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Reversible phenotypic flexibility allows organisms to better match phenotypes to prevailing environmental conditions and may produce fitness benefits. Costs and constraints of phenotypic flexibility may limit the capacity for flexible responses but are not well understood nor documented. Costs could include expenses associated with maintaining the flexible system or with generating the flexible response. One potential cost of maintaining a flexible system is an energetic cost reflected in the basal metabolic rate (BMR), with elevated BMR in individuals with more flexible metabolic responses. We accessed data from thermal acclimation studies of birds where BMR and/or M sum (maximum cold-induced metabolic rate) were measured before and after acclimation, as a measure of metabolic flexibility, to test the hypothesis that flexibility in BMR (ΔBMR), M sum (ΔM sum ), or metabolic scope (M sum  - BMR; ΔScope) is positively correlated with BMR. When temperature treatments lasted at least three weeks, three of six species showed significant positive correlations between ΔBMR and BMR, one species showed a significant negative correlation, and two species showed no significant correlation. ΔM sum and BMR were not significantly correlated for any species and ΔScope and BMR were significantly positively correlated for only one species. These data suggest that support costs exist for maintaining high BMR flexibility for some bird species, but high flexibility in M sum or metabolic scope does not generally incur elevated maintenance costs., (© 2023. The Author(s).)

We investigated the role of mitochondrial function in the avian thermoregulatory response to a cold environment. Using black-capped chickadees (Poecile atricapillus) acclimated to cold (-10°C) and thermoneutral (27°C) temperatures, we expected to observe an upregulation of pectoralis muscle and liver respiratory capacity that would be visible in mitochondrial adjustments in cold-acclimated birds. We also predicted that these adjustments would correlate with thermogenic capacity (Msum) and basal metabolic rate (BMR). Using tissue high-resolution respirometry, mitochondrial performance was measured as respiration rate triggered by proton leak and the activity of complex I (OXPHOSCI) and complex I+II (OXPHOSCI+CII) in the liver and pectoralis muscle. The activity of citrate synthase (CS) and cytochrome c oxidase (CCO) was also used as a marker of mitochondrial density. We found 20% higher total CS activity in the whole pectoralis muscle and 39% higher total CCO activity in the whole liver of cold-acclimated chickadees relative to that of birds kept at thermoneutrality. This indicates that cold acclimation increased overall aerobic capacity of these tissues. Msum correlated positively with mitochondrial proton leak in the muscle of cold-acclimated birds while BMR correlated with OXPHOSCI in the liver with a pattern that differed between treatments. Consequently, this study revealed a divergence in mitochondrial metabolism between thermal acclimation states in birds. Some functions of the mitochondria covary with thermogenic capacity and basal maintenance costs in patterns that are dependent on temperature and body mass., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2022. Published by The Company of Biologists Ltd.)

It is generally assumed that small birds improve their shivering heat production capacity by developing the size of their pectoralis muscles. However, some studies have reported an enhancement of thermogenic capacity in the absence of muscle mass variation between seasons or thermal treatments. We tested the hypothesis that an increase in muscle mass is not a prerequisite for improving avian thermogenic capacity. We measured basal (BMR) and summit (M sum ) metabolic rates of black capped chickadees (Poecile atricapillus) acclimated to thermoneutral (27 °C) and cold (-10 °C) temperatures and obtained body composition data from dissections. Cold acclimated birds consumed 44% more food, and had 5% and 20% higher BMR and M sum , respectively, compared to individuals kept at thermoneutrality. However, lean dry pectoralis and total muscle mass did not differ between treatments, confirming that the improvement of thermogenic capacity did not require an increase in skeletal muscle mass. Nevertheless, within temperature treatments, M sum was positively correlated with the mass of all measured muscles, including the pectoralis. Therefore, for a given acclimation temperature individuals with large muscles do benefit from muscle size in term of heat production but improving thermogenic capacity during cold acclimation likely requires an upregulation of cell functions.

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.

