恒温脊椎动物最大持续能量代谢率及其研究进展

研究最大持续代谢率具有重要的理论和实践意义,本文概述了近年来国外对脊椎动物最大持续代谢率研究的成果,概括介绍了限制最大持续代谢率的四个假说：（1）食物限制假说（food limit hypothesis) ,(2)外周限制假说（peripheral limit hypotheisis),(3)中心限制假说（central limit hypothesis)和（4）对称性形态构成假说（symmorphosis hypothesis),分析了静止代谢率随持续代谢率的增加而增加的机理,讨论了最大持续代谢率对动物的生态进化意义,最后指出了最大持续代谢率研究中可能的发展前景。     

中华蟾蜍静止代谢率与高耗能器官大小的关系

摄食增量假说认为,持续能量支出与静止代谢率正相关,且需要较大的高耗能器官(心脏、肝脏、肾脏和小肠等)与其匹配.因此,该假说预测静止代谢率与高耗能器官质量正相关.该预测在脊椎动物种间研究中得到普遍支持,但在种内研究中的结论尚不一致.在外温动物中的种内验证尤其缺乏.本研究以中华蟾蜍Bufo gargari-zans为研究对...     

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。     

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。     

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.     

不同温度下黑线仓鼠应对食物短缺的能量学对策

食物资源的不确定性是动物在自然环境中面临的重要挑战之一."代谢率转换"假说认为,动物应对食物短缺的能量学策略在于降低代谢率以减少能量支出.然而在不同环境温度下非冬眠小型哺乳动物应对食物短缺的"代谢率转换"策略,尚不明确.为探究这一问题,将成年雄性黑线仓鼠在低温(5.0℃)、室温(21.0℃)和高温(32.5℃)下断食处...     

植物群落物种共存机制:负密度制约假说

负密度制约假说主要描述由于资源竞争、有害生物侵害(比如病原微生物、食草动物捕食)等, 同种个体之间发生的相互损害行为; 它主要强调同种个体之间的相互作用, 解释自然群落物种共存的机理; 负密度制约机制主要在小尺度上降低群落内同种个体生长率, 同时提高个体死亡率, 从而为其他物种的生存提供空间和资源, 促进物种共存。目前负密度制约假说的检验研究主要侧重密度制约、距离制约、群落补偿效应等三个方面。最近, 研究者又探讨了近缘物种之间由于对相似资源的竞争所产生的负效应, 扩展了负密度制约假说, 进而提出异群保护假说和谱系多样性制约假说。负密度制约假说引起生态学家长久的探讨和关注, 世界范围内大尺度森林动态样地的建立, 又为探索不同尺度上密度制约效应的研究提供了条件。然而目前的研究仍然存在不足, 比如负密度制约假说的检验受到其他因素的干扰、区域研究不平衡等。因此, 生态学家们仍然怀疑负密度制约效应调节群落物种共存的重要性, 但是目前的研究还没有发现否定负密度制约假说的充分证据。     

光照条件、植株冠层结构和枝条寿命的关系——以桂花和水杉为例

根据衰老理论的代谢率假说,生物寿命与其代谢率有关,个体大小相同的生物体,在质量较好的微生境中通常比较差生境中具有更高的代谢速率。因此,生物体在资源供给较差的生境中通常比资源供给较充足的生境中具有更长的寿命。枝条是木本植物植冠构建的基本单元之一,如果枝条遵循代谢率假说,则可推测在光照较好环境下的植物枝条或小枝将比其在遮荫环境下具有更短的寿命,即枝条寿命与光照条件成反比。以常绿物种桂花(Osmanthus fragrans)和落叶物种水杉(Metasequoiaglyptostroboides)为研究对象,通过测量不同光照环境下,植株大小(株高和胸径)、冠层深度、冠层轮廓(冠层深度/冠层宽度)、相对冠层宽度(冠层宽度/植株高度)以及植株凋落枝条寿命等性状,探讨了光照条件对成年植株冠层形态结构和植株枝条寿命的影响。调查发现:1)枝条的寿命在遮荫条件下显著高于全光照条件下,与理论预测吻合;2)随遮荫程度增加,植株冠层深度和冠层轮廓增加,相对冠层宽度减小;3)枝条的平均寿命与植株冠层深度和冠层轮廓成正比,与植株相对冠层宽度成反比。这表明光照条件可能通过改变植株冠层结构来影响枝条寿命。未来需要进一步研究枝条生物量分配、叶片光合能力和呼吸速率在不同生活型物种之间的差异,以便更全面的理解枝条寿命与生境质量之间的关系。     

日粮水平对鲇鱼日总代谢,特殊动力作用和活动代谢的影响

在２５℃不同日粮水平（饥饿─４％体重／ｄ）条件下,采用封闭式呼吸仪测定了３１尾鲇鱼（体重７２．９─１３３．３ｇ）的日总代谢率,然后,将鱼限制在呼吸室中的限动笼内测定了其中２０尾鲇鱼的静止日总代谢率,并计算出了鲇鱼的特殊动力作用（ＳＤＡ）和活动代谢率。日总代谢率和活动代谢率都随日粮水平的升高呈“Ｖ”形变化,分别在约１％和２％体重／ｄ的日粮水平时最低,ＳＤＡ的能量支出占摄入能量的２２．１４％。     

