黑线仓鼠繁殖输出与基础代谢率的关系

为了解黑线仓鼠繁殖输出与基础代谢率(BMR)的关系,阐明最大持续能量收支(SusMR)的限制水平, 揭示哺乳期能量收支对策,本文测定了哺乳期黑线仓鼠的体重、摄食量、BMR 和身体组成,以及哺乳期的胎仔数、胎仔重和泌乳能量支出(MEO)。结果显示,黑线仓鼠哺乳期体重降低了15.0 ± 0.8% , 摄食量显著增加, 哺乳高峰期平均摄食量为13.9 ± 0.3 g /d, 摄入能为222.1 ± 5.3 kJ/ d, 比哺乳初期增加121% , 比对照组高288% ;哺乳高峰期MEO 为62.4 ± 2.3 kJ/ d, 哺乳末期BMR 为49.7 ± 1.1 kJ/ d; 断乳时平均胎仔数4.7 ± 0.2、窝胎仔重50.5 ±1.6 g; 哺乳末期BMR 比对照组增加48% ,BMR 与消化系统各器官的相关性高于对照组; BMR 与胎仔数、胎仔重、乳腺重量和MEO 显著正相关。结果表明:初次繁殖的黑线仓鼠哺乳期SusMR 限制为4.47 ×BMR, 在自身维持和繁殖输出之间采取了“权衡分配”的原则,通过体重降低以减少BMR 的增加幅度, 从而有利于繁殖输出。     

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。     

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。     

小鼠基础代谢率与繁殖输出和繁殖期末器官的关系

以封闭式流体压力呼吸计测定KM小鼠(Mus musculus)的基础代谢率(BMR);采用残差分析和Pearson相关分析检验BMR与繁殖输出、内脏器官的相关性。哺乳末期BMR显著高于繁殖前,繁殖前BMR与繁殖输出不相关,但哺乳末期BMR与体重、摄食量、胎仔数和胎仔重、内脏器官和消化道显著正相关;与消化道器官的相关性高于其他内脏器官。研究结果支持"哺乳期较高的BMR有利于消化系统增强消化和吸收能力,以增加能量摄入用于繁殖输出"的假设。     

剃毛对大绒鼠持续能量摄入的影响

持续能量摄入(Sus EI)的上限对于哺乳期母体的生存非常重要。先前的研究表明大绒鼠Eothenomys miletus Sus EI的限制可能受到外周限制假说,而不支持热散失假说。为了再次验证这2个假说,本研究在哺乳早期对大绒鼠剃毛,测定剃毛组和对照组的体质量、食物摄入量、静止代谢率、胎仔数、胎仔质量和泌乳输出。结果表明:剃毛可以显著增加大绒鼠的食物摄入量和静止代谢率,但是对于胎仔数和泌乳输出没有影响。所有结果不支持热散失假说,而支持外周限制假说,说明大绒鼠在哺乳期的Sus EI可能受到乳腺泌乳能力的限制。暗示在泌乳高峰期Sus EI的限制在不同物种间的反应可能是不一样的。     

随机饥饿和重喂食对小鼠能量代谢和行为的影响

为阐明能量代谢和行为的可塑性对动物适应食物资源变化的意义,将成年雄性KM 小鼠随机饥饿驯化4
周,再重喂食驯化4 周。采用食物平衡法测定摄食量、封闭式流体压力呼吸计测定基础代谢率(BMR) 和非颤
抖性产热(NST)、观察法测定行为。随机饥饿使摄食量、消化道重量显著增加,BMR 和NST 显著降低。与对照
组相比,饥饿组休息行为显著增加,活动显著降低。重喂食后,上述指标均恢复到对照组水平,表现出显著的
可塑性。研究结果表明,动物适应难以预测的食物资源短缺的主要策略包括:增加自由取食期间的摄食量;降
低BMR、NST 和活动行为,从而保存身体贮存的能量。能量代谢和活动行为在较短的时间尺度内表现出显著的
可塑性对小鼠适应不可预测的食物资源短缺的应激环境具有重要意义。     

