Ontogenetic body-mass scaling of resting metabolic rate covaries with species-specific metabolic level and body size in spiders and snakes

According to common belief, metabolic rate usually scales with body mass to the 3/4-power, which is considered by some to be a universal law of nature. However, substantial variation in the metabolic scaling exponent (b) exists, much of which can be related to the overall metabolic level (L) of various taxonomic groups of organisms, as predicted by the recently proposed metabolic-level boundaries (MLB) hypothesis. Here the MLB hypothesis was tested using data for intraspecific (ontogenetic) body-mass scaling of resting metabolic rate in spiders and boid snakes. As predicted, in both animal groups b varies mostly between 2/3 and 1, and is significantly negatively related to L. L is, in turn, negatively related to species-specific body mass (M_m: estimated as the mass at the midpoint of a scaling relationship), and as a result, larger species tend to have steeper metabolic scaling slopes (b) than smaller species. After adjusting for the effects of M_m, b and L are still negatively related, though significantly only in the spiders, which exhibit a much wider range of L than the snakes. Therefore, in spiders and snakes the intraspecific scaling of metabolic rate with body mass itself scales with interspecific variation in both metabolic level and body mass.     

Body temperature and metabolic rate during natural hypothermia in endotherms

During daily torpor and hibernation metabolic rate is reduced to a fraction of the euthermic metabolic rate. This reduction is commonly explained by temperature effects on biochemical reactions, as described by Q ₁₀ effects or Arrhenius plots. This study shows that the degree of metabolic suppression during hypothermia can alternatively be explained by active downregulation of metabolic rate and thermoregulatory control of heat production. Heat regulation is fully adequate to predict changes in metabolic rate, and Q ₁₀ effects are not required to explain the reduction of energy requirements during hibernation and torpor.Abbreviations BMR basal metabolic rate - BW body weight - C thermal conductance - C_HL thermal conductance as derived from HL - C_HP thermal conductance as derived from HP - HL heat loss - HP heat production - MR metabolic rate - RQ respiratory quotient - T_a ambient temperature - T_b body temperature     

The effect of temperature on the pattern of torpor in a marsupial hibernator

Physiological variables of torpor are strongly temperature dependent in placental hibernators. This study investigated how changes in air temperature affect the duration of torpor bouts, metabolic rate, body temperature and weight loss of the marsupial hibernator Burramys parvus (50 g) in comparison to a control group held at a constant air temperature of 2°C. The duration of torpor bouts was longest (14.0±1.0 days) and metabolic rate was lowest (0.033±0.001 ml O₂·g^-1·h^-1) at2°C. At higher air temperatures torpor bouts were significantly shorter and the metabolic rate was higher. When air temperature was reduced to 0°C, torpor bouts also shortened to 6.4±2.9 days, metabolic rate increased to about eight-fold the values at 2°C, and body temperature was maintained at the regulated minimum of 2.1±0.2°C. Because air temperature had such a strong effect on hibernation, and in particular energy expenditure, a change in climate would most likely increase winter mortality of this endangered species.Abbreviationst STP standard temperature and pressure - T _a air temperature - T _b body temperature - VO₂ rate of oxygen consumption     

Activity affects intraspecific body-size scaling of metabolic rate in ectothermic animals

Metabolic rate is commonly thought to scale with body mass (M) to the 3/4 power. However, the metabolic scaling exponent (b) may vary with activity state, as has been shown chiefly for interspecific relationships. Here I use a meta-analysis of literature data to test whether b changes with activity level within species of ectothermic animals. Data for 19 species show that b is usually higher during active exercise (mean ± 95% confidence limits = 0.918 ± 0.038) than during rest (0.768 ± 0.069). This significant upward shift in b to near 1 is consistent with the metabolic level boundaries hypothesis, which predicts that maximal metabolic rate during exercise should be chiefly influenced by volume-related muscular power production (scaling as M ¹). This dependence of b on activity level does not appear to be a simple temperature effect because body temperature in ectotherms changes very little during exercise.     

