Shivering thermogenesis in leg and breast muscles of galliform chicks and nestlings of the domestic pigeon.

We studied the ontogeny of shivering thermogenesis in breast and leg muscles of precocial galliforms (domestic fowl, grey partridge, and Japanese quail) and the altricial domestic pigeon using electromyography (EMG) and indirect calorimetry. Galliforms were able to increase heat production by shivering in leg muscles at the youngest age studied (1-2 d). Pectorals contributed to heat production from days 7-10 onward, but in the partridge and especially in the fowl, shivering by the pectorals was weaker than in the quail. In the pigeon, shivering began in pectorals and legs at 2 and 4 d of age, respectively, and pectorals had clearly the predominant role in thermogenesis. Despite the early beginning of electrical signs of shivering, significant thermogenesis did not appear in the pigeon before the age of 6 d. All galliforms shivered in bursts, like pigeons aged 2-4 d. From the age of 6 d onward, continuous shivering became predominant in the pigeon. In pectorals of 2-6-d-old pigeons, shivering did not increase linearly during decreasing ambient temperature, as in other muscles and species, but started abruptly, at full intensity. Furthermore, in 2-4-d-old pigeons, cooling induced movement activity in legs. The median frequency of shivering EMGs varied (1) with maturation of the muscle, (2) with size of the adult bird, and (3) between altricials and precocials.     

Effect of photoperiod and melatonin on cold resistance,thermoregulation and shivering/nonshivering thermogenesis in Japanese quail

Summary The effect of photoperiod and melatonin treatment on cold resistance and thermogenesis of quails was studied. The birds were acclimated for 8 weeks to short day (8L:16D) or long day (16L:8D) conditions, and 8 of 16 quails in each group were implanted with melatonin capsules. One group of quails was maintained outside in an aviary during winter. Oxygen consumption ( ) body temperature (T _b, recorded with temperature transmitters) and shivering (integrated pectoral EMG) were recorded continuously, and samples of heart rate and breathing rate were picked up when ambient temperature was decreased stepwise from 27 down to –75 °C. Heat production maximum (HP_max), cold limit, lower critical temperature, basal metabolic rate (BMR) and thermal conductance were determined.The results show that short day, cold and melatonin treatment improved cold resistance and thermal insulation of quils when compared with quails acclimated to long day conditions. An increase in HP_max was induced only by melatonin treatment. The results suggest that the acclimatization of quails is under control of the pineal gland.The linear increase of shivering intensity with at moderate cold load shows that shivering is the primary source for thermoregulatory heat production in the quail. AtT _a's below –40 °C shivering remained constant although , heart rate and breathing rate continued to increase with increasing cold load. This could indicate the existence of a nonshivering thermogenesis in birds. Unlike to mammals, this non-shivering thermogenesis in birds would serve as secondary source of heat supporting shivering thermogenesis in severe coldAbbreviations BMR basal metabolic rate - ECG electrocardiogram - EMG electromyogram - NST nonshivering thermogenesis - SMR standard metabolic rate     

Shivering thermogenesis in the neonatal pig

(I) Shivering intensity and metabolic rate were determined in Large White pigs aged 2, 24, 48 h and 5 d, at temperatures ranging from thermoneutrality (36°C) to cold (20°C). (2) Shivering is the main heat producing mechanism, the absence of nonshivering thermogenesis being implied by both the absence of delay between the onset of shivering (S_tt) and the increase in metabolic rate (L_ct) and by the linearity of the relationship between metabolic rate and shivering intensity in the cold. (3) For a comparable thermal demand, shivering intensity decreased with age whereas cold induced heat production remained constant, which suggests that the thermogenic efficiency of shivering is improved during the first 5 days of life.     

Alterations in thyroid hormone physiology induced by temperature and feeding in newly hatched chickens

