Variation In Basal Metabolic Rate Research Articles

Metabolic Rate and Oxidative Stress as a Risk Factors in the Development of Colorectal Cancer.

There is growing evidence that the body's energy expenditures constitute a significant risk factor for the development of most deadly diseases, including cancer. Our aim was to investigate the impact of basal metabolic rate (BMR) on the growth and progression of colorectal cancer (CRC). To do so, we used a unique model consisting of three lines of laboratory mice (Mus musculus) artificially selected for high (HBMR) and low (LBMR) basal metabolic rate and randomly bred individuals (non-selected, NSBMR). The experimental individuals were implanted with human colorectal cancer cells DLD-1. The variation in BMR between the lines allowed for testing the impact of whole-body metabolism on oxidative and antioxidant parameters in the liver throughout the cancerogenesis process. We investigated the dependence between metabolic values, reactive oxygen species (ROS) levels, and Kelch-like ECH-associated protein 1-based E3 ligase complexes (Keap1) gene activity in these animals. We found that the HBMR strain had a higher concentration of oxidative enzymes compared to the LBMR and NSBMR. Furthermore, the growth rate of CRC tumors was associated with alterations in the levels of oxidative stress enzymes and Keap1 expression in animals with a high metabolic rate. Our results indicate that a faster growth and development of CRC line DLD-1 is associated with enzymatic redox imbalance in animals with a high BMR.

Predation can shape the cascade interplay between heterothermy, exploration and maintenance metabolism underhigh food availability.

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.

Prehatching temperatures drive inter-annual cohort differences in great tit metabolism

Basal metabolic rate (BMR) constitutes the lowest metabolic rate in a resting animal and is, therefore, considered to reflect the energetic cost of maintenance in endotherms. BMR is a reversible plastic trait that changes with environmental and ecological circumstances, albeit being heritable and susceptible to selection. Inter-individual variation within populations of small birds is substantial, and while many of the drivers of such variation have been identified, many remain unexplained. We studied winter BMR variation of juveniles over a 15-year period in a wild population of great tits Parus major at the northern border of their distribution. BMR during winter consistently changed between years, even after controlling for environmental factors, suggestive of a non-reversible developmental plasticity shaping the adult metabolic phenotype. BMR in cohorts of wintering great tits varied among winters as a response to minimum ambient temperatures experienced early in life, during the prehatching period. This developmental plasticity might be adaptive if temperatures experienced by growing embryos would metabolically prime them to an environment that they will likely encounter in future life. However, in line with a more unpredictable future climate, the risk of phenotype-environment mismatch is likely to lead to certain cohorts being poorly adapted to prevailing winter conditions, resulting in wider annual fluctuations in population size.

Morphological and physiological correlates of among- individual variation in basal metabolic rate in two passerine birds

Basal metabolic rate (BMR) has been shown to be a highly phenotypic flexibility trait within species. A significant proportion of an individual's energy budget is accounted for by BMR, hence among-individual variation in this trait may affect other energetic processes, as well as fitness. In this study, we measured BMR, organ mass, mitochondrial respiration capacities and cytochrome c oxidase (COX) activities in muscle and liver and circulating levels of plasma triiodothyronine (T3) in Chinese bulbuls (Pycnonotus sinensis) and Eurasian tree sparrows (Passer montanus). Our results showed that heart and kidney mass was positively correlated with BMR in Chinese bulbuls, whereas liver and kidney mass was positively correlated with BMR in Eurasian tree sparrows. Regarding metabolic biochemical markers of tissues, state 4 respiration and COX activity in the muscles of the Chinese bulbuls was correlated with BMR, while state 4 respiration in the muscle and liver was correlated with BMR in Eurasian tree sparrows. T3 was significantly and positively correlated with BMR in Chinese bulbuls and Eurasian tree sparrows. Consistent with the above results, our findings suggest that T3 levels play an important role in modulating BMR in Chinese bulbuls and Eurasian tree sparrows. Moreover, individual variation in BMR can be explained partly by morphological and physiological mechanisms.