A bioenergetic study of a benthic nematode,Plectus palustris de man 1880, throughout its life cycle

Summary Respiration rates of the bacterivorous freshwater nematode Plectus palustris were measured during the whole life cycle of the species and for animals grown at two food densities. Covariance analysis showed that small, but significant differences exist in the elevation of the respiration rate—body weight regressions (R=aW ^b, in nl O₂/ind·h and g wet weight) for different food densities. At a food density of 6–9·10⁸ bacterial cells/ml the level of respiration is 14% lower compared to rates of animals cultured at a ten times higher food density. However, the allometric function, R-aW ^b, adequately describes the relationship of respiration and body weight only during the larval growth phase and for young females, while respiration rates of newly hatched larvae and mature females at maximal egg production have lower metabolic rates. Cumulated metabolic costs to attain a certain age, size and stage of development have been determined and are used in a subsequent paper (Schiemer et al., 1979) to calculate the energy budgets of the species.     

Effects of temperature and body size on radiocaesium retention in brown trout, Salmo trutta

1. The elimination rate of radiocaesium in brown trout Salmo trutta L. was determined in the laboratory at four water temperatures (range 4.4–15.6°C). In the experiments three or four homogenous size-groups of fish (mean weights 23–496 g) were studied at each temperature. 2. The brown trout received acute oral doses of ¹³⁴Cs and were killed at intervals for radioactivity counting. The retention versus time curves were composed of two distinct exponential components. The long-lived component was quantitatively the most important for retention of radiocaesium. Elimination rate increased with increasing water temperature and decreased with increasing body weight. 3. The biological half-life of ¹³⁴Cs (T_b, days) was related to fresh body weight (W, g) and water temperature (t, °C) by the equation: T_b= 290 ×W°.¹⁷⁶× e-°.^106×t. The elimination rate of Cs could be predicted from weight-specific metabolic rate as given by Elliott's equations for brown trout.     

Summer acclimatization in the short-tailed field vole,Microtus agrestis

We investigated the changes that occurred in basal and noradrenaline-induced metabolic rate, body temperature and body mass in short-tailed field voles,Microtus agrestis, during exposure to naturally increasing photoperiod and ambient temperature. These parameters were first measured in winter-acclimatized voles (n=8) and then in the same voles which had been allowed to seasonally acclimatize to photoperiod and ambient temperature (6 months later). Noradrenaline induced metabolic rate, basal metabolic rate and nonshivering thermogenesis were significantly higher in winter-acclimatized compared to summer-acclimatized voles. There was a significant positive relationship between basal metabolic rate and noradrenaline-induced metabolic rate. Body mass was significantly higher in summer-acclimatized compared to winter-acclimatized voles. There was a significant positive relationship between body mass and noradrenaline-induced metabolic rate in both winter-acclimalized and summer-acclimatized voles; however, there was no relationship between basal metabolic rate and body mass in either seasonal group of voles. Body temperature after measurements of basal metabolic rate was not significantly different in the seasonal cohorts of voles. However, body temperature was significantly higher in winter-acclimatized compared to summer-acclimatized voles after injection of noradrenaline. Previously we have found that a long photoperiod was not a sufficient stimulus to reduce thermogenic capacity in winter-acclimatized voles during cold exposure, since basal metabolic rate increased to compensate for a reduction in regulatory nonshivering thermogenesis. Here we found that a combination of increased ambient temperature and photoperiod did significantly reduce thermogenic capacity in winter-acclimatized voles. This provided evidence that the two aspects of non-shivering thermogenesis, obligatory and regulatory, are stimulated by different exogenous cues. Summer acclimatization in the shorttailed field vole is manifest as a significant decrease in both basal and noradrenaline-induced metabolic rate, combined with a significant increase in body mass.Abbreviations ANCOV A analysis of covariance - BAT brown adipose tissue - BM body mass - BMR basal metabolic rate - NST non-shivering thermogenesis - NA noradrenaline - V the maximum V recorded following mass specific injection of noradrenaline - V the maximum V recorded following mass specific injection of saline - T _a ambient temperature - T _b rectal body temperature - T _1c lower critical temperature - UCP uncoupling protein - V oxygen consumption     

