繁殖经历对黑线仓鼠哺乳期能量收支的影响

为探讨繁殖经历与哺乳期最大持续能量收支的关系,对连续4 次繁殖的黑线仓鼠哺乳期的能量收支情况进行了测定。结果显示:1)不同繁殖组哺乳高峰期的摄食量、泌乳能量支出(MEO)、胎仔数和胎仔重差异不显著,静止代谢率(RMR)、非颤抖性产热(NST)、褐色脂肪组织(BAT)线粒体细胞色素c 氧化酶(COX)活性、血清甲状腺激素(T3 、T4 )和催乳素水平也无明显变化;2)摄食量与MEO、胎仔重和RMR 呈显著正相关。结果表明,不同繁殖经历的黑线仓鼠主要通过降低产热和增加能量摄入来满足哺乳高峰期的能量需求;哺乳期最大持续代谢率(SusMR)可能受乳腺组织泌乳能力的限制,与“外周限制假说” 的预测一致,不支持“中心限制假说”;SusMR 限制因素和哺乳期能量收支策略可能与繁殖经历无关。     

小型啮齿动物的繁殖能量代价

繁殖是动物向后代传递和保持遗传信息的方式。因此繁殖的意义是显而易见的,但也需要付出代价。主要代价是能量需求增加。在对小家鼠繁殖能量需求的研究中发现,能量摄入在妊娠期只是稍微增加,而在哺乳期则急剧增加。尽管在妊娠期增加的幅度很小,但这可能反映了消化道和发育的胎儿之间在动物腹中的空间竞争,从而可能使能量摄入受到限制进而影响到繁殖过程。哺乳期间,能量摄入急剧增加,在哺乳后期达到高峰并趋于稳定。对野生鼠的研究也表明,野生鼠妊娠期和哺乳期的能量摄入模式与小家鼠是基本相同的,这样我们在小家鼠研究工作中的发现就具有更普遍的适应意义。对哺乳后期能量摄入的限制机制研究至少已经进行了15 年。能量摄入受消化道消化能力的限制(中心限制假说)或者受乳腺泌乳能力限制(外周限制假说) 的假说,都不能合理地解释一些现有的结果。我们提出了一个新的假说,即能量摄入可能受啮齿动物散热能力的限制(热耗散限制假说)。很久以来,一直认为散热能力是对大型哺乳动物哺乳的一个限制因素,但它在小型啮齿动物中的意义尚不清楚。传统观点认为,啮齿动物哺乳期对褐色脂肪组织产热水平的调节是为了重新分配能量以满足哺乳所需;但现在看来,实际上可能是动物为了避免体温过高而降低其基本的产热水平。我们在这个领域已经有了一些进展,但要利用这些知识来理解即使很简单的生活史权衡等问题也还有很多的工作需要做。     

暖温经历对雌性黑线仓鼠能量代谢、产热和体脂含量的影响

能量代谢的生理调节是小型哺乳动物应对不同环境温度的重要策略之一,为探讨暖温下代谢产热在体重和体脂适应性调节中的作用和机理,本研究将雌性黑线仓鼠(Cricetulus barabensis)暴露于暖温(30°C)1个月、3个月和4个月,测定体重、摄入能、代谢产热、体脂含量、褐色脂肪组织(BAT)细胞色素c氧化酶(COX)活性和解偶联蛋1 (UCP1) mRNA表达等。结果显示,暖温对黑线仓鼠体重无显著影响,但使脂肪含量显著增加。与室温组相比(21°C),暖温组消化率显著升高,但摄入能和消化能显著降低;暖温下非颤抖性产热(NST)显著降低,脑、肝脏和心脏COX活性、BAT COX活性和UCP1 mRNA的表达显著下调。结果表明,暖温下降低代谢产热补偿了能量摄入的减少,机体处于正能量平衡状态,是脂肪含量显著增加的主要原因之一。脑、肝脏、心脏和BAT代谢活性降低是代谢产热降低的主要机制,与脂肪累积有关。     

