Seasonal metabolic acclimatization in mountain chickadees and juniper titmice

Mountain chickadees and juniper titmice from northern Utah were examined to determine metabolic and body-composition characteristics associated with seasonal acclimatization. These species use behavioral adaptations and nocturnal hypothermia, which reduce energetic costs. These adjustments could reduce the need for extensive metabolic adjustments typically found in small passerines that overwinter in cold regions. In addition, these species live at higher altitudes, which may also decrease metabolic acclimatization found in birds. Winter birds tolerated colder test temperatures than summer birds. This improved cold tolerance was associated with an increase in maximal thermogenic capacity or summit metabolism (M(sum)). Winter M(sum) exceeded summer M(sum) by 26.1% in chickadees and 16.2% in titmice. Basal metabolic rates (BMR) were also significantly higher in winter birds compared with summer birds. Pectoralis wet muscle mass increased 33.3% in chickadees and 24.1% in titmice in winter and paralleled the increased M(sum) and BMR. Dry mass of contour plumage increased in winter for both species and was associated with decreased thermal conductance in winter chickadees compared to summer chickadees. Chickadees and titmice show metabolic acclimatization similar to other temperate species.     

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

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

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

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

Shorebirds' seasonal adjustments in thermogenic capacity are reflected by changes in body mass: how preprogrammed and instantaneous acclimation work together

Phenotypic flexibility in shorebirds has been studied mainly in the context of adjustments to migration and to quality of food; little is known on how birds adjust their phenotype to harsh winter conditions. We showed earlier that red knot (Calidris canutus islandica) can acclimate to cold by elevating body mass. This goes together with larger pectoral muscles, i.e., greater shivering machinery, and thus, better thermogenic capacity. Here, we present results of a yearlong experiment with indoor captive knots to determine whether this strategy is part of their natural seasonal phenotypic cycle. We maintained birds under three thermal regimes: constant cold (5 °C), constant thermoneutrality (25 °C) and natural seasonal variation between these extremes (9-22 °C). Each month we measured variables related to the birds' endurance to cold and physiological maintenance [body mass, thickness of pectoral muscles, summit metabolic rate (M(sum)), food intake, gizzard size, basal metabolic rate (BMR)]. Birds from all treatments expressed synchronized and comparable variation in body mass in spite of thermal treatments, with a 17-18% increase between the warmest and coldest months of the year; which appeared regulated by an endogenous driver. In addition, birds living in the cold exhibited a 10% higher average body mass than did those maintained at thermoneutrality. Thickness of the pectoral muscle tracked changes in body mass in all treatments and likely contributed to greater capacity for shivering in heavier birds. Consequently, M(sum) was 13% higher in cold-acclimated birds compared to those experiencing no thermoregulation costs. However, our data also suggest that part of maximal heat production comes from nonshivering processes. Birds facing cold conditions ate up to 25% more food than did birds under thermoneutral conditions, yet did not develop larger gizzards. Seasonal variation in BMR followed changes in body mass, probably reflecting changes in mass of metabolically active tissues. Just as cold-exposed birds, red knots in the variable treatment increased body mass in winter, thereby improving cold endurance. During summer, however, they maintained a lower body mass and thermogenic capacity compared to cold-exposed birds, similar to individuals kept at thermoneutrality. We conclude that red knots acclimate to seasonal variations in ambient temperature by modulating body mass, combining a preprogrammed increase in mass during winter with a capacity for fine-tuning body mass and thermogenic capacity to temperature variations.     