Preliminary studies on the respiratory energy loss of a spider,Lycosa pseudoannulata

To elucidate the basic food requirement of spiders, the important polyphagous predators of rice-plant insect pests, an attempt was made to measure the respiratory energy loss of fasting spiders, Lycosa pseudoannulata. Relationship between fresh (y) and dry (x) weights of spiders inhabiting the bottom layer of the rice-plant community was represented by the following allometric equation:y=0.428x^0.872. The carbon dioxide production by previously fed and unfed females under the dark at 29°C 100% R. H. was measured by a titration technique. The relationship between fresh body weight and CO₂ production by unfed animals could be represented by the equation M=aW^b, M being the CO₂ output per individual per day and W the fresh body weight. The constant b, which determines the slope of curve, was 0.808. Respiration of the adult female with 100 mg fresh weight was 1.155±0.250 mg CO₂/100 g fresh weight/day or 48.69 mg CO₂/g dry weight/day. This value corresponds to 35.81 cal/g fresh weight/day or 150.94 cal/g dry weight/day. Supposing the calorific content of spiders to be 5820 cal/g dry weight, rate of the respiratory energy loss to total energy of the body was estimated to be 2.60%. This rate did not strongly contradict with the loss of fresh body weight before and after the measurement. The metabolic rate showed remarkable fluctuation with changing food supply. The CO₂ production of starved individuals decreased to 83.63±16.34% as compared with individuals which were fed before the measurement.     

Effects of temperature,swimming speed and body mass on standard and active metabolic rate in vendace (<Emphasis Type="Italic">Coregonus albula</Emphasis>)

This study gives an integrated analysis of the effects of temperature, swimming speed and body mass on standard metabolism and aerobic swimming performance in vendace (Coregonus albula (L.)). The metabolic rate was investigated at 4, 8 and 15°C using one flow-through respirometer and two intermittent-flow swim tunnels. We found that the standard metabolic rate (SMR), which increased significantly with temperature, accounted for up to 2/3 of the total swimming costs at optimum speed (U _opt), although mean U _opt was high, ranging from 2.0 to 2.8 body lengths per second. Net swimming costs increased with swimming speed, but showed no clear trend with temperature. The influence of body mass on the metabolic rate varied with temperature and activity level resulting in scaling exponents (b) of 0.71–0.94. A multivariate regression analysis was performed to integrate the effects of temperature, speed and mass (AMR = 0.82M ^0.93 exp(0.07T) + 0.43M ^0.93 U ^2.03). The regression analysis showed that temperature affects standard but not net active metabolic costs in this species. Further, we conclude that a low speed exponent, high optimum speeds and high ratios of standard to activity costs suggest a remarkably efficient swimming performance in vendace.     

Cube law, condition factor and weight–length relationships: history, meta-analysis and recommendations

This study presents a historical review, a meta‐analysis, and recommendations for users about weight–length relationships, condition factors and relative weight equations. The historical review traces the developments of the respective concepts. The meta‐analysis explores 3929 weight–length relationships of the type W = aL^b for 1773 species of fishes. It shows that 82% of the variance in a plot of log a over b can be explained by allometric versus isometric growth patterns and by different body shapes of the respective species. Across species median b = 3.03 is significantly larger than 3.0, thus indicating a tendency towards slightly positive‐allometric growth (increase in relative body thickness or plumpness) in most fishes. The expected range of 2.5 < b < 3.5 is confirmed. Mean estimates of b outside this range are often based on only one or two weight–length relationships per species. However, true cases of strong allometric growth do exist and three examples are given. Within species, a plot of log a vs b can be used to detect outliers in weight–length relationships. An equation to calculate mean condition factors from weight–length relationships is given as K_mean = 100aL^b?3. Relative weight W_rm = 100W/(a_mL^bm) can be used for comparing the condition of individuals across populations, where a_m is the geometric mean of a and b_m is the mean of b across all available weight–length relationships for a given species. Twelve recommendations for proper use and presentation of weight–length relationships, condition factors and relative weight are given.     

