The pace-of-life syndrome (POLS) hypothesis specifies that closely related species or populations experiencing different ecological conditions should differ in a suite of metabolic, hormonal and immunity traits that have coevolved with the life-history particularities related to these conditions. Surprisingly, two important dimensions of the POLS concept have been neglected: (i) despite increasing evidence for numerous connections between behavioural, physiological and life-history traits, behaviours have rarely been considered in the POLS yet; (ii) the POLS could easily be applied to the study of covariation among traits between individuals within a population. In this paper, we propose that consistent behavioural differences among individuals, or personality, covary with life history and physiological differences at the within-population, interpopulation and interspecific levels. We discuss how the POLS provides a heuristic framework in which personality studies can be integrated to address how variation in personality traits is maintained within populations.

The study of phenotypic evolution should be an integrative endeavor that combines different approaches and crosses disciplinary and phylogenetic boundaries to consider complex traits and organisms that historically have been studied in isolation from each other. Analyses of individual variation within populations can act to bridge studies focused at the levels of morphology, physiology, biochemistry, organismal performance, behavior, and life history. For example, the study of individual variation recently facilitated the integration of behavior into the concept of a pace-of-life syndrome and effectively linked the field of energetics with research on animal personality. Here, we illustrate how studies on the pace-of-life syndrome and the energetics of personality can be integrated within a physiology-performance-behavior-fitness paradigm that includes consideration of ecological context. We first introduce key concepts and definitions and then review the rapidly expanding literature on the links between energy metabolism and personality traits commonly studied in nonhuman animals (activity, exploration, boldness, aggressiveness, sociability). We highlight some empirical literature involving mammals and squamates that demonstrates how emerging fields can develop in rather disparate ways because of historical accidents and/or particularities of different kinds of organisms. We then briefly discuss potentially interesting avenues for future conceptual and empirical research in relation to motivation, intraindividual variation, and mechanisms underlying trait correlations. The integration of performance traits within the pace-of-life-syndrome concept has the potential to fill a logical gap between the context dependency of selection and how energetics and personality are expected to interrelate. Studies of how performance abilities and/or aspects of Darwinian fitness relate to both metabolic rate and personality traits are particularly lacking.

The domestic dog has undergone extensive artificial selection resulting in an extreme diversity in body size, personality, life-history, and metabolic traits among breeds. Here we tested whether proactive personalities (high levels of activity, boldness, and aggression) are related to a fast "pace of life" (high rates of growth, mortality, and energy expenditure). Data from the literature provide preliminary evidence that artificial selection on dogs (through domestication) generated variations in personality traits that are correlated with life histories and metabolism. We found that obedient (or docile, shy) breeds live longer than disobedient (or bold) ones and that aggressive breeds have higher energy needs than unaggressive ones. These correlations could result from either human preference for particular trait combinations or, more likely, correlated responses to artificial selection on personality. Our results suggest the existence of a general pace-of-life syndrome arising from the coevolution of personality, metabolic, and life-history traits.

1 Personality is highly relevant to ecology and the evolution of fast–slow metabolic and life-history strategies. One of the most important personality traits is exploratory behaviour, usually measured on an animal introduced to a novel environment (e.g. open-field test). 2 Here, we use a unique comparative dataset on open-field exploratory behaviour of muroid rodents to test a key assumption of a recent evolutionary model, i.e. that exploration thoroughness is positively correlated to age at first reproduction (AFR). We then examine how AFR and exploratory behaviour are related to basal metabolic rate (BMR). 3 Inter-specific variation in exploratory behaviour was positively correlated with AFR. Both AFR and exploration behaviour were negatively correlated with BMR. These results remained significant when taking phylogeny into account. 4 We suggest that species occupying unproductive and unpredictable environments simultaneously benefit from high exploration, low BMR and delayed AFR because exploration increases the likelihood of finding scarce resources, whereas low BMR and delayed reproduction enhance survival during frequent resources shortages. 5 This study provides the first empirical evidence for a link between personality, life-history, phylogeny and energy metabolism at the inter-specific level. The superficial-thorough exploration continuum can be mapped along the fast–slow metabolic and life-history continua.

Understanding the impacts of activity on energy balance is crucial. Increasing levels of activity may bring diminishing returns in energy expenditure because of compensatory responses in non-activity energy expenditures^1–3. This suggestion has profound implications for both the evolution of metabolism and human health. It implies that a long-term increase in activity does not directly translate into an increase in total energy expenditure (TEE) because other components of TEE may decrease in response–energy compensation. We used the largest dataset compiled on adult TEE and BEE (N = 1,754) of people living normal lives to find that energy compensation by a typical human averages 28% due to reduced BEE; this suggests that only 72% of the extra calories we burn from additional activity translate into extra calories burned that day. Moreover, the degree of energy compensation varied considerably between people of different body composition. This association between compensation and adiposity could be due to among-individual differences in compensation: people who compensate more may be more likely to accumulate body fat. Alternatively, the process might occur within individuals: as we get fatter, our body might compensate more strongly for the calories burned during activity, making losing fat progressively more difficult. Determining the causality of the relationship between energy compensation and adiposity will be key to improving public health strategies regarding obesity.