树麻雀代谢率和器官重量在季节驯化中表型的可塑性变化

动物能量代谢的生理生态特征与物种的分布和丰富度密切相关,基础代谢率(BMR)是内温动物能量预算的重要组成部分。北温带的小型鸟类,通过增加产热来适应低温环境。增加BMR的基础之一是中心器官(代谢机器)发生明显的变化。本研究中我们测定了树麻雀(Passermontanus)的BMR、体重和各器官的重量,分析了麻雀各器官的季节性变化及与BMR的关系。方差分析表明:麻雀的BMR存在明显的季节性变化,在冬季和秋季较高。麻雀内部器官的变化同样有明显的季节性,冬季和秋季麻雀的肝脏、心脏、肌胃、小肠、直肠和整体消化道的重量,都有明显的增加。相关分析表明:麻雀的BMR与肝脏、心脏和消化道等内部器官存在明显的相关性。我们的结果验证了“中心限制假说”,即麻雀体内存在着与BMR相关的“代谢机器”,中心器官是提高麻雀BMR的基础之一。     

Seasonal acclimatization of metabolism in Eurasian tree sparrows (Passer montanus)

Acclimatization to winter conditions is an essential prerequisite for survival of small passerines of the northern temperate zone. Changes in photoperiod, ambient temperature and food availability trigger seasonal acclimatization in physiology and behavior of many birds. In the present study, seasonal adjustments in several physiological, hormonal, and biochemical markers were examined in wild-captured Eurasian tree sparrows (Passer montanus) from the Heilongjiang Province in China. In winter sparrows had higher body mass and basal metabolic rate (BMR). Consistently, the dry mass of liver, heart, gizzard, small intestine, large intestine and total digestive tract were higher in winter than in that in summer. The contents of mitochondrial protein in liver, and state-4 respiration and cytochrome c oxidase (COX) activity in liver and muscle increased significantly in winter. Circulating level of serum triiodothyronine (T₃) was significantly higher in winter than in summer. Together, these data suggest that tree sparrows mainly coped with cold by enhancing thermogenic capacities through increased organ masses and heightened activity of respiratory enzymes activities. The results support the view that prominent winter increases in BMR are manifestations of winter acclimatization in tree sparrows and that seasonal variation in metabolism in sparrows is similar to that in other small temperate-wintering birds.     

Seasonal thermoregulation in the burrowing parrot (Cyanoliseus patagonus)

Birds exposed to seasonal environments are faced with the problem of maintaining thermogenic homoeostasis. Previous studies have established that birds native to the Holarctic increase their Resting Metabolic Rate at different ambient temperatures (RMR_Ta) and Basal Metabolic Rate (BMR) in winter as an adaptation to cold temperature since winters are more severe, while their non-Holarctic counterparts generally decrease their winter BMR as an energy saving mechanism during unproductive and dry winter months. In this study, we examined seasonal thermoregulation in the burrowing parrot (Cyanoliseus patagonus), a colonial psittacine native to the Patagonian region of Argentina, a region with an unpredictable environment. We found significantly higher mass specific RMR_Ta and BMR in summer than in winter. Both summer and winter BMR of the species fell within the predicted 95% confident interval for a parrot of its size. Body mass was significantly higher in winter than in summer. The burrowing parrot had broad thermo-neutral zones in winter and summer. The circadian rhythm of core body temperature (T_b) of burrowing parrots was not affected by season, showing that this species regulated its T_b irrespective of season. These results suggest that the burrowing parrots' seasonal thermoregulatory responses represent that of energy conservation which is important in an unpredictable environment.     

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

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

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

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

     

Seasonal variation in thermal energetics of the Australian owlet-nightjar (Aegotheles cristatus)

Many birds living in regions with seasonal fluctuations in ambient temperatures (T_a) typically respond to cold by increasing insulation and adjusting metabolic rate. Seasonal variation in thermal physiology has not been studied for the Caprimulgiformes, an order of birds that generally have basal metabolic rates (BMR) lower than predicted for their body mass. We measured the metabolic rate and thermal conductance of Australian owlet-nightjars (Aegotheles cristatus) during summer and winter using open-flow respirometry. Within the thermoneutral zone (TNZ; 31.3 to 34.8 °C), there was no seasonal difference in BMR or thermal conductance (C), but body temperature was higher in summer- (38.2 ± 0.3 °C) than winter-acclimatized (37.1 ± 0.5 °C) birds. Below the TNZ, resting metabolic rate (RMR) increased linearly with decreasing T_a, and RMR and C were higher for summer- than winter-acclimatized birds. The mean mass-specific BMR of owlet-nightjars (1.27 mL O₂ g^− 1 h^− 1) was close to the allometrically predicted value for a 45 g Caprimulgiformes, but well below that predicted for birds overall. These results suggest that owlet-nightjars increase plumage insulation to cope with low winter T_a, which is reflected in the seasonal difference in RMR and C below the TNZ, rather than adjusting BMR.     