高原鼠兔褐色脂肪组织及肝脏产热的季节性变化

非颤抖性产热(nonshtverlng thermogenesis, NST)是小哺乳动物在低温环境下增加产热的有效途径(Foster等1978.Can JPhyslol,56:110~122)。NST主要产生于褐色脂肪组织(BAT),所有影响BAT结构与功能的因素均可影响NST能力(Cannon等,1993．Life in the cold．369～379)。 高原鼠兔(Ochotona curzoniae)为高寒草甸生态系统中非冬眠的小哺乳类动物,其生态学特征已被进行了广游酊探^的研究．王德华等(1990。兽类学报10(1)：40～53; 1993．兽类学报13(4)：271～276)对高原鼠兔NST能力,BAT组织成分及线粒体蛋白的季节性变化进行了研究。本文在此工作的基础上。试图从细胞承千上进一步探讨高原鼠兔产热的季节性变化及适应意义。     

黑线仓鼠自身生存和繁殖输出间的权衡不受温度影响

  为阐明野生小型哺乳动物哺乳期能量收支对策,深入理解最大持续能量摄入（SusEI）限制的因素和机理,测定了不同环境温度下（21℃、30℃和5℃）与哺育不同胎仔数（自然胎仔数Con、减少Minus和增加胎仔数Plus）的黑线仓鼠哺乳期体重、摄食量、基础代谢率（BMR）、非颤抖性产热（NST）,以及褐色脂肪组织（BAT）细胞色素C氧化酶（COX）活性、血清T₃、T₄和催乳素水平。结果显示,哺乳期体重显著降低,摄食量显著增加,21℃和30℃组间差异不显著。最大持续摄食量约为14g/d,Minus组比Con和Plus组低20.3%和18.6%。温度对摄食量的影响显著,5℃下摄食量达16g/d,比21℃和30℃组高14%（P<0.05）。Con和Minus组胎仔数维持稳定,而Plus组胎仔数显著降低,断乳时Con和Plus组胎仔数差异不显著。Minus组胎仔重显著低于Con和Plus组。断乳时Minus组平均幼体体重比Con和Plus组分别高17.9%和24.9%（P<0.05）。5℃下BMR、NST、COX活性、血清T₃、T₄和催乳素水平显著高于21℃和30℃,而21℃和30℃组间差异不显著。结果表明：黑线仓鼠SusEI水平为5×BMR,低温下可通过增加能量摄入应对代谢产热的能量支出,在自身维持和繁殖输出之间采取了“权衡分配”的能量学策略,研究结果支持热耗散限制假说,也符合外周限制假说的预测。     

冷驯化对中缅树鼩褐色脂肪组织适应性产热的影响

目的探讨环境温度对中缅树鼩体重、褐色脂肪组织(BAT)产热活性及解偶联蛋白1含量的影响,为建立树鼩肥胖模型提供理论依据。方法将40只体重相似的成年中缅树鼩随机分为5组(每组8只):对照组(0 d),置于(25±1)℃,12 L/12 D条件下饲养;以及置于(5±1)℃,12 L/12 D条件下分别驯化7、14、21 d和28d组。驯化结束后测定动物的体重、非颤抖性产热(NST)、褐色脂肪组织重量以及解偶联联蛋白1(UCP1)的含量。结果与对照组(0 d)相比,冷驯化组中缅树鼩的体重、NST、BAT重量以及UCP1含量都显著增加,BAT颜色也明显加深,冷驯化28 d后,体重增加了26.32%,NST增加了20.65%,BAT重量增加了53.85%,UCP1含量增加了43%。UCP1含量与BAT重量和NST显著正相关。结论中缅树鼩可能通过冷驯化诱导BAT组织增生和UCP1表达上调,从而增强BAT产热活力以增加能量支出,推测BAT可能作为以能量学途径治疗肥胖的靶器官。     

