Animal Personality Traits Research Articles

Repeatable social network node-based metrics across populations and contexts in a passerine.

Behavioural traits are considered animal personality traits when individuals differ consistently in their expression across time and across context. Here, we test this idea on three metrics derived from social interaction networks (strength, betweenness and closeness). Using experimental data from house sparrows in captive populations, and observational data from house sparrows in a wild population, we show that all three metrics consistently exhibit repeatability across both study populations and two methods of recording interactions. The highest repeatability values were estimated in male-only captive groups, whereas repeatabilities estimated in single-sex networks subsetted from mixed-sex groups showed no sex specificity. We also show that changes in social group composition led to a decrease in repeatability for up to six months. This work provides substantial and generalizable support for the notion that social network node-based metrics can be considered animal personalities. Our work suggests that social network traits may be heritable and thus could be selected for.

More exploratory house mice judge an ambiguous situation more negatively.

Exploration tendency, one of the most investigated animal personality traits, may be driven by either positive (when seeking interesting information) or negative (to reduce the uncertainty of the environment) affective/emotional profiles. To disentangle the valence of the affective state associated with exploration trait, we applied a judgment bias test to evaluate the animals' responses in an ambiguous situation, allowing an assessment of their affective state or mood. Experiments were carried out in male house mice (Mus musculus) of wild origin. Individual differences in exploration tendency were assessed by repeated open field and novel object tests. To evaluate the animals' judgment bias, we trained the subjects for 8days in a 3-arm maze to discriminate between two extreme locations (outer arms: either positively reinforced with sugary water or less-positively reinforced with plain water), in terms of a shorter latency to approach the positively reinforced arm. After this learning criterion was reached, we repeatedly tested their responses to an ambiguous location (intermediate arm). The latencies to approach and consume the ambiguous reward were highly repeatable over the 3days of testing; hence individuals expressed a stable judgment bias. Most importantly, more exploratory animals showed a more negative judgment bias, which supports the hypothesis that a higher exploration tendency was associated with a negative affective state. Further studies should investigate whether exploration in different situations might be due to distinct affective states.

The genetic basis of animal behavioural diversity in natural populations.

Individual differences in animal behaviour influence ecological and evolutionary processes. Much behavioural variation has a heritable component, suggesting that genetics may play a role in its development. Yet, the study of the mechanistic description linking genes to behaviour in nature remains in its infancy, and such research is considered a challenge in contemporary biology. Here, we performed a literature review and meta-analysis to assess trends in analytical approaches used to investigate the relationship between genes and behaviour in natural systems, specifically candidate gene approaches, quantitative trait locus (QTL) mapping, and genome-wide association studies (GWAS). We aimed to determine the efficacy and success of each approach, while also describing which behaviours and species were examined by researchers most often. We found that the majority of QTL mapping and GWAS results revealed a significant or suggestive effect (Zr=0.3 [95% CI: 0.25:0.35] and Zr=0.39 [0.33:0.46], respectively) between the trait of interest and genetic marker(s) tested, while over half of candidate gene accounts (Zr=0.16 [0.11:0.21]) did not find a significant association. Approximately a third of all study estimates investigated animal personality traits; though, reproductive and migratory behaviours were also well-represented. Our findings show that despite widespread accessibility of molecular approaches given current sequencing technologies, efforts to elucidate the genetic basis of behaviour in free-ranging systems has been limited to relatively few species. We discuss challenges encountered by researchers, and recommend integration of novel genomic methods with longitudinal studies to usher in the next wave of behavioural genomic research.

Long‐term repeatability and stability of three personality traits in meadow voles

AbstractConsistency of animal personality traits is expected. Mounting evidence suggests that animal personalities will change with time and major life events. Research into this phenomenon is often limited to span only short periods of time in the animals’ life or have few measures across their life. We measured Exploration, Activity and Boldness of litters of meadow voles, Microtus pennsylvanicus, in a pair of novel object tests six times over a year, representing a large proportion of the lifespan of the vole. Our null hypothesis was that the personality traits would not change, and our first alternative hypothesis was that personality traits would change equally for all individuals as they aged. Neither was supported. The data support our third hypothesis that traits would change and change unequally. Activity and Boldness were consistent across time. However, Exploratory behaviour increased with the vole's age. The supported models show that personality traits change consistently for related individuals, but the litters differ in their change with age. This suggests that meadow vole life history affects how personality traits develop, but not necessarily their magnitudes. We also tested the repeatability of traits across the testing intervals. We discovered that the repeatability of these three personality traits varied across time, depending on the trait. Activity was repeatable equally across ages. Exploration became more repeatable and Boldness became less so. Thus, personality traits may be more malleable than supposed, especially over long periods of time.

