Phenotypic plasticity in the scaling of avian basal metabolic rate

Many birds exhibit short-term, reversible adjustments in basal metabolic rate (BMR), but the overall contribution of phenotypic plasticity to avian metabolic diversity remains unclear. The available BMR data include estimates from birds living in natural environments and captive-raised birds in more homogenous, artificial environments. All previous analyses of interspecific variation in BMR have pooled these data. We hypothesized that phenotypic plasticity is an important contributor to interspecific variation in avian BMR, and that captive-raised populations exhibit general differences in BMR compared to wild-caught populations. We tested this hypothesis by fitting general linear models to BMR data for 231 bird species, using the generalized least-squares approach to correct for phylogenetic relatedness when necessary. The scaling exponent relating BMR to body mass in captive-raised birds (0.670) was significantly shallower than in wild-caught birds (0.744). The differences in metabolic scaling between captive-raised and wild-caught birds persisted when migratory tendency and habitat aridity were controlled for. Our results reveal that phenotypic plasticity is a major contributor to avian interspecific metabolic variation. The finding that metabolic scaling in birds is partly determined by environmental factors provides further support for models that predict variation in scaling exponents, such as the allometric cascade model.     

Physiological responses in rufous-collared sparrows to thermal acclimation and seasonal acclimatization

A large number of physiological acclimation studies assume that flexibility in a certain trait is both adaptive and functionally important for organisms in their natural environment; however, it is not clear how an organism’s capacity for temperature acclimation translates to the seasonal acclimatization that these organisms must accomplish. To elucidate this relationship, we measured BMR and TEWL rates in both field-acclimatized and laboratory-acclimated adult rufous-collared sparrows (Zonotrichia capensis). Measurements in field-acclimatized birds were taken during the winter and summer seasons; in the laboratory-acclimated birds, we took our measurements following 4 weeks at either 15 or 30°C. Although BMR and TEWL rates did not differ between winter and summer in the field-acclimatized birds, laboratory-acclimated birds exposed to 15°C exhibited both a higher BMR and TEWL rate when compared to the birds acclimated to 30°C and the field-acclimatized birds. Because organ masses seem to be similar between field and cold-acclimated birds whereas BMR is higher in cold-acclimated birds, the variability in BMR cannot be explained completely by adjustments in organ masses. Our findings suggest that, although rufous-collared sparrows can exhibit thermal acclimation of physiological traits, sparrows do not use this capacity to cope with minor to moderate fluctuations in environmental conditions. Our data support the hypothesis that physiological flexibility in energetic traits is a common feature of avian metabolism.     

Effects of short-term acclimation on thermoregulatory responses of the rock kestrel, Falco rupicolus

The effects of a short-term acclimation period on basal metabolic rate (BMR) and resting metabolic rate (RMR) were measured in captive-bred Rock Kestrels (Falco rupicolus). Birds were exposed to winter conditions (pre-acclimation) in a semi-natural environment before they were acclimated for a period of 3 weeks at a constant temperature of 25 °C and a constant light:dark cycle (12:12 h) (post-acclimation). After acclimation the kestrels showed changes in RMR, BMR and the width of the thermoneutral zone. There was inter- and intra-individual phenotypic plasticity in BMR and RMR both pre- and post-acclimation. However, more inter-individual variation was seen after acclimation. This study concurs with recent suggestions that phenotypic plasticity in BMR is prevalent in avian physiology, and thus a single-species-specific BMR value may not be representative. Furthermore, comparative avian studies of BMR need to account for phenotypic plasticity.     

Intra-Seasonal Flexibility in Avian Metabolic Performance Highlights the Uncoupling of Basal Metabolic Rate and Thermogenic Capacity

Stochastic winter weather events are predicted to increase in occurrence and amplitude at northern latitudes and organisms are expected to cope through phenotypic flexibility. Small avian species wintering in these environments show acclimatization where basal metabolic rate (BMR) and maximal thermogenic capacity (M_SUM) are typically elevated. However, little is known on intra-seasonal variation in metabolic performance and on how population trends truly reflect individual flexibility. Here we report intra-seasonal variation in metabolic parameters measured at the population and individual levels in black-capped chickadees ( Poecile atricapillus ). Results confirmed that population patterns indeed reflect flexibility at the individual level. They showed the expected increase in BMR (6%) and M_SUM (34%) in winter relative to summer but also, and most importantly, that these parameters changed differently through time. BMR began its seasonal increase in November, while M_SUM had already achieved more than 20% of its inter-seasonal increase by October, and declined to its starting level by March, while M_SUM remained high. Although both parameters co-vary on a yearly scale, this mismatch in the timing of variation in winter BMR and M_SUM likely reflects different constraints acting on different physiological components and therefore suggests a lack of functional link between these parameters.     

