Oxygen supply and the placenta limit thermogenic responses in fetal sheep

The aim of this study was to assess the individual effects of cooling, increased oxygenation, and umbilical cord occlusion on nonshivering thermogenesis in utero. A cooling coil was placed around eight fetal sheep of 132-145 days gestation; thermistors were placed in the fetal esophagus and maternal iliac artery, vascular catheters and a tracheal catheter were inserted, and a snare was placed loosely around the umbilical cord. The next day cold water was circulated through the coil for 5 h. During the 1st h of cooling alone, fetal core temperature fell 2.79 degrees C, but indexes of brown fat activity increased only slightly. After ventilation with O2, plasma free fatty acid concentration (FFA) rose 7.4-fold to 244 +/- 42 mu eq/l, glycerol concentration rose fourfold to 376 +/- 85 microM, and the difference between brown fat and core temperature widened to 0.60 +/- 0.10 degrees C. Ventilation with N2-enriched air did not evoke similar responses. After snaring the umbilical cord while ventilation was continued, FFA rose to 554 +/- 95 mu eq/l, glycerol rose to 684 +/- 76 microM, and the temperature difference widened to 0.77 +/- 0.13 degrees C. Whole-body O2 consumption peaked at 19.6 ml.min-1.kg-1 of fetal tissue. We conclude that fetal thermogenic responses are limited in part by O2 delivery to brown fat and are augmented by occlusion of the umbilical cord.     

Changes in plasma adenosine during simulated birth of fetal sheep

Adenosine is known to inhibit nonshivering thermogenesis in adult brown fat. These experiments were undertaken to test whether fetal adenosine, normally present in high concentrations, suppresses lipolysis in utero and then falls after birth, permitting thermogenesis to begin. To test this hypothesis, we measured fetal plasma adenosine concentration [ADO] using high-performance liquid chromatography in 11 fetal sheep at 135-140 days gestation during simulated birth. During an initial control period, fetal [ADO] averaged 1.9 +/- 0.3 microM, about four times maternal [ADO] (0.4 +/- 0.1 microM, P less than 0.001). The fetus was then cooled by circulating cold water through a plastic coil encircled about the fetal torso. One hour later, when fetal core temperature had decreased 2.3 degrees C, fetal [ADO] averaged 2.8 +/- 0.5 microM, a 50% increase (P less than 0.05), while thermogenesis remained inactive. Next the fetal lungs were ventilated with O2 to raise arterial Po2 to greater than or equal to 150 Torr. Fetal [ADO] decreased only slightly, and thermogenic responses were modest. Finally, the umbilical cord was occluded. Fetal [ADO] decreased rapidly and 60 min later averaged 1.1 +/- 0.2 microM, 40% below initial control (P less than 0.05) and 57% below the previous period (P less than 0.001). As [ADO] fell, strong thermogenic responses became apparent, as indicated by seven- to eightfold increases in plasma glycerol (P less than 0.001) and a doubling in fetal O2 consumption (P less than 0.001). These results are consistent with the hypothesis that high fetal [ADO] inhibits thermogenesis before birth but then decreases after cord occlusion, allowing thermogenesis to begin.(ABSTRACT TRUNCATED AT 250 WORDS)     

Suppressive action of endogenous adenosine on ovine fetal nonshivering thermogenesis

Ball, Karen T., Tania R. Gunn, Peter D. Gluckman, and GordonG. Power. Suppressive action of endogenous adenosine on ovinefetal nonshivering thermogenesis. J. Appl.Physiol. 81(6): 2393-2398, 1996.Nonshiveringthermogenesis is not initiated when the fetal sheep is cooled in uterobut appears to require the removal of an inhibitor of placental originat birth. To test whether adenosine is such an inhibitor, we examinedthe effect of the adenosine antagonist theophylline on the initiationof nonshivering thermogenesis during sequential cooling, ventilation, and umbilical cord occlusion in utero. Theophylline (18 mg/kg bolus and0.6 mg ^· kg^{1 ·} min¹thereafter) was infused for 90 min before and 90 min after cord occlusion. Theophylline enhanced the nonshivering thermogenic freefatty acid (FFA) and glycerol responses before cord occlusion, raisingFFA concentrations 99% to 415 ± 60 µeq/l(P < 0.01) and glycerol levels 87%to 526 ± 135 µmol/l (P < 0.05). These FFA (P < 0.001) andglycerol (P < 0.05) concentrationswere significantly greater than the corresponding period during thebirth-simulation control. Umbilical cord occlusion did not alter FFAlevels but induced a 41% rise in glycerol concentrations to 774 ± 203 µmol/l (P < 0.05). Theincreases in nonshivering thermogenic indexes after the administrationof the adenosine-receptor antagonist suggest that the quiescent stateof ovine fetal brown adipose tissue may result, in part, from the tonicinhibitory actions of adenosine and that a decrease in adenosineconcentrations enhances nonshivering thermogenesis. However, thefurther rise after umbilical cord occlusion suggests that at least oneother inhibitor of placental origin inhibits nonshivering thermogenesisbefore birth.

