Individual Variation In Behaviour Research Articles

Not so fast: Paradoxically increased variability in the glucose tolerance test due to food withdrawal in continuous glucose-monitored mice

ObjectiveThis study was performed to determine the effect of fasting on reproducibility of the glucose tolerance test. Due to individual variation in animal feeding behaviors, fasting animals prior to metabolic and behavioral experiments is widely held to reduce inter-subject variation in glucose and metabolic parameters of preclinical rodent models. Reducing variability is especially important for studies where initial metabolite levels can influence the magnitude of experimental interventions, but fasting also imposes stress that may distort the variables of interest. One such intervention is the glucose tolerance test (GTT) which measures the maximum response and recovery following a bolus of exogenous glucose. We sought to investigate how fasting affects the response of individual mice to a GTT. MethodsUsing simultaneous continuous glucose monitoring (CGM) and indirect calorimetry, we quantified blood glucose, physical activity, body temperature, metabolic rates, and food consumption levels on a minute-to-minute basis in adult male mice for 4 weeks. We tested the effects of a 4-h or 18-h fast on the GTT to examine the effect of food withdrawal in light or dark photoperiods. Studies were also performed with 4-h fasting in additional mice without implanted CGM probes. ResultsContrary to our expectations, a 4-h fast during the light photoperiod promotes a paradoxical increase in inter-animal variation in metabolic rate, physical activity, body temperature, glycemia, and glucose tolerance. This hyperglycemic and hyper-metabolic phenotype promotes increased corticosterone levels and is consistent with a behavioral stress response to food deprivation, even in well-fed mice. We find that mice undergoing an 18-h fast entered torpor, a hibernation-like state. In addition to low body temperature and metabolic rate, torpor is also associated with glucose levels 56 mg/dl lower than those seen in mice with ad libitum access to food. Moreover, the time spent in torpor affects the response to a GTT. ConclusionOur results suggest fasting mice before glucose tolerance testing, and perhaps other experiments, can have the opposite of the intended effect where fasting can increase, rather than decrease, experimental variability.

Biobehavioral susceptibility for obesity in childhood: Behavioral, genetic and neuroimaging studies of appetite

Modern food environments are conducive to overeating and weight gain, but not everyone develops obesity. One reason for this may be that individuals differ in appetitive characteristics, or traits, that manifest early in life and go on to influence their behavioral susceptibility to gain and maintain excess weight. Classic studies showing that eating behavior in children can be measured by behavioral paradigms such as tests of caloric compensation and eating in the absence of hunger inspired the development and validation of psychometric instruments to assess appetitive characteristics in children and infants. A large body of evidence now suggests that food approach traits increase obesity risk, while food avoidant traits, such as satiety responsiveness, decrease obesity risk. Twin studies and genetic association studies have demonstrated that appetitive characteristics are heritable, consistent with a biological etiology. However, family environment factors are also influential, with mounting evidence suggesting that genetic and environmental risk factors interact and correlate with consequences for child eating behavior and weight. Further, neuroimaging studies are revealing that individual differences in responses to visual food cues, as well as to small tastes and larger amounts of food, across a number of brain regions involved in reward/motivation, cognitive control and other functions, may contribute to individual variation in appetitive behavior. Growing evidence also suggests that variation on psychometric measures of appetite is associated with regional differences in brain structure, and differential patterns of resting state functional connectivity. Large prospective studies beginning in infancy promise to enrich our understanding of neural and other biological underpinnings of appetite and obesity development in early life, and how the interplay between genetic and environmental factors affects appetitive systems. The biobehavioral susceptibility model of obesity development and maintenance outlined in this narrative review has implications for prevention and treatment of obesity in childhood.

The influence of flow on movement of a headwater specialist in an intermittent urban headwater stream