The seasonal variation of basal metabolic rate is related to the expression of torpor among small mammals

A variety of responses to climate seasonality have evolved by small mammals, including adjustments of the basal rate of metabolism (BMR) and the use of daily or seasonal torpor (here referred to as short-bout and long-bout torpor). The seasonal variation of their BMR is known to depend mainly on the concurrent variation of body mass, but it should also be affected by structural and functional changes occurring within the body that could depend on the expression of torpor. Thus it was hypothesized that BMR seasonality is related to the expression of torpor at an interspecific level. Seasonal BMR and body mass data were gathered from the literature and phylogenetic comparative analyses were done to test this hypothesis among mammals of less than 1 kg. BMR seasonality (dBMR) was quantified as the log-transformed ratio of the mean whole-animal BMR reported for the period P2 (autumn-winter) over that for the period P1 (spring-summer). Predictors were the seasonal body mass adjustment (dm), mean body mass (m) and torpor expression (TO, a three-level factor: no torpor, short-bout torpor, long-bout torpor). The seasonal variation of BMR was significantly related to dm but also to TO. Accounting for dm, species expressing long-bout torpor, but not those entering short-bout torpor, collectively exhibited a lower dBMR than species not entering torpor. Fat storage and use by species entering long-bout torpor, alone, could not explain their lower dBMR, as the TO:dm interaction was not significant. The low dBMR of species entering long-bout torpor may result from their collective tendency to down-regulate more strongly costly visceral organs during P2. The dBMR of the different TO categories overlapped appreciably, which highlights our still limited knowledge of the BMR seasonality among small mammals.

Significance of variation in basal metabolic rate in laboratory mice for translational experiments

The basal metabolic rate (BMR) accounts for 60–70% of the daily energy expenditure (DEE) in sedentary humans and at least 50% of the DEE in laboratory mice in the thermoneutral zone. Surprisingly, however, the significance of the variation in the BMR is largely overlooked in translational research using such indices as physical activity level (PAL), i.e., the ratio of DEE/BMR. In particular, it is unclear whether emulation of human PAL in mouse models should be carried out within or below the thermoneutral zone. It is also unclear whether physical activity within the thermoneutral zone is limited by the capacity to dissipate heat generated by exercise and obligatory metabolic processes contributing to BMR. We measured PAL and spontaneous physical activity (SPA) in laboratory mice from two lines, divergently selected towards either high or low level of BMR, and acclimated to 30 °C (i.e., the thermoneutral zone), 23 or 4 °C. The mean PAL did not differ between both lines in the mice acclimated to 30 °C but became significantly higher in the low BMR mouse line at the lower ambient temperatures. Acclimation to 30 °C reduced the mean locomotor activity but did not affect the significant difference observed between the selected lines. We conclude that carrying out experiments within the thermoneutral zone can increase the consistency of translational studies aimed at the emulation of human energetics, without affecting the variation in physical activity correlated with BMR.

Thomas H. Kunz

Thomas H. Kunz

Individual (Co)variation in Resting and Maximal Metabolic Rates in Wild Mice

Basal metabolic rate (BMR) represents the lowest level of aerobic metabolism in a resting, postabsorptive endotherm as measured within the thermoneutral zone. By contrast, maximal metabolic rate (V˙O2max) reflects the upper limit of aerobic metabolism achieved during intensive exercise. As BMR and V˙O2max define the boundaries of the possible levels of aerobic metabolism expressed by a normothermic individual, a key question is whether BMR and V˙O2max are correlated. In the present study, we took repeated paired measurements of thermoneutral resting metabolic rate (RMRt) and V˙O2max on 165 white-footed mice (Peromyscus leucopus). Over a single summer (May–October), repeatability (R ± SE) was low but statistically significant (P<0.05) for both RMRt and V˙O2max (R=0.15±0.08 for RMRt; R=0.27±0.12 for V˙O2max). Willingness to run during the forced-exercise trials was also significantly repeatable (R=0.16±0.08). At the residual level (within individual), RMRt and V˙O2max tended to be positively correlated (re=0.23±0.11, P=0.051), suggesting the presence of correlated phenotypic plasticity. By contrast, RMRt and V˙O2max were significantly negatively correlated at the among-individual level (rind=−0.84±0.40). To the extent that variation in RMRt reflects variation in BMR, the negative among-individual correlation does not corroborate the idea that a costly metabolic machinery is needed to support a high V˙O2max. Future research should investigate the (genetic) relationship between RMRt (and BMR) and other energetically expensive behaviors and activities to better understand how energy is allocated within individuals.

