RESUMEN

Maintenance metabolism as the minimum energy expenditure needed to maintain homeothermy (a high and stable body temperature, T b), reflects the magnitude of metabolic machinery and the associated costs of self-maintenance in endotherms (organisms able to produce heat endogenously). Therefore, it can interact with most, if not all, organismal functions, including the behavior-fitness linkage. Many endothermic animals can avoid the costs of maintaining homeothermy and temporally reduce T b and metabolism by entering heterothermic states like torpor, the most effective energy-saving strategy. Variations in BMR, behavior, and torpor use are considered to be shaped by food resources, but those conclusions are based on research studying these traits in isolation. We tested the effect of ecological contexts (food availability and predation risk) on the interplay between the maintenance costs of homeothermy, heterothermy, and exploration in a wild mammal-the yellow-necked mouse. We measured maintenance metabolism as basal metabolic rate (BMR) using respirometry, distance moved (exploration) in the open-field test, and variation in T b (heterothermy) during short-term fasting in animals captured at different locations of known natural food availability and predator presence, and with or without supplementary food resources. We found that in winter, heterothermy and exploration (but not BMR) negatively correlated with natural food availability (determined in autumn). Supplementary feeding increased mouse density, predation risk and finally had a positive effect on heterothermy (but not on BMR or exploration). The path analysis testing plausible causal relationships between the studied traits indicated that elevated predation risk increased heterothermy, which in turn negatively affected exploration, which positively correlated with BMR. Our study indicates that adaptive heterothermy is a compensation strategy for balancing the energy budget in endothermic animals experiencing low natural food availability. This study also suggests that under environmental challenges like increased predation risk, the use of an effective energy-saving strategy predicts behavioral expression better than self-maintenance costs under homeothermy.

RESUMEN

Metabolic physiology and animal behaviour are often considered to be linked, positively or negatively, according to either the performance or allocation models. Performance seems to predominate over allocation in natural systems, but the constraining environmental context may reveal allocation limitations to energetically expensive behaviours. Habitat disturbance, such as the large-scale fire that burnt wetlands of Biebrza National Park (NE Poland), degrades natural ecosystems. It arguably reduces food and shelter availability, modifies predator-prey interactions, and poses a direct threat for animal survival, such as that of the wetland specialist root vole Microtus oeconomus. We hypothesized that fire disturbance induces physiology-behaviour co-expression, as a consequence of changed environmental context. We repeatedly measured maintenance and exercise metabolism, and behavioural responses to the open field, in a root voles from post-fire and unburnt locations. Highly repeatable maintenance metabolism and distance moved during behavioural tests correlated positively, but relatively labile exercise metabolism did not covary with behaviour. At the same time, voles from a post-fire habitat had higher maintenance metabolism and moved shorter distances than voles from unburnt areas. We conclude there is a prevalence of the performance mechanism, but simultaneous manifestation of context-dependent allocation constraints of the physiology-behaviour covariation after disturbance. The last occurs at the within-individual level, indicating the significance of behavioural plasticity in the context of environmental disturbance.

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RESUMEN

For the last two decades, behavioral physiologists aimed to explain a plausible covariation between energetics and personality, predicted by the "pace-of-life syndrome" (POLS) hypothesis. However, the results of these attempts are mixed with no definitive answer as to which of the two most acknowledged models "performance" or "allocation" predicts covariation between consistent among-individual variation in metabolism and repeatable behavior (animal personality). The general conclusion is that the association between personality and energetics is rather context-dependent. Life-history, behavior, and physiology as well as its plausible covariation can be considered a part of sexual dimorphism. However, up to now, only a few studies demonstrated a sex-specific correlation between metabolism and personality. Therefore, we tested the relationships between physiological and personality traits in a single population of yellow-necked mice Apodemus flavicollis in the context of a plausible between-sexes difference in this covariation. We hypothesized that the performance model will explain proactive behavior in males and the allocation model will apply to females. Behavioral traits were determined using the latency of risk-taking and the open field tests, whereas the basal metabolic rates (BMR) was measured using indirect calorimetry. We have found a positive correlation between body mass-adjusted BMR and repeatable proactive behavior in male mice, which can support the performance model. However, the females were rather consistent mainly in avoidance of risk-taking that did not correlate with BMR, suggesting essential differences in personality between sexes. Most likely, the lack of convincing association between energetics and personality traits at the population level is caused by a different selection acting on the life histories of males and females. This may only result in weak support for the predictions of the POLS hypothesis when assuming that only a single model explaining the link between physiology and behavior operates in males and females. Thus, there is a need to consider the differences between sexes in behavioral studies to evaluate this hypothesis.

RESUMEN

Small mammals that are specialists in homeothermic thermoregulation reduce their self-maintenance costs of normothermy to survive the winter. By contrast, heterothermic ones that are considered generalists in thermoregulation can lower energy expenditure by entering torpor. It is well known that different species vary the use of their strategies to cope with harsh winters in temperate zones; however, little is still known about the intraspecific variation within populations and the associated external and internal factors. We hypothesized that yellow-necked mice Apodemus flavicollis decrease their resting metabolic rate (RMR) from autumn to winter, and then increase it during spring. However, since the alternative for seasonal reduction of RMR could be the development of heterothermy, we also considered the use of this strategy. We measured body mass (m b), RMR, and body temperature (T b) of mice during 2 consecutive years. In the 1st year, mice decreased whole animal RMR in winter, but did not do so in the 2nd year. All mice entered torpor during the 2nd winter, whereas only a few did so during the first one. Mice showed a continuous increase of m b, which was steepest during the 2nd year. The relationship between RMR and m b varied among seasons and years most likely due to different mouse development stages. The m b gain at the individual level was correlated positively with RMR and heterothermy. This indicates that high metabolism in winter supports the growth of smaller animals, which use torpor as a compensatory mechanism. Isotope composition of mice hair suggests that in the 1st year they fed mainly on seeds, while in the 2nd, they likely consumed significant amounts of less digestible herbs. The study suggests that the use of specialist or generalist thermoregulatory strategies can differ with environmental variation and associated differences in developmental processes.

RESUMEN

Theoretical modelling predicts that the thermoregulatory strategies of endothermic animals range from those represented by thermal generalists to those characteristic for thermal specialists. While the generalists tolerate wide variations in body temperature (Tb), the specialists maintain Tb at a more constant level. The model has gained support from inter-specific comparisons relating to species and population levels. However, little is known about consistent among-individual variation within populations that could be shaped by natural selection. We studied the consistency of individual heterothermic responses to environmental challenges in a single population of yellow-necked mice (Apodemus flavicollis), by verifying the hypothesis that Tb variation is a repeatable trait. To induce the heterothermic response, the same individuals were repeatedly food deprived for 24 h. We measured Tb with implanted miniaturised data loggers. Before each fasting experiment, we measured basal metabolic rate (BMR). Thus, we also tested whether individual variation of heterothermy correlates with individual self-maintenance costs, and the potential benefits arising from heterothermic responses that should correlate with body size/mass. We found that some individuals clearly entered torpor while others kept Tb stable, and that there were also individuals that showed intermediate thermoregulatory patterns. Heterothermy was found to correlate negatively with body mass and slightly positively with the BMR achieved 1-2 days before fasting. Nonetheless, heterothermy was shown to be highly repeatable, irrespective of whether we controlled for self-maintenance costs and body size. Our results indicate that specialist and generalist thermoregulatory phenotypes can co-exist in a single population, creating a heterothermy continuum.

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