The roles of energetics,water economy,foraging behavior,and geothermal refugia in the distribution of the bat,Macrotus californicus

Summary Energy metabolism, thermoregulation, and water flux ofMacrotus californicus, the most northerly representative of the Phyllostomidae, were studied in the laboratory using standard methods, and energy metabolism and water fluxes were studied in the field using the doubly labelled water method together with a time budget. Daily energy expenditures of free-living bats averaged 22.8 kJ during the winter study period. Approximately 60% of this was allocated to resting metabolism costs while in the primary roosts (22 h/day).Macrotus californicus is unable to use torpor. The thermoneutral zone (TNZ) in this species is narrow (33 to 40 °C) and metabolic rate increased rapidly as ambient temperature decreased below the TNZ. Basal metabolic rate was 1.25 ml O₂/g·h, or 24 J/g·h. Total thermal conductance below the TNZ. was 1.8 mW/g·°C, similar to values measured for other bats. Evaporative water loss showed a hyperbolic increase with increasing ambient temperature, and was approximately 1% of total body mass/h in the TNZ. The success of these bats as year-round residents in deserts in the southwestern United States is probably not due to special physiological adaptations, but to roosting and foraging behavior. They use geothermally-heated winter roost sites (stable year-round temperatures of approximately 29 °C) which minimize energy expenditures, and they have an energetically frugal pattern of foraging that relies on visual prey location. These seem to be the two major factors which have allowedM. californicus to invade the temperate zone.Abbreviations BMR basal metabolic rate - FMR field metabolic rate - T _a ambient temperature - T _b body temperature - T _lc,T _uc lower and upper critical temperature, respectively - TBW total body water - TNZ thermoneutral zone     

Metabolism,thermogenesis and daily rhythm of body temperature in the wood lemming,Myopus schisticolor

Wood lemmings (Myopus schisticolor) were captured during their autumnal migration in September and October. The animals were maintained at 12°C and under 12L:12D photoperiod. Basal metabolic rate and thermogenic capacity of the wood lemming were studied. Basal metabolic rate was 3.54 ml O₂·g^-1·h^-1, which is 215–238% of the expected value. The high basal metabolic rate seems to be typical of rodents living in high latitudes. The body temperature of the wood lemming was high (38.0–38.8°C), and did not fluctuate much during the 24-h recording. The high basal metabolic rate and the high body temperature are discussed with regard to behavioural adaptation to a low-quality winter diet. Thermogenic capacity, thermal insulation and non-shivering thermogenesis of the wood lemming displayed higher values than expected: 53.0 mW·g^-1, 0.53 mW·g^-1·C^-1 and 53.2 mW·g^-1, respectively. Brown adipose tissue showed typical thermogenic properties, although its respiratory property was fairly low, but mitochondrial protein content was high compared to other small mammals. The 24-h recording of body temperature and motor activity did not reveal whether the wood lemming is a nocturnal animal. Possibly, the expression of a circadian rhythm was masked by peculiar feeding behaviour. It is concluded that the wood lemming is well adapted to living in cold-temperature climates.Abbreviations BAT brown adipose tissue; bm, body mass - BMR basal metabolic rate - C conductance - Cox cytochrome-c-oxidase - HP heat production - HP_max maximum heat production - M metabolism - NA noradrenaline - NST non-shivering thermogenesis - NST_max maximum non-shivering thermogenesis - RMR resting metabolic rate - RQ respiratory quotient - T _a anibient temperature - T _b body temperature - T _lc lower critical temperature - UCP uncoupling protein - vO₂ oxygen consumption - vO_{2 max} maximum oxygen consumption     