Individual exploratory responses are not repeatable across time or context for four species of food-storing corvid

Exploration is among one of the most studied of animal personality traits (i.e., individual-level behavioural responses repeatable across time and contexts). However, not all species show clear evidence of this personality trait, and this is particularly so for members of the Corvidae family. We assessed the exploratory behaviour of four food-caching corvid species: pinyon jays (Gymnorhinus cyanocephalus), Clark’s nutcrackers (Nucifraga columbiana), California scrub jays (Aphelocoma californica), and black-billed magpies (Pica hudsonia). Contextual repeatability was assessed through examining behavioural measures during the Novel Environment task and the Novel Object task, whereas temporal repeatability was assessed by examining changes in these measures over repeated trials. Our results suggest that, for corvids, an individual’s exploratory behaviour was not repeatable across contexts or over time. Hence, we found no evidence that exploration constitutes a personality trait for these species of corvid. We did find differences in exploratory behaviour, at a species level, that may be explained by relative reliance on cached food.

Intraspecific variation in boldness and exploration shapes behavioral responses to stress in Galápagos sea lion pups

Behavioral plasticity allows individual organisms to alter behavioral traits in response to environmental or state-dependent conditions. Structured animal personality traits often influence the degree of between-individual variation in plasticity within a population, especially as it relates to an individual’s behavioral coping style towards disturbances. Here, we investigated whether boldness and exploration measured during novel object tests in young Galapagos sea lion pups were associated with individual behavioral responses towards capture. We found that only state-related traits, such as sex and body condition, and not personality affected baseline capture responses (struggle and escape responses after release). Male pups and those that maintained good body condition tended to show initially low struggle and escape behavior. These responses, however, were not highly repeatable within individuals during the study period. Different personality types greatly differed in how they modulated their behavior over time, and few sea lions utilized fixed strategies. Unexpectedly, bolder but less explorative individuals habituated quickest to capture, while shy and explorative pups increased responses over time. Pups found to be bolder but less explorative in novel object tests also moved between fewer sites within their habitat based on observational data. This may suggest that these individuals may be less sensitive to disturbances by increasing familiarity within a limited home range. Our results illustrate that individual pups may differentially assess risk and adjust behavior to cope with disturbance within their early life environment. Galapagos sea lions are an endangered pinniped species which live within a habitat characterized by a large degree of environmental uncertainty. We hypothesized that, depending on their personality, Galapagos sea lion pups may show varying degrees of behavioral plasticity in response to a dynamic early life environment. We show that bold pups and those that were less explorative during novel object tests and within their natural habitat were more likely to habituate quickly to stressful experiences (i.e., a capture and handling regime). We postulate that a familiarity with the physical and social environment in addition to personality traits may play an important role in how individual sea lion pups assess and cope with early life risks.

Personality predicts foraging site fidelity and trip repeatability in a marine predator.

Animal populations are often comprised of both foraging specialists and generalists. For instance, some individuals show higher foraging site fidelity (spatial specialization) than others. Such individual differences in degree of specialization can persist over time‐scales of months or even years in long‐lived animals, but the mechanisms leading to these different individual strategies are not fully understood.There is accumulating evidence that individual variation in foraging behaviour is shaped by animal personality traits, such as boldness. Despite this, the potential for boldness to drive differences in the degree of specialization is unknown.In this study, we used novel object tests to measure boldness in black‐legged kittiwakes (Rissa tridactyla) breeding at four colonies in Svalbard and deployed GPS loggers to examine their at‐sea foraging behaviour. We estimated the repeatability of foraging trips and used a hidden Markov model to identify locations of foraging sites in order to quantify individual foraging site fidelity.Across the breeding season, bolder birds were more repeatable than shy individuals in the distance and range of their foraging trips, and during the incubation period (but not chick rearing), bolder individuals were more site‐faithful. Birds exhibited these differences while showing high spatial similarity in foraging areas, indicating that site selection was not driven by personality‐dependent spatial partitioning.We instead suggest that a relationship between boldness and site fidelity may be driven by differences in behavioural flexibility between bold and shy individuals. Together, these results provide a potential mechanism by which widely reported individual differences in foraging specialization may emerge.