Functional and structural changes in aorta of mice divergently selected for basal metabolic rate

Cardiovascular diseases (CVD) are one of the most common causes of mortality likely genetically linked to the variation in basal metabolic rate (BMR). A robust test of the significance of such association may be provided by artificial selection experiments on animals selected for diversification of BMR. Here we asked whether genetically determined differences in BMR correlate with anatomical shift in endothelium structure and if so, the relaxation and contraction responses of the aorta in mice from two lines of Swiss-Webster laboratory mice (Mus musculus) divergently selected for high or low BMR (HBMR and LBMR lines, respectively). Functional and structural study of aorta showed that a selection for divergent BMR resulted in the between-line difference in diastolic aortic capacity. The relaxation was stronger in aorta of the HBMR mice, which may stem from greater flexibility of aorta mediated by higher activity of Ca2+-activated K+ channels. Structural examination also indicated that HBMR mice had significantly thicker aorta's middle layer compared to LBMR animals. Such changes may promote arterial stiffness predisposing to cardiovascular diseases. BMR-related differences in the structure and relaxation ability of aortas in studied animals may be reminiscent of potential risk factors in the development of CVD in humans.

Testosterone does not mediate variation in basal metabolic rate and activity in relation to reproductive condition and photoperiod in white-footed mice (Peromyscus leucopus).

The photoperiodic response of many temperate zone rodents, including white-footed mice (Peromyscus leucopus), is a heritable life-history trait with underlying physiological variation. Previous studies of intact male P. leucopus utilized two wild-derived bidirectional selection lines, a short photoperiod responsive (R) line selected for reproductive suppression in short-day conditions (SD) and a nonresponsive (NR) line selected for reproductive maturity in SD. NR mice in SD had greater food intake, but also higher levels of locomotor activity, and basal metabolic rate (BMR) than R mice. We hypothesized that testosterone may be a key mediator of this metabolic difference, as it is likely to be significantly reduced in R SD mice. Male P. leucopus from either line in SD were castrated and given either an implant containing testosterone (T) or a sham control (C). They were then tested for variation in metabolic rate and activity in SD, thermoneutral conditions. T mice had significantly higher levels of food intake, testosterone, and seminal vesicle dry weight than C mice. Seminal vesicle dry weight was significantly and positively correlated with average testosterone level, indicating an effect of the T implants. There was no statistically significant difference among treatment groups in BMR and average daily metabolic rate, suggesting that differences in testosterone alone are not the cause of differences in metabolic rate between selection lines.

A novel polymorphism in the fatty acid desaturase 2 gene (Fads2): A possible role in the basal metabolic rate.

Fatty acyl composition of cell membrane lipids, particularly the abundance of highly unsaturated docosahexaenoic fatty acid (22:6n-3, DHA), is likely to be an important predictor of basal metabolic rate (BMR). Our study was performed using two lines of laboratory mice divergently selected for either high or low BMR. We describe a novel single nucleotide polymorphism in the Fads2 gene encoding Δ6-desaturase, a key enzyme in the metabolic pathways of polyunsaturated fatty acids (PUFAs). The allele frequencies of Fads2 were significantly different in both lines of mice. The analysis of genetic distances revealed that the genetic differentiation between the two studied lines developed significantly faster at the Fads2 locus than it did at neutral loci. Such a pattern suggests that the Fads2 polymorphism is related to the variation in BMR, i.e. the direct target of selection. The Fads2 polymorphism significantly affected abundance of several PUFAs; however, the differences in PUFA composition between lines were compatible with the difference in frequency of Fads2 alleles only for DHA. We hypothesize that the polymorphism in the Fads2 gene affects the BMR through modification of DHA abundance in cell membranes. This may be the first example of a significant link between a polymorphism in a gene responsible for fatty acyl composition and variation in BMR.

Influence of nutritional status on basal metabolic rates among rural agriculturalists of Ngilo-Ilo, East Java.