Avian basal metabolic rate (BMR) and summit metabolic rate (M sum ) vary in parallel during cold acclimation and acclimatization, which implies a functional link between these variables. However, evidence suggests that these parameters may reflect different physiological systems acting independently. We tested this hypothesis in white-throated sparrows (Zonotrichia albicollis) acclimated to two temperatures (-8° and 28°C) and two diets (0% and 30% cellulose). We expected to find an uncoupling of M sum and BMR where M sum , a measure of maximal shivering heat production, would reflect muscle and heart mass variation and would respond only to temperature, while BMR would reflect changes in digestive and excretory organs in response to daily food intake, responding to both temperature and diet. We found that the gizzard, liver, kidneys, and intestines responded to treatments through a positive relationship with food intake. BMR was 15% higher in cold-acclimated birds and, as expected, varied with food intake and the mass of digestive and excretory organs. In contrast, although M sum was 19% higher in cold-acclimated birds, only heart mass responded to temperature (+18% in the cold). Pectoral muscles did not change in mass with temperature but were 8.2% lighter on the cellulose diet. Nevertheless, M sum varied positively with the mass of heart and skeletal muscles but only in cold-acclimated birds. Our results therefore suggest that an upregulation of muscle metabolic intensity is required for cold acclimation. This study increases support for the hypothesis that BMR and M sum reflect different physiological systems responding in parallel to constraints associated with cold environments.

In birds, acclimation and acclimatization to temperature are associated with changes in basal (BMR), summit (Msum) and maximal (MMR) metabolic rates but little is known about the rate at which species adjust their phenotype to short-term temperature variations. Our aims were (1) to determine the pattern of metabolic adjustments following a rapid temperature change, (2) to determine whether performance varies at similar rates during exposure to warm or cold environments, and (3) to determine if BMR, Msum and MMR change at comparable rates during thermal acclimation. We measured these parameters in white-throated sparrows (Zonotrichia albicollis), black-capped chickadees (Poecile atricapillus), and snow buntings (Plectrophenax nivalis) after acclimation to 10 °C (day 0) and on the 4th and 8th days of acclimation to either -5 or 28 °C. Birds changed their metabolic phenotype within 8 days with patterns differing among species. Sparrows expressed the expected metabolic increases in the cold and decreases at thermoneutrality while performance in chickadees and buntings was not influenced by temperature but changed over time with inverse patterns. Our results suggest that BMR varies at comparable rates in warm and cold environments but changes faster than Msum and MMR, likely due to limitations in the rate of change in organ size and function. They also suggest that maximal metabolic capacity is lost faster in a warm environment than it is gained in a cold environment. With the expected increase in temperature stochasticity at northern latitudes, a loss of thermogenic capacity during warm winter days could, therefore, be detrimental if birds are slow to readjust their phenotype with the return of cold days.

Reaction norms reflect an organisms' capacity to adjust its phenotype to the environment and allows for identifying trait values associated with physiological limits. However, reaction norms of physiological parameters are mostly unknown for endotherms living in natural conditions. Black-capped chickadees (Poecile atricapillus) increase their metabolic performance during winter acclimatization and are thus good model to measure reaction norms in the wild. We repeatedly measured basal (BMR) and summit (Msum) metabolism in chickadees to characterize, for the first time in a free-living endotherm, reaction norms of these parameters across the natural range of weather variation. BMR varied between individuals and was weakly and negatively related to minimal temperature. Msum varied with minimal temperature following a Z-shape curve, increasing linearly between 24°C and -10°C, and changed with absolute humidity following a U-shape relationship. These results suggest that thermal exchanges with the environment have minimal effects on maintenance costs, which may be individual-dependent, while thermogenic capacity is responding to body heat loss. Our results suggest also that BMR and Msum respond to different and likely independent constraints.

Abstract Small avian species wintering at northern latitudes typically show increases in basal metabolic rate (BMR) and maximal thermogenic capacity (Msum). Those are widely assumed to reflect changes in body composition, with enlargement of digestive and excretory organs resulting in elevated winter BMR and larger body muscles driving the increase in Msum. Using free-living black-capped chickadees (Poecile atricapillus) as our model species, we investigated seasonal changes in body composition and tested for relationships between mass variations of body organs and variability of both BMR and Msum. Our results confirmed the expected winter increase in mass of body muscles and cardiopulmonary organs (heart + lungs) and showed that 64% of the observed Msum variations throughout the year were explained by changes in these organs. In contrast, we found little support for an effect of the digestive organs (gizzard + intestines) on BMR seasonal changes. Instead, this variable was mainly influenced by variations in mass of body muscles and excretory organs (liver + kidney), explaining up to 35% of its variability.

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 (MSUM) 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 (Poecileatricapillus). Results confirmed that population patterns indeed reflect flexibility at the individual level. They showed the expected increase in BMR (6%) and MSUM (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 MSUM had already achieved more than 20% of its inter-seasonal increase by October, and declined to its starting level by March, while MSUM remained high. Although both parameters co-vary on a yearly scale, this mismatch in the timing of variation in winter BMR and MSUM likely reflects different constraints acting on different physiological components and therefore suggests a lack of functional link between these parameters.