     

Initiation of pulmonary gas exchange by fetal sheep in utero

The role of umbilical cord occlusion in the initiation of breathing at birth was investigated using unanesthetized fetal sheep that were provided with access to a tracheal supply of hyperoxic air. Near-term fetuses were studied in utero to eliminate extraneous sensory stimuli. Gasping movements began 1.4 +/- 0.1 min after cord occlusion. Breathing was irregular for several minutes before continuous breathing (greater than or equal to 40 min-1) began 6 +/- 1 min after cord occlusion (n = 10). Arterial PO2 rose significantly from 18 +/- 2 mmHg before occlusion and was 115 +/- 15 mmHg immediately before cord release at 15 or 30 min. Breathing continued even during high-voltage electrocortical activity. Cord release caused the breathing rate to decrease from 77 +/- 13 min-1 during the last 5 min of cord occlusion to 5 +/- 3 min-1 10 min after cord release (P less than 0.002; n = 7). Results indicate the change from placental to lung gas exchange can occur in the absence of sensory and thermal changes normally present at birth and that the transition is reversible.     

Developmental and adult acclimation effects of ambient temperature on temperature regulation of mice selected for high and low levels of nest-building

Summary Genetic and environmental components of adaptation to cold inMus musculus were assessed in a study of the effects of selective breeding for behavioral temperature regulation (indexed by high and low levels of nest-building), rearing mice from birth in the cold, and cold acclimation of adult animals, on thermoregulatory traits. Mice from the eleventh selected generation of a high-nesting line maintained higher resting metabolic rates and body temperatures, while at the same time consuming less food when compared with mice from the low-nesting line (Table 1). High-nesting mice were also more discriminating in their temperature preference when placed on a thermal gradient. Thus, common genetic loci must influence a variety of energy conservation measures important for survival in the cold, including insulative nest-building, metabolic efficiency, and optimum microhabitat selection.Rearing mice at 5°C from birth until 70 days of age resulted in permanent increases in nonshivering thermogenesis, weight of interscapular brown adipose tissue, and core body temperature when compared to mice raised at 22°C (Table 1). These greater heat production capacities were accompanied by consumption of more food. Cold acclimation of adults at 5°C for 3 weeks similarly increased measures of thermogenic capacity (nonshivering thermogenesis and interscapular brown adipose tissue) as well as food consumption, when compared to the effects of warm acclimation, but differed from the effects of cold-rearing in that while resting metabolic rates were elevated, no significant differences in body temperature were found (Table 1).Sex differences were also noted for most of the thermoregulatory measures, with the lighter females scoring higher on thermal preference, resting metabolic rate, nonshivering thermogenesis, brown fat, and food consumption.In general, these results suggest that a more precise partitioning of the genetic and environmental factors which influence thermoregulatory traits in mammals could eventually result in a better understanding of the differences which exist between acclimated and acclimatized animals.     

Plasma free fatty acid levels during cold-induced and noradrenaline-induced nonshivering thermogenesis in the Djungarian hamster

Summary In Djungarian hamsters the cold-induced thermoregulatory heat production was preceeded and accompanied by an increase in the plasma level of free fatty acids. In warm-acclimated hamsters this increase was found more pronounced (0.85 to 1.48 mM) than in cold-acclimated hamsters (0.64 to 0.88 mM). Noradrenaline-induced thermogenesis at thermoneutrality provoked a similar increase in the free fatty acid level. Inhibition of nonshivering thermogenesis during cold exposure by propranolol abolished the increase in free fatty acids completely. The surgical removal of brown adipose tissue proportionately reduced the increase in free fatty acids. This indicates that the rise in plasma free fatty acids is functionally related to nonshivering thermogenesis and originates from brown adipose tissue.     