Seasonal adjustments in body mass and thermogenesis in Mongolian gerbils (<Emphasis Type="Italic">Meriones unguiculatus</Emphasis>): the roles of short photoperiod and cold

Seasonal adjustments in body mass and thermogenesis are important for the survival of small mammals during acclimatization in the temperate zone. To determine the contributions of short photoperiod and cold temperatures to seasonal changes in thermogenesis and body mass in Mongolian gerbils (Meriones unguiculatus), body mass, basal metabolic rate (BMR), nonshivering thermogenesis (NST), energy intake and energy digestibility were determined in seasonally acclimatized and laboratory acclimated animals. Body mass showed significant seasonal changes and decreased to a minimum in winter. Both BMR and NST increased in winter, and these changes were mimicked by exposing animals to short photoperiod or cold temperatures in the animal house. Digestible energy intake also increased significantly in winter, and also during exposure of housed animals to both short photoperiod and cold. These results suggest that Mongolian gerbils overcome winter thermoregulatory challenges by increasing energy intake and thermogenesis, and decreasing body mass to reduce total energy requirements. Short photoperiod and cold can serve as effective environmental cues during seasonal acclimatization.     

Effects of helium/oxygen and temperature on aerobic metabolism in the marsupial sugar glider, Petaurus breviceps

Helox (79% helium and 21% oxygen) has often been used for thermobiological studies, primarily because helium is thought to be metabolically inert and to produce no adverse effects other than increasing heat loss. However, these assumptions have been questioned. As basal metabolic rate (BMR) represents maintenance energy requirements for vital body functions, potential physiological effects of helox should be reflected in changes of BMR. In this study, sugar gliders were subjected to both air and helox atmospheres over a wide range of T(a)'s, including the thermoneutral zone (TNZ), to determine (1) whether helox has any influence other than on heat loss and (2) the maximum heat production (HP(max)) and thermal limits of this species. Although thermal conductance in the TNZ increased in helox, BMR was similar in air and helox (0.55+/-0.07 and 0.57+/-0.06 mL g(-1) h(-1), respectively). The TNZ in helox, however, was shifted upwards by about 3 degrees C. Below the TNZ, sugar gliders were able to withstand an effective temperature of -24.7+/-7.3 degrees C with an HP(max) of 3.14+/-0.36 mL g(-1) h(-1). The low effective temperature tolerated by sugar gliders shows that they are competent thermoregulators despite their apparent lack of functional brown fat. Similarities of BMRs in air and helox suggest that the effect of helox is restricted to an increase of heat loss, and, consequently, helox represents a useful tool for thermal physiologists. Moreover, the lack of increase of BMR in helox despite an increase in thermal conductance of sugar gliders suggests that BMR is not a function of body surface.     