Environment, migratory tendency, phylogeny and basal metabolic rate in birds

Basal metabolic rate (BMR) represents the minimum maintenance energy requirement of an endotherm and has far-reaching consequences for interactions between animals and their environments. Avian BMR exhibits considerable variation that is independent of body mass. Some long-distance migrants have been found to exhibit particularly high BMR, traditionally interpreted as being related to the energetic demands of long-distance migration. Here we use a global dataset to evaluate differences in BMR between migrants and non-migrants, and to examine the effects of environmental variables. The BMR of migrant species is significantly higher than that of non-migrants. Intriguingly, while the elevated BMR of migrants on their breeding grounds may reflect the metabolic machinery required for long-distance movements, an alternative (and statistically stronger) explanation is their occupation of predominantly cold high-latitude breeding areas. Among several environmental predictors, average annual temperature has the strongest effect on BMR, with a 50% reduction associated with a 20 degrees C gradient. The negative effects of temperature variables on BMR hold separately for migrants and non-migrants and are not due their different climatic associations. BMR in migrants shows a much lower degree of phylogenetic inertia. Our findings indicate that migratory tendency need not necessarily be invoked to explain the higher BMR of migrants. A weaker phylogenetic signal observed in migrants supports the notion of strong phenotypic flexibility in this group which facilitates migration-related BMR adjustments that occur above and beyond environmental conditions. In contrast to the findings of previous analyses of mammalian BMR, primary productivity, aridity or precipitation variability do not appear to be important environmental correlates of avian BMR. The strong effects of temperature-related variables and varying phylogenetic effects reiterate the importance of addressing both broad-scale and individual-scale variation for understanding the determinants of BMR.     

Seasonal variation in the metabolism-temperature relation of House Sparrows (Passer domesticus) in KwaZulu-Natal,South Africa

Many birds exhibit considerable phenotypic flexibility in metabolism to maintain thermoregulation or to conserve energy. This flexibility usually includes seasonal variation in metabolic rate. Seasonal changes in physiology and behavior of birds are considered to be a part of their adaptive strategy for survival and reproductive success. House Sparrows (Passer domesticus) are small passerines from Europe that have been successfully introduced to many parts of the world, and thus may be expected to exhibit high phenotypic flexibility in metabolic rate. Mass specific Resting Metabolic Rate (RMR) and Basal Metabolic Rate (BMR) were significantly higher in winter compared with summer, although there was no significant difference between body mass in summer and winter. A similar, narrow thermal neutral zone (25–28 °C) was observed in both seasons. Winter elevation of metabolic rate in House Sparrows was presumably related to metabolic or morphological adjustments to meet the extra energy demands of cold winters. Overall, House Sparrows showed seasonal metabolic acclimatization similar to other temperate wintering passerines. The improved cold tolerance was associated with a significant increase in VO₂ in winter relative to summer. In addition, some summer birds died at 5 °C, whereas winter birds did not, further showing seasonal variation in cold tolerance. The increase in BMR of 120% in winter, compared to summer, is by far the highest recorded seasonal change so far in birds.     

The allometry of avian basal metabolic rate: good predictions need good data

Basal metabolic rate (BMR) is often predicted by allometric interpolation, but such predictions are critically dependent on the quality of the data used to derive allometric equations relating BMR to body mass (Mb). An examination of the metabolic rates used to produce conventional and phylogenetically independent allometries for avian BMR in a recent analysis revealed that only 67 of 248 data unambiguously met the criteria for BMR and had sample sizes with n>/=3. The metabolic rates that represented BMR were significantly lower than those that did not meet the criteria for BMR or were measured under unspecified conditions. Moreover, our conventional allometric estimates of BMR (W; logBMR=-1.461+0.669logMb) using a more constrained data set that met the conditions that define BMR and had n>/=3 were 10%-12% lower than those obtained in the earlier analysis. The inclusion of data that do not represent BMR results in the overestimation of predicted BMR and can potentially lead to incorrect conclusions concerning metabolic adaptation. Our analyses using a data set that included only BMR with n>/=3 were consistent with the conclusion that BMR does not differ between passerine and nonpasserine birds after taking phylogeny into account. With an increased focus on data mining and synthetic analyses, our study suggests that a thorough knowledge of how data sets are generated and the underlying constraints on their interpretation is a necessary prerequisite for such exercises.     