Effect of constant and fluctuating temperatures on resting and active oxygen consumption of toads,Bufo boreas

Summary The relations of standard and active rates of oxygen consumption to body temperature (T_b) were tested in montane Bufo b. boreas and lowland Bufo boreas halophilus acclimated to constant T _b of 10, 20, or 30° C or to a fluctuating cycle of 5–30° C. Standard metabolic rates (SMR) of boreas acclimated to 30° C and halophilus acclimated to 10° C show pronounced regions of thermal independence but all other standard and active metabolic rates of groups acclimated to other thermal regimes are thermally sensitive. The SMR of both subspecies acclimated to the 5–30° C cycle are more thermally sensitive than those of similar individuals acclimated to constant T _b. In cases where the relation between SMR and T _b is linear for both halophilus and boreas at the same acclimation temperature, the slope and Q₁₀ of the relation for boreas are significantly higher than those of halophilus. Acclimation had little or no effect on the active metabolic rates of either subspecies. The relation between SMR and T _b of boreas maintained under field conditions (Carey, 1979) is matched only by those of individuals from the same population acclimated to 20° C.     

Gene or Size: Metabolic Rate and Body Temperature in Obese Growth Hormone‐Deficient Dwarf Mice

Objective: SMA1 mice carry a missense mutation in the growth hormone gene that leads to semidominant dwarfism and obesity. In this study, the basic thermal and metabolic properties of SMA1 mice were examined to detect metabolic alterations that can support the accretion of excess fat. Research Methods and Procedures: Basal and resting metabolic rates (RMRs) in wild‐type and SMA1 (sma1/+ and sma1/sma1) mice were determined by indirect calorimetry. Body temperature (T_b) was recorded using intraperitoneally implanted temperature‐sensitive transmitters, and body composition was determined by DXA. Results: SMA1 mice have proportionally lower basal and resting metabolic rates, higher body mass (BM)‐specific RMRs, and a higher lower critical temperature, and display a decrease in T_b by 0.4 °C in sma1/+ and 0.9 °C in sma1/sma1. Discussion: The analysis of gene effects on BM and energy expenditure in mouse mutants must consider the appropriate allometric relationship between BM and metabolic rate. With the exception of T_b, all metabolic alterations observed in SMA1 reflect reduced size.     

Warm-up rates during arousal from torpor in heterothermic mammals: physiological correlates and a comparison with heterothermic insects

Summary This study examines the relationship between warm-up rate, body mass, metabolic rate, thermal conductance and normothermic body temperature in heterothermic mammals during arousal from torpor. Predictions based on the assumption that the energetic cost of arousal has been minimised are tested using data for 35 species. The observation that across-species warm-up rate correlates negatively with body mass is confirmed using a comparative technique which removes confounding effects due to the non-independence of species data due to shared common ancestry. Mean warm-up rate during arousal correlates negatively with basal metabolic rate and positively with the temperature difference through which the animal warms, having controlled for other factors. These results suggest that selection has operated to minimise the overall energetic, cost of warm-up. In contrast, peak warm-up rate during arousal correlates positively with peak metabolic rate during arousal, and negatively with thermal conductance, when body mass has been taken into account. These results suggest that peak warm-up rate is more sensitive to the fundamental processes of heat generation and loss. Although heterothermic marsupials have lower normothermic body temperatures and basal metabolic rates, marsupials and heterothermic eutherian mammals do not differ systematically in warm-up rate. Pre-flight warm-up rates in one group of endothermic insects, the bees, are significantly higher than predictions based on rates of arousal of a mammal of the same body mass.Abbreviations BMR basal metabolic rate - ICM independent comparisons method - MWR mean warm-up rate - PMR peak metabolic rate - PWR peak·warm-up rate - Tb_activity body temperature during activity - Tb_torpor body temperature during torpor - T _arousal increase in body temperature during arousal     