AbstractFish movement in freshwater streams is often tied to patterns of flow, especially when the stream is intermittent, flashy, or some combination thereof. Relationships between flow and fish movement are exacerbated in small urban streams in impervious surface‐dominated watersheds that lead to extreme flow conditions. Here we examined the response of Creek Chub (Semotilus atromaculatus), a common pioneer fish species in eastern North America, to flow events of various sizes in a small intermittent urban headwater stream. Our primary goals were to determine (1) the level of Creek Chub movement occurring in an intermittent headwater stream, (2) the influence of flow on movement direction and magnitude, (3) the degree of variation in individual movement behaviour, and (4) mortality rates in the system. The study used PIT tagging and telemetry surveys to monitor Creek Chub movement over a six‐month period. We found that individual Creek Chub in our study system were generally mobile (71%), but usually moved short distances only; however, a few individuals moved much farther, with a maximum single movement of 521 m. Moderate or major flow events increased the probability and magnitude of Creek Chub movement, but there was no detectable asymmetry towards upstream or downstream directionality. Movement was variable among individual fish and fish were able to respond to large and small flow events to move among habitats. Finally, we observed relatively high, but episodic, mortality in our study site, potentially related to pollution events. Evidence of elevated mortality points to the potential importance of movement in highly disturbed streams, as it may enable recolonization after local extirpation.

Disparate behavioral types in wild and reared juveniles of gilthead seabream

Fish differ consistently in behavior within the same species and population, reflecting distinct behavioral types (BTs). Comparing the behavior of wild and reared individuals provides an excellent opportunity to delve into the ecological and evolutionary consequences of BTs. In this work, we evaluated the behavioral variation of wild and reared juvenile gilthead seabreams, Sparus aurata, a highly relevant species for aquaculture and fisheries. We quantified behavioral variation along the five major axes of fish behavioral traits (exploration-avoidance, aggressiveness, sociability, shyness-boldness, and activity) using standardized behavioral tests and a deep learning tracking algorithm for behavioral annotation. Results revealed significant repeatability in all five behavior traits, suggesting high consistency of individual behavioral variation across the different axes in this species. We found reared fish to be more aggressive, social and active compared to their wild conspecifics. Reared individuals also presented less variance in their aggressiveness, lacking very aggressive and very tame individuals. Phenotypic correlation decomposition between behavioral types revealed two different behavioral syndromes: exploration-sociability and exploration-activity. Our work establishes the first baseline of repeatability scores in wild and reared gilthead seabreams, providing novel insight into the behavior of this important commercial species with implications for fisheries and aquaculture.

Individual variation in preference behavior in sailfin fish refines the neurotranscriptomic pathway for mate preference.

Social interactions can drive distinct gene expression profiles which may vary by social context. Here we use female sailfin molly fish (Poecilia latipinna) to identify genomic profiles associated with preference behavior in distinct social contexts: male interactions (mate choice) versus female interactions (shoaling partner preference). We measured the behavior of 15 females interacting in a non-contact environment with either two males or two females for 30 min followed by whole-brain transcriptomic profiling by RNA sequencing. We profiled females that exhibited high levels of social affiliation and great variation in preference behavior to identify an order of magnitude more differentially expressed genes associated with behavioral variation than by differences in social context. Using a linear model (limma), we took advantage of the individual variation in preference behavior to identify unique gene sets that exhibited distinct correlational patterns of expression with preference behavior in each social context. By combining limma and weighted gene co-expression network analyses (WGCNA) approaches we identified a refined set of 401 genes robustly associated with mate preference that is independent of shoaling partner preference or general social affiliation. While our refined gene set confirmed neural plasticity pathways involvement in moderating female preference behavior, we also identified a significant proportion of discovered that our preference-associated genes were enriched for 'immune system' gene ontology categories. We hypothesize that the association between mate preference and transcriptomic immune function is driven by the less well-known role of these genes in neural plasticity which is likely involved in higher-order learning and processing during mate choice decisions.

Boldness and physiological variation in round goby populations along their Baltic Sea invasion front

The round goby (Neogobius melanostomus) is a fish native to the Ponto-Caspian region that is highly invasive through freshwater and brackish habitats in northern Europe and North America. Individual behavioural variation appears to be an important factor in their spread, for example a round goby's personality traits can influence their dispersal tendency, which may also produce variation in the behavioral composition of populations at different points along their invasion fronts. To further analyze the drivers of behavioral variation within invasive round goby populations, we focused on two populations along the Baltic Sea invasion front with closely comparable physical and community characteristics. Specifically, this study measured personality within a novel environment and predator response context (i.e., boldness), and directly analyzed links between individuals’ personality traits and their physiological characteristics and stress responses (i.e., blood cortisol and lactate, brain neurotransmitters). In contrast to previous findings, the more recently established population had similar activity levels but were less bold in response to a predator cue than the older population, which suggests that behavioral compositions within our study populations may be more driven by local environmental conditions rather than being a result of personality-biased dispersal. Furthermore, we found that both populations showed similar physiological stress responses, and there also appeared to be no detectable relationship between physiological parameters and behavioral responses to predator cues. Instead, body size and body condition were important factors influencing individual behavioral responses. Overall, our results reinforce the importance of boldness traits as a form of phenotypic variation in round goby populations in the Baltic Sea. We also highlight the importance of these traits for future studies specifically testing for effects of invasion processes on phenotypic variation in the species. Nonetheless, our results also highlight that the physiological mechanisms underpinning behavioural variation in these populations remain unclear.