Estimation of heritability and repeatability of resting metabolic rate in birds, with free-living pied flycatchers Ficedula hypoleuca (Aves: Passeriformes) as an example

Estimates of a trait heritability and repeatability can get at an idea of its usefulness for being an individual characteristic and its ability to change under selection pressure. Heritability and repeatability of energetic parameters still poorly studied in birds. The most important physiological characteristic of homoiotherms is resting metabolic rate (RMR), which, in the absence of productive processes, does not exceed basal metabolic rate (BMR). We estimated BMR repeatability in free-living pied flycatchers in Moscow Region (55 degrees 44' N, 36 degrees 51' E; 1992-2008) and Tomsk (56 degrees 20' N, 84 degrees 56' E; 2008-2009) populations over intervals from 40 days to 3 years. In Moscow Region population, BMR repeatability amounted to tau = 0.34 +/- 0.10 (n=80) if measured over 1 year interval, tau = 0.60 +/- 0.15 (n=19) if measured over 2 years interval, and tau = 0.85 +/- 0.13 (n=6) if measured over 3 years interval providing that consecutive BMR measurements were done in the same period of reproductive season. In Tomsk population, BMR repeatability, measured over 1 year interval, amounted to tau = 0.49 +/- 0.11 (n=50). Repeatability is a measure of a trait constancy and sets the upper limit of its heritability. To estimate RMR heritability, cross-fostering experiments have been conducted in 2003-2005 with flycatchers of Moscow Region population. RMR of chicks positively correlated with BMR of their biological fathers, whereas such correlation in metabolic rates between chicks and their foster fathers was absent. The RMR heritability estimate turned out to be h2 = 0.43 +/- 0.17 (n=210). The obtained estimates of heritability and repeatability of fundamental energetic traits are rather high for physiological features. This suggests the existence of a potential for direct selection on BMR and evolutionary stable diversity of avian populations with regard to basal metabolic rate.     

Disentangling environmental drivers of metabolic flexibility in birds: the importance of temperature extremes versus temperature variability

Examining physiological traits across large spatial scales can shed light on the environmental factors driving physiological variation. For endotherms, flexibility in aerobic metabolism is especially important for coping with thermally challenging environments and recent research has shown that aerobic metabolic scope [the difference between maximum thermogenic capacity (M_sum) and basal metabolic rate (BMR)] increases with latitude in mammals. One explanation for this pattern is the climatic variability hypothesis, which predicts that flexibility in aerobic metabolism should increase as a function of local temperature variability. An alternative explanation is the cold adaptation hypothesis, which predicts that cold temperature extremes may also be an important driver of variation in metabolic scope. To determine the thermal drivers of aerobic metabolic flexibility in birds, we combined data on metabolic scope from 40 bird species sampled across a range of environments with several indices of local ambient temperature. Using phylogenetically‐informed analyses, we found that minimum winter temperature was the best predictor of variation in avian metabolic scope, outperforming all other thermal variables. Additionally, M_sum was a better predictor of latitudinal patterns of metabolic scope than BMR, with species inhabiting colder environments exhibiting increased M_sum over their counterparts in warmer environments. Taken together, these results suggest that cold temperature extremes drive latitudinal patterns of metabolic scope via selection for enhanced thermogenic performance in cold environments, supporting the cold adaptation hypothesis. Temperature extremes may therefore be an important selective pressure driving macrophysiological trends of aerobic performance in endotherms.     

Individual variation and repeatability of basal metabolism in the bank vole, Clethrionomys glareolus

Basal metabolic rate (BMR) is a fundamental energetic trait and has been measured in hundreds of birds and mammals. Nevertheless, little is known about the consistency of the population-average BMR or its repeatability at the level of individual variation. Here, we report that average mass-independent BMR did not differ between two generations of bank voles or between two trials separated by one month. Individual differences in BMR were highly repeatable across the one month interval: the coefficient of intraclass correlation was 0.70 for absolute log-transformed values and 0.56 for mass-independent values. Thus, BMR can be a meaningful measure of an individual physiological characteristic and can be used to test hypotheses concerning relationships between BMR and other traits. On the other hand, mass-independent BMR did not differ significantly across families, and the coefficient of intraclass correlation for full sibs did not differ from zero, which suggests that heritability of BMR in voles is not high.     