Starvation decreases behavioral consistency in a Neotropical harvestman

Consistent inter-individual differences in behavior have been shown in several animal groups, ranging from vertebrates to invertebrates. One of the most studied personality traits in animals is boldness, which is the tendency to expose to risky situations. Theory proposes that individuals’ state (e.g., body energy) would influence the expression of personality traits. In this study, we tested if boldness levels of the harvestman Mischonyx cuspidatus (measured as duration of death feigning/freezing after simulated predator attack) differ between two different states, namely sated and food deprived. We also tested if the degree of repeatability in boldness is affected by the individual state. We found no differences in the levels of boldness expressed by M. cuspidatus when comparing between different conditions (sated and food deprived) at a population level. However, we found that individuals showed more consistency in boldness when sated relative to a food-deprived condition. Finally, we suggest new avenues for future studies addressing personality in harvestmen.

Exploratory behavior undergoes genotype-age interactions in a wild bird.

Animal personality traits are often heritable and plastic at the same time. Indeed, behaviors that reflect an individual's personality can respond to environmental factors or change with age. To date, little is known regarding personality changes during a wild animals' lifetime and even less about stability in heritability of behavior across ages. In this study, we investigated age‐related changes in the mean and in the additive genetic variance of exploratory behavior, a commonly used measure of animal personality, in a wild population of great tits. Heritability of exploration is reduced in adults compared to juveniles, with a low genetic correlation across these age classes. A random regression animal model confirmed the occurrence of genotype–age interactions (G×A) in exploration, causing a decrease in additive genetic variance before individuals become 1 year old, and a decline in cross‐age genetic correlations between young and increasingly old individuals. Of the few studies investigating G×A in behaviors, this study provides rare evidence for this phenomenon in an extensively studied behavior. We indeed demonstrate that heritability and cross‐age genetic correlations in this behavior are not stable over an individual's lifetime, which can affect its potential response to selection. Because G×A is likely to be common in behaviors and have consequences for our understanding of the evolution of animal personality, more attention should be turned to this phenomenon in the future work.

Links between personality, early natal nutrition and survival of a threatened bird.

There is growing recognition that variation in animal personality traits can influence survival and reproduction rates, and consequently may be important for wildlife population dynamics. Despite this, the integration of personality research into conservation has remained uncommon. Alongside the establishment of personality as an important source of individual variation has come an increasing interest in factors affecting the development of personality. Recent work indicates the early environment, including natal nutrition, may play a stronger role in the development of personality than previously thought. In this study, we investigated the importance of three personality metrics (activity, boldness and acclimation time) for estimating survival of a threatened species, the hihi (Notiomystis cincta), and evaluated the influence of early natal nutrition on those metrics. Our results showed that boldness (as measured from a one-off cage test) had a positive effect on the probability of juvenile hihi surviving to adulthood. There was also a tendency for juveniles that received carotenoid supplementation in the nest to be bolder than those that did not, suggesting that the early environment had some influence on the expression of boldness in juvenile hihi. Linking the development of personality traits with ultimate effects on vital rates may benefit conservation management, as it could enable developmentally targeted management interventions. To our knowledge, this study is the first to identify potential linkages between early natal nutrition, personality and fitness in a wild-living population. This article is part of the theme issue 'Linking behaviour to dynamics of populations and communities: application of novel approaches in behavioural ecology to conservation'.

Social cues affect quantitative genetic variation and covariation in animal personality traits.