Mid‐summer fish behavior in a high‐latitude twilight zone

AbstractThe behavior of the mesopelagic fish Benthosema glaciale was studied at 60°N in mid‐summer. We hypothesized that diel vertical migration (DVM) is constrained by short and dusk nights (surface illumination > 10−2 μmol m−2 s−1) and that individuals are active at depth during the long summer days. Submerged echosounders provided high‐resolution data throughout the water column. During the day, a part of the population ascended toward the increasing daylight. Short vertical relocations were followed by minutes of vertical inactivity. Swimming included horizontal turns and loops associated with the vertical steps. Normal DVM was initiated ~ 4 h before sunset and reflected independent individual decisions. The fish initially ascended stepwise but switched to mostly straight upwards swimming attaining 3–4 cm s−1. Their vertical speed was faster than the slow ascent of isolumes and even the deepest living fish potentially could reach upper layers shortly after sunset. However, many individuals aborted their ascent and returned to depth before the darkest time of the night, while others returned downward closer to sunrise. The daytime swimming and individual variability in diel migration behavior have implications for encounters with prey and predators in the twilight zone and the biological carbon pump. A principal conclusion is that mesopelagic fishes can modify their behavior and migration patterns to suit a wide range of changing conditions.

Studying individual-level interactions can transform our understanding of avian mixed-species flocks

Abstract Avian mixed-species flocks are ubiquitous across habitats and a model for studying how heterospecific sociality influences the behavior and composition of animal communities. Here, we review the literature on mixed-species flocks and argue that a renewed focus on individual-level interactions among flock members can transform our understanding of this iconic, avian social system. Specifically, we suggest that an individual perspective will further our understanding of (1) how inter- and intraspecific variation in flock participation links to fitness costs and benefits, (2) the implications of familiarity between individuals in structuring mixed-species flock communities, and (3) how social roles within mixed-species flocks are related to social behavior within and across species. We summarize studies that use an individual perspective in each of these areas and discuss knowledge from conspecific social behavior to posit more broadly how individuals may shape mixed-species flocks. We encourage research approaches that incorporate individual variation in traits, relationships, and social roles in their assessment of mixed-species flocking dynamics. We propose that the analysis of individual variation in behavior will be particularly important for explicitly identifying fitness outcomes that led to the evolution of mixed-species flocks, which in turn affect community structure and resilience.

Mouse spontaneous behavior reflects individual variation rather than estrous state

Behavior is shaped by both the internal state of an animal and its individual behavioral biases. Rhythmic variation in gonadal hormones during the estrous cycle is a defining feature of the female internal state, one that regulates many aspects of sociosexual behavior. However, it remains unclear whether estrous state influences spontaneous behavior and, if so, how these effects might relate to individual behavioral variation. Here, we address this question by longitudinally characterizing the open-field behavior of female mice across different phases of the estrous cycle, using unsupervised machine learning to decompose spontaneous behavior into its constituent elements.1,2,3,4 We find that each female mouse exhibits a characteristic pattern of exploration that uniquely identifies it as an individual across many experimental sessions; by contrast, estrous state only negligibly impacts behavior, despite its known effects on neural circuits that regulate action selection and movement. Like female mice, male mice exhibit individual-specific patterns of behavior in the open field; however, the exploratory behavior of males is significantly more variable than that expressed by females both within and across individuals. These findings suggest underlying functional stability to the circuits that support exploration in female mice, reveal a surprising degree of specificity in individual behavior, and provide empirical support for the inclusion of both sexes in experiments querying spontaneous behaviors.