Maximum ingested food size in captive strepsirrhine primates: Scaling and the effects of diet

Little is known about ingested food size (V_b) in primates, even though this variable has potentially important effects on food intake and processing. This study provides the first data on V_b in strepsirrhine primates using a captive sample of 17 species. These data can be used for generating and testing models of feeding energetics. Strepsirrhines are of interest because they are hypometabolic and chewing rate and daily feeding time do not show a significant scaling relationship with body size. Using melon, carrot, and sweet potato we found that maximum V_b scales isometrically with body mass and mandible length. Low dietary quality in larger strepsirrhines might explain why V_b increases with body size at a greater rate than does resting metabolic rate. Relative to body size, V_b is large in frugivores but small in folivores; furthermore scaling slopes are higher in frugivores than in folivores. A gross estimate of dietary quality explains much of the variation in V_b that is not explained by body size. Gape adaptations might favor habitually large bites for frugivores and small ones for folivores. More data are required for several feeding variables and for wild populations. Am J Phys Anthropol 142:625–635, 2010. © 2010 Wiley‐Liss, Inc.     

Reduction of metabolism during hibernation and daily torpor in mammals and birds: temperature effect or physiological inhibition?

Summary The present study addresses the controversy of whether the reduction in energy metabolism during torpor in endotherms is strictly a physical effect of temperature (Q₁₀) or whether it involves an additional metabolic inhibition. Basal metabolic rates (BMR; measured as oxygen consumption, ), metabolic rates during torpor, and the corresponding body temperatures (T _b) in 68 mammalian and avian species were assembled from the literature (n=58) or determined in the present study (n=10). The Q₁₀ for change in between normothermia and torpor decreased from a mean of 4.1 to 2.8 with decreasingT _b from 30 to <10°C in hibernators (species that show prolonged torpor). In daily heterotherms (species that show shallow, daily torpor) the Q₁₀ remained at a constant value of 2.2 asT _b decreased. In hibernators with aT _b<10°C, the Q₁₀ was inversely related to body mass. The increase of mass-specific metabolic rate with decreasing body mass, observed during normothermia (BMR), was not observed during torpor in hibernators and the slope relating metabolic rate and mass was almost zero. In daily heterotherms, which had a smaller Q₁₀ than the hibernators, no inverse relationship between the Q₁₀ and body mass was observed, and consequently the metabolic rate during torpor at the sameT _b was greater than that of hibernators. These findings show that the reduction in metabolism during torpor of daily heterotherms and large hibernators can be explained largely by temperature effects, whereas a metabolic inhibition in addition to temperature effects may be used by small hibernators to reduce energy expenditure during torpor.Abbreviation BMR basal metabolic rate     

Physiological ecology of frillneck lizards in a seasonal tropical environment

The frillneck lizard, Chlamydosaurus kingii, is a conspicuous component of the fauna of the wetdry tropics of northern Australia during the wet season, but it is rarely seen in the dry season. Previous studies have demonstrated that during the dry season the field metabolic rate (FMR) is only about one-quarter of the wet-season rate, and one factor involved in this seasonal drop is a change in the behavioural thermoregulation of the species such that lower body temperatures (T _bs) are selected during dry-season days. Here we examine other factors that could be responsible for the seasonal change in FMR: standard metabolic rates (SMR) and activity. Samples from stomach flushing revealed that the lizards in the dry season continued to feed, but the volume of food was half as much as in the wet season. SMR in the laboratory was 30% less in the dry season. During the dry season, the energy expended by the lizards is 60.4 kJ kg^-1 day^-1 less than during the wet season. Combining laboratory and field data, we determined the relative contribution of the factors involved in this energy savings: 10% can be attributed to lower nighttime T _b, 12% is attributable to lower daytime T _b, 12% is attributable to decreased metabolism, and the remaining 66% is attributable to other activities (including e.g. locomotion, reproductive costs, digestion). Calculations indicate that if FMR did not drop in the dry season the lizards would not survive on the observed food intake during this season. Seasonal analysis of blood plasma and urine indicated an accumulation of some electrolytes during the dry season suggesting modest levels of water stress.     