秋季迁徙中北美戴菊的寒冷耐受能力及最大代谢产热

由于金冠戴菊冬季的分布地与红玉冠戴菊相比更偏北,生活的环境更加寒冷,人们认为金冠戴菊对寒冷的耐受力更强。然而,有关红玉冠戴菊和金冠戴菊两者之间对寒冷的耐受性和最大产热能力的直接证据尚无报道。在美国南达科他秋季鸟类迁徙季节,作者采用冷暴露氦氧混合气体(79%氦和21%氧),对红玉冠戴菊和金冠戴菊的寒冷耐受能力和最大代谢率(最大冷诱导代谢,Msux)进行了测定。结果显示:金冠戴菊对低温的耐受能力高于红玉冠戴菊,由于金冠戴菊和红玉冠戴菊的体重和热传导差异不显著,表明对低温的耐受能力的差异不是体重和热传导引起的;而金冠戴菊的最大代谢产热(2.51ml±0.32mlO2min-1,n=11)明显高于雄性(2.27ml±0.25mlO2min-1,n=13)和雌性(2.05ml±0.18mlO2min-1,n=13)的红玉冠戴菊,表明最大代谢的差异可能导致对寒冷的耐受能力不同。相对于红玉冠戴菊,金冠戴菊有良好的耐寒冷能力,与其在冬季分布更北相一致,并与其高的代谢产热能力有关,此模式和许多雀形目鸟类在季节性驯化中增加对寒冷的耐受能力相一致。     

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

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

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

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

黑线仓鼠体重和能量代谢的季节性变化

为阐明小型哺乳动物体重和能量代谢的季节性变化以及生理调节机制,将黑线仓鼠驯化于自然环境下12个月,测定其体重、能量收支、身体组织器官和血清瘦素水平的季节性变化。黑线仓鼠能量摄入和支出的季节性变化显著,冬季摄入能、基础代谢率(BMR)、非颤抖性产热(NST)显著高于夏季。体重季节性变化不显著,但身体组织器官重量呈现显著的季节性变化,冬季肝脏、心脏、肾脏以及消化道重量显著高于夏季。体脂含量夏季最高,冬季最低,冬季显著低于夏、秋和春季(P <0.01)。血清瘦素水平的季节性变化显著,夏季瘦素水平比秋、冬季分别高88.2% 和52.4% (P <0.05)。结果表明,黑线仓鼠体重维持季节性稳定,与“调定点假说”的预测不同;但脂肪含量和血清瘦素季节性变化显著,符合该假说。夏季血清瘦素升高具有抑制能量摄入的作用,冬季血清瘦素可能是促进代谢产热的重要因子,瘦素对能量代谢和体重的调节作用与气候的季节性变化有关。       

Phenotypic flexibility of body composition associated with seasonal acclimatization in passerine birds

Improved winter cold tolerance is widespread among small birds overwintering in cold climates and is associated with improved shivering endurance and elevated summit metabolic rate (M_sum). Phenotypic flexibility resulting in elevated M_sum could result from either increased skeletal muscle mass (perhaps with support from similar adjustments in “nutritional organs”) and/or cellular metabolic intensity. We investigated seasonal changes in body composition of three species of passerine birds resident in cold winter climates, all of which show large seasonal variations in M_sum (>25%); white-breasted nuthatch (Sitta carolinensis), black-capped chickadee (Poecile atricapillus), and house sparrow (Passer domesticus). All three species displayed significant winter increases in pectoralis and heart masses, and supracoracoideus mass also increased in winter chickadees. Gizzard mass increased in winter for all three species, but masses of other nutritional organs did not vary consistently with season. These data suggest that winter increases in pectoralis and heart masses are important contributors to elevated thermogenic capacity and cold tolerance, but seasonal variation in nutritional organ masses, other than gizzard, which is likely associated with dietary changes, are not universally associated with seasonal phenotypes. The winter increases in pectoralis and heart masses are consistent with data from other small passerines showing marked seasonal changes in cold tolerance and support the Variable Maximum Model of seasonal phenotypic flexibility, where physiological adjustments that promote improved cold tolerance, also result in elevated M_sum.     