The social environment is expected to have substantial effects on behavior, and as a consequence, its heritability and evolvability. We investigated these effects by exposing Australian field crickets (Teleogryllus oceanicus) to either silence or recordings of male acoustic sexual signals. We used a combined pedigree and full-sib/half-sib breeding design to estimate the repeatability, heritability, and evolvability of behaviors related to boldness, exploration, and activity. All behaviors measured were significantly repeatable in both social environments. Additionally, most behaviors showed significant heritabilities in the two environments. We found no difference in repeatabilities between the silent and the acoustic environment but did find significant differences in the heritabilities and evolvabilities between these environments. There was a high degree of similarity between the phenotypic covariance matrices across the two environments, while the genotypic covariance matrices were highly dissimilar. Reflecting this, we found significant genotype-by-environment interactions for most of the behaviors. Lastly, we found that the repeatable aspect of behavior ("personality") was significantly heritable for most behaviors, but that these heritabilities were higher in the acoustic than in the silent environment. We conclude that the social environment can have a significant impact on the heritability and evolvability of behavior, and argue that evolutionary inferences from phenotypic studies should be made with caution.

Don't poke the bear: using tracking data to quantify behavioural syndromes in elusive wildlife

Animal personality traits and the emergence of behavioural syndromes, i.e. between-individual correlation of behaviours, are commonly quantified from behavioural observations in controlled environments. Subjecting large and elusive wildlife to controlled test situations is, however, rarely possible, suggesting that ecologists should exploit alternative measures of behaviours for quantifying differences between individuals. Our goal was to test whether movement and space use data can be used to quantify behavioural syndromes in the wild. We quantified six behaviours from GPS and dual motion sensor tracking devices of 46 adult female brown bears followed in southcentral Sweden over the summer and early autumn. As well as daily travel distance, an indicator for activity, and daily displacement, an indicator for exploration, we quantified four behaviours that increase a bear's likelihood of encountering humans and could thus serve as indicators for boldness: diurnality, selection for roads and selection for two open habitat types, bogs and clearcuts, with low lateral cover. We tested (1) whether behaviours showed repeatable between-individual variation (animal personality) and (2) whether behaviours were correlated between individuals and thus formed a behavioural syndrome. Repeatability of behaviours ranged from 0.16 to 0.61 confirming between-individual variation in movement, activity and space use. A multivariate mixed model revealed significant positive correlations between travel distance, displacement and diurnality, suggesting the existence of an activity–exploration and potentially partial boldness syndrome in our bear population. Selection for exposed or human-frequented habitats were uncorrelated with the activity–exploration syndrome and with each other, albeit there was a trend for stronger road avoidance by bears that readily used clearcuts. We show that large tracking data sets can be used to quantify between-individual correlation in spatial behaviours. We suggest that delineating behavioural types from wildlife tracking data will be of increasing interest because of the importance of animal personality for ecological processes, wildlife conservation and human–wildlife coexistence.

Experimental evolution of personality traits: open-field exploration in bank voles from a multidirectional selection experiment

Evolution of complex physiological adaptations could be driven by natural selection acting on behavioral traits. Consequently, animal personality traits and their correlation with physiological traits have become an engaging research area. Here, we applied a unique experimental evolution model—lines of bank voles selected for (A) high exercise-induced aerobic metabolism, (H) ability to cope with low-quality herbivorous diet, and (P) intensity of predatory behavior, that is, traits shaping evolutionary path and diversity of mammals—and asked how the selection affected the voles’ personality traits, assessed in an open field test. The A- and P-line voles were more active, whereas the H-line voles were less active, compared those from unselected control lines (C). H-line voles moved slower but on more meandering trajectories, which indicated a more thorough exploration, whereas the A- and P-line voles moved faster and on straighter trajectories. A-line voles showed also an increased escape propensity, whereas P-line voles tended to be bolder. The remarkable correlated responses to the selection indicate a common genetic underlying mechanism of behavioral and physiological traits, and support the paradigm of evolutionary physiology built around the concept of correlated evolution of behavior and physiology.

