Summit Metabolism Research Articles

The role of the heart in the evolution of aerobic performance.

Aerobic metabolism underlies vital traits such as locomotion and thermogenesis, and aerobic capacity influences fitness in many animals. The heart is a key determinant of aerobic capacity, but the relative influence of cardiac output versus other steps in the O2 transport pathway remains contentious. In this Commentary, we consider this issue by examining the mechanistic basis for adaptive increases in aerobic capacity (thermogenic V̇O2,max; also called summit metabolism) in deer mice (Peromyscus maniculatus) native to high altitude. Thermogenic V̇O2,max is increased by acclimation to cold hypoxia (simulating high-altitude conditions), and high-altitude populations generally have greater V̇O2,max than their low-altitude counterparts. This plastic and evolved variation in V̇O2,max is associated with corresponding variation in maximal cardiac output, along with variation in other traits across the O2 pathway (e.g. arterial O2 saturation, blood haemoglobin content and O2 affinity, tissue O2 extraction, tissue oxidative capacity). By applying fundamental principles of gas exchange, we show that the relative influence of cardiac output on V̇O2,max depends on the O2 diffusing capacity of thermogenic tissues (skeletal muscles and brown adipose tissues). Functional interactions between cardiac output and blood haemoglobin content determine circulatory O2 delivery and thus affect V̇O2,max, particularly in high-altitude environments where erythropoiesis can increase haematocrit and blood viscosity. There may also be functional linkages between cardiac output and tissue O2 diffusion due to the role of blood flow in determining capillary haematocrit and red blood cell flux. Therefore, the functional interactions between cardiac output and other traits in the O2 pathway underlie the adaptive evolution of aerobic capacities.

Metabolic adjustments to winter severity in two geographically separated great tit (Parus major) populations.

Understanding the potential limits placed on organisms by their ecophysiology is crucial for predicting their responses to varying environmental conditions. A main hypothesis for explaining avian thermoregulatory mechanisms is the aerobic capacity model, which posits a positive correlation between basal (basal metabolic rate [BMR]) and summit (Msum) metabolism. Most evidence for this hypothesis, however, comes from interspecific comparisons, and the ecophysiological underpinnings of avian thermoregulatory capacities hence remain controversial. Indeed, studies have traditionally relied on between-species comparisons, although, recently, there has been a growing recognition of the importance of intraspecific variation in ecophysiological responses. Therefore, here, we focused on great tits (Parus major), measuring BMR and Msum during winter in two populations from two different climates: maritime-temperate (Gontrode, Belgium) and continental (Zvenigorod, Russia). We tested for the presence of intraspecific geographical variation in metabolic rates and assessed the predictions following the aerobic capacity model. We found that birds from the maritime-temperate climate (Gontrode) showed higher BMR, whereas conversely, great tits from Zvenigorod showed higher levels of Msum. Within each population, our data did not fully support the aerobic capacity model's predictions. We argued that the decoupling of BMR and Msum observed may be caused by different selective forces acting on these metabolic rates, with birds from the continental-climate Zvenigorod population facing the need to conserve energy for surviving long winter nights (by keeping their BMR at low levels) while simultaneously being able to generate more heat (i.e., a high Msum) to withstand cold spells.

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 (Msum) 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 Msum (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 Msum 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, Msum 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.

Phenotypic flexibility of metabolic rate and evaporative water loss does not vary across a climatic gradient in an Afrotropical passerine bird.

Small birds inhabiting northern temperate and boreal latitudes typically increase metabolic rates during cold winters or acclimation to low air temperatures (Taccl). Recent studies suggest considerable variation in patterns of seasonal metabolic acclimatization in birds from subtropical and tropical regions with milder winters, but there remains a dearth of acclimation studies investigating metabolic flexibility among lower-latitude birds. We used short-term thermal acclimation experiments to investigate phenotypic flexibility in basal metabolic rate (BMR), thermoneutral evaporative water loss (EWL) and summit metabolism (Msum) in three populations of white-browed sparrow-weavers (Plocepasser mahali) along a climatic and aridity gradient. We allocated individuals to one of three Taccl treatments (5, 20 and 35°C; n=11 per population per Taccl) for 28 days, and measured post-acclimation BMR, EWL and Msum using flow-through respirometry. Our data reveal the expected pattern of lower BMR and EWL (∼12% and 25% lower, respectively) in birds at Taccl=35°C compared with cooler Taccl treatments, as observed in previous acclimation studies on subtropical birds. We found no variation in the reaction norms of BMR and EWL among populations in response to acclimation, suggesting previously documented differences in seasonal BMR acclimatization are the result of phenotypic flexibility. In contrast to higher-latitude species, Msum did not significantly vary in response to thermal acclimation. These findings support the idea that factors other than enhancing cold tolerance may be driving patterns of metabolic variation in subtropical birds.