温度和增加胎仔数对大绒鼠哺乳期能量代谢的影响

为了阐明温度和增加胎仔数对大绒鼠Eothenomys miletus哺乳期能量代谢的影响,本研究测定了在不同温度条件下增加胎仔数(比正常胎仔数多),大绒鼠的摄食量、胎仔数、胎仔质量、静止代谢率、非颤抖性产热和乳腺质量。结果表明:不同胎仔数对大绒鼠的摄食量、胎仔质量和产热能力没有影响。低温增加了母体的摄食量和产热能力,断奶时低温组的胎仔质量显著低于常温组。低温组和常温组大绒鼠的乳腺质量差异无统计学意义。研究结果说明低温可以增加大绒鼠摄食量,但是对乳腺质量没有影响,表明大绒鼠哺乳期的持续能量摄入上限可能受到乳腺分泌乳汁的限制,支持外周限制假说。     

随机限食和重喂食小鼠能量收支和生长发育的可塑性

为阐明能量收支和生长发育的可塑性对动物适应食物资源变化的作用和意义,将断乳后的雄性KM小鼠40只随机限食4周,再重喂食4周。采用封闭式流体压力呼吸计测定基础代谢率(BMR)。限食使摄食量显著增加,BMR和活动行为降低,胴体和生殖腺重量显著降低。重喂食后上述指标均恢复到对照组水平,表现出显著的可塑性变化。结果表明,KM小鼠能通过摄食量、BMR、活动行为和身体组成的可塑性调节以适应难以预测的食物资源变化。     

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。     

黑线仓鼠断乳后能量代谢和脂肪累积的适应性调节

小型哺乳动物能量代谢和脂肪累积的适应性调节是其应对自然环境变化的主要能量学策略,但在不同的生活史阶段,脂肪组织适应性调节的特征和能量机理尚不清楚。为探讨不同繁殖阶段能量代谢和脂肪累积的变化及其内分泌机理,本文测定了黑线仓鼠哺乳期和断乳后摄食量、脂肪重量,以及血清瘦素水平、下丘脑瘦素受体（Ob-Rb）和相关神经肽的基因表达。结果显示,哺乳高峰期黑线仓鼠的脂肪重量几乎降低至零,断乳后显著增加;与非繁殖对照组相比,皮下脂肪、肾周脂肪与腹腔脂肪重量分别增长了1.5倍、37.1倍和1.9倍。断乳后摄食量、血清瘦素水平显著高于非繁殖对照组,Ob-Rb基因表达显著下调,而促食与抑食神经肽的基因表达均未发生显著变化。哺育不同胎仔数的黑线仓鼠在断乳后能量摄入、静止代谢率、身体组分未出现显著差异。研究表明,在不同的繁殖阶段脂肪累积呈现显 著的适应性调节,瘦素抵抗是断乳后脂肪累积补偿性增长的重要内分泌机制之一。这对迅速恢复脂肪累积,以应对将来的能量需求增加或者食物资源短缺的环境,进而提高自身的适合度具有重要意义。     

Large litter size increases maternal energy intake but has no effect on UCP1 content and serum-leptin concentrations in lactating Brandt’s voles (<Emphasis Type="Italic">Lasiopodomys brandtii</Emphasis>)

Lactation is the most energetically demanding period in the female mammal's life. We measured maternal energy intake, uncoupling protein 1 (UCP1) content in brown adipose tissue (BAT), serum-leptin concentration, and litter growth in lactating Brandt's voles (Lasiopodomys brandtii) with different litter sizes. Litter mass was positively related to litter size but there was no difference in pup mass at birth. Maternal gross energy intake at peak lactation was positively correlated with litter size and litter mass. Maternal resting metabolic rate (RMR) was positively correlated with litter mass, but not with litter size. No significant differences were detected in body-fat mass, serum-leptin concentration, or UCP1 in lactating voles with different litter sizes. Serum-leptin concentration was negatively correlated with energy intake during lactation. Our data did not support the hypothesis that there is a trade-off in energy allocation between maternal maintenance and offspring growth in lactating Brandt's voles, but support the idea that if the mothers with ten pups should have less energy available for their maintenance than mothers raising fewer pups. Also, leptin is probably not the only factor that induces the high energy intake in mothers with large litter sizes, although it was involved in the regulation of energy intake during lactation.     