Previous research has shown that tropical island populations have reduced basal metabolic rates (BMR) associated with the joint influences of heat stress and undernutrition. This study examines variation in BMR among an indigenous population of Indonesia, and compares these data with those collected from earlier studies in Indonesia. Anthropometric dimensions and BMR were measured on a sample of 35 Indonesian adults (28 men, 7 women) from the rural village of Ngilo-Ilo, East Java. Mean measured BMRs (±SD) were 1433 ± 344 kcal/d in men and 1256 ±257 kcal/d in women, and were not significantly different from estimates using the FAO/WHO/UNU predictive questions. Underweight individuals (BMIs <18.5 kg/m2 ) had BMRs that were 7.6% below predicted levels, while those with BMIs ≥18.5 kg/m2 had BMRs that were 8.0% above predicted levels (P < .01). Underweight individuals also had significantly higher respiratory quotients (RQ = .94 vs. .89; P < .05), suggesting lower levels of fat oxidation. Compared to data from previous studies (1929-1979), men of the Ngilo-Ilo sample had similar BMIs (19.8 vs. 19.2 kg/m2 ), but higher BMRs, after adjusting for age and body weight (+2.1% vs. -5.6%; P < .05). Among the agriculturalists of Ngilo-Ilo, measured BMRs were low, but not significantly different from those predicted by the FAO/WHO/UNU equations. Among subjects of this sample and from earlier studies, poorer physical nutritional status was associated with reduced BMRs. These results suggest that chronic energy stress has consistently shaped metabolic function among Indonesian rural populations.

Energetic trade-offs and feedbacks between behavior and metabolism influence correlations between pace-of-life attributes

Correlations between behavioral, physiological, and morphological traits linked to life history have been given the label “pace-of-life syndrome” (POLS), hypothesized to arise through variation in the resolution of a trade-off between present and future reproduction. However, other trade-offs over energy allocation may also have effects and influence the present-future trade-off. We analyzed an optimality model of basal metabolic rate (BMR) across variation in food availability and two types of mortality. The model contained three major features: (1) feedback between activity and energy acquisition, (2) links between BMR and the use of energy for other traits, and (3) allocation trade-offs between BMR and all other traits, between activity and defense, and between defense against activity-related risk and activity-independent risk. The model produced an intermediate optimal BMR that was usually highest at an intermediate level of food availability. Food availability and both types of mortality risk interacted to influence the exact value of optimal BMR. Trait correlations expected in the POLS existed under some environmental conditions, but these correlations flipped sign under different conditions and were not always strong. Our model reproduces trait correlations consistent with the POLS, but also generated a “sloppy” syndrome with considerable non-POLS-like variation. In addition, among-individual, non-adaptive variation in BMR produced adjustments of the other traits. These fit a best-of-a-bad job strategy, and the adjustments further weakened trait correlations. The results emphasize that variation in resources and mortality risk creates a diversity of correlation structures. This complexity means the POLS is likely to be a variable construct. Many attributes important for reproduction and survival are associated. Such associations may arise through common physiological processes and correlated selection. We modeled metabolic rate within a system in which foraging behavior both depended on and mediated the acquisition of resources necessary for metabolism, while energy was allocated among multiple attributes. Variation in several environmental variables (food availability and two types of mortality risk) influenced basal metabolic rate, activity, and defenses against mortality risk. This variation affected the correlations between the traits in complex ways. When basal metabolic rate was non-optimal, evolution of the allocation of energy to other traits partially compensated, but this further eroded consistent trait correlations. Our results indicate that complexity in how energy is acquired and used can potentially disrupt trait correlations normally associated with the pace-of-life syndrome.

The performing animal: causes and consequences of body remodeling and metabolic adjustments in red knots facing contrasting thermal environments.

Using red knots (Calidris canutus) as a model, we determined how changes in mass and metabolic activity of organs relate to temperature-induced variation in metabolic performance. In cold-acclimated birds, we expected large muscles and heart as well as improved oxidative capacity and lipid transport, and we predicted that this would explain variation in maximal thermogenic capacity (Msum). We also expected larger digestive and excretory organs in these same birds and predicted that this would explain most of the variation in basal metabolic rate (BMR). Knots kept at 5°C were 20% heavier and maintained 1.5 times more body fat than individuals kept in thermoneutral conditions (25°C). The birds in the cold also had a BMR up to 32% higher and a Msum 16% higher than birds at 25°C. Organs were larger in the cold, with muscles and heart being 9-20% heavier and digestive and excretory organs being 21-36% larger than at thermoneutrality. Rather than the predicted digestive and excretory organs, the cold-induced increase in BMR correlated with changes in mass of the heart, pectoralis, and carcass. Msum varied positively with the mass of the pectoralis, supracoracoideus, and heart, highlighting the importance of muscles and cardiac function in cold endurance. Cold-acclimated knots also expressed upregulated capacity for lipid transport across mitochondrial membranes [carnitine palmitoyl transferase (CPT)] in their pectoralis and leg muscles, higher lipid catabolism capacity in their pectoralis muscles [β-hydroxyacyl CoA-dehydrogenase (HOAD)], and elevated oxidative capacity in their liver and kidney (citrate synthase). These adjustments may have contributed to BMR through changes in metabolic intensity. Positive relationships among Msum, CPT, and HOAD in the heart also suggest indirect constraints on thermogenic capacity through limited cardiac capacity.