Context-dependent regulation of pectoralis myostatin and lipid transporters by temperature and photoperiod in dark-eyed juncos

A prominent example of seasonal phenotypic flexibility is the winter increase in thermogenic capacity (=summit metabolism, ) in small birds, which is often accompanied by increases in pectoralis muscle mass and lipid catabolic capacity. Temperature or photoperiod may be drivers of the winter phenotype, but their relative impacts on muscle remodeling or lipid transport pathways are little known. We examined photoperiod and temperature effects on pectoralis muscle expression of myostatin, a muscle growth inhibitor, and its tolloid-like protein activators (TLL-1 and TLL-2), and sarcolemmal and intracellular lipid transporters in dark-eyed juncos Junco hyemalis. We acclimated winter juncos to four temperature (3 °C or 24 °C) and photoperiod [short-day (SD) = 8L:16D; long-day (LD) = 16L:8D] treatments. We found that myostatin, TLL-1, TLL-2, and lipid transporter mRNA expression and myostatin protein expression did not differ among treatments, but treatments interacted to influence lipid transporter protein expression. Fatty acid translocase (FAT/CD36) levels were higher for cold SD than for other treatments. Membrane-bound fatty acid binding protein (FABPpm) levels, however, were higher for the cold LD treatment than for cold SD and warm LD treatments. Cytosolic fatty acid binding protein (FABPc) levels were higher on LD than on SD at 3 °C, but higher on SD than on LD at 24 °C. Cold temperature groups showed upregulation of these lipid transporters, which could contribute to elevated Msum compared to warm groups on the same photoperiod. However, interactions of temperature or photoperiod effects on muscle remodeling and lipid transport pathways suggest that these effects are context-dependent.

Seasonal Metabolic Acclimatization Varies in Direction and Magnitude among Populations of an Afrotropical Passerine Bird.

Avian metabolic responses demonstrate considerable diversity under fluctuating environmental conditions, a well-studied example being the seasonal upregulation of basal metabolic rate (BMR) and summit metabolism (Msum) in temperate species experiencing harsh winters. Fewer studies have examined seasonal metabolic acclimatization in subtropical or tropical species. We investigated seasonal metabolic variation in an Afrotropical ploceid passerine, the white-browed sparrow-weaver (Plocepasser mahali; ∼47 g), at three sites along a climatic gradient of approximately 7°C in winter minimum air temperature (Ta). We measured Msum (n ≥ 10 per site per season) in a helox atmosphere, BMR of the same birds at thermoneutrality (Ta ≈ 30°C), and resting metabolic rates at 5°C ≤ Ta ≤ 20°C. Patterns of seasonal adjustments in BMR varied among populations in a manner not solely related to variation in seasonal Ta extremes, ranging from BMR ∼52% higher in winter than in summer to no seasonal difference. Greater cold tolerance was found in a population at a colder desert site, manifested as higher Msum (∼25% higher) and lower helox temperature at cold limit values compared with a milder, mesic site. Our results lend support to the idea that greater variance in the pattern of seasonal metabolic responses occurs in subtropical and tropical species compared with their temperate-zone counterparts and that factors other than Ta extremes (e.g., food availability) may be important in determining the magnitude and direction of seasonal metabolic adjustments in subtropical birds.

Solar Radiation during Rewarming from Torpor in Elephant Shrews: Supplementation or Substitution of Endogenous Heat Production?