Limits to sustainable energy budget during lactation in the striped hamster (Cricetulus barabensis) raising litters of different size

A female animal appears to approach an upper limit to the rate of sustained energy intake/metabolic rate (SusEI/MR) during lactation. However, different species of animals may respond differently to the sustainable limit. Here, we measured energy budget during lactation in female striped hamsters raising litters of natural size (Con), and females whose litter size was manipulated during early lactation to support fewer or more pups (minus pups, MP or plus pups, PP). The striped hamsters significantly decreased their body mass and increased food intake from early to late lactation; and MP females had lower weight loss and food intake than the control and PP females. Litter size of the PP group decreased significantly over the period of lactation, and pups were weaned at a similar weight to that of the controls. MP females supported a significantly lower litter mass throughout lactation compared with the control and PP females, but during late lactation the pups from the MP group were significantly heavier. Resting metabolic rate (RMR) did not differ significantly between the three groups and the gross energy intake during peak lactation was 5.0×, 4.2× and 5.0 × RMR for the control, MP and PP females, respectively. Female striped hamsters reached a plateau in food intake at around 14 g/d during peak lactation, which might signify a limit of SusEI at 5.0 × RMR. However, it was not possible to determine whether the limitation on SusEI was imposed centrally by the capacity of the gastrointestinal tract to process food, peripherally by the capacity of the mammary gland to produce milk, or by the capacity of animals to dissipate heat.     

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。     

Reproductive effort during gestation and lactation by Richardson's ground squirrels

Summary Reproductive effort by yearling and older female Richardson's ground squirrels was studied over a 4-year period in southern Alberta by obtaining serial weight records from marked individuals to compare the mother's mass at critical points in the annual cycle (emergence from hibernation, estrus, parturition, and litter emergence) with her litter's mass at birth and weaning. Yearlings weighed only 80% of older adults at emergence from hibernation, but they mated at the same time as older females, attained adult mass coincident with pregnancy, and weaned litters that were not significantly smaller in size or mass than those of older females. Age and maternal mass were weak predictors of litter size and litter mass. Of the net increase in mass of the combined mother-litter unit during gestation, over half (60% of 139 g for yearlings; 52% of 127 g for older females) was attributable to an increase in the mother's own mass, whereas during lactation almost all of the net increase (93% of 545 g for yearlings; 96% of 567 g for older females) was attributable to an increase in the litter's mass. On a daily basis, deposition of mass in the litter was 6 times greater during lactation than gestation. On average, neonates weighed 2.3% (6.5 g) of maternal mass at birth and 23.1% (81 g) at emergence from the natal burrow; offspring masses at birth and at emergence were significantly negatively correlated with litter size. On average, litters weighed 16.3% (48 g) of maternal mass at birth and 157.5% (578 g) at emergence from the natal burrow. Compared with other hibernating sciurids, Richardson's ground squirrels have a similar offspring mass relative to maternal mass both at birth and at emergence from the natal burrow. However, because of the large litter size (typically 6–8), absolute reproductive effort, measured either as litter mass at birth or at natal emergence, is large for the body size of the species.     

The functional significance of individual variation in basal metabolic rate

Basal metabolic rate (BMR) was established as a common reference point allowing comparable measures across different individuals and species. BMR is often regarded as a minimal rate of metabolism compatible with basic processes necessary to sustain life. One confusing aspect, however, is that BMR is highly variable, both within and between species. A potential explanation for this variability is that while individuals with high BMRs may suffer the disadvantage of having to feed for longer to cover the extra energy demands, this may be offset by advantages that accrue because of the high metabolic rate. One suggested advantage is that high levels of BMR are a consequence of maintaining a morphology that permits high rates of the maximal sustained rate of metabolism (SusMR)--the rate of metabolism that can be sustained for days or weeks. We have been studying the energetics of MF1 laboratory mice during peak lactation to investigate this idea. In this article, we review some of our work in connection with three particular predictions that derive from the hypothesised links among morphology, basal metabolism, and sustained metabolic rate. By comparing groups of individuals, for example, lactating and nonlactating individuals, the patterns that emerge are broadly consistent with the hypothesis that BMR and SusMR are linked by morphology. Lactating mice have bigger organs connected with energy acquisition and utilisation, greater resting metabolic rates in the thermoneutral zone, called RMRt (approximately equivalent to BMR), and high sustainable rates of maximal energy intake. However, when attempts are made to establish these relationships across individuals within lactating mice, the associations that are anticipated are either absent or very weak and depend on shared variation due to body mass. At this level there is very little support for the suggestion that variation in RMRt (and thus BMR) is sustained by associations with SusMR.