How low can you go? An adaptive energetic framework for interpreting basal metabolic rate variation in endotherms.

Adaptive explanations for both high and low body mass-independent basal metabolic rate (BMR) in endotherms are pervasive in evolutionary physiology, but arguments implying a direct adaptive benefit of high BMR are troublesome from an energetic standpoint. Here, we argue that conclusions about the adaptive benefit of BMR need to be interpreted, first and foremost, in terms of energetics, with particular attention to physiological traits on which natural selection is directly acting. We further argue from an energetic perspective that selection should always act to reduce BMR (i.e., maintenance costs) to the lowest level possible under prevailing environmental or ecological demands, so that high BMR per se is not directly adaptive. We emphasize the argument that high BMR arises as a correlated response to direct selection on other physiological traits associated with high ecological or environmental costs, such as daily energy expenditure (DEE) or capacities for activity or thermogenesis. High BMR thus represents elevated maintenance costs required to support energetically demanding lifestyles, including living in harsh environments. BMR is generally low under conditions of relaxed selection on energy demands for high metabolic capacities (e.g., thermoregulation, activity) or conditions promoting energy conservation. Under these conditions, we argue that selection can act directly to reduce BMR. We contend that, as a general rule, BMR should always be as low as environmental or ecological conditions permit, allowing energy to be allocated for other functions. Studies addressing relative reaction norms and response times to fluctuating environmental or ecological demands for BMR, DEE, and metabolic capacities and the fitness consequences of variation in BMR and other metabolic traits are needed to better delineate organismal metabolic responses to environmental or ecological selective forces.

Basal metabolic rate in free-living tropical birds: the influence of phylogenetic, behavioral, and ecological factors.

The majority of our knowledge of avian energetics is based on studies of birds from temperate and high latitudes. Using the largest existing sample of wild-caught Old World tropical species, we showed that birds from Southern Vietnam had lower basal metabolic rate (BMR) than temperate species. The strongest dissimilarity between tropical and temperate species was the low scaling exponent in the allometric relation between BMR and body mass in tropical birds (the regression slope was 0.573). The passerine migrants to temperate and high latitudes had higher BMR than tropical sedentary passerines. Body mass alone accounted for 93% of the variation in BMR (body mass ranged from 5 to 252 g). Contrary to some other studies, we did not find evidence besides the above mentioned that phylogeny, taxonomy, behavior, or ecology have a significant influence on BMR variation among tropical birds.

Understanding evolutionary variation in basal metabolic rate: An analysis in subterranean rodents

Understanding how evolutionary variation in energetic metabolism arises is central to several theories in animal biology. Basal metabolic rate (BMR) –i.e., the minimum rate of energy necessary to maintain thermal homeostasis in endotherms– is a highly informative measure to increase our understanding, because it is determined under highly standardized conditions. In this study we evaluate the relationship between taxa- and mass-independent (residual) BMR and ten environmental factors for 34 subterranean rodent species. Both conventional and phylogenetically informed analyses indicate that ambient temperature is the major determinant of residual BMR, with both variables inversely correlated. By contrast, other environmental factors that have been shown to affect residual BMR in endotherms, such as habitat productivity and rainfall, were not significant predictors of residual BMR in this group of species. Then, the results for subterranean rodents appear to support a central prediction of the obligatory heat model (OHM), which is a mechanistic model aimed to explain the evolution of residual BMR. Specifically, OHM proposes that during the colonization of colder environments, individuals with greater masses of metabolically expensive tissues (and thus with greater BMR) are favored by natural selection due to the link between greater masses of metabolically expensive tissues and physiological capacities. This way, natural selection should establishes a negative correlation between ambient temperature and both internal organ size and residual BMR.