The energy cost of egg production in passerine birds has typically been estimated to be 45%-60% of basal metabolic rate (BMR), but this is based on theoretical models using data on energy content of eggs and reproductive tissue; there are still very few empirical data on egg production costs. In this study, we directly measured resting metabolic rate (RMR) in egg-laying female European starlings (Sturnus vulgaris) over 3 yr. We compared these data with RMR of nonbreeding and chick-rearing birds and with estimated energy expenditure generated from a typical energy content model by using empirically derived data from body composition analysis for this species. We found marked variation in RMR between years and between reproductive stages, which complicates comparisons among breeding stages for the assessment of relative egg production costs. On the basis of this method, RMR during egg laying varied from +74% to -13% of nonbreeding RMR and from +20% to -7% of chick-rearing RMR. We therefore used an alternate approach: measuring changes in RMR through the complete cycle of follicle development and ovulation. The increase in RMR from the beginning of prelaying to the six-follicle stage (before first ovulation) when birds have a complete developing follicle hierarchy was 22.4%. This value is still much lower than that estimated from our energy content model. We discuss conceptual problems associated with the theoretical energy content approach but also suggest, on the basis of earlier work done in our lab, that the measured increase in RMR might still underestimate the actual cost of egg production if birds reallocate energy between different physiological systems.

Rising global temperatures are expected to increase reproductive costs for wildlife as greater thermoregulatory demands interfere with reproductive activities. However, predicting the temperatures at which reproductive performance is negatively impacted remains a significant hurdle. Using a thermoregulatory polygon approach, we derived a reproductive threshold temperature for an Arctic songbird—the snow bunting (Plectrophenax nivalis). We defined this threshold as the temperature at which individuals must reduce activity to suboptimal levels (i.e. less than four-time basal metabolic rate) to sustain nestling provisioning and avoid overheating. We then compared this threshold to operative temperatures recorded at high (82° N) and low (64° N) Arctic sites to estimate how heat constraints translate into site-specific impacts on sustained activity level. We predict buntings would become behaviourally constrained at operative temperatures above 11.7°C, whereupon they must reduce provisioning rates to avoid overheating. Low-Arctic sites had larger fluctuations in solar radiation, consistently producing daily periods when operative temperatures exceeded 11.7°C. However, high-latitude birds faced entire, consecutive days when parents would be unable to sustain required provisioning rates. These data indicate that Arctic warming is probably already disrupting the breeding performance of cold-specialist birds and suggests counterintuitive and severe negative impacts of warming at higher latitude breeding locations. [ABSTRACT FROM AUTHOR]

Resident temperate passerines adjust their phenotypes to cope with winter constraints, with peak performance in metabolic traits typically occurring during the coldest months. However, it is sparsely known whether cold-adapted northern species make similar adjustments when faced with variable seasonal environments. Life in near-constant cold could be associated with limited flexibility in traits underlying cold endurance. We investigated this by tracking individual physiological changes over five consecutive winters in snow buntings (Plectrophenax nivalis), an Arctic-breeding migratory passerine typically confronted with nearly constant cold. Buntings were held in an outdoor aviary and exposed to seasonal temperature variation typical of temperate zone climates. We measured phenotypic changes in body composition (body, fat, and lean mass, pectoralis muscle thickness), oxygen transport capacity (hematocrit), metabolic performance (basal metabolic rate [BMR] and summit metabolic rate [ M sum]), thermogenic endurance (time to reach M sum), and cold tolerance (temperature at M sum). Snow buntings showed flexibility in functions underlying thermogenic capacity and cold endurance comparable to that observed in temperate resident passerines wintering at similar latitudes. Specifically, they increased body mass (13%), fat mass (246%), hematocrit (23%), pectoralis muscle thickness (8%), and M sum (27%). We also found remarkable cold tolerance in these birds, with individuals reaching M sum in helox at temperatures equivalent to less than −90°C in air. However, in contrast with resident temperate passerines, lean mass decreased by 12%, and there was no clear increase in maintenance costs (BMR). Our results show that the flexibility of traits underlying thermal acclimatization in a cold-adapted northern species is comparable to that of temperate resident species living at lower latitudes and is therefore not limited by life in near-constant cold. [ABSTRACT FROM AUTHOR]