In the chicken the transition of a poikilotherm to a homeotherm reaction upon cold exposure takes place in the perinatal period between pipping and hatching. However, newly hatched chicks cannot maintain their body temperature within narrow limits after cold exposure. The fact that relatively little attention was payed on the role of thyroid hormones in the thermoregulatory reaction to cold of young chicks was probably due to the hypothetically long latention time that was thought to be necessary to bring about changes in secretory activity by cold stimulation. However, more recently, rapid changes (within hours) of thyroid hormone concentrations upon cold exposure were described in the chickens and the quail. In this study, changes in circulating T3 and T4 concentrations upon cold exposure of young chicks during the first two weeks were followed, that means during the period wherein NST (non-shivering thermogenesis), if it exists at all, should be progressively replaced by ST (shivering thermogenesis). Because of the importance of feeding condition on thyroid hormone levels, the experiments were carried out with and without a preceeding fasting period. In all experiments a short-term cold exposure of young chickens (1-11 days) fed ad lib decreased T3 but increased T4 levels while a reversed picture was found after short cold exposure of the fasted animals. However, after prolonged cold stimulus (15 degrees C) of young chickens fed ad lib, plasma T3 was also significantly elevated over that of controls whereas T4 levels returned to normal values. A prolonged warm treatment (37 degrees C) of young chickens fed ad lib resulted in significantly lower T3 and higher T4 concentrations. After a prolonged cold treatment no differences in T4 or T3 response upon TRH were found whereas the warm treatment abolished these responses upon TRH. However, a cold treatment at the stage of incubation during which the hypothalamo-hypophyseal control of thyroid function is established (dag 10-14) enhanced the T4 response to TRH with a long lasting effect extending to the posthatch period. Since T3 is thought to be the active form of thyroid hormones with regard to thermopoiesis we have studied more specifically the effect of blocking peripheral conversion of T4 on thermoregulatory abilities in young chicks and the influence of temperature treatment on monodeiodination capacity. The lower rectal temperatures following the interference with the peripheral monodeiodination of T4, the effect being more pronounced at the lower ambient temperature, are indicative for a preponderant role of T3 on thermogenesis.(ABSTRACT TRUNCATED AT 400 WORDS)     

Shivering and non-shivering thermogenesis in a marsupial,the eastern barred bandicoot (Perameles gunnii)

We investigated the metabolic rate of the Tasmanian marsupial, the eastern barred bandicoot, Perameles gunnii, before and after acclimation to cold temperature (5 °C) for a 2-week period. Although body temperature did not change significantly, we observed a significant increase in the metabolic rate (MR) when measured at 5 °C before and after cold acclimation. Nor-epinephrine had a significant effect on the metabolic rate when measured in the thermoneutral zone and when measured at 5 °C after cold acclimation; however, there was no significant increase when measured at 5 °C before cold acclimation. Nor-epinephrine also resulted in a small but significant decrease in body temperature. Electromyography (EMG) measurements were obtained before and after cold acclimation during shivering. Shivering decreased after two weeks of cold exposure indicating that the bandicoot had acclimated to that temperature. Nor-epinephrine (NE) significantly reduced shivering before but not after cold acclimation. The metabolic rate and shivering decreased in the adult eastern barred bandicoot after acclimation at 5 °C and nor-epinephrine had similar effects to cold acclimation. Our findings of minor changes in thermal conductance suggest that insulation differences were unlikely explanations for our results. These experiments indicate that this marsupial is able to increase its heat production by non-shivering thermogenesis.     

Does cold acclimation induce nonshivering thermogenesis in juvenile birds? Experiments with Pekin ducklings and Japanese quail chicks

The capability to produce heat in cold by nonshivering thermogenesis (NST) was studied in Pekin ducklings and Japanese quail chicks acclimated to cold for 3 weeks using indirect calorimetry (oxygen consumption) and electromyography from breast (M. pectoralis) and leg muscles (quails: M. gastrocnemius; ducklings: M. gastrocnemius, M. iliofibularis). Respiration of muscles in vitro was studied by measuring cytochrome c oxidase activity. In both species, cold acclimation induced clear morphometric and physiological changes, but no clear evidence of nonshivering thermogenesis. This was evident because increased shivering at least in one muscle coincided with increased oxygen consumption. In ducklings, however, amplitudes of shivering EMGs were low (<30 μV) in all muscles studied in both the control and cold-acclimated groups. Ducklings reacted to cold mainly by means of increasing body weight (1796 g in control, 2095 g in cold-acclimated) and circulatory changes. Acclimation did not change oxygen consumption either in vivo or in vitro. In quails, in addition to increased body weight (78.1 g control, 89.9 g cold-acclimated), improved insulation and metabolic adaptation to cold (increased respiration in vivo and in M. pectoralis in vitro) was also utilized. In Japanese quail chicks, 3 weeks of cold acclimation does not seem to induce NST, while in Pekin ducklings the existence of NST could not be totally excluded because of weak overall shivering activity. Accepted: 13 July 2000     