Differential associations of mentally-active and passive sedentary behaviours and physical activity with putative cognitive decline in healthy individuals and those with bipolar disorder: Findings from the UK Biobank cohort

ObjectivePhysical activity confers protection against age-related cognitive dysfunction, but less is known about whether cognition is impacted by sedentary behaviour, specifically its mentally-active and passive forms. Both physical inactivity and sedentary behaviour are highly prevalent in bipolar disorder (BD), yet if and how they relate to the disorder's cognitive symptoms remains unclear. In this study, we explored whether individual variation in physical activity and sedentary behaviour was directly associated with cognition and/or moderated the extent of decrements in cognitive performance occurring as a function of age in people with BD versus psychiatrically-healthy controls. MethodsRelevant UK Biobank data were available for 1074 BD patients and 59,653 psychiatrically-healthy controls, aged 40–70. Age, mentally-active (computer use) and passive sedentary behaviour (TV viewing), physical activity, diagnosis, and their interactions were regressed on a global cognitive score. ResultsCognition was inversely associated with physical activity (β = −0.003, CI: −0.003, −0.003) and mentally-passive sedentary behaviour (β = −0.03, CI: −0.04, −0.02) and positively associated with mentally-active sedentary behaviour (β = 0.13, CI: 0.12, 0.15). The latter association was stronger in the BD group. Mentally-active sedentary behaviour moderated the age-cognition association in both BD patients and controls (β = 0.01, CI: 0.008, 0.01), such that age-related decrements in cognition were more apparent in those who engaged in less mentally-active sedentary behaviours compared to those who engaged in more. ConclusionsMentally-active sedentary behaviour may not only attenuate age-related cognitive dysfunction, but may be important for cognition irrespective of age, particularly in those with BD. These findings support the notion that intellectual stimulation is a protective factor conferring cognitive benefits, even in the context of a sedentary lifestyle.

Behavioural responses are associated with mortality and mobility after reintroduction in the endangered Yellow Cardinal

Reintroductions are an important management tool for the conservation of threatened species. Personality or temperament of individuals is likely to influence survival in the reintroduction success of released individuals. We studied behavioural traits related to personality in the globally endangered Yellow Cardinal Gubernatrix cristata and analysed whether these traits were associated with the outcome of a reintroduction programme in Argentina. We found that exploratory behaviour and anti‐predator responses were correlated in Yellow Cardinals. Individuals with higher activity levels in the presence of a predator had a lower risk of mortality following release. Also, less neophobic individuals with increased exploratory behaviour moved farther away from the release site. High activity levels in terms of a predatory response could outweigh the cost of higher exposure to predators and may be beneficial in decreasing predation probability. We recommend the enhancement of predatory responses before release, especially in individuals with inherent low mobility. Long‐term monitoring of future releases will help understand how individual variation in behaviour influences reintroduction success in this species.

Temporal repeatability of behaviour in a lizard: implications for behavioural syndrome studies

It is well established that, across taxa, individuals within populations exhibit consistent differences in their behaviour across time and/or contexts. Further, the functional coupling of traits may result in the formation of a behavioural syndrome. Despite extensive evidence on the existence of consistent among-individual differences in behaviour and behavioural syndromes in the animal realm, these findings are predominately based upon short-term assessments, leading to questions regarding their stability over longer periods. Understanding if these estimates are temporally stable would allow predictions of individual behaviour to be made using short-term repeated measures. Here, we used 57 adult male delicate skinks (Lampropholis delicata) to evaluate the stability of behavioural variation observed both among (animal personality and behavioural plasticity) and within individuals (behavioural predictability), as well as behavioural syndromes, across short (four weeks) and long (five months) timeframes. To do so, we repeatedly assayed activity, exploration, and boldness five times per each individual. Overall, our study revealed complex patterns of behavioural variation and trait (co)variation over time. Activity was always repeatable across time intervals, whereas behavioural differences among individuals in exploration and boldness were not consistent. Yet a behavioural syndrome between activity and exploration was detected at both shorter and longer temporal scales, suggesting that syndrome structure in these traits does not vary as a function of time. Our findings indicate that, at least for some traits (e.g. activity) and studies, short-term measures may be adequate in serving as a proxy for long-term variation in individual behaviour, and to reveal the existence of behavioural syndromes at the population level.

Influence of natal habitat preference on habitat selection during extra-home range movements in a large ungulate.