Small resident bird species living at northern latitudes increase their metabolism in winter, and this is widely assumed to improve their chances of survival. However, the relationship between winter metabolic performance and survival has yet to be demonstrated. Using capture-mark-recapture,we followed a population of free-living black-capped chickadees (Poecile atricapillus) over 3 yr and evaluated their survival probability within and among winters. We also measured the size-independent body mass (Ms), hematocrit (Hct), basal metabolic rate (BMR), and maximal thermogenic capacity (Msum) and investigated how these parameters influenced survival within and among winters. Results showed that survival probability was high and constant both within (0.92) and among (0.96) winters. They also showed that while Ms, Hct, and BMR had no significant influence, survival was positively related to Msum—following a sigmoid relationship— within but not among winter. Birds expressing an Msum below 1.26W(i.e., similar to summer levels) had a !50% chance of survival, while birds with anMsumabove 1.35Whad at least a 90% chance of surviving through the winter. Our data therefore suggest that black-capped chickadees that are either too slow or unable to adjust their phenotype from summer to winter have little chances of survival and thus that seasonal upregulation of metabolic performance is highly beneficial. This study is the first to document in an avian system the relationship between thermogenic capacity and winter survival, a proxy of fitness. [ABSTRACT FROM AUTHOR]

ABSTRACT Resident passerines inhabiting high latitude environments are faced with strong seasonal changes in thermal conditions and energy availability. Summit metabolic rate (maximal metabolic rate elicited by shivering during cold exposure: Msum) and basal metabolic rate (BMR) vary in parallel among seasons and increase in winter due to cold acclimatization, and these adjustments are thought to be critical for survival. Wintering individuals expressing consistently higher Msum and BMR could therefore be seen as better performers with higher chances of winter survival than those exhibiting lower metabolic performance. In this study, we calculated repeatability to evaluate temporal consistency of body mass, BMR and Msum within and across three consecutives winters in black-capped chickadees ( Poecile atricapillus). We found that body mass was significantly repeatable both within and across winters (R 0.51-0.90). BMR (R 0.29-0.47) was only repeatable within winter while Msum was repeatable both among (R 0.33-0.49) and within winters (R 0.33-0.49) with the magnitude and significance of repeatability in both variables depending on the year and whether they were corrected for body mass or body size. The patterns of repeatability observed among years also differed between the two variables. Our findings suggest that the relative ranking of individuals in winter metabolic performance is affected by local ecological conditions and can change within relatively short periods of time. J. Exp. Zool. 323A: 179-190, 2015. © 2015 Wiley Periodicals, Inc. [ABSTRACT FROM AUTHOR]

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). 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) 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 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, 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. [ABSTRACT FROM AUTHOR]

Interest in phenotypic flexibility has increased dramatically over the last decade, but flexibility during reproduction has received relatively little attention from avian scientists, despite its possible impact on fitness. Because most avian species maintain atrophied reproductive organs when not active, reproduction in birds requires major tissue remodeling in preparation for breeding. Females undergo rapid (days) recrudescence and regression of their reproductive organs at each breeding attempt, while males grow their organs ahead of time at a much slower rate (weeks) and may maintain them at maximal size throughout the breeding season. Reproduction is associated with significant metabolic costs. Egg production leads to a 22%-27% increase in resting metabolic rate (RMR) over non-reproductive values. This is partly due to the activity of the oviduct, an organ that may allow females to adjust reproductive investment by modulating egg size and quality. In males, gonadal recrudescence may lead to a 30% increase in RMR, but the data are inconsistent and general conclusions regarding energetic costs of reproduction in males will require more research. Recent studies on captive female zebra finches describe the impacts of these costs on daily energy budgets and highlight the strategies used by birds to maintain their investment in reproduction when energy is limited. Whenever possible, birds use behavioral flexibility as a first means of saving energy. Decreasing locomotor activity saves energy during challenges such as egg production or exposure to cold temperatures and is an efficient way to buffer variation in individual daily energy budgets. However, when behavioral flexibility is not possible, birds must rely on flexibility at the physiological level to meet energy demands. In zebra finches breeding in the cold, this results in a reduced pace of laying, likely due to down-regulation of both reproductive and non-reproductive function, allowing females to defend minimal egg size and maintain reproductive success. More research involving a range of species in captive and free-living conditions is needed to determine how phenotypic flexibility during tissue remodeling and early reproductive investment translates to natural conditions and affects fitness. [ABSTRACT FROM AUTHOR]