Substrate preference in aminophylline-stimulated thermogenesis

The present study investigated the suitability of different substrates on aminophylline (AMPY)-induced thermogenesis in rats during cold exposure. Feeding of distilled water 60 min prior to cold exposure in two-day fasted rats resulted in the lowest total heat production and final body temperature in both saline- and AMPY-treated groups. Feeding of 5 ml Intralipid (2 Kcal/ml), a triglyceride mixture, did not improve thermogenesis beyond the control levels. However, feeding of isocaloric substitutes of sucrose elevated significantly the total thermogenesis by 7.9% and 7.4% and final body temperature by 2.23 and 1.61 degrees C, respectively, in saline- and AMPY-treated groups. The increase in thermogenesis by sucrose is not due to its thermic effect. It is concluded that sucrose, in combination with AMPY, may be of value in improving resistance to cold.     

Effects of fasting and aminophylline on norepinephrine-stimulated non-shivering thermogenesis

In an attempt to further elucidate the mechanisms of fasting-depressed maximum thermogenesis and cold tolerance, norepinephrine (NE)-stimulated non-shivering thermogenesis (NST) in cold-acclimated rats was used as a functional index of possible alterations in adrenergic efficacy after fasting. Fasting decreased the magnitude of maximum NE-Stimulated NST by 18.2% [6.87±0.47 Kcal (Kg^.75.min)^?1 well-fed vs. 5.81±0.39 Kcal (Kg^.75.min)^?1 fasted], but the apparent adrenergic binding affinity was not affected [Ke=0.43 μg NE min^?1 well-fed vs 0.55 μg NE min^?1 fasted]. Pretreatment with aminophylline [15 mg Kg^?1, i.p.], a phosphodiesterase inhibitor, restored the fasting-depressed NE-stimulated NST to the fed level. The results suggest that the depression of maximum thermogenesis after fasting is not due to changes in adrenergic binding characteristics but to alteration in cAMP production/degradation, resulting in decreased substrate mobilization for thermogenesis.     

Ontogeny of thermoregulatory mechanisms in king penguin chicks (Aptenodytes patagonicus)

The rapid maturation of thermoregulatory mechanisms may be of critical importance for optimising chick growth and survival and parental energy investment under harsh climatic conditions. The ontogeny of thermoregulatory mechanisms was studied in growing king penguin chicks from hatching to the full emancipation observed at 1 month of age in the sub-Antarctic area (Crozet Archipelago). Newly hatched chicks showed small, but significant regulatory thermogenesis (21% rise in heat production assessed by indirect calorimetry), but rapidly became hypothermic. Within a few days, both resting (+32%) and peak (+52%) metabolic rates increased. The first week of life was characterised by a two-fold rise in thermogenic capacity in the cold, while thermal insulation was not improved. During the second and third weeks of age, thermal insulation markedly rose (two-fold drop in thermal conductance) in relation to down growth, while resting heat production was slightly reduced (-13%). Shivering (assessed by electromyography) was visible right after hatching, although its efficiency was limited. Thermogenic efficiency of shivering increased five-fold with age during the first weeks of life, but there was no sign of non-shivering thermogenesis. We conclude that thermal emancipation of king penguin chicks may be primarily determined by improvement of thermal insulation after thermogenic processes have become sufficiently matured. Both insulative and metabolic adaptations are required for the rapid ontogeny of thermoregulation and thermal emancipation in growing king penguin chicks.     

Effects of fasting on heat balance and nonshivering thermogenesis in febrile adult guinea pigs

1. 1. The calorigenic responses of 60-h fasted (F) and control (C) guinea pigs to E. coli endotoxin (5 μg·kg⁻¹ i.v.) injection were compared at T_a = 25°C.

2. 2. In contrast to fed (C) guinea pigs, the F group showed significantly lower calorigenic response.

3. 3. The brown adipose tissue (BAT) thermogenesis judge by the BAT temperature did not change significantly. Shivering activity was not different from that in the C group.

4. 4. The results conclude that fasting attenuates endotoxin-induced fever and this attenuation is due to suppression of nonshivering thermogenesis (NST) in guinea pigs.

5. 5. The results from direct calorimetry indicate that endotoxin injection evoked a prominent increase in heat production, while the changes in heat loss do not have an important role.

Shivering in the cold: from mechanisms of fuel selection to survival.