Seasonal acclimation of bank voles and wood mice: nonshivering thermogenesis and thermogenic properties of brown adipose tissue mitochondria

Summary Seasonal acclimation of nonshivering thermogenesis and brown adipose tissue was studied in wild bank voles (Clethrionomys glareolus), yellow necked field mice and wood mice (Apodemus flavicollis, A. sylvaticus). Both, voles and mice increased their capacity for nonshivering thermogenesis during winter. Thermogenic properties of brown fat (cytochrome c oxidase activity, mitochondrial protein content, GDP-binding of brown fat mitochondria) showed similar changes during seasonal acclimation;Clethrionomys andApodemus spp. both showed lowest thermogenic properties in the summer during August, a rapid increase during fall, and highest levels of thermogenic activity in the winter months. With regard to changes in body weight and brown fat mass these species show different strategies for seasonal acclimation. InClethrionomys a reduction of body mass in the winter was found, both in the wild population as well as in individual animals housed in the laboratory.A. flavicollis showed a reduction of body weight during fall, whereasA. sylvaticus maintained a constant body mass throughout the year. Brown fat mass and cellularity increased in theApodemus spp. during winter, in parallel with the thermogenic properties of brown fat, whereas inClethrionomys brown fat mass and cellularity remained seasonally constant. These species live in the same habitat and were trapped in the same area. It is concluded that seasonal improvements of in vivo and in vitro thermogenesis are very similar in these species, although the physiological basis for this improvement is different inClethrionomys andApodemus.Abbreviations BAT brown adipose tissue - BMR basal metabolic rate (resting metabolic rate at thermoneutrality) - BW body weight - COX cytochrome c oxidase - GDP guanosine diphosphate - MP mitochondrial protein - NA noradrenaline - NST nonshivering thermogenesis - NSTcap NST capacity (NST maximum minus BMR) - T _a ambient temperature     

Effect of exercise training on the disappearance of cold adaptability in rats

Following the transfer of cold-adapted rats to a warm environment at 25 degrees C, enhanced nonshivering thermogenesis and enlarged interscapular brown adipose tissue (BAT) decreased gradually and reached a steady state after 4 weeks of de-adaptation. Animals that were exercised in the process of de-adaptation, however, showed no decrease in enhanced nonshivering thermogenesis, but did show a decrease in BAT weight as compared with sedentarily de-adapted animals. Triiodothyronine (T3), the physiologically most active thyroid hormone, was at a higher plasma level in cold-adapted rats than in de-adapted animals with or without exercise loads. Although the resting level of T3 in running-trained rats was not higher than that in sedentary rats, some fluctuations of T3 level were observed during running.     

Absence of adaptive nonshivering thermogenesis in a marsupial,the fat-tailed dunnart (<Emphasis Type="Italic">Sminthopsis crassicaudata</Emphasis>)

The presence of nonshivering thermogenesis in marsupials is controversially debated. Survival of small eutherian species in cold environments is crucially dependent on uncoupling protein 1 (UCP1)-mediated, adaptive nonshivering thermogenesis that is executed in brown adipose tissue. In a small dasyurid marsupial species, the fat-tailed dunnart (Sminthopsis crassicaudata), an orthologue of UCP1 has been recently identified which is upregulated during cold exposure resembling adaptive molecular adjustments of eutherian brown adipose tissue. Here, we tested for a thermogenic function of marsupial brown adipose tissue and UCP1 by evaluating the capacity of nonshivering thermogenesis in cold-acclimated dunnarts. In response to an optimal dosage of noradrenaline, cold-acclimated dunnarts (12°C) showed no additional recruitment of noradrenaline-induced maximal thermogenic capacity in comparison to warm-acclimated dunnarts (24°C). While no differences in body temperature were observed between the acclimation groups, basal metabolic rate was significantly elevated after cold acclimation. Therefore, we suggest that adaptive nonshivering thermogenesis does not occur in this marsupial species despite the cold recruitment of oxidative capacity and UCP1 in the interscapular fat deposit. In conclusion, the ancient UCP orthologue in marsupials does not contribute to the classical nonshivering thermogenesis, and may exhibit a different physiological role.     