Does basal metabolic rate predict weight gain?

Some previous studies have indicated that a low basal metabolic rate (BMR) is an independent predictor of future weight gain, but low rates of follow-up and highly select populations may limit the ability to generalize the results. We assessed whether adults with a low BMR gain more weight than do adults with a high BMR who are living in a typical Western environment. We extracted BMR, body-composition, demographic, and laboratory data from electronic databases of 757 volunteers who were participating in our research protocols at the Mayo Clinic between 1995 and 2012. Research study volunteers were always weight stable, had no acute illnesses and no confounding medication use, and were nonsmokers. The top and bottom 15th percentiles of BMR, adjusted for fat-free mass (FFM), fat mass, age, and sex, were identified. Follow-up electronic medical record system data were available for 163 subjects, which allowed us to determine their subsequent weight changes for ≥3 y (mean: ∼9.7 y). By definition, the BMR was different in the high-BMR group (2001 ± 317 kcal/d; n = 86) than in the low-BMR group (1510 ± 222 kcal/d; n = 77), but they were comparable with respect to age, body mass index, FFM, and fat mass. Rates of weight gain were not greater in the bottom BMR group (0.3 ± 1.0 kg/y) than in the top BMR group (0.5 ± 1.5 kg/y) (P = 0.17). Adults with low BMRs did not gain more weight than did adults with high BMRs, implying that habitual differences in food intake or activity counterbalance variations in BMR as a risk factor for weight gain in a typical Western population.

Basal Metabolic Rate of the Black-Faced Sheathbill (Chionis minor): Intraspecific Variation in a Phylogenetically Distinct Island Endemic.

Metabolic rate is a fundamental characteristic of all organisms. It covaries most significantly with activity, body mass, seasonality, and temperature. Nonetheless, substantial additional variation in metabolic rate, especially either resting rate or basal rate, is associated with a range of factors including phylogenetic position, ecological distinctiveness, range position, and diet. Understanding this variation is a key goal of physiological ecology. The black-faced sheathbill is a phylogenetically distinct, high-latitude, island-endemic bird occurring exclusively on several archipelagos in the southern Indian Ocean. Here we examined the idea that the unique phylogenetic position and ecology of the black-faced sheathbill may lead to a basal metabolic rate (BMR) different from that predicted by its body mass. When compared with BMR data available for all birds and a subset of island species, it was clear that the BMR of the black-faced sheathbill on subantarctic Marion Island, estimated at 15°C using indirect calorimetry (2.370 ± 0.464 W, mean ± SD; n = 22), for a group of birds with a mean mass of 459 ± 64 g, is no different from that expected based on body mass. However, variation in BMR, associated with habitat use and diet, even when correcting for variation in mass, was found. Sheathbills foraging year-round in comparatively resource-rich king penguin colonies have a higher BMR (2.758 ± 0.291 W, n = 12) than sheathbills that split their foraging between rockhopper penguin colonies and the intertidal zone (2.047 ± 0.303 W, n = 10), which are poorer in resources. Because these populations coexist at relatively small spatial extents (the entire island is 290 km(2)), other factors seem unlikely as causes of this variation.

Fluctuating selection on basal metabolic rate.

BMR (Basal metabolic rate) is an important trait in animal life history as it represents a significant part of animal energy budgets. BMR has also been shown to be positively related to sustainable work rate and maximal thermoregulatory capacity. To this date, most of the studies have focused on the causes of interspecific and intraspecific variation in BMR, and fairly little is known about the fitness consequences of different metabolic strategies. In this study, we show that winter BMR affects local survival in a population of wild blue tits (Cyanistes caeruleus), but that the selection direction differs between years. We argue that this fluctuating selection is probably a consequence of varying winter climate with a positive relation between survival and BMR during cold and harsh conditions, but a negative relation during mild winters. This fluctuating selection can not only explain the pronounced variation in BMR in wild populations, but will also give us new insights into how energy turnover rates can shape the life-history strategies of animals. Furthermore, the study shows that the process of global warming may cause directional selection for a general reduction in BMR, affecting the general life-history strategy on the population level.