Many small mammals bask in the sun during rewarming from heterothermy, but the implications of this behaviour for their energy balance remain little understood. Specifically, it remains unclear whether solar radiation supplements endogenous metabolic thermogenesis (i.e., rewarming occurs through the additive effects of internally-produced and external heat), or whether solar radiation reduces the energy required to rewarm by substituting (i.e, replacing) metabolic heat production. To address this question, we examined patterns of torpor and rewarming rates in eastern rock elephant shrews (Elephantulus myurus) housed in outdoor cages with access to either natural levels of solar radiation or levels that were experimentally reduced by means of shade cloth. We also tested whether acclimation to solar radiation availability was manifested via phenotypic flexibility in basal metabolic rate (BMR), non-shivering thermogenesis (NST) capacity and/or summit metabolism (Msum). Rewarming rates varied significantly among treatments, with elephant shrews experiencing natural solar radiation levels rewarming faster than conspecifics experiencing solar radiation levels equivalent to approximately 20% or 40% of natural levels. BMR differed significantly between individuals experiencing natural levels of solar radiation and conspecifics experiencing approximately 20% of natural levels, but no between-treatment difference was evident for NST capacity or Msum. The positive relationship between solar radiation availability and rewarming rate, together with the absence of acclimation in maximum non-shivering and total heat production capacities, suggests that under the conditions of this study solar radiation supplemented rather than substituted metabolic thermogenesis as a source of heat during rewarming from heterothermy.

Global patterns of seasonal acclimatization in avian resting metabolic rates

The adjustment of resting metabolic rates represents an important component of avian seasonal acclimatization, with recent studies revealing substantial differences between summer and winter in birds from a wide range of latitudes. We compared seasonal variation in basal metabolic rate (BMR) and summit metabolism (M sum) between temperate and tropical/subtropical latitudes, and examined correlations with latitude and temperature. The direction and magnitude of seasonal adjustments in BMR are broadly related to temperature and latitude, but are significantly more variable among tropical and subtropical species compared to those inhabiting temperate zones. Winter adjustments in BMR among subtropical species, when expressed relative to summer values, range from decreases of approximately 35 % to increases of more than 60 %, whereas the majority of temperate-zone species show increases in BMR during winter. Relatively few seasonal M sum data exist for tropical/subtropical species, but those that are available involve responses ranging from winter decreases to increases of similar magnitude to those characteristic of many temperate-zone species. Recent studies also highlight the substantial variation in seasonal adjustments that may occur within species, and reiterate the need for further investigations of the relative roles of environmental variables such as temperature and food availability as determinants of seasonal metabolic variation.

Summit metabolic rate exhibits phenotypic flexibility with migration, but not latitude in a neotropical migrant, Parkesia noveboracensis

Physiology–life history interactions suggest that birds living with a fast ‘pace-of-life’ should have higher metabolic capacity to provision higher reproductive activity. Previous work supports this, but does not consider migration. We measured summit metabolism (\(\dot{V}{\text{O}}_{{ 2\;{\text{summit}}}}\)) in Northern Waterthrush (Parkesia noveboracensis) while wintering in the Republic of Panama, migrating northwards through eastern North America, and while breeding in northeastern North America. \(\dot{V}{\text{O}}_{{ 2\;{\text{summit}}}}\) is similar between breeding and overwintering (non-migratory) and is significantly elevated in migration. These data suggest that migration is a driver of phenotypic flexibility in these birds and that migration, like winter survival, may be an important determinant of connections between life history and physiology.

Reaction norms in natural conditions: how does metabolic performance respond to weather variations in a small endotherm facing cold environments?

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.

Characterization of muscle plasticity in a neotropical migrant, Dumetella carolinesis (gray catbird) (878.2)

This study focuses on how avian migration is facilitated by changes in organismal metabolism, muscle structure, and muscle metabolism. These specific migratory changes were examined across the annual cycle on‐an organismal level by investigating whole body oxidative capacity (ability to use energy) and, on a tissue level by characterizing muscle tissue size and oxidative capacity. In order to quantify whole body oxidative capacity, summit metabolism was measured to determine the peak oxygen consumed by each individual. Birds were euthanized and the pectoralis (flight muscle) and gastrocnemius (leg muscle) were removed and weighed. Mitochondrial content of the muscle was quantified by measuring enzyme activity via citrate synthase and cytochrome oxidase assays to determine oxidative capacity of the muscles. Summit metabolism and flight muscle mass were greatest during southward migration compared to temperate breeding and tropical wintering. The Gray Catbird displays phenotypic plasticity during migration compared to non‐migratory periods suggesting that augmentation of the flight muscle and increased metabolism are necessary for migration.Grant Funding Source: Supported by NSF #IOS‐1257455 to PJS

Relative roles of temperature and photoperiod as drivers of metabolic flexibility in dark-eyed juncos