Evidence for a contribution by brown adipose tissue to the development of fever in the young rabbit

This study was undertaken to determine if brown adipose tissue was involved in heat production during fever produced by S. abortus equi (1 micrograms) in unanesthetized rabbits aged 19-26 days. The fever (0.9-1.6 degrees C) occurred after a delay of 20-30 min and was frequently biphasic. Radiolabelled microspheres for measuring tissue blood flow were injected intraventricularly into three groups of animals: rabbits not given pyrogen, rabbits in which the febrile response to pyrogen was developing, and rabbits in which the febrile response had peaked. Blood flow to brown fat deposits and other organs was calculated from the fractional distribution of the microspheres and the recovery of microspheres in a reference arterial blood sample. At the fever peak, blood flow to brown fat was not significantly different (p greater than 0.05) from the control value (0.9 +/- 0.2), but during the rising phase of the fever the flow increased significantly (p less than 0.01) to 2.6 +/- 0.4 mL min-1 g-1. The blood flow to muscles of the forelimbs and hind limbs was also increased significantly (p less than 0.05) during the rising phase of the fever. No significant change in blood flow to other organs or tissues was found during the rising phase of the fever. These results indicate that both nonshivering as well as shivering thermogenesis contribute to heat production during development of fever in the young rabbit. However, nonshivering thermogenesis was not involved in the maintenance of the elevated body temperature after the fever had peaked.     

Increased contribution of brown adipose tissue to nonshivering thermogenesis in the Djungarian hamster during cold-adaptation

Summary The effect of cold-adaptation was investigated on the brown adipose tissue of Djungarian hamsters. Animals maintained at 23°C and 16 hours light per day (controls) were exposed to 5°C. The wet weight of the total brown fat is reduced by some 40% within 4 days of coldexposure, as a result of extensive triacylglycerol depletion of the tissue with no reduction in DNA; the tissue mass remains constant unde persistent cold influence. The total amount of tissue mitochondria is doubled by 24 h and increases by a factor of 3 under persistent cold-stimulus, the specific respiratory capacity of the organelles remaining unchanged. The amount of 32 kDa regulatory protein per mg mitochondrial protein quantified from high-affinity GDP-binding, is increased by a factor of 2.7 after 21 days of cold-adaptation; a 9-fold increment is found of the total mitochondrial GDB-binding capacity.Comparison of nonshivering thermogenesis and the maximal thermogenic capacity of brown fat, estimated from the maximal respiration of the isolated mitochondria and the total amount of mitochondria in the tissue, suggests that brown fat may contribute about 20% to the whole-body nonshivering thermogenesis in warm-adapted controls and 45% in cold-adapted hamsters. The estimated increase in thermogenic capacity of the tissue in response to 21 days of cold-adaptation corresponds to the increase in nonshivering thermogenesis, suggesting a central thermoregulatory role of brown fat during cold-adaptation.Abbreviation FCCP carbonylcyanide-p-trifluoromethoxyphenylhydrazone     

自愿转轮运动对雄性长爪沙鼠体重和能量代谢的影响

运动是影响动物能量平衡和体重变化的重要因素之一。为研究自愿转轮运动对体重和能量代谢的影响,我们监测了8 周自愿转轮运动过程中,雄性长爪沙鼠的体重、能量摄入、消化率、静止代谢率(RMR) 和非颤抖性产热(NST)的变化,以及8 周后的体脂含量、血清甲状腺激素(T3、T4 )和瘦素(leptin)水平的变化等。结果发现,8 周自愿转轮运动增加了长爪沙鼠的体重和能量摄入以及血清瘦素水平(血清瘦素浓度比对照组高27% ),但对消化率、RMR 和褐色脂肪组织的线粒体蛋白浓度等没有明显影响。尽管体脂含量和血清T₃ 和T₄ 没有显著差异,但运动组体脂含量比对照组高33%,血清T₃ 和T₄ 水平分别比对照组低10% 和38%。血清瘦素浓度与体脂重量呈正相关。因此,自动转轮运动并没有降低动物的体重和体脂含量,但瘦素和甲状腺素在雄性长爪沙鼠能量代谢和能量平衡中的作用尚需进一步确定。     

Brown fat UCP1 is not involved in the febrile and thermogenic responses to IL-1beta in mice