In cold-exposed adult humans, significant or lethal decreases in body temperature are delayed by reducing heat loss via peripheral vasoconstriction and by increasing rates of heat production via shivering thermogenesis. This brief review focuses on the mechanisms of fuel selection responsible for sustaining long-term shivering thermogenesis. It provides evidence to explain large discrepancies in fuel selection measurements among shivering studies, and it proposes links between choices in fuel selection mechanism and human survival in the cold. Over the last decades, a number of studies have quantified the contributions of carbohydrate (CHO) and lipid to total heat generation. However, the exact contributions of these fuels still remain unclear because of large differences in fuel selection measurements even at the same metabolic rate. Recent advances on the mechanisms of fuel selection during shivering provide some plausible explanations for these discrepancies between shivering studies. This new evidence indicates that muscles can sustain shivering over several hours using a variety of fuel mixtures achieved by modifying diet (changing the size of CHO reserves) or by changing muscle fiber recruitment (increasing or decreasing the recruitment of type II fibers). From a practical perspective, how does the choice of fuel selection mechanism affect human survival in the cold? Based on a glycogen-depletion model, estimates of shivering endurance show that, whereas the oxidation of widely different fuel mixtures does not improve survival time, the selective recruitment of fuel-specific muscle fibers provides a substantial advantage for cold survival. By combining fundamental research on fuel metabolism and applied strategies to improve shivering endurance, future research in this area promises to yield important new information on what limits human survival in the cold.     

Participation of breast and leg muscles in shivering thermogenesis in young turkeys and guinea fowl

Turkey (Meleagris gallopavo) and guinea fowl (Numida meleagris) chicks (0–27 days posthatch) were exposed to decreasing or increasing ambient temperatures. Root mean square electromyographic activity of musculus pectoralis (m. pect.) and musculus iliotibialis (m. iliot.) was recorded simultaneously with O₂ consumption and CO₂ production. From both muscles, relative mass, water fraction and fibre type were determined. M. iliot. participated in shivering from hatching onwards. The relationship between its root mean square electromyographic activity and ambient temperature resembled that of metabolic rate and ambient temperature, and the shivering threshold temperature was indistinguishable from the lower critical temperature. This suggests that the leg muscles are major contributors to shivering thermogenesis. M. pect. participated in shivering only at days 6–20 in turkeys and at days 6–10 in guinea fowl. Both water fraction and histological analysis indicated that m. pect. was less developed than m. iliot. at hatching. We hypothesize that a minimal level of maturity is required before a muscle can participate in shivering, which is probably represented by a water fraction of about 0.85. Both species recruited the aerobic leg muscles first; the anaerobic breast muscle was recruited only when the rate of mass-specific heat loss was high. Accepted: 20 March 1997     

Ontogeny of cardiac rate regulation and brown fat thermogenesis in golden hamsters (Mesocricetus auratus)

It was shown previously in infant rats (Rattus norvegicus) that the ability to produce heat in the cold using brown adipose tissue (BAT) is closely related to the ability to maintain cardiac rate. When the limits of BAT thermogenesis were exceeded, interscapular temperature (which reflects the temperature of the interscapular BAT depot) and cardiac rate fell together. As an extension of this earlier study, the relation between BAT thermogenesis and cardiac rate was examined here in the golden hamster (Mesocricetus auratus), a species whose young do not exhibit BAT thermogenesis until the end of the 2nd week postpartum. It was found that 3 to 12-day-old hamsters were unable to increase shivering or nonshivering thermogenesis in the cold and exhibited decreases in cardiac rate that proceeded in lock-step with decreases in interscapular temperature. In contrast, as the thermogenic capability of hamsters increased after 12 days of age, cardiac rate was maintained within narrow limits across a wide range of air temperatures. These results support the hypothesis that heat produced by BAT helps to warm the heart and thus aids in the maintenance of cardiac rate during cold exposure. Accepted: 16 August 1997     

Effects of fasting on maximum thermogenesis in temperature-acclimated rats

To further investigate the limiting effect of substrates on maximum thermogenesis in acute cold exposure, the present study examined the prevalence of this effect at different thermogenic capabilities consequent to cold- or warm-acclimation. Male Sprague-Dawley rats (n=11) were acclimated to 6, 16 and 26C, in succession, their thermogenic capabilities after each acclimation temperature were measured under helium-oxygen (21% oxygen, balance helium) at –10C after overnight fasting or feeding. Regardless of feeding conditions, both maximum and total heat production were significantly greater in 6>16>26C-acclimated conditions. In the fed state, the total heat production was significantly greater than that in the fasted state at all acclimating temperatures but the maximum thermogenesis was significant greater only in the 6 and 16C-acclimated states. The results indicate that the limiting effect of substrates on maximum and total thermogenesis is independent of the magnitude of thermogenic capability, suggesting a substrate-dependent component in restricting the effective expression of existing aerobic metabolic capability even under severe stress.     