Cortisol during adolescence organises personality traits and behavioural syndromes

Despite the growing evidence for the importance of developmental experiences shaping consistent individual differences in behaviour and physiology, the role of endocrine factors underlying the development and maintenance of such differences across multiple traits, remains poorly understood. Here, we investigated how an experimental manipulation of circulating glucocorticoids during early adolescence affects behavioural and physiological variation and covariation later in life in the precocial cavy (Cavia aperea). Plasma cortisol concentrations were experimentally elevated by administering cortisol via food for 3 weeks. Struggle docility, escape latency, boldness, exploration and social behaviour were then tested three times after individuals attained sexual maturity. In addition, blood samples were taken repeatedly to monitor circulating cortisol concentrations. Exogenous cortisol affected mean trait expression of plasma cortisol levels, struggle docility and escape latency. Repeatability of cortisol and escape latency was increased and repeatability of struggle docility tended to be higher (approaching significance) in treated individuals. Increased repeatability was mainly caused by an increase of among-individual variance. Correlations among docility, escape latency and cortisol were stronger in treated animals compared to control animals. These results suggest that exposure to elevated levels of cortisol during adolescence can alter animal personality traits as well as behavioural syndromes. Social and risk-taking traits showed no correlation with cortisol levels and were unaffected by the experimental manipulation, indicating behavioural modularity. Taken together, our data highlight that cortisol can have organising effects during adolescence on the development of personality traits and behavioural syndromes, adding to the increasing evidence that not only early life but also adolescence is an important sensitive period for behavioural development.

The number of neurons in specific amygdala regions is associated with boldness in mink: a study in animal personality.

Conspecifics vary consistently in their behavioural responses towards environment stimuli such as exposure to novel objects; ethologists often refer to this variability as animal personality. The neurological mechanisms underlying animal personality traits remain largely unknown, but linking the individual variation in emotional expression to brain structural and neurochemical factors is attracting renewed interest. While considerable research has focused on hormonal and neurotransmitter effects on behavioural responses, less is known about how individual variation in the number of specific neuron populations contributes to individual variation in behaviour. The basolateral amygdala (BLA) and the central nuclei of the amygdala (CeA) mediate emotional processing by regulating behavioural responses of animals in a potentially threatening situation. As such, these structures are good candidates for evaluating the relationship between neuronal populations and behavioural traits. We now show that individual American mink (Neovison vison) reacting more boldly towards novelty have more neurons in the BLA than do their more timid conspecifics, suggesting that a developmental pattern of the number of amygdala neurons can influence behavioural traits of an adult animal. Furthermore, post hoc correlations revealed that individuals performing with higher arousal, as reflected by their frequency of startle behaviour, have more CeA neurons. Our results support a direct link between the number of neurons in amygdala regions and aspects of animal personality.

Does the use of a multi-trait, multi-test approach to measure animal personality yield different behavioural syndrome results?

Abstract A multi-trait, multi-test approach to investigate the convergence or discrimination between behavioural tests putatively targeting the same or different animal personality traits has been recommended, yet whether and how the approaches affect the identification of behavioural syndrome(s), the suite of correlated personality traits, requires investigation. Here, we used behavioural measures collected from five commonly used behavioural tests targeting three personality traits, evaluated their convergence/discrimination through exploratory factor analysis (EFA), and then explored whether the identification of syndrome changed based on the approach we used to quantify personality traits. Our results indicated that tests presumably targeting the same personality trait actually measured distinct behavioural aspects. Syndrome defined using correlation changed due to how we identified personality traits, but not when using structural equation models (SEMs). Overall, this study emphasizes that it is critical to clarify the approach and terms we use for ‘personality traits’ in the field of animal personality.

An evolutionary framework outlining the integration of individual social and spatial ecology.

Behaviour is the interface between an organism and its environment, and behavioural plasticity is important for organisms to cope with environmental change. Social behaviour is particularly important because sociality is a dynamic process, where environmental variation influences group dynamics and social plasticity can mediate resource acquisition. Heterogeneity in the ecological environment can therefore influence the social environment. The combination of the ecological and social environments may be interpreted collectively as the "socioecological environment," which could explain variation in fitness. Our objective was to outline a framework through which individual social and spatial phenotypes can be integrated and interpreted as phenotypes that covary as a function of changes in the socioecological environment. We propose the socioecological environment is composed of individual behavioural traits, including sociality and habitat selection, both of which are repeatable, potentially heritable and may reflect animal personality traits. We also highlight how ecological and social niche theory can be applied to the socioecological environment framework, where individuals occupy different socioecological niches. Individual sociality and habitat selection are also density-dependent, and theory predicts that density-dependent traits should affect reproduction, survival, and therefore fitness and population dynamics. We then illustrate the proximate links between sociality, habitat selection and fitness as well as the ultimate, and possibly adaptive, consequences associated with changes in population density. The ecological, evolutionary and applied implications of our proposed socioecological environment framework are broad and changes in density could influence individual fitness and population dynamics. For instance, human-induced environmental changes can influence population density, which can affect the distribution of social and spatial phenotypes within a population. In summary, we outline a conceptual framework that incorporates individual social and spatial behavioural traits with fitness and we highlight a range of ecological and evolutionary processes that are likely associated with the socioecological environment.

