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。     

繁殖和运动对小型兽类褐色脂肪组织产热的影响

综述了近年来国内外9学者对小型兽类的繁殖、运动与能量平衡和褐色脂肪组织（BAT）产热关系的研究,大多数学者认为繁殖（尤其是哺乳）和运动能促进动物摄食量的增加,而降低BAT产热。这说明小型兽类为满足繁殖和运动过程中高能量的需求,除了大幅度增加能量摄入之入,还采取降低BAT产热以节约非哺乳能量消耗的策略。     

小型哺乳动物的持续能量收支限制研究进展

综述了小型哺乳动物持续能量收支限制的研究概况和进展。最大持续能量收支在决定物种的地理分布、生存适应、繁殖成功等方面都具有重要意义,但在许多条件下受到限制。食物的丰富度,或者动物自身的摄食、消化和吸收能力似乎不是主要限制因素。持续能量收支可能被外周组织和器官消耗能量的能力限制,即"外周限制"假说;或者机体的散热能力所限制,即"热耗散限制"假说。动物也可能通过衡量季节性繁殖投资的价值,实现最大繁殖输出,即"季节性投资"假说。尽管这些假说得到了一些研究的证实,但仍未阐明持续能量收支限制的机理。本文对相关研究的发展方向进行了展望。     

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。     

高蛋白食物和繁殖对布氏田鼠能量摄入和产热的影响

为探讨高蛋白食物和繁殖对布氏田鼠食物摄入和产热等特征的效应,将成年雌性布氏田鼠分为非繁殖对照食物组、非繁殖高蛋白组、繁殖对照食物组和繁殖高蛋白组。对照食物蛋白含量为17.7%,高蛋白食物的蛋白含量为36.6%。实验过程中测定动物的体重、食物摄入量、静止代谢率(RMR)、身体成分、内脏器官重量、褐色脂肪组织(BAT)解偶联蛋白1(UCP1)含量、血清瘦素和催乳素水平等。结果发现:高蛋白食物明显抑制布氏田鼠的体重,但动物在妊娠期和哺乳期,这种抑制作用消失。高蛋白食物明显抑制非繁殖组动物的干物质摄入、摄入能和消化能,但对哺乳期动物没有影响。高蛋白食物提高了非繁殖期和繁殖期动物的消化率,降低了血清瘦素浓度,但仅提高了繁殖期动物肾脏的重量,而降低了盲肠的重量。RMR、UCP1含量和血清催乳素浓度等则不受高蛋白食物的影响。繁殖期动物的体重、能量摄入、RMR和血清催乳素浓度等均高于非繁殖动物。这些结果表明,食物蛋白含量可影响布氏田鼠的能量代谢和产热特征等,且在繁殖期和非繁殖期有不同的反应方式。     

棕色田鼠消化道形态变化与能量需求的关系

为研究能量代谢与消化道形态结构变化及其某些生活史特征之间的关系，采用食物平衡法、耗氧量测定、形态测量和组织学方法，分别测定了雄性、非繁殖雌性及哺乳雌性棕色田鼠(Mirotus mandarinus)的摄食量、每日消化能、静止代谢率和胃肠器官长度、重量以及肠道各段管径和黏膜厚度。结果表明：哺乳雌鼠摄食量、每日消化能、静止代谢率高于非繁殖雌鼠和雄鼠，且消化道各器官有最大的重量、管径和黏膜厚度。由此可见，哺乳雌鼠能量需求增加，促使消化道形态结构进行一些有益的调节。棕色田鼠在哺乳期代谢率增加时，仅有消化道器官重量、黏膜厚度及小肠管径的明显变化，这可能与其哺乳期较长，胎仔数较少，生后生长缓慢等生活史特征有关。同时也说明在未受到十分严峻的能量胁迫的情况下，动物并不需付出昂贵的代价去增加消化器官的长度。     