环境温度和繁殖经历对黑线仓鼠哺乳期能量收支的影响

为探讨环境温度和繁殖经历对黑线仓鼠哺乳期能量收支的影响,将连续3次繁殖的黑线仓鼠暴露于温度梯度降低的条件下(30—0℃,1℃/4d),使初次、第2和3次繁殖的动物分别暴露于30—20℃、20—10℃、10—0C℃,测定了哺乳期能量收支。与初次繁殖的动物相比,第3次繁殖组动物的摄食量显著增加,静止代谢率、非颤抖性产热、褐色脂肪组织细胞色素c氧化酶(COX)活性和血清T3水平显著增加,而断乳时胎仔重显著降低。结果表明:(1)低温下繁殖的黑线仓鼠处于负能量平衡,在自身维持和哺育后代的能量分配之间存在权衡,低温下产热增加,繁殖输出减少;(2)黑线仓鼠可能感知环境温度的变化,在连续降低温度的条件下降低繁殖投资,符合"季节性投资假说"的预测。     

光照和温度对根田鼠褐色脂肪组织产热能力的影响

测定了根田鼠在室内驯化2周后褐色脂肪组织（BAT）的蛋白含量、线粒体蛋白含量和细胞色素C氧化酶活性的变化。动物进行如下4种处理:23℃,16L:8D;23℃,8L：16D;5℃,16L：8D和5℃,8L:16D.。结果表明,短光照可刺激根田鼠的BAT线粒体细胞色素C氧化酶的活性增加;低温驯化可导致根田鼠的BAT线粒体蛋白含量增加;短光照加低温可进一步促进酶的活性增加。结合个体水平的产热特点,说明根田鼠在产热能力的季节驯化过程中,光周期是一主要的季节调节信号,环境温度进一步增强光照的作用,环境温度和光周期共同作用以诱导其产热能力等方面的生理调节。研究结果支持了环境温度和光周期共同作用以调节动物产热变化的学说。     

同步遥测分析大鼠棕色脂肪产热与体温昼夜节律变化的时间关系

目的：同步遥测棕色脂肪组织（BAT）产热与体核温度昼夜律变化的时间曲线,分析二者昼夜节律变化的时间关系。方法：实验用成年雄性SD大鼠,在22℃环境温度下,明暗时间各12h,昼光时间为06：00h-18：00h,同步无线遥测体核温度（TC）、BAT温度（T队T）、腋窝温度（Tax）和动物活动的昼夜节律变化。结果：①在昼光中,TBAT较TC低0．67％,而在暗光中二者则相似。大鼠从昼光进入暗光时,TBAT升高的速率较TC升高速率快,开始上升的时间较TC提前8min;而从暗光进入昼光时,TBAT开始下降的时间则较TC提前4min。②Tax的昼夜节律幅度与TC相似,但无论动物在明光期或暗光期中,Tax均低于同步测量的TC。③从昼光期转入暗光期时,动物的行为活动出现增加反应先于TBAT和TC开始上升的时间。结论：实验结果证明,在暗光期中大鼠TC升高与BAT产热增加有关,说明BAT昼夜节律性产热的变化在维持体温昼夜生理节律中有重要的作用。     

冷驯化条件下高山姬鼠能量对策的初步研究

为进一步了解横断山区小型哺乳动物的能量代谢对策,本文主要研究了横断山的外来物种——高山姬鼠(Apodemus chevrieri)持续冷暴露28d的能量变化。结果表明,随着冷驯化时间延长,体重和体温降低;摄入能、消化能和可代谢能增加,至21d后这3项指标又下降;BMR和NST增加,并且28d后它们分别是对照组的187.19%和155.97%;(NST-BMR)/BMR比值7d时增加到1.02,21d后下降至0.53,并保持稳定。因此,在持续冷暴露下,高山姬鼠采取适当降低体重和体温、增加能量摄入、增加BMR和NST产热等途径来维持能量代谢平衡,但是NST在产热中的作用逐渐降低。这种能量模式很可能代表了一种外来小型啮齿动物对于在横断山生存独特的能量适应策略。     

Adaptive character of metabolism in Eothenomys miletus in Hengduan Mountains region during cold acclimation