Seasonal variation in body composition in an Afrotropical passerine bird: increases in pectoral muscle mass are,unexpectedly, associated with lower thermogenic capacity

Phenotypic flexibility in avian metabolic rates and body composition have been well-studied in high-latitude species, which typically increase basal metabolic rate (BMR) and summit metabolism (M_sum) when acclimatized to winter conditions. Patterns of seasonal metabolic acclimatization are more variable in lower-latitude birds that experience milder winters, with fewer studies investigating adjustments in avian organ and muscle masses in the context of metabolic flexibility in these regions. We quantified seasonal variation (summer vs winter) in the masses of organs and muscles frequently associated with changes in BMR (gizzard, intestines and liver) and M_sum (heart and pectoral muscles), in white-browed sparrow-weavers (Plocepasser mahali). We also measured pectoral muscle thickness using a portable ultrasound system to determine whether we could non-lethally estimate muscle size. A concurrent study measured seasonal changes in BMR and M_sum in the same population of sparrow-weavers, but different individuals. There was no seasonal variation in the dry masses of the gizzard, intestines or liver of sparrow-weavers, and during the same period, BMR did not vary seasonally. We found significantly higher heart (~ 18% higher) and pectoral muscle (~ 9% higher) dry mass during winter, although ultrasound measurements did not detect seasonal changes in pectoral muscle size. Despite winter increases in pectoral muscle mass, M_sum was ~ 26% lower in winter compared to summer. To the best of our knowledge, this is the first study to report an increase in avian pectoral muscle mass but a concomitant decrease in thermogenic capacity.

     

Ecological and evolutionary physiology of desert birds: a progress report

The adaptive significance of mechanisms of energy and water conservation among species of desert rodents, which avoid temperature extremes by remaining within a burrow during the day, is well established. Conventional wisdom holds that arid-zone birds, diurnal organisms that endure the brunt of their environment, occupy these desert climates because of the possession of physiological design features common to all within the class Aves. We review studies that show that desert birds may have evolved specific features to deal with hot desert conditions including: a reduced basal metabolic rate (BMR) and field metabolic rate (FMR), and lower total evaporative water loss (TEWL) and water turnover (WTO).Previous work on the comparative physiology of desert birds relied primarily on information gathered on species from the deserts of the southwestern U.S., which are semi-arid habitats of recent geologic origin. We include data on species from Old World deserts, which are geologically older than those in the New World, and place physiological responses along an aridity axis that includes mesic, semi-arid, arid, and hyperarid environments.The physiological differences between desert and mesic birds that we have identified using the comparative method could arise as a result of acclimation to different environments, of genetic change mediated by selection, or both. We present data on the flexibility of BMR and TEWL in Hoopoe Larks that suggest that phenotypic adjustments in these variables can be substantial. Finally, we suggest that linkages between the physiology of individual organism and its life-history are fundamental to the understanding of life-history evolution.     

Seasonal effects on metabolism and thermoregulation abilities of the Red-winged Starling (Onychognathus morio)

Basal metabolic rate (BMR) of birds is beginning to be viewed as a highly flexible physiological trait influenced by environmental fluctuations, and in particular changes in ambient temperatures (T_a). Southern Africa is characterized by an unpredictable environment with daily and seasonal variation. This study sought to evaluate the effects of seasonal changes in T_a on mass-specific resting metabolic rate (RMR), BMR and body temperature (T_b) of Red-winged Starlings (Onychognathus morio). They have a broad distribution, from Ethiopia to the Cape in South Africa and are medium-sized frugivorous birds. Metabolic rate (VO₂) and T_b were measured in wild caught Red-winged Starlings after a period of summer and winter acclimatization in outdoor aviaries. RMR and BMR were significantly higher in winter than summer. Body mass of Starlings was significantly higher in winter compared with summer. The increased RMR and BMR in winter indicate improved ability to cope with cold and maintenance of a high T_b. These results show that the metabolism of Red-winged Starlings are not constant, but exhibit a pronounced seasonal phenotypic flexibility with maintenance of a high T_b.     