Natal habitat preference induction (NHPI) occurs when animals exhibit a preference for new habitat that is similar to that which they experienced in their natal environment, potentially leading to post-dispersal success. While the study of NHPI is typically focused on post-settlement home ranges, we investigated how this behavior may manifest during extra-home range movements (EHRMs), both to identify exploratory prospecting behavior and assess how natal habitat cues may influence path selection before settlement. We analyzed GPS collar relocation data collected during 79 EHRMs made by 34 juvenile and subadult white-tailed deer (Odocoileus virginianus) across an agricultural landscape with highly fragmented forests in Illinois, USA. We developed a workflow to measure multidimensional natal habitat dissimilarity for each EHRM relocation and fit step-selection functions to evaluate whether natal habitat similarity explained habitat selection along movement paths. Across seasons, selection for natal habitat similarity was generally weak during excursive movements, but strong during dispersals, indicating that NHPI is manifested in dispersal habitat selection in this study system and bolstering the hypothesis that excursive movements differ functionally from dispersal. Our approach for extending the NHPI hypothesis to behavior during EHRMs can be applied to a variety of taxa and can expand our understanding of how individual behavioral variation and early life experience may shape connectivity and resistance across landscapes.

The link between animal personality and habitat selection in males of the Neotropical poison frog Allobates femoralis

Abstract Environmental variation plays a key role in the evolution and maintenance of animal personality. Individuals with different personality types might exhibit different habitat preferences. Alternatively, variation in individual behaviour across space could arise as a plastic adaptation to distinct habitats. Our study aims to investigate if habitat choice is influenced by an individual’s personality. We assessed individual levels of activity, boldness, and exploration in male poison frogs, and performed a habitat choice test under controlled laboratory conditions. Individuals were consistent in their behaviours, but all tested frogs chose the complex over the simple habitat. Individuals that were characterized as bold and very explorative also showed more movements between the two different habitats in the choice test. These results indicate that personality measured in a highly standardized artificial setup, such as a novel environment test, indeed can reflect boldness and exploration related behaviours measured in a more naturalistic setup.

A test of attribute normalization via a double decoy effect

We report a “Double Decoy” experiment designed to separate two competing accounts of the asymmetric dominance effect. The experiment places an additional decoy alternative within the range of existing alternatives, which should leave choice behaviour unaltered if attributes are weighted by their range. Instead, we observe a decrease in the relative proportion of targets chosen, particularly for subjects who exhibited an initial decoy effect. We also observe considerably more variation in individual behaviour than expected. We therefore consider an alternative theory in which attributes values are compared with diminishing sensitivity (via divisive normalization) and assess its performance in an additional discrete choice experiment previously used in the discrete choice literature. We find that divisive normalization captures behaviour better than range normalization and the linear additive Logit model typically used in applied settings. We therefore propose divisive normalization as both a neuro-computational explanation for context effects and a useful empirical tool for applied researchers.

Individual variability in habitat selection by aquatic insects is driven by taxonomy rather than specialisation

Habitat selection, the choice of a habitat based on its perceived quality, is a key mechanism structuring freshwater communities. To date, individual variability in habitat selection has been neglected, and specialisation has never been considered in this type of studies. We examined the individual differences in the habitat selection of backswimmers (Notonectidae) and diving beetles (Dytiscidae). From each family, we selected one habitat generalist able to coexist with fish (Notonecta glauca, Dytiscus marginalis), and one species specialised to fishless habitats (Notonecta obliqua, Acilius sulcatus). We performed a mesocosm experiment quantifying the consistency in individuals’ decisions in response to fish and vegetation structure, in relation to sex and specialisation. Neither the overall pattern of preferences nor consistency in individuals’ decisions differed between specialists and generalists or between the sexes, but both were consistent within families. At the population level, backswimmers preferred fishless pools with submersed and floating macrophytes, while diving beetles showed no clear preferences. Individual decisions of backswimmers were consistent and likely driven by conspecific/heterospecific attraction. In diving beetles, individual decisions were primarily density-dependent. Our results reinforce the significance of habitat selectivity for aquatic community assembly, while suggesting a range of mechanisms driving variability in individual behaviour.