Environmental factors play an important role in the seasonal adaptation of body mass and thermogenesis in small, wild mammals. The purpose of the present study was to test the hypothesis that ambient temperature was a cue to trigger the seasonal adjustments in body mass, energy intake, uncoupling protein 1 (UCP1) in brown adipose tissue (BAT), and other biochemical characteristics of Eothenomys miletus during 49 days of cold exposure. Our data demonstrated that cold acclimation induced a remarkable decrease in body mass, a significant increase in energy intake and metabolic rate, and high expression of UCP1 in BAT of E. miletus. Biochemical characteristics of BAT and liver respiration were also increased following cold acclimation. These data suggest that E. miletus reduced the body mass and increased energy intake and expenditure under cold acclimation. Increased expression of UCP1 was potentially involved in the regulation of energy metabolism and thermogenic capacity following cold acclimation.     

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).     

黑线仓鼠的BMR个体差异及其应对高脂食物的能量学对策

为探讨高脂食物对小型哺乳动物能量代谢的影响及其与基础代谢率(Basal metabolic rate, BMR)的关系,将成年雌性黑线仓鼠(Cricetulus barabensis)分为高、低BMR组,每组再随机分为低脂、高脂食物组,驯化6周后,测定体重、摄入能和代谢率,以及消化酶活力、褐色脂肪组织(Brown adipose tissue, BAT)和主要内脏器官与肌肉的细胞色素c氧化酶(Cytochrome c oxidase, COX)活性、解偶联蛋白(Uncoupling protein, UCP) mRNA表达等。结果显示,高脂食物对高、低BMR组动物体重均无显著影响。与低脂食物组相比,高脂食物组的摄食量、摄入能和消化能显著下降,小肠脂肪酶活力显著增强,消化率明显增加,但高、低BMR组的组间差异不显著。夜间代谢水平显著高于昼间,高脂食物使高BMR组的夜间代谢率显著升高。BAT、肌肉和内脏器官COX活性不受高脂食物的影响,高、低BMR组的组间差异也不显著。高脂食物组仅肝脏UCP2表达显著上调。结果表明,能量摄入和消化系统形态及功能的可塑性调节是黑线仓鼠应对高脂食物的主要策略;黑线仓鼠的代谢率具有显著的昼夜节律,既受高脂食物的影响,也与动物自身的BMR水平有关,但UCP表达具有组织特异性,这可能不是导致BMR个体差异的因素。     

Adaptive Activation of Thyroid Hormone and Energy Expenditure

The mechanisms by which thyroid hormone accelerates energy expenditure are poorly understood. In the brown adipose tissue (BAT), activation of thyroid hormone by type 2 iodothyronine deiodinase (D2) has been known to play a role in adaptive energy expenditure during cold exposure in human newborns and other small mammals. Although BAT is not present in significant amounts in normal adult humans, recent studies have found substantial amounts of D2 in skeletal muscle, a metabolically relevant tissue in humans. This article reviews current biological knowledge about D2 and adaptive T3 production and their roles in energy expenditure.     

Effects of exercise training on brown adipose tissue thermogenesis in ovariectomized obese rats

The effect of exercise training on brown adipose tissue (BAT) thermogenesis was studied by measuring cytochrome oxidase activity, as a marker of mitochondrial abundance, mitochondrial guanosine-5'-diphosphate (GDP) binding, as an indicator of thermogenic activity and oxygen consumption in BAT in ovariectomized (OVX) obese rats and sham-operated rats. Six-week exercise training significantly suppressed body weight gain in OVX rats to the level of sedentary control rats, although food intake in exercise trained OVX rats increased more than in the sedentary OVX rats. Exercise training increased cytochrome oxidase activity, mitochondrial GDP binding and oxygen consumption in BAT in OVX rats, which were reduced in a sedentary condition, as well as in the control rats. These results suggest that exercise training potentiates BAT thermogenesis, which may contribute to the reduction of body weight in OVX obese rats.     