Basal metabolic rate, food intake, and body mass in cold- and warm-acclimated Garden Warblers

We address the question of whether physiological flexibility in relation to climate is a general feature of the metabolic properties of birds. We tested this hypothesis in hand-raised Garden Warblers (Sylvia borin), long-distance migrants, which normally do not experience great temperature differences between summer and winter. We maintained two groups of birds under cold and warm conditions for 5 months, during which their body mass and food intake were monitored. When relatedness (siblings vs. non-siblings) of the experimental birds was taken into account, body mass in cold-acclimated birds was higher than in warm-acclimated birds. BMR, measured at the end of the 5-month temperature treatment, was also higher in the cold- than the warm-acclimated group. Migrant birds thus seem to be capable of the same metabolic cold-acclimation response as has been reported in resident birds. The data support the hypothesis that physiological flexibility is a basic trait of the metabolic properties of birds.     

Phenotypic flexibility in basal metabolic rate is associated with rainfall variability among populations of rufous-collared sparrow

Phenotypic flexibility in metabolic rates allows organisms to reversibly adjust their energy flow to meet challenges imposed by a variable environment. In turn, the food habits hypothesis (FHH) predicts that species or populations adjust their basal metabolic rate (BMR) according to the diet attributes such as food abundance or predictability. Desert ecosystems represent a temporally heterogeneous environment because of low rain pulse predictability, which is also associated with temporal variation in food resources. In the present study, we investigated the relationship between the magnitude of BMR flexibility in response to dietary acclimation and the inter-annual rainfall variability in three populations of rufous-collared sparrows. Specifically we addressed the question of whether birds from a desert environment are more flexible in BMR than those from non-desert habitats. We found a positive trend between BMR flexibility and the inter-annual rainfall variability. In fact, dietary treatments had a significant effect only in desert birds, a result that also supported the FHH. Our study confirms the existence of phenotypic variation in response to environmental conditions among populations, and also highlights the importance of considering the circumstances in which phenotypic flexibility evolves and the specific environmental cues that induce their expression.     

Latitudinal patterns in rodent metabolic flexibility

Macrophysiology is defined as the study of variation in physiological traits-including physiological trait flexibility-over large geographical and temporal scales, and the ecological implications of this variation. A classic example of a macrophysiological trend is the one emerging from the climatic variability hypothesis, which states that as the range of climatic fluctuation experienced by terrestrial animals increases with latitude, individuals at higher latitudes should be more plastic than individuals inhabiting lower latitudes. In this context, we evaluate the correlation between absolute metabolic scope during cold exposure (an instantaneous measure of metabolic flexibility) and different geographic and climatic variables for 48 rodent species. Conventional and phylogenetic informed analyses indicated a positive correlation between metabolic scope and geographic latitude. These findings, together with previous reports on latitudinal pattern in phenotypic flexibility, suggest that an increase in physiological flexibility with latitude may hold for many phenotypic traits.     

Tough decisions: Reproductive timing and output vary with individuals' physiology,behavior and past success in a social opportunistic breeder

This article is part of a Special Issue "SBN 2014".Photoperiod and the hormonal response it triggers are key determinants of reproductive timing in birds. However, other cues and physiological traits may permit flexibility in the timing of breeding and perhaps facilitate adaptation to global change. Opportunistic breeders are excellent models to study the adaptive significance of this flexibility, especially at the individual level. Here, we sought to quantify whether particular male physiological and behavioral traits were linked to reproductive timing and output in wild-derived zebra finches. We repeatedly assessed male stress-induced corticosterone levels (CORT), basal metabolic rate (BMR), and activity before releasing them into outdoor aviaries and quantifying each pair's breeding timing, investment, and output over a seven-month period. Despite unlimited access to food and water, the colony breeding activity occurred in waves, probably due to interpair social stimulations. Pairs adjusted their inter-clutch interval and clutch size to social and temperature cues, respectively, but only after successful breeding attempts, suggesting a facultative response to external cues. When these effects were controlled for statistically or experimentally, breeding intervals were repeatable within individuals across reproductive attempts. In addition, males' first laying date and total offspring production varied with complex interactions between pre-breeding CORT, BMR and activity levels. These results suggest that no one trait is under selection but that, instead, correlational selection acts on hormone levels, metabolism, and behavior. Together our results suggest that studying inter-individual variation in breeding strategy and their multiple physiological and behavioral underpinnings may greatly improve our understanding of the mechanisms underlying the evolution of breeding decisions.     