Seasonal phenotypic flexibility in small birds produces a winter phenotype with elevated maximum cold-induced metabolic rates (=summit metabolism, Msum). Temperature and photoperiod are candidates for drivers of seasonal phenotypes, but their relative impacts on metabolic variation are unknown. We examined photoperiod and temperature effects on Msum, muscle masses and activities of key catabolic enzymes in winter dark-eyed juncos (Junco hyemalis). We randomly assigned birds to four treatment groups varying in temperature (cold=3°C; warm=24°C) and photoperiod [short day (SD)=8 h:16 h light:dark; long day (LD)=16 h:8 h light:dark] in a two-by-two design. We measured body mass (Mb), flight muscle width and Msum before and after 3 and 6 weeks of acclimation, and flight muscle and heart masses after 6 weeks. Msum increased for cold-exposed, but not for warm-exposed, birds. LD birds gained more Mb than SD birds, irrespective of temperature. Flight muscle size and mass did not differ significantly among groups, but heart mass was larger in cold-exposed birds. Citrate synthase, carnitine palmitoyl transferase and β-hydroxyacyl Co-A dehydrogenase activities in the pectoralis were generally higher for LD and cold groups. The cold-induced changes in Msum and heart mass parallel winter changes for small birds, but the larger Mb and higher catabolic enzyme activities in LD birds suggest photoperiod-induced changes associated with migratory disposition. Temperature appears to be a primary driver of flexibility in Msum in juncos, but photoperiod-induced changes in Mb and catabolic enzyme activities, likely associated with migratory disposition, interact with temperature to contribute to seasonal phenotypes.

Basal metabolic rate, maximum thermogenic capacity and aerobic scope in rodents: interaction between environmental temperature and torpor use

When torpid animals arouse and warm up to restore normal body temperature (T(b)), they produce heat at levels that can reach up to 10 times basal metabolic rate (BMR), close to the cold-induced summit metabolism (VO(2)-sum). Because torpor is an adaptation aimed at conserving energy over periods of low ambient temperature (T(a)) and food availability, selective forces that have led to the evolution of torpor may have simultaneously favoured high thermogenic capacity (i.e. VO(2)-sum) relative to the maintenance costs (i.e. BMR), hence a higher factorial aerobic scope (FAS; the ratio of VO(2)-sum to BMR). My objective was to test this adaptive hypothesis using a phylogenetically informed comparative approach with data on BMR and VO(2)-sum in rodents. I found a strong negative correlation between FAS and the average of the daily minimum T(a) (T(min)) in species using torpor, which was due to differential effects of T(a) on BMR (but not VO(2)-sum) in species that use torpor compared with species that do not. In addition, FAS was negatively correlated with the lowest torpid T(b) in a subset of nine species. These results suggest that in species using torpor, selective forces may have acted to maximize the efficiency of thermogenic capacity (VO(2)-sum) relative to maintenance costs (BMR), resulting in an increasing FAS with decreasing T(a).

Phenotypic flexibility in body mass, basal metabolic rate and summit metabolism in southern red bishops (Euplectes orix): Responses to short term thermal acclimation

Avian basal metabolic rate (BMR) and summit metabolism (Msum) vary within individuals in response to seasonal acclimatization in free-ranging birds, and thermal acclimation under laboratory conditions. We examined relationships between acclimation air temperature (Tacc) and body mass (Mb), BMR and Msum in female southern red bishops (Euplectes orix) from a relatively mild coastal site and a seasonally colder, inland site. Following acclimation for 21days to Tacc=10, 22 or 35°C, Mb, BMR and Msum were all significantly and negatively related to Tacc. The significant relationship between BMR and Tacc did not remain after Mb was included as a covariate, whereas that between Msum and Tacc did. A subsequent reverse acclimation protocol, where bishops were acclimated to a second Tacc value and then re-acclimated to the first, revealed that short-term changes in Mb and BMR were completely reversible, but changes in Msum were only partially reversible. Following the reverse-acclimation protocol, metabolic expansibility (Msum/BMR) varied significantly with air temperature, being greatest at Tacc=35°C. Our data suggest that the intraspecific variation in seasonal metabolic variation previously reported for this species is at least partly driven by factors other than temperature.

Does low daily energy expenditure drive low metabolic capacity in the tropical robin, Turdus grayi?