Thermogenic capabilities of the opossum Monodelphis domestica when warm and cold acclimated: similarities between American and Australian marsupials

1. Monodelphis domestica is a small marsupial mammal from South America. Its thermogenic abilities in the cold were determined when the opossums were both warm (WA) and cold (CA) acclimated. Maximum heat production of M. domestica was obtained at low temperatures in helium-oxygen. 2. Basal metabolic rate (BMR) in the WA animals was 3.2 W/kg and mean body temperature was 32.6 degrees C at 30 degrees C. These values were lower than those generally reported for marsupials. Nevertheless, these M. domestica showed considerable metabolic expansibility in response to cold. Sustained (summit) metabolism was 8-9 times BMR, while peak metabolism was 11-13 times BMR. These maximum values were equal to, or above, those expected in small placentals. 3. Cold acclimation altered the thermal responses of M. domestica, particularly in warm TaS. However, summit metabolism was not significantly increased; nor did M. domestica show a significant thermogenic response to noradrenaline, which in many small placentals elicits non-shivering thermogenesis. The thermoregulatory responses of this American marsupial were, in most aspects, similar to those of Australian marsupials. This suggests that the considerable thermoregulatory abilities of marsupials are of some antiquity.     

Seasonal variation in metabolic rate of a medium-sized frugivore, the Knysna Turaco (Tauraco corythaix)

Many seasonal thermoregulation studies have been conducted on Holarctic birds that live in predictable, highly seasonal climates with severe winters. However, relatively few studies have been conducted on their southern hemisphere Afrotropical counterparts that encounter less predictable climates with milder winters. These latter birds are expected to conserve energy in winter by downregulating their metabolic rates. Therefore in this study, metabolic rate was measured during summer and winter in Knysna Turaco, Tauraco corythaix (Musophagiformes, Musophagidae) (c. 310 g), a non-passerine, in order to test whether there is energy conservation in winter. No overall significant differences in resting metabolic rates over a range of ambient temperatures were observed between winter and summer. However, whole-organism basal metabolic rates were 18.5% higher (p=0.005) in winter than in summer (210.83±15.97 vs. 186.70±10.52 O₂ h⁻¹). Knysna Turacos had broad thermoneutral zones ranging from 20 to 28 °C in winter and 10 to 30 °C in summer. These results suggest that Knysna Turacos show seasonal thermoregulatory responses that represent cold defense rather than energy conservation, which is contrary to what was expected.     

Seasonal variations in basal metabolic rate, lower critical temperature and responses to temporary starvation in the arctic fox (Alopex lagopus) from Svalbard

Metabolic rates of four resting, post-absorptive male adult summer- and winter-adapted captive arctic foxes (Alopex lagopus) were recorded. Basal metabolic rates (BMR) varied seasonally with a 36% increase from winter to summer, while body mass was reduced by 17% in the same period. The lower critical temperature (T _1c) of the winter-adapted arctic fox was estimated to −7°C, whereas T _lc during summer was 5°C. The similarity of these values, which are much higher than hitherto assumed (e.g. Scholander et al. 1950b), is mainly due to a significantly (P<0.05) lower BMR in winter than in summer. Body core (stomach) temperature was stable, even at ambient temperatures as low as −45°C, but showed a significant (P<0.05) seasonal variation, being lower in winter (39.3±0.33°C) than in summer (39.8±0.16°C). The thermal conductivity of arctic fox fur was the same during both seasons, whereas the thermal conductance in winter was lower than in summer. This was reflected in an increase in fur thickness of 140% from summer to winter, and in a reduced metabolic response to ambient temperatures below T _lc in winter. Another four arctic foxes were exposed to three periods of forced starvation, each lasting 8 days during winter, when body mass is in decline. No significant reduction in mass specific BMR was observed during the exposure to starvation, and respiratory quotient was unchanged at 0.73±0.02 during the first 5 days, but dropped significantly (P<0.05) to 0.69±0.03 at day 7. Locomotor activity and body core (intraperitoneal) temperature was unaltered throughout the starvation period, but body mass was reduced by 18.5±2.1% during these periods. Upon re-feeding, locomotor activity was significantly (P<0.05) reduced for about 6 days. Energy intake was almost doubled, but stabilised at normal levels after 11 days. Body mass increased, but not to the level before the starvation episodes. Instead, body mass increased until it reached the reduced body mass of ad libitum fed control animals. This indicates that body mass in the arctic fox is regulated according to a seasonally changing set point.