In this paper we show how animal personality could explain some of the large inter-individual variation in resting metabolic rate (MR) and explore methodological and functional linkages between personality and energetics. Personality will introduce variability in resting MR measures because individuals consistently differ in their stress response, exploration or activity levels, all of which influence MR measurements made with respirometry and the doubly-labelled water technique. Physiologists try to exclude these behavioural influences from resting MR measurements, but animal personality research indicates that these attempts are unlikely to be successful. For example, because reactive animals ''freeze'' when submitted to a stress, their MR could be classified as ''resting'' because of immobility when in fact they are highly stressed with an elevated MR. More importantly, recent research demonstrating that behavioural responses to novel and highly artificial stimuli are correlated with both behaviour and fitness under more natural circumstances calls into question the wisdom of excluding these behavioural influences on MR measurements. The reason that intra-specific variation in resting MR are so weakly correlated with daily energy expenditure (DEE) and fitness, may be that the latter two measures fully incorporate personality while the former partially excludes its influence. Because activity, exploration, boldness and aggressiveness are energetically costly, personality and metabolism should be correlated and physiological constraints may underlie behavioural syndromes. We show how physiological ecologists can better examine behavioural linkages between personality and metabolism, as required to better understand the physiological correlates of personality and the evolutionary consequences of metabolic variability.Although inter-individual variation in phenotypic traits is omnipresent, it has historically been considered to be noise superimposed on the evolutionarily important signal, the population mean. Recently, however, researchers from a broad array of ecological sub-disciplines Á population biology (Bolnick et al. 2003), epidemiology (Lloyd-Smith et al. 2005), endocrinology (Williams 2008), behavioural ecology (Wilson et al. 1994, Sih et al. 2004a, Réale et al. 2007, and physiology (Bennett 1987, this paper) Á have begun to consider inter-individual variation as an important ecological and evolutionary characteristic of wild populations. The burgeoning field of animal personality seeks to explain the maintenance of variation in numerous behavioural traits, including exploration, boldness, activity and stress response among others, by examining their fitness in a variety of ecological, developmental, and demographic contexts. In this forum, we argue that personality may explain some of the large observed variation in rates of energy metabolism in animals and we explore potential synergies between personality and metabolism research. Energy metabolism: the mystery of intra-specific variationEnergy is the c...

Hibernation is widely regarded as an adaptation to seasonal energy shortage, but the actual influence of energy availability on hibernation patterns is rarely considered. Here we review literature on the costs and benefits of torpor expression to examine the influence that energy may have on hibernation patterns. We first establish that the dichotomy between food- and fat-storing hibernators coincides with differences in diet rather than body size and show that small or large species pursuing either strategy have considerable potential scope in the amount of torpor needed to survive winter. Torpor expression provides substantial energy savings, which increase the chance of surviving a period of food shortage and emerging with residual energy for early spring reproduction. However, all hibernating mammals periodically arouse to normal body temperatures during hibernation. The function of these arousals has long been speculated to involve recovery from physiological costs accumulated during metabolic depression, and recent physiological studies indicate these costs may include oxidative stress, reduced immunocompetence, and perhaps neuronal tissue damage. Using an optimality approach, we suggest that trade-offs between the benefits of energy conservation and the physiological costs of metabolic depression can explain both why hibernators periodically arouse from torpor and why they should use available energy to minimize the depth and duration of their torpor bouts. On the basis of these trade-offs, we derive a series of testable predictions concerning the relationship between energy availability and torpor expression. We conclude by reviewing the empirical support for these predictions and suggesting new avenues for research on the role of energy availability in mammalian hibernation.

To predict the consequences of human-induced global climate change, we need to understand how climate is linked to biogeography. Energetic constraints are commonly invoked to explain animal distributions, and physiological parameters are known to vary along distributional gradients. But the causal nature of the links between climate and animal biogeography remain largely obscure. Here we develop a bioenergetic model that predicts the feasibility of mammalian hibernation under different climatic conditions. As an example, we use the well-quantified hibernation energetics of the little brown bat (Myotis lucifugus) to parameterize the model. Our model predicts pronounced effects of ambient temperature on total winter energy requirements, and a relatively narrow combination of hibernaculum temperatures and winter lengths permitting successful hibernation. Microhabitat and northern distribution limits of M. lucifugus are consistent with model predictions, suggesting that the thermal dependence of hibernation energetics constrains the biogeography of this species. Integrating projections of climate change into our model predicts a pronounced northward range expansion of hibernating bats within the next 80 years. Bioenergetics can provide the simple link between climate and biogeography needed to predict the consequences of climate change.

By advancing spring leaf flush and ensuing food availability, climatic warming results in a mismatch between the timing of peak food supply and nestling demand, shifting the optimal time for reproduction in birds. Two populations of blue tits (Parus caeruleus) that breed at different dates in similar, but spatially distinct, habitat types in Corsica and southern France provide a unique opportunity to quantify the energetic and fitness consequences when breeding is mismatched with local productivity. As food supply and demand become progressively mismatched, the increased cost of rearing young pushes the metabolic effort of adults beyond their apparent sustainable limit, drastically reducing the persistence of adults in the breeding population. We provide evidence that the economics of parental foraging and limits to sustainable metabolic effort are key selective forces underlying synchronized seasonal breeding and long-term shifts in breeding date in response to climatic change.

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