vary across individuals and that are reasonably stable across time [2].

Unpredictable animals: individual differences in intraindividual variability (IIV)

When an individual is repeatedly observed or tested in the same context, it does not always express the same behaviour. Intraindividual variability (IIV) refers to the short-term, unpredictable, reversible variation in behaviour that often occurs in this situation. Although individual differences in IIV have been well documented in humans, this topic has been virtually ignored by researchers studying other animals. Here, we review evidence from humans and animals that IIV can vary in important ways across individuals (e.g. as a function of age or prior experience) and that individual differences in IIV may be related to differences in performance. However, most statistical models currently used to study individual differences in behaviour in animals rely on the assumption that IIV does not vary across individuals. Using ‘boldness’ data for hermit crabs, Pagurus bernhardus, and Ward’s damselfish, Pomacentrus wardi, we show how to measure IIV when behaviour systematically changes over a series of observations (e.g. as a result of habituation), and how to avoid the adverse effects of censored data on estimates of IIV. After controlling for systematic changes in behaviour over time, we observed strong, significant individual differences in IIV in both species. That is, some individuals were much more predictable in the same situation than were others. We conclude by discussing proximate and ultimate factors that might have contributed to interindividual variation in IIV in these species, and the implications of our findings for methods currently used to study individual differences in behaviour in animals.

FROM INDIVIDUALS TO POPULATIONS: PREY FISH RISK-TAKING MEDIATES MORTALITY IN WHOLE-SYSTEM EXPERIMENTS

Recent research suggests that the behavior of individuals under risk of pre- dation could be a key link between individual behavior and population and community dynamics. Yet existing theory remains largely untested at large spatial and temporal scales. We manipulated food available to age-0 rainbow trout while at risk of cannibalism, in a replicated factorial whole-lake experiment, to test whether the trade-off between growth and mortality rates is mediated by foraging activity by young fish under predation risk. We found that this trade-off exists for young fish at the whole-system scale, and that food- dependent behavioral variation has large mortality consequences. In high-food lakes, age- 0 trout spent less time moving, fewer individuals swam continuously, and those swimming continuously swam at slower speeds relative to those in low-food lakes. Age-0 trout also used deep, risky habitats less when food was abundant. This lower activity, combined with avoidance of risky habitats, coincided with 68% higher survival in high-food lakes. If general, this trade-off may be a key mechanism linking individual behavior to population- level processes in size-structured populations.

Ontogeny of energy allocation reveals selective pressure promoting risk-taking behaviour in young fish cohorts

Given limited food, prey fishes in a temperate climate must take risks to acquire sufficient reserves for winter and/or to outgrow vulnerability to predation. However, how can we distinguish which selective pressure promotes risk-taking when larger body size is always beneficial? To address this question, we examined patterns of energy allocation in populations of age-0 trout to determine if greater risk-taking corresponds with energy allocation to lipids or to somatic growth. Trout achieved maximum growth rates in all lakes and allocated nearly all of their acquired energy to somatic growth when small in early summer. However, trout in low-food lakes took greater risks to achieve this maximal growth, and therefore incurred high mortality. By late summer, age-0 trout allocated considerable energy to lipids and used previously risky habitats in all lakes. These results indicate that: (i) the size-dependent risk of predation (which is independent of behaviour) promotes risk-taking behaviour of age-0 trout to increase growth and minimize time spent in vulnerable sizes; and (ii) the physiology of energy allocation and behaviour interact to mediate growth/mortality trade-offs for young animals at risk of predation and starvation.

Staying Cool: Behavioral Thermoregulation during Summer by Young-of-Year Brook Trout in a Lake

Abstract Thermal habitat selection and behavior by young-of-year brook trout Salvelinus fontinalis was studied in a lake in central Ontario, Canada. In May, trout foraged actively within 2 m of shore in the warmest water available (∼15°C). In early June, trout foraged near the bottom within 4 m of shore, where bottom water temperatures were near, or at, the upper thermal tolerance, for trout, of 20°C. In July, when ambient water temperatures ranged from 23°C to 27°C, trout lay on the bottom in the coldest water available (18–20°C) in discrete areas 3–8 m from shore. Flow rate of cold groundwater accounted for 87% of the variance in trout density in these areas, and the data suggest that a minimum flow rate of 125 mL·m−2·min−1 is required for trout to take up station. When trout were displaced from holding positions, sites with greater groundwater flow were more quickly reoccupied by trout than sites with lower flow. Experimentally created trough-like depressions at these sites attracted higher densities o...