Metabolic and ventilatory acclimatization to cold stress in house sparrows (Passer domesticus)

Passerines that overwinter in temperate climates undergo seasonal acclimatization that is characterized by metabolic adjustments that may include increased basal metabolic rate (BMR) and cold-induced summit metabolism (M(sum)) in winter relative to summer. Metabolic changes must be supported by equivalent changes in oxygen transport. While much is known about the morphology of the avian respiratory system, little is known about respiratory function under extreme cold stress. We examined seasonal variation in BMR, M(sum), and ventilation in seasonally acclimatized house sparrows from Wisconsin. BMR and M(sum) increased significantly in winter compared with summer. In winter, BMR increased 64%, and M(sum) increased 29% over summer values. The 64% increase in winter BMR is the highest recorded for birds. Metabolic expansibility (M(sum)/BMR) was 9.0 in summer and 6.9 in winter birds. The metabolic expansibility of 9.0 in summer is the highest yet recorded for birds. Ventilatory accommodation under helox cold stress was due to changes in breathing frequency (f), tidal volume, and oxygen extraction efficiency in both seasons. However, the only significant difference between summer and winter ventilation measures in helox cold stress was f. Mean f in helox cold stress for winter birds was 1.23 times summer values.     

Intraspecific correlations of basal and maximal metabolic rates in birds and the aerobic capacity model for the evolution of endothermy

The underlying assumption of the aerobic capacity model for the evolution of endothermy is that basal (BMR) and maximal aerobic metabolic rates are phenotypically linked. However, because BMR is largely a function of central organs whereas maximal metabolic output is largely a function of skeletal muscles, the mechanistic underpinnings for their linkage are not obvious. Interspecific studies in birds generally support a phenotypic correlation between BMR and maximal metabolic output. If the aerobic capacity model is valid, these phenotypic correlations should also extend to intraspecific comparisons. We measured BMR, M(sum) (maximum thermoregulatory metabolic rate) and MMR (maximum exercise metabolic rate in a hop-flutter chamber) in winter for dark-eyed juncos (Junco hyemalis), American goldfinches (Carduelis tristis; M(sum) and MMR only), and black-capped chickadees (Poecile atricapillus; BMR and M(sum) only) and examined correlations among these variables. We also measured BMR and M(sum) in individual house sparrows (Passer domesticus) in both summer, winter and spring. For both raw metabolic rates and residuals from allometric regressions, BMR was not significantly correlated with either M(sum) or MMR in juncos. Moreover, no significant correlation between M(sum) and MMR or their mass-independent residuals occurred for juncos or goldfinches. Raw BMR and M(sum) were significantly positively correlated for black-capped chickadees and house sparrows, but mass-independent residuals of BMR and M(sum) were not. These data suggest that central organ and exercise organ metabolic levels are not inextricably linked and that muscular capacities for exercise and shivering do not necessarily vary in tandem in individual birds. Why intraspecific and interspecific avian studies show differing results and the significance of these differences to the aerobic capacity model are unknown, and resolution of these questions will require additional studies of potential mechanistic links between minimal and maximal metabolic output.     

Dominant black-capped chickadees pay no maintenance energy costs for their wintering status and are not better at enduring cold than subordinate individuals

Winter requires physiological adjustments in northern resident passerines. Cold acclimatization is generally associated with an increase in physiological maintenance costs, measured as basal metabolic rate (BMR), and cold endurance, reflected by summit metabolic rate (M _sum). However, several northern species also form social groups in winter and a bird’s hierarchical position may influence the size of its metabolically active organs as well as its BMR. Winter metabolic performance in these species may therefore reflect a complex set of adjustments to both seasonal climatic variations and social environment. We studied the effect of social status on parameters of cold acclimatization (body mass, size of fat reserves and pectoral muscles, BMR and M _sum) in free-living black-capped chickadees (Poecile atricapillus). Birds that were structurally large and heavy for their body size, mostly dominant individuals, carried more fat reserves and had larger pectoral muscles. However, social status had little effect on metabolic performance in the cold. Indeed, M _sum was independent of social rank while mass-corrected BMR was slightly lower in dominant individuals, likely due to a statistical dilution effect caused by large metabolically inactive fat reserves. BMR and M _sum, whether considered in terms of whole-animal values, corrected for body mass or body size were nevertheless correlated, suggesting a functional link between these metabolic components. Our results therefore indicate that the energy cost of social dominance is not a generalized phenomenon in small wintering birds.