Previous experiments showed reduction of basal metabolic rate (BMR) in birds facing energetic challenges. We alternately exposed two groups of red knots (Calidris canutus) to either 6 h or 22 h of food availability for periods of 22 d. Six h of access to food led to a 6%-10% loss of body mass over the first 8 d, with nearly all of the birds' daily energy expenditures supported by body nutrient stores during the first 2 d. Birds responded by increasing feeding behavior and food intake, but the response was slow. There were no gains of mass before day 15, which suggests a digestive bottleneck and a period of physiological adjustment. Food-restricted birds exhibited decreases in pectoral-muscle thickness and BMR in association with a loss of body mass. Although a decrease in BMR saves energy, savings represented only 2%-7% of the daily energy spent in excess of that acquired during the deficit period. Red knots did not downregulate mass-independent BMR. On the bases of recent independent findings and the pattern of mass gain observed when food access was switched from 6 h to 22 h, we suggest that these birds routinely maintain nutrient stores as a buffer against periods of energy shortages, thereby precluding the need for downregulation of mass-independent BMR. [ABSTRACT FROM AUTHOR]

Basal metabolic rate (BMR) in animals is interpreted as reflecting the size and metabolic intensity of energy-consuming tissues. However, studies investigating relationships between the mass of specific organs and interindividual variation in BMR have produced inconsistent patterns with regard to which organs have the largest impact on BMR variation. Because of the known flexibility in organ mass and metabolic intensity within individual organs, relationships between BMR and body-composition variables are bound to be context specific. Altricial nestlings are excellent models to illustrate this phenomenon because of the extreme variation in body composition occurring during growth. Using European starlings at three age classes, we studied changes in body composition together with its effect on variation in resting metabolic rate (RMR) in order to highlight the context-specific nature of these relationships. Our data suggest a transition in metabolic costs during growth in starling nestlings. During the linear phase of growth, energy is mainly consumed by tissue-synthesis processes, with fast-growing organs having a large influence on RMR. In the plateau phase of growth, the energy expenditure is transferred to functional costs, with high-intensity organs having a predominant effect on RMR variation. Our data illustrates the context-specific nature of organ mass-metabolic rate correlations, which complicates inter- and intraspecific comparisons of BMR. In the future, such comparisons must be done while taking the physiological state of the study animal into account. [ABSTRACT FROM AUTHOR]

Even though molt involves both endocrine and energetic changes in bird bodies, this study is among the first to combine assessments of energy costs together with thyroid hormone variations in molting birds. Individual shorebirds (red knots Calidris canutus islandica) were measured while in full summer and winter plumage as well as during peak of molt. Molt was associated with a 9.8% increase in average mass-independent basal metabolic rate (BMR) above nonmolting levels. Individual plasma levels of thyroxine (T4) were correlated with individual rate of body feather renewal, confirming that T4 is related to body molt but also showing that it is potentially regulating its rate. Across seasons, mass-independent average heat loss measured as conductance gradually declined with conductance during molt falling between measured values for summer and winter. During the molting period, however, body molting rate was positively correlated with thermal conductance, indicating that for a given ambient temperature below thermoneutrality, the fastest molting birds were losing more body heat. Across seasons, triiodothyronine (T3), a hormone typically upregulated in response to a cold stimulus, was correlated with individual thermal conductance and BMR. We suggest that the increased heat loss of fast-molting birds leads to a cold-acclimatization response that may be partly responsible for the elevated BMR measured during molt. This could be mediated through a stimulatory effect of T3 on BMR in response to increased heat loss. Our interpretation is supported by a positive relationship between the individual changes in conductance and the change in BMR from summer to the molting period. [ABSTRACT FROM AUTHOR]

In the calidrine sandpiper red knot (Calidris canutus), the weeks preceding takeoff for long-distance migration are characterized by a rapid increase in body mass, largely made up of fat but also including a significant proportion of lean tissue. Before takeoff, the pectoral muscles are known to hypertrophy in preparation for endurance flight without any specific training. Because birds facing cold environments counterbalance heat loss through shivering thermogenesis, and since pectoral muscles represent a large proportion of avian body mass, we asked the question whether muscle hypertrophy in preparation for long-distance endurance flight would induce improvements in thermogenic capacity. We acclimated red knots to different controlled thermal environments: 26°C, 5°C, and variable conditions tracking outdoor temperatures. We then studied within-individual variations in body mass, pectoral muscle size (measured by ultrasound), and metabolic parameters [basal metabolic rate (BMR) and summit metabolic rate (Msum)] throughout a 3-mo period enclosing the migratory gain and loss of mass. The gain in body mass during the fattening period was associated with increases in pectoral muscle thickness and thermogenic capacity independent of thermal acclimation. Regardless of their thermal treatment, birds showing the largest increases in body mass also exhibited the largest increases in Msum. We conclude that migratory fattening is accompanied by thermoregulatory side effects. The gain of body mass and muscle hypertrophy improve thermogenic capacity independent of thermal acclimation in this species. Whether this represents an ecological advantage depends on the ambient temperature at the time of fattening. [ABSTRACT FROM AUTHOR]