Inhibitory control of shivering and nonshivering thermogenesis by preoptic warm-sensitive neurons

To investigate the characteristics of efferent projections from the preoptic area for the control of shivering and nonshivering thermogenesis, we tested the effects of thermal stimulation and injecting excitatory substances into the preoptic area on shivering and nonshivering thermogenesis in anesthetized rats. Preoptic warming and injection of glutamate suppressed shivering at ambient temperatures of 15–21°C. Likewise, preoptic warming and d,l-homocysteic acid injection suppressed nonshivering thermogenesis elicited by electrical stimulation of the ventromedial hypothalamic nucleus. Inhibitory signals from warm-sensitive neurons, thus, contribute a larger efferent signal for heat production than do signals from cold-sensitive neurons.     

Metabolic changes associated with sustained 48-Hr shivering thermogenesis in the newborn pig

Metabolic changes associated with sustained 48-hr shivering thermogenesis were studied in piglets maintained at 34 (thermoneutrality) or 25°C (cold) between 6 and 54 hr of life. Despite their high shivering activity and elevated heat production, cold-exposed piglets exhibited a slightly lower rectal temperature than thermoneutral animals (-1.1°C; P < 0.01) at the end of the treatment. The enhancement of heat production and shivering activity were associated with a decrease in muscle glycogen (− 47%; P < 0.05) and total lipid content (− 23%; P < 0.05), a reduction of blood lactate levels (P < 0.05) and an enhancement of muscle cytochrome oxidase activity (+20%;P < 0.05), which suggests that muscle oxidative potential was increased by cold exposure. Potential for capturing lipids (lipoprotein lipase activity) was also higher in the redrhomboideus muscle (+ 71%; P < 0.01) and lower in adipose tissue (−58%; P < 0.01) of the cold-exposed piglets. Measurements performed at the mitochondrial level show no changes inrhomboideus muscle, but respiratory capacities (state IV and FCCP-stimulated respiration) and intermyofibrillar mitochondria oxidative and phosphorylative (creatine kinase activity) capacities were enhanced inlongissimus dorsi muscle (P < 0.05). These changes may contribute to provide muscles with nonlimiting amount of readily oxidable substrates and ATP necessary for shivering thermogenesis. A rise in plasma norepinephrine levels was also observed during the second day of cold exposure (P < 0.05).     

Shivering modulation in humans: Effects of rapid changes in environmental temperature

During cold exposure, increase in heat production is produced via the activation of shivering thermogenesis and nonshivering thermogenesis, the former being the main contributor to compensatory heat production in non-acclimatized humans. In rats, it has been demonstrated that shivering thermogenesis is modulated solely by skin thermoreceptors but this modulation has yet to be investigated in humans. The aim of this study was to determine if cold-induced shivering in humans can be modulated by cutaneous thermoreceptors in conditions where increases in heat loss can be adequately compensated by increases in thermogenic rate. Using a liquid-conditioned suit, six non-acclimatized men were exposed to cold (6 °C) for four 30 min periods, each of them separated by 15 min of heat exposure (33 °C). Core temperature remained stable throughout exposures whereas skin temperatures significantly decreased by 12% in average during the sequential cold/heat exposures compared to baseline (p<0.0001). Shivering intensity and metabolic rate increased significantly during 6 °C exposures (3.3±0.7% MVC, 0.40±0.0 L O₂/min, respectively) and were significantly reduced during 33 °C exposure (0.5±0.1% MVC, 0.25±0.0 L O₂/min; p<0.005 for both). Most importantly, shivering could be quickly and strongly inhibited during 33 °C exposure although skin temperature often remained below baseline values. In conclusion, under compensatory conditions, cutaneous thermoreceptors appear to be a major modulator of the shivering response in humans and seem to react rapidly to changes in the microclimate right next to the skin and to skin temperature.     