Temperate and tropical birds possess divergent life history strategies. Physiological parameters including energy metabolism correlate with the life history such that tropical species with a slower 'pace of life' have lower resting and maximal metabolic rates than temperate congeners. To better understand the physiological mechanisms underlying these differences, we investigated the relationship of metabolic capacity, muscle oxidative capacity and activity patterns to variation in life history patterns in American robins (Turdus migratorius), while resident in central North America and Clay-colored robins (Turdus grayi) resident in Panama. We measured summit metabolism [Formula: see text] in birds from both tropical and temperate habitats and found that the temperate robins have a 60% higher metabolic capacity. We also measured the field metabolic rate (FMR) of free-living birds using heart rate (HR) telemetry and found that temperate robins' daily energy expenditure was also 60% higher. Thus, [Formula: see text] and FMR both reflect life history differences between the species. Further, both species operate at a nearly identical ~50% of their thermogenic capacity throughout a given day. As a potential mechanism to explain differences in activity and metabolic capacity, we ask whether oxidative properties of flight muscle are altered in accordance with life history variation and found minimal differences in oxidative capacity of skeletal muscle. These data demonstrate a close relationship between thermogenic capacity and daily activity in free-living birds. Further, they suggest that the slow pace of life in tropical birds may be related to the maintenance of low activity rather than functional capacity of the muscle tissue.

Summit metabolism and metabolic expansibility in Wahlberg's epauletted fruit bats (Epomophorus wahlbergi): seasonal acclimatisation and effects of captivity

Summit metabolism (M sum), the maximum rate of resting metabolic thermogenesis, has been found to be broadly correlated with climatic variables and the use of heterothermy in some endotherms. Far less is known about M sum and metabolic expansibility [ME, the ratio of M sum to basal metabolic rate (BMR)] in bats compared with many other endotherm taxa. We measured BMR and M sum during winter and summer in captive and wild populations of a pteropodid from the southern subtropics, Wahlberg's epauletted fruit bat (Epomophorus wahlbergi) in Pretoria, South Africa. The M sum of fruit bats ranged from 5.178 ± 0.611 W (captive, summer) to 6.006 ± 0.890 W (captive, winter), and did not vary significantly between seasons. In contrast, BMR decreased by 17-25% in winter. The combination of seasonally stable M sum but flexible BMR resulted in ME being significantly higher in winter than in summer, ranging from 7.24 ± 1.49 (wild, summer) to 13.11 ± 2.14 (captive, winter). The latter value is well above the typical mammalian range. Moreover, both M sum and ME were significantly higher in captive bats than in wild individuals; we speculate this represents a phenotypic response to a reduction in exercise-associated heat production while in captivity. Our data for E. wahlbergi, combined with those currently available for other chiropterans, reveal that M sum in bats is highly variable compared with allometrically expected values for other mammals.

Seasonal Metabolic Variation in Two Populations of an Afrotropical Euplectid Bird

Many birds exhibit seasonal phenotypic flexibility in basal metabolic rate (BMR) and summit metabolism (M(sum)), but very little information is available for species from subtropical latitudes or for conspecific populations from sites that vary in climate. We measured body mass (M(b)), BMR, and M(sum) in summer and winter in two populations of the southern red bishop Euplectes orix, a passerine that is widespread in southern Africa. One population occurs at a comparatively warmer coastal site (mean daily minimum air temperature [T(a)] in midwinter, 8.3°C) and the other at a colder inland site (mean daily minimum T(a) in midwinter, -2.8°C). Bishops from both populations significantly increased M(b) in winter. However, seasonal metabolic adjustments differed considerably between the populations. The inland population significantly increased BMR by approximately 58%, mass-specific BMR by approximately 31%, and M(sum) by approximately 15% in winter, although mass-specific M(sum) did not change significantly. In contrast, the coastal population showed no significant seasonal change in BMR and significant winter reductions in mass-specific BMR (~15%), M(sum) (~8%), and mass-specific M(sum) (~15%). The interpopulation differences in the magnitude and direction of seasonal mass-specific BMR changes are qualitatively consistent with global patterns, although the increase shown by the inland population is larger than expected. Our data reveal that avian seasonal metabolic adjustments can vary greatly within subtropical species depending on the climatic conditions experienced by the birds, and our findings reiterate the need to better understand metabolic flexibility in species that inhabit lower latitudes.