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。     

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。     

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。     

剃毛和增加胎仔数对KM 小鼠哺乳期能量收支的影响

为探讨最大持续能量收支限制的因素和生理机理,本文测定了增加胎仔数和背部剃毛的KM 小鼠的哺乳
期摄食量和繁殖输出。基础代谢率(BMR)以封闭式流体压力呼吸计测定。结果发现,增加胎仔数对哺乳期体
重、摄食量、热传导率、BMR 和胎仔重的影响不显著;与非剃毛对照组相比,剃毛使哺乳高峰期日平均摄食量
增加了13.8% (P<0.001)、BMR 增加了18.1% (P< 0.01)、热传导增加了30.8% (P< 0.01),但剃毛对体
重、胎仔数和胎仔重的影响不显著;胎仔数与断乳时胎仔重显著正相关,与幼体平均体重显著负相关。BMR 与
母体体重和胎仔重显著正相关。这些结果表明,KM 小鼠哺乳期能量收支受到了限制,剃毛显著增强了KM 小鼠
的散热能力,但未影响繁殖输出;支持“外周限制假说”,不符合“热耗散限制假说”。     

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.     

Energy metabolism and the evolution of reproductive suppression in the human female

Reproduction places severe demands on the energy metabolism in human females. When physical work entails higher energy expenditure, not enough energy will be left for the support of the reproductive processes and temporal suppression of the reproductive function is expected. While energy needed for reproduction may be obtained by increases in energy intake, utilization of fat reserves, or reallocation of energy from basal metabolism, several environmental or physiological constraints render such solutions unlikely. For human ancestors increases in energy intake were limited by availability of food, by labor of food preparation and by metabolic ceilings to energy assimilation. Energy stored as fat may support only a fraction of the requirements for reproduction (especially lactation). Effects of intense physical activity on basal metabolism may also interfere with fat accumulation during pregnancy. Finally, the female physiology may experience demands on increasing the basal metabolism as a consequence of physical activity and, at the same time, on decreasing the basal metabolism, when energy to support the ongoing pregnancy or lactation is inadequate. The resulting metabolic dilemmas could constitute a plausible cause for the occurrence of reproductive suppression in response to physical activity. It is, therefore, likely that allocating enough energy to the reproductive processes during periods when energy expenditure rises may be difficult due to physiological and bioenergetic constraints. Females attempting pregnancy in such conditions may compromise their lifetime reproductive output. A reproductive suppression occurring in low energy availability situations may thus represent an adaptive rather then a pathological response.     

Limits to sustained energy intake IX: a review of hypotheses

Several lines of evidence indicate that animals in the wild may be limited in their maximal rates of energy intake by their intrinsic physiology rather than food availability. Understanding the limits to sustained energy intake is important because this defines an envelope within which animals must trade-off competing activities. In the first part of this review, we consider the initial ideas that propelled this area and experimental evidence connected with them. An early conceptual advance in this field was the idea that energy intake could be centrally limited by aspects of the digestive process, or peripherally limited at the sites of energy utilisation. A model system that has been widely employed to explore these ideas is lactation in small rodents. Initial studies in the late 1980s indicated that energy intake might be centrally limited, but work by Hammond and colleagues in the 1990s suggested that it was more likely that the limits were imposed by capacity of the mammary glands, and other works tended to support this view. This consensus, however, was undermined by studies that showed milk production was higher in mice at low temperatures, suggesting that the capacity of the mammary gland is not a limiting factor. In the second part of the review we consider some additional hypotheses that might explain these conflicting data. These include the heat dissipation limits hypothesis, the seasonal investment hypothesis and the saturated neural control hypothesis. Current evidence with respect to these hypotheses is also reviewed. The limited evidence presently available does not unambiguously support any one of them. Limits to sustained energy intake VIII: Król et al. (2003) J Exp Biol 206:4283-4291.     