The activity of brown adipose tissue (BAT), a site of nonshivering metabolic thermogenesis, has been reported to increase after interleukin (IL)-1beta/lipopolysaccharide injection. To clarify the possible contribution of BAT thermogenesis to whole body febrile response, we investigated febrile and thermogenic response to IL-1beta using mice deficient in uncoupling protein-1 (UCP1), a key molecule for BAT thermogenesis. In wild-type (WT) mice, IL-1beta injection (5 microg/kg ip) increased body temperature (+1.82 degrees C at 20 min), decreased physical activity (-37% at 1 h), and produced a slight and insignificant rise (+15% at 1 h) in oxygen consumption (Vo(2)). Vo(2) dependent on metabolic thermogenesis (DeltaVO2 thermogenesis) calculated by correcting the effect of physical activity was increased after IL-1beta injection (726 +/- 200 ml x h(-1) x kg(-1) at 1 h). Almost the same responses were observed in UCP1-deficient mice, showing 638 +/- 87 ml x h(-1) x kg(-1) of DeltaVO2 thermogenesis at 1 h. In contrast, CL316,243, a selective activator of BAT thermogenesis, increased body temperature, decreased physical activity, and produced a significant rise in Vo2 in WT mice, showing 1,229 +/- 35 ml x h(-1) x kg(-1) of DeltaVO2 thermogenesis at 1 h. These changes were not observed in UCP1-deficient mice. These results, conflicting with a previously proposed idea of a role of BAT in fever, suggest a minor contribution of BAT thermogenesis to IL-1beta-induced fever. In support of this, we found no effect of IL-1beta on triglyceride content and UCP1 mRNA level in BAT, in contrast with apparent effects of CL316,243.     

High body weight associated with impaired nonshivering thermogenesis but improved glucose tolerance in Mongolian gerbils (Meriones unguiculatus)

Overweight and obesity correspond with metabolic syndromes, such as glucose intolerance and type 2 diabetes. The objective of this study was to determine whether decreased thermogenesis mass and glucose intolerance are directly related to changes in body mass in Mongolian gerbils. High body weight gerbils displayed increase in total body fat mass especially epididymal fat pad, and decrease in nonshivering thermogenesis, as indicated by depressed mitochondrial protein content and uncoupling protein-1 content in brown adipose tissue. No variations of sirtuin 1 and subunit IV of cytochrome oxidase expression were found in brown adipose tissue and skeletal muscle between the two groups. High body weight gerbils showed increased serum leptin and insulin concentrations but surprisingly increased glucose tolerance, suggesting a difference from other obese species in the regulation of glucose metabolism. Serum leptin levels were negatively correlated with UCP1 content in BAT and positively correlated with energy intake and insulin concentration. Our data suggest that leptin may be involved in thermogenesis regulation, insulin secretion and glucose metabolism in HBW gerbils.     

Photoperiod and thermoregulation in vertebrates: body temperature rhythms and thermogenic acclimation

Evidence has recently begun to accumulate that photoperiodic responses of mammals and birds may affect the control of energy balance and thermoregulation. Exposure to short photoperiod can lower the set point for body temperature regulation in birds and mammals, as well as the voluntarily selected body temperature in ectothermic lizards. This decrease is accompanied by a reorganization of circadian or ultradian rhythms of body temperature, particularly an increase in periods spent at rest with minimum body temperatures. Short photoperiod is also used as an environmental cue for induction of seasonal torpor or facilitation of hibernation. During winter, cold tolerance of small mammals is improved by an increase of nonshivering thermogenesis in brown fat. Thermogenic capacity of brown fat (respiratory enzymes, mitochondria, uncoupling protein) is enhanced in response to short photoperiod. This response is mediated via an increase in the activity of sympathetic innervation in brown fat. Moreover, an exposure to short photoperiod prior to low temperatures may act in preparing brown fat for facilitated thermogenesis during acclimation to cold. This shows that photoperiodic control not only affects energy balance indirectly via the control of reproduction or body mass, but may directly interact with central control of thermoregulation and may influence the process of acclimatization.     

Effects of photoperiod on energy budgets and thermogenesis in Mongolian gerbils (Meriones unguiculatus)

(1)

To investigate the role of photoperiod on the regulation of energy budgets and thermogenesis in Mongolian gerbils, body mass (BM), body fat mass (BFM), basal metabolic rate (BMR), nonshivering thermogenesis (NST), gross energy intake (GEI), mitochondrial cytochrome c oxidase (COX) activity and uncoupling protein1 (UCP1) content of brown adipose tissue (BAT), and serum tri-iodothyronine (T₃), thyroxine (T₄) and leptin levels were measured.     