Seasonality of peak metabolic rate in non‐migrant tropical birds

Birds that are year‐round residents of temperate and tropical regions have divergent life histories. Tropical birds have a slower ‘pace of life’, one characteristic of which includes lower peak metabolic rate and daily activity levels. Temperate resident birds are faced with seasonal variation in thermogenic demand. This challenge is met with seasonally increased peak metabolic rate during winter. These thermogenic demands are much lower in birds that are year‐round tropical residents. By measuring peak (summit) metabolic rate in tropical and temperate resident bird species during summer and winter, we asked whether tropical birds exhibit seasonality in peak metabolic rate, and if the direction of seasonality differs between tropical and temperate species. We measured summit metabolism in seven tropical and one temperate species during the winter and during the summer breeding season to test the hypothesis that summit metabolism of tropical residents would change seasonally. We consider whether metabolic seasonality is associated with breeding season for tropical species. We found that summit metabolism was significantly greater during the summer for most tropical residents, while the temperate resident matched several previous reports with higher summit metabolism in winter. We conclude that metabolic seasonality occurs in tropical residents and differs from temperate residents, suggesting that breeding during the summer may be driving relatively higher metabolism as compared to winter thermogenesis in temperate birds.

Acclimatization of seasonal energetics in northern cardinals (Cardinalis cardinalis) through plasticity of metabolic rates and ceilings

Northern cardinals (Cardinalis cardinalis) are faced with energetically expensive seasonal challenges that must be met to ensure survival, including thermoregulation in winter and reproductive activities in summer. Contrary to predictions of life history theory that suggest breeding metabolic rate should be the apex of energetic effort, winter metabolism exceeds that during breeding in several temperate resident bird species. By examining whole-animal, tissue and cellular function, we ask whether seasonal acclimatization is accomplished by coordinated phenotypic plasticity of metabolic systems. We measured summit metabolism (V(O(2),sum)), daily energy expenditure (DEE) and muscle oxidative capacity under both winter (December to January) and breeding (May to June) conditions. We hypothesize that: (1) rates of energy utilization will be highest in the winter, contrary to predictions based on life history theory, and (2) acclimatization of metabolism will occur at multiple levels of organization such that birds operate with a similar metabolic ceiling during different seasons. We measured field metabolic rates using heart rate telemetry and report the first daily patterns in avian field metabolic rate. Patterns of daily energy use differed seasonally, primarily as birds maintain high metabolic rates throughout the winter daylight hours. We found that DEE and V(O(2),sum) were significantly greater and DEE occurred at a higher fraction of maximum metabolic capacity during winter, indicating an elevation of the metabolic ceiling. Surprisingly, there were no significant differences in mass or oxidative capacity of skeletal muscle. These data, highlighting the importance of examining energetic responses to seasonal challenges at multiple levels, clearly reject life history predictions that breeding is the primary energetic challenge for temperate zone residents. Further, they indicate that metabolic ceilings are seasonally flexible as metabolic effort during winter thermoregulation exceeds that of breeding.

Chronic cold acclimation increases thermogenic capacity, non-shivering thermogenesis and muscle citrate synthase activity in both wild-type and brown adipose tissue deficient mice

The purpose of this study was to determine whether chronic cold exposure would increase the aerobic capacity of skeletal muscle in UCP-dta mice, a transgenic line lacking brown adipose tissue (BAT). Wild type and UCP-dta mice were acclimated to either warm (23°C), or cold (4°C) conditions. Cold increased muscle oxidative capacity nearly equivalently in wild-type and UCP-dta mice, but did not affect the respiratory function of isolated mitochondria. Summit metabolism (V˙O2summit) and norepinephrine-induced thermogenesis (V˙O2NST) were significantly lower in UCP-dta mice relative to wild-type mice regardless of temperature treatment, but both were significantly higher in cold relative to warm acclimated mice. BAT mass was significantly higher in the cold relative to warm acclimated wild-type mice, but not in cold acclimated UCP-dta mice. BAT citrate synthase activity was lower in transgenic animals regardless of acclimation temperature and BAT citrate synthase activity per depot was significantly higher only in the cold acclimated wild-type mice. Muscle citrate synthase activity was increased in both genotypes. As defects in muscle oxidative function have been observed with obesity and type 2 diabetes, these results suggest that chronic cold exposure is a useful intervention to drive skeletal muscle oxidative capacity in mouse models of obesity.