Indication of a personality trait in dairy calves and its link to weight gain through automatically collected feeding behaviours

Farm animal personality traits are of interest since they can help predict individual variation in behaviour and productivity. However, personality traits are currently inferred using behavioural tests which are impractical outside of research settings. To meet the definition of a personality trait, between-individual differences in related behaviours must be temporally as well as contextually stable. In this study, we used data collected by computerised milk feeders from 76 calves over two contexts, pair housing and group housing, to test if between-individual differences in feeding rate and meal frequency meet the definition for a personality trait. Results show that between-individual differences in feeding rate and meal frequency were related, and, for each behaviour, between-individual differences were positively and significantly correlated across contexts. In addition, feeding rate and meal frequency were positively and significantly associated with weight gain. Together, these results indicate the existence of a personality trait which positions high meal frequency, fast drinking, fast growing calves at one end and low meal frequency, slow drinking, and slow growing calves at the other. Our results suggest that data already available on commercial farms could be harnessed to establish a personality trait.

Weak evidence for personality and behavioural syndrome in Teleogryllus commodus crickets

ABSTRACT Individuals of the same species and even within the same populations often vary in their behavioural responses to environmental cues. Individual variation in behaviour can be consistent across time (personality) and may also be correlated across contexts (behavioural syndromes). Here, we assessed whether Australian black field crickets (Teleogryllus commodus) display consistent inter-individual variation in four different contexts: general activity, exploration in a novel environment, boldness after a predation-risk event, and aggression towards a conspecific (fighting). We then assessed whether these four behaviours are linked, forming a behavioural syndrome. We found weak evidence for consistent individual differences in general activity, latency to touch novel objects (i.e. proxy of boldness), and the outcome of a fight. However, none of the repeatable behaviours were correlated across contexts, indicating that there is no behavioural syndrome linking these three particular behaviours. These results suggest that these behavioural traits might be regulated by different biological processes.

Background matching explains repeatable individual variation in the defence strategies of a stick insect

Visually matching the background environment is an important means by which many animals avoid predation. Yet natural habitats are rarely uniform over space and time. A key question is whether prey behaviour depends on an individual's perceived vulnerability. Using the crowned stick insect, Onchestus rentzi , as a model, we asked whether individuals' willingness to take risks, and their decision to react or to remain motionless when facing a predator threat, is explained by their appearance matching the background and/or variation in their life history traits. Over 4 consecutive weeks, we assayed individuals' tendencies to take risks and exhibit defensive displays in response to predation risk. We also measured individuals' body reflectance relative to their background (i.e. contrast) and traits related to life history that might influence risk-taking behaviours, including reproductive traits in both males and females. Individuals differed consistently from each other in their behaviour over time, with risk-prone individuals engaging more in defensive displays than risk-averse animals. Although we found no effects of body reflectance per se on individual variation in behaviour, we observed a strong positive correlation between defensive behaviours and reflectance relative to the background (i.e. contrast), revealing a functional link between behavioural variation and perceived vulnerability that is independent of life history traits. • Does prey behaviour depend on an individual's perceived vulnerability? • We test whether background matching affects the defence strategies of stick insects. • Individuals differ consistently in their defensive and risk-taking behaviours. • Individuals display stronger defensive behaviours when background matching is low. • Variation in life history traits does not explain variation in defence strategies.

Social context mediates the expression of a personality trait in a gregarious lizard

The social environment is a key factor that influences behavioural traits across a wide array of species. Yet, when investigating individual differences in behaviour, studies tend to measure animals in isolation from other conspecifics—even in social species. Surprisingly, whether behavioural traits measured in isolation are predictive of individual-level behaviour when in social groups is still poorly understood. Here, we repeatedly measured risk-taking behaviour (i.e. boldness; 741 total trials) in both the presence and absence of conspecifics in a social lizard, the delicate skink (Lampropholis delicata). Further, we manipulated food availability during group trials to test whether the effect of the social environment on risk-taking behaviour was mediated by competition over resources. Using 105 lizards collected from three independent populations, we found that individual risk-taking behaviour was repeatable when measured in either social isolation or within groups both with and without food resources available. However, lizards that were bolder during individual trials were not also bolder when in groups, regardless of resource availability. This was largely driven by individual differences in social behavioural plasticity, whereby individual skinks responded differently to the presence of conspecifics. Together, this resulted in a rank order change of individual behavioural types across the social conditions. Our results highlight the importance of the social environment in mediating animal personality traits across varying levels of resource availability. Further, these findings suggest that behavioural traits when measured in isolation, may not reflect individual variation in behaviour when measured in more ecologically realistic social groups.