Serum leptin, energy budget, and thermogenesis in striped hamsters exposed to consecutive decreases in ambient temperatures

Leptin has been found to be a direct participant in the regulation of both energy intake and energy expenditure in small mammals showing seasonal declines in body mass (M(b)) and fat mass, but its roles in an animal exhibiting seasonally increased thermogenesis and unchanged M(b) remain unclear. Serum leptin levels, energy budget, and thermogenesis were measured in striped hamsters exposed to consecutive decreases in ambient temperatures ranging from 23° to -23°C. Cold-exposed hamsters had significant increases in gross energy intake (GEI), the rate of basal metabolism, nonshivering thermogenesis, and activity of cytochrome c oxidase (COX) in brown adipose tissue (BAT), compared with control hamsters, indicating a cold-induced elevation of thermogenesis. Body mass and fat content were decreased in cold-exposed animals, and serum leptin levels were increased in hamsters exposed to temperatures of -8°C and below in inverse proportion to body fat content. Serum leptin levels were positively correlated with GEI and BAT COX activity in cold-exposed hamsters, but no such relationships were observed in control animals. These findings suggest that cold-exposed hamsters increase food consumption to meet the energy requirements for increased BAT thermogenesis. The increases in serum leptin levels are likely involved in increased thermogenesis in hamsters under cold stress. Cold-exposed hamsters may become leptin resistant, which is associated with impaired regulation of food intake. This new natural model of leptin resistance may also provide insight into the dynamic long-term control of energy homeostasis for animals that do not exhibit seasonal decline in M(b).     

Only UCP1 can mediate adaptive nonshivering thermogenesis in the cold.

Adaptive nonshivering thermogenesis may have profound effects on energy balance and is therefore therefore is a potential mechanism for counteracting the development of obesity. The molecular basis for adaptive nonshivering thermogenesis has remained a challenge that sparked acute interest with the identification of proteins (UCP2, UCP3, etc.) with high-sequence similarity to the original uncoupling protein-1 (UCP1), which is localized only in brown adipose tissue. Using UCP1-ablated mice, we examined whether any adaptive nonshivering thermogenesis could be recruited by acclimation to cold. Remarkably, by successive acclimation, the UCP1-ablated mice could be made to subsist for several weeks at 4C during which they had to constantly produce heat at four times their resting levels. Despite these extreme requirements for adaptive nonshivering thermogenesis, however, no substitution of shivering by any adaptive nonshivering thermogenic process occurred. Thus, although the existence of, for example, muscular mechanisms for adaptive nonshivering thermogenesis has recurrently been implied, we did not find any indication of such thermogenesis. Not even during prolonged and enhanced demand for extra heat production was any endogenous hormone or neurotransmitter able to recruit any UCP1-independent adaptive nonshivering thermogenic process in muscle or in any other organ, and no proteins other than UCP1-not even UCP2 or UCP3-therefore have the ability to mediate adaptive nonshivering thermogenesis in the cold.     

温度和高脂食物对黑线仓鼠代谢产热和体脂累积的影响

能量代谢的适应性调节是小型哺乳动物应对环境季节性变化的主要策略之一。为探讨不同温度下动物在代谢产热能量支出与脂肪累积之间的权衡策略,以成年雄性黑线仓鼠为研究对象开展了3 个实验:实验1 将动物驯化于高脂和低脂食物;实验2 将动物暴露于低温(5℃)和暖温(30℃);实验3 将饲喂高脂食物的动物暴露于低温。以食物平衡法测定摄食量、摄入能和消化率,以开放式氧气分析仪测定代谢产热,以索氏抽提法测定脂肪含量。结果发现,取食高脂食物的黑线仓鼠摄食量显著减少,但脂肪累积显著增加;暖温下摄食量显著减少,但体脂含量显著增加,低温下摄食量显著升高,但体脂含量显著减少;饲喂高脂食物的黑线仓鼠在低温下摄入能显著增加,非颤抖性产热增强,但体脂含量显著降低。结果表明高脂食物对黑线仓鼠体脂累积的影响与环境温度有关,低温诱导脂肪动员,暖温促进脂肪贮存;低温下黑线仓鼠增加能量摄入不能完全补偿用于产热的能量支出,导致脂肪动员增加;暖温下代谢产热降低是脂肪累积的主要因素;与能量摄入相比代谢产热的能量支出在体脂累积的适应性变化中发挥更重要的作用。