Personality in the chimpanzees of Gombe National Park

Researchers increasingly view animal personality traits as products of natural selection. We present data that describe the personalities of 128 eastern chimpanzees (Pan troglodytes schweinfurthii) currently living in or who lived their lives in the Kasekela and Mitumba communities of Gombe National Park, Tanzania. We obtained ratings on 24 items from an established, reliable, well-validated questionnaire used to study personality in captive chimpanzee populations. Ratings were made by former and present Tanzanian field assistants who followed individual chimpanzees for years and collected detailed behavioral observations. Interrater reliabilities across items ranged from acceptable to good, but the personality dimensions they formed were not as interpretable as those from captive samples. However, the personality dimensions corresponded to ratings of 24 Kasekela chimpanzees on a different questionnaire in 1973 that assessed some similar traits. These correlations established the repeatability and construct validity of the present ratings, indicating that the present data can facilitate historical and prospective studies that will lead to better understanding of the evolution of personality in chimpanzees and other primates.

Repeatability and consistency of individual behaviour in juvenile and adult Eurasian harvest mice

Knowledge on animal personality has provided new insights into evolutionary biology and animal ecology, as behavioural types have been shown to affect fitness. Animal personality is characterized by repeatable and consistent between-individual behavioural differences throughout time and across different situations. Behavioural repeatability within life history stages and consistency between life history stages should be checked for the independence of sex and age, as recent data have shown that males and females in some species may differ in the repeatability of behavioural traits, as well as in their consistency. We measured the repeatability and consistency of three behavioural and one cognitive traits in juvenile and adult Eurasian harvest mice (Micromys minutus). We found that exploration, activity and boldness were repeatable in juveniles and adults. Spatial recognition measured in a Y Maze was only repeatable in adult mice. Exploration, activity and boldness were consistent before and after maturation, as well as before and after first sexual contact. Data on spatial recognition provided little evidence for consistency. Further, we found some evidence for a litter effect on behaviours by comparing different linear mixed models. We concluded that harvest mice express animal personality traits as behaviours were repeatable across sexes and consistent across life history stages. The tested cognitive trait showed low repeatability and was less consistent across life history stages. Given the rising interest in individual variation in cognitive performance, and in its relationship to animal personality, we suggest that it is important to gather more data on the repeatability and consistency of cognitive traits.

Does Individual Variation in Male Mate Choice Copying Reflect Differences in Social Responsiveness?

During mate choice copying (MCC), individuals evaluate the attractiveness of potential mating partners while observing if they are preferred or rejected by other individuals. Numerous studies have examined female MCC, while its role during male mate choice received far less attention. Using western mosquitofish (Gambusia affinis) as our study organism, we asked if individual focal males would spend more time associating with the previously rejected female of a pair of stimulus females (a large- and a small-bodied female) after they had observed another (model) male consort the rejected female. We used the resulting ‘copying scores’ to test if individual variation in MCC can be explained as part of a larger behavioural syndrome (i.e., social responsiveness). In one experiment, we assessed focal males’ copying tendencies and then tested them in another context of socially-dependent mate choice (audience effects). Males with high copying tendencies were also more likely to respond to rival presence by showing audience effects (concealment of mating preferences). In another experiment, we assessed copying tendencies and behavioral correlates of boldness, activity and sociability (shoaling tendencies) of the focal males. All three traits were correlated, but did not predict male copying tendencies. Altogether, our results suggest that male MCC may indeed be part of a social responsiveness syndrome (explaining correlated responses in two contexts of socially-dependent mate choice), but this syndrome seems to be independent of standard behavioral parameters widely used to quantify animal personality traits.