Energy turnover in European hares is centrally limited during early,but not during peak lactation

We investigated metabolizable energy intake (MEI) and milk energy output in European hares throughout gestation and lactation in females raising three young, i.e., close to maximum litter size in this precocial species. We hypothesized that herbivorous hares may face a central limitation of energy turnover during lactation, imposed by maximum capacity of the gastrointestinal tract. Females were provided with low-energy or high-energy diets, either continually, or during lactation only. Unexpectedly, females on either diet reached identical peak MEIs (>6 times BMR) during late lactation, with females on low-energy diet increasing food intake proportionally. Thus, we reject our hypothesis that in lactating hares, peak MEI is centrally limited. During early lactation, MEI and milk transfer was, however, significantly impaired in females on the low-energy diet, indicating a temporal central limitation due to a time-lag caused by the readjustment of energy intake capacity. Importantly, irrespective of the diet, females significantly increased peak MEI late in the breeding season. Consequently, earlier in the season, when energy reserves are still high, energy throughput was not limited by physiological constraints at all. We conclude that extreme MEI may have fitness costs, and that females maximize lifetime reproductive success by actively down-regulating MEI whenever possible.     

Metabolic compensation during high energy output in fasting, lactating grey seals (Halichoerus grypus): metabolic ceilings revisited

Lactation is the most energetically expensive period for female mammals and is associated with some of the highest sustained metabolic rates (SusMR) in vertebrates (reported as total energy throughput). Females typically deal with this energy demand by increasing food intake and the structure of the alimentary tract may act as the central constraint to ceilings on SusMR at about seven times resting or standard metabolic rate (SMR). However, demands of lactation may also be met by using a form of metabolic compensation such as reducing locomotor activities or entering torpor. In some phocid seals, cetaceans and bears, females fast throughout lactation and thus cannot offset the high energetic costs of lactation through increased food intake. We demonstrate that fasting grey seal females sustain, for several weeks, one of the highest total daily energy expenditures (DEE; 7.4 x SMR) reported in mammals, while progressively reducing maintenance metabolic expenditures during lactation through means not explained by reduction in lean body mass or behavioural changes. Simultaneously, the energy-exported in milk is progressively increased, associated with increased lipoprotein lipase activity in the mammary gland, resulting in greater offspring growth. Our results suggest that females use compensatory mechanisms to help meet the extraordinary energetic costs of lactation. Additionally, although the concepts of SusMR and ceilings on total DEE may be somewhat different in fasting lactating species, our data on phocid seals demonstrate that metabolic ceilings on milk energy output, in general, are not constrained by the same kind of peripheral limitations as are other energy-consuming tissues. In phocid seals, the high ceilings on DEE during lactation, coupled with metabolic compensation, are undoubtedly important factors enabling shortened 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。     

Demographics of the European hare (Lepus europaeus) in the Mediterranean climate zone of Australia

Demographic parameters of the European hare (Lepus europaeus) in southern Australia were investigated by dissecting hares shot by hunters during each month of the year. Gender, body weight, age, sucking, lactation, weight of the abdominal alimentary canal, weight of the left peri-renal fat body, pregnancy status, presence and counts of placental scars, litter size, and stage of gestation were recorded. From those data, growth rates, age at weaning, age and weight at puberty, date of conception, projected birth date, recruitment, survivorship, and the relationships between lactation and fat stores and alimentary capacity were determined.Fecundity of the southern Australian hares followed the seasonal pattern reported for northern hemisphere populations. However, output was lower per female and particularly per older female. Females began breeding at an earlier age such that recruitment into the southern Australian population was more dependent on females in their first year of life than on older females. Growth rates were comparable with European rates. Although high chill factors were apparently associated with higher leveret mortality, there was paradoxically higher overall mortality during the spring-early summer period of higher plant growth than in the late summer–winter period of lower plant growth and more extreme weather conditions. Fat was accumulated during pregnancy and would act as a buffer against the possibility of inadequate food availability during lactation, but hares increased the capacity of the alimentary canal during lactation and presumably with it their ability to assimilate energy to meet the demands of lactation.