Normal thyroid thermogenesis but reduced viability and adiposity in mice lacking the mitochondrial glycerol phosphate dehydrogenase

The mitochondrial glycerol phosphate dehydrogenase (mGPD) is important for metabolism of glycerol phosphate for gluconeogenesis or energy production and has been implicated in thermogenesis induced by cold and thyroid hormone treatment. mGPD in combination with the cytosolic glycerol phosphate dehydrogenase (cGPD) is proposed to form the glycerol phosphate shuttle, catalyzing the interconversion of dihydroxyacetone phosphate and glycerol phosphate with net oxidation of cytosolic NADH. We made a targeted deletion in Gdm1 and produced mice lacking mGPD. On a C57BL/6J background these mice showed a 50% reduction in viability compared with wild-type littermates. Uncoupling protein-1 mRNA levels in brown adipose tissue did not differ between mGPD knockout and control pups, suggesting normal thermogenesis. Pups lacking mGPD had decreased liver ATP and slightly increased liver glycerol phosphate. In contrast, liver and muscle metabolites were normal in adult animals. Adult mGPD knockout animals had a normal cold tolerance, normal circadian rhythm in body temperature, and demonstrated a normal temperature increase in response to thyroid hormone. However, they were found to have a lower body mass index, a 40% reduction in the weight of white adipose tissue, and a slightly lower fasting blood glucose than controls. The phenotype may be secondary to consequences of the obligatory production of cytosolic NADH from glycerol metabolism in the mGPD knockout animal. We conclude that, although mGPD is not essential for thyroid thermogenesis, variations in its function affect viability and adiposity in mice.     

Daily rhythm of oxygen consumption and thermoregulatory responses in some European winter- or summer-acclimatized finches at different ambient temperatures

The oxygen consumption of European finches, the siskin (Carduelis spinus), the brambling (Fringilla montifringilla), the bullfinch (Pyrhulla pyrhulla), the greenfinch (Carduelis chloris) and the hawfinch (Coccothraustes coccothraustes), was recorded continuously while ambient temperature was decreased stepwise from +30 down to-75°C. The oxygen consumption, body temperature (telemetrically), and shivering (integrated pectoral electromyography) of greenfinches were measured simultaneously at ambient temperatures between +30 and-75°C. Maximum heat production, cold limit, lower critical temperature, basal metabolic rate and thermal conductance (of the greenfinch) were determined. The diurnal variation of oxygen consumption of siskins and greenfinches was recorded at thermoneutrality and below the thermoneutral zone in winter- and summer-acclimatized birds. The diurnal variation of body temperature and thermal conductance of greenfinches were also determined. The diurnal variation of heat production was not seasonal or temperature dependent in the siskin and in the greenfinch. Nocturnal reduction of oxygen consumption saved 15–33% energy in the siskin and greenfinch. Body temperature of the greenfinch was lowered by 2.5–3.4°C. The nocturnal reduction of thermal conductance in the greenfinch was 39–48%. The basal metabolic rate was lowest in the largest bird (hawfinch) and highest in the smallest bird (siskin). The values were in the expected range. The heat production capacity of finches in winter was 4.7 times basal metabolic rate in the siskin, 4.2 times in the brambling, 3.5 times in the greenfinch and 2.9 times in the bullfinch and hawfinch. The heat production capacity of the siskin and greenfinch was not significantly lower in summer. The cold limit temperatures (°C) in winter were-61.2 in the siskin,-41.3 in the greenfinch,-37.0 in the bullfinch,-35.7 in the brambling and-28.9 in the hawfinch. The cold limit was 14.3°C higher in summer than in winter in the siskin and 8.7°C in the greenfinch. Thermal insulation of the greenfinch was significantly better in winter than in summer. The shivering of the greenfinch increased linearly when ambient temperature was decreased down to-40°C. Maintenance of shivering was coincident with season. In severe cold integrated pectoral electromyography did not correlate with oxygen consumption as expected. The possible existence of non-shivering thermogenesis in birds is discussed. It is concluded that the acclimatization of European finches is primarily metabolic and only secondly affected by insulation.Abbreviations AAT avian adipose tissue - bm body mass - BMR basal metabolic rate - C _t thermal conductance - EMG electromyogram - HP heat production - HP _max maximum heat production - MR metabolic rate - NST non-shivering thermogenesis - RMR resting metabolic rate - RQ respiratory quotient - T _a ambient temperature - T _b body temperature - T _c colonic temperature - T _1c lower critical temperature - TNZ thermoneutral zone - T _st shivering threshold temperature - V oxygen consumption