Metabolic and cardiovascular adaptations in the western chipmunks, genusEutamias

Summary The metabolic and cardiac responses to temperature were studied in two species (four subspecies) of western chipmunks (genusEutamias), inhabiting boreal and alpine environments. A specially designed (Fig. 1) implantable biopential radiotransmitter was used to measure heart rate in unrestrained animals. The estimated basal metabolic rates (EBMR) were 1.78 (E. minimus borealis), 1.64 (E. m. oreocetes), 1.50 (E. m. operarius), and 1.69 ml O₂ g^–1 h^–1 (E. amoenus luteiventris), or 839, 752, 698, and 628 ml O₂ kg^–0.75 h^–1, respectively, for the four subspecies (Table 1). The two alpine species (E.m.or. andE.m.op.) had significantly lower EBMR than both of their boreal counterparts. The EBMR from all animals are 120–135% of the predicted values based on body weights of the animals. The thermal neutral zone for the four subspecies ranged from 23.5 to 32°C and the minimum thermal conductances were 0.113, 0.111, 0.112 and 0.112 ml O₂ g^–1 h^–1 °C^–1, respectively, or 54.4, 54.0, 50.4 and 52.1 ml O₂ kg^–0.75 h^–1 °C^–1, respectively (Fig. 2). No interspecific diffence in conductance was observed. These values are 72 to 85% of their weight specific values. The body temperature ranged between 35.0 and 39.5°C and was usually maintained between 36 and 38°C in all subspecies between ambient temperatures of 3 and 32°C. The estimated basal heart rates were 273, 296, 273 and 264 beats/min, respectively, for the four subspecies, 49–55% of their predicted weight specific values. The resultant oxygen pulses (metabolic rate/heart rate) were 5.49, 4.50, 4.48 and 5.56×10^–3 ml O₂/beat, respectively, which are 2 to 2.4 times their weight specific values (Table 2).The observed reduction of basal heart rate without the corresponding decreases of basal metabolic rate and body temperature indicate sufficient compensatory increases in stroke volume and/or A-V oxygen difference at rest. Such cardiovascular modifications provide extra reserves when demand for aerobic metabolism rises during bursts of activity typically observed in the western chipmunk.Abbreviations A-V arterio-venous - EBMR estimated basal metabolic rate (ml O₂ g^–1 h^–1) - HR heart rate (beats/min) - MR metabolic rate (ml O₂ g^–1 h^–1) - OP oxygen pulse (ml O₂/heart beat) - T_a, T_b ambient and body temperature (°C)     

Evidence of facultative daytime hypothermia in a small passerine wintering at northern latitudes

The use of hypothermia as a means to save energy is well documented in birds. This energy‐saving strategy is widely considered to occur exclusively at night in diurnally active species. However, recent studies suggest that facultative hypothermia may also occur during the day. Here, we document the use of daytime hypothermia in foraging Black‐capped Chickadees Poecile atricapillus wintering in eastern Canada. We measured the body temperature (T_b) of 126 individuals (plus 48 repeated measures) during a single winter and related values to ambient temperature (T_a) at the time of capture. We also tested whether daytime hypothermia was correlated with the size of body reserves (residuals of mass on structural size and fat score) and levels of metabolic performance (basal metabolic rate and maximum thermogenic capacity). We found that T_b of individual birds was lower when captured at low T_a, reaching values as low as 35.5 °C in actively foraging individuals. T_b was unrelated to metabolic performance or measures of body reserves. Therefore, daytime hypothermia does not result from individuals being unable to maintain T_b during cold spells or to a lack of body reserves. Our data also demonstrated a high level of individual variation in the depth of hypothermia, the causes of which remain to be explored.