Lactation in the tammar wallaby (Macropus eugenii). II. Intake of milk components and maternal allocation of energy

The intake of milk components (total solids, carbohydrate, protein, lipid, energy) by suckling tammar wallabies ( Macropus eugenii ), from peak lactation to independence, was measured using a double-isotope dilution technique and chemical analysis of milk samples. The time of peak intake of milk solids (day 256 of lactation or 1126 g of offspring weight) was similar to that for whole milk. Peak intake of carbohydrate occurred earlier than this (235 days) and peak intakes of protein and lipid occurred later (262 days and 266 days, respectively). Intake of gross energy peaked at 262 days and represented a maternal yield of about 207 KJ.Kg^-0.75.d^-1. This is much lower than peak lactational energy yields in most other mammals, but the duration of lactation is longer in tammars than in other mammals. Total output of energy in milk by tammar mothers was 63 MJ, and this would require an intake of about 98 MJ of metabolizable energy in food. This requirement, which is equivalent to 21 MJ/kg of maternal weight, is similar to those calculated for sheep and cattle, suggesting that there are not large differences between marsupial and placental herbivores in terms of weight-related allocation of energy to reproduction. We suggest that a strategy of minimizing the peak energetic demand of lactation may be an important adaptation for a small, primarily grazing mammal due to size-related physiological constraints on elevating herbage intake and the increasing risk from predators when grazing time is increased.     

Uncoupling proteins and thermoregulation.

Energy balance in animals is a metabolic state that exists when total body energy expenditure equals dietary energy intake. Energy expenditure, or thermogenesis, can be subcategorized into groups of obligatory and facultative metabolic processes. Brown adipose tissue (BAT), through the activity of uncoupling protein 1 (UCP1), is responsible for nonshivering thermogenesis, a major component of facultative thermogenesis in newborn humans and in small mammals. UCP1, found in the mitochondrial inner membrane in BAT, uncouples energy substrate oxidation from mitochondrial ATP production and hence results in the loss of potential energy as heat. Mice that do not express UCP1 (UCP1 knockouts) are markedly cold sensitive. The recent identification of four new homologs to UCP1 expressed in BAT, muscle, white adipose tissue, brain, and other tissues has been met by tremendous scientific interest. The hypothesis that the novel UCPs may regulate thermogenesis and/or fatty acid metabolism guides investigations worldwide. Despite several hundred publications on the new UCPs, there are a number of significant controversies, and only a limited understanding of their physiological and biochemical properties has emerged. The discovery of UCP orthologs in fish, birds, insects, and even plants suggests the widespread importance of their metabolic functions. Answers to fundamental questions regarding the metabolic functions of the new UCPs are thus pending and more research is needed to elucidate their physiological functions. In this review, we discuss recent findings from mammalian studies in an effort to identify potential patterns of function for the UCPs.     

Different physiological roles of serum leptin in the regulation of energy intake and thermogenesis between pregnancy and lactation in primiparous Brandt's voles (Lasiopodomys brandtii)

Reproduction, especially lactation, is associated with major metabolic adaptive changes. In this study, we investigated the metabolic changes and the roles of leptin during different periods of reproduction in primiparous Brandt's voles (Lasiopodomys brandtii). Energy intake, thermogenic capacity and serum leptin levels were examined in non-reproductive, mid pregnant, late pregnant, early lactating and peak lactating voles. Voles increased body mass by nearly 70% during late pregnancy compared to the non-breeding controls. The increase in body mass was mainly due to the increase in body fat mass which increased by 56%, and the growth of the reproductive tissues and digestive organs. Lactating voles decreased body fat by nearly 27% at peak lactation compared to the controls, and 53% compared to late pregnant voles. At the same time they increased food intake significantly. Uncoupling protein 1 (UCP1) content in brown adipose tissue (BAT) decreased significantly at peak lactation. Serum leptin increased significantly in the mid pregnancy, at a time when there was no increase in body fat, and remained at this high level in late pregnancy. Leptin levels decreased after parturition and reached a nadir at peak lactation. Serum leptin was negatively correlated with energy intake during lactation, but not during pregnancy. These data suggest that Brandt's voles adjust energy intake, thermogenic capacity and body reserves to match the high energy demands for reproduction. Hyperleptinemia, without decreased energy intake suggests a state of leptin resistance during pregnancy, and hypoleptinemia during lactation might act as a signal to stimulate energy intake.