横断山脉大绒鼠最大代谢率的季节性差异

最大代谢率对于动物的生存、繁殖和分布具有关键作用。实验用开放式呼吸仪测定了大绒鼠(Eothenomys miletus)夏季和冬季的冷诱导最大代谢率和运动最大代谢率。大绒鼠夏季的冷诱导最大代谢率O2为(7.24±0.61)ml/(g.h),运动最大代谢率O2为(7.69±0.59)ml/(g.h);冬季的冷诱导最大代谢率O2为(8.61±0.42)ml/(g.h),运动最大代谢率O2为(7.51±0.51)ml/(g.h)。冷诱导最大代谢率冬夏之间具有差异显著,而运动最大代谢率则差异不显著。结果表明:栖息于横断山脉的大绒鼠由于受到低温的胁迫,导致其冷诱导最大代谢率季节性变化较大,而运动最大代谢率则相对稳定。     

The quantitative genetics of sustained energy budget in a wild mouse

We explored how morphological and physiological traits associated with energy expenditure over long periods of cold exposure would be integrated in a potential response to natural selection in a wild mammal, Phyllotis danwini. In particular, we studied sustained energy expenditure (SusMR), the rate of expenditure fueled by concurrent energy intake, basal metabolic rate (BMR), and sustained metabolic scope (SusMS = SusMR/BMR), a measure of the reserve for sustained work. We included the masses of different central processing organs as an underlying factor that could have a mechanistic link with whole animal traits. Only the liver had heritability statistically different from zero (0.73). Physiological and morphological traits had high levels of specific environmental variance (average 70%) and postnatal common environmental variance (average 30%) which could explain the low heritabilities estimates. Our results, (1) are in accordance with previous studies in mammals that report low heritabilities for metabolic traits (SusMR, BMR, SusMS), (2) but not completely with previous ones that report high heritabilities for morphological traits (masses of central organs), and (3) provide important evidence of the relevance of postnatal common environmental variance to sustained energy expenditure.     

A seasonal difference of daily energy expenditure in a free-living subterranean rodent,the silvery mole-rat (Heliophobius argenteocinereus; Bathyergidae)

In seasonal climatic regimes, animals have to deal with changing environmental conditions. It is reasonable to expect that seasonal changes are reflected in animal overall energetics. The relation between daily energy expenditure (DEE) and seasonally variable ecological determinants has been studied in many free-living small mammals; however with inconsistent results. Subterranean mammals, i.e. fossorial (burrowing) mammals which live and forage underground, live in a seasonally and diurnally thermally stable environment and represent a suitable model to test seasonality in DEE in respect to seasonal changes, particularly those in soil characteristics and access to food supply. Both factors are affected by seasonal rainfall and are supposed to fundamentally determine activity of belowground dwellers. These ecological constraints are pronounced in some tropical regions, where two distinct periods, dry and rainy seasons, regularly alternate. To explore how a tropical mammal responds to an abrupt environmental change, we determined DEE, resting metabolic rate (RMR) and sustained metabolic scope (SusMS) in a solitary subterranean rodent, the silvery mole-rat, at the end of dry season and the onset of rainy season. Whereas RMR did not differ between both periods, mole-rats had 1.4 times higher DEE and SusMS after the first heavy rains. These findings suggest that rainfall is an important environmental factor responsible for higher energy expenditure in mole-rats, probably due to increased burrowing activity. SusMS in the silvery mole-rat is comparable to values in other bathyergids and all bathyergid values rank among the lowest SusMS found in endothermic vertebrates.     

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.     

Energetic costs of activity by lizards in the field

1. The available data related to the activity energetics of lizards in the field were collated with respect to three indices of activity energetics: the ecological cost of transport (ECT) expressed as a percentage of the total energy, the proportion of total energy used in all forms of activity (%AR), and the sustained metabolic scope (SusMS), defined as the ratio of the total energy expenditure to the total resting metabolism.
2. The ECT values of lizards ranged from 3 to 36% with five of 11 species having values >20%. The percentage AR ranged from 23 to 80% for lizards during active seasons, with most species having values > 50%. The SusMS ranged from 1·1 to 5·1.
3. Values of ECT are higher for lizards than for mammals, in part because the costs of maintenance metabolism are higher in mammals.
4. The percentage AR and SusMS values of mammals are higher than those of lizards.
5. It follows from the previous two points that the proportion of the total daily energy that is expended in non-locomotory activities is disproportionately higher in mammals compared with lizards.
6. The energy expended in locomotion is a significant portion of the energy budget of lizards. This is generally true for all seasons in which there is activity.     

Seasonal variation in the metabolic rate and body composition of female grey seals: fat conservation prior to high-cost reproduction in a capital breeder?

Many animals rely on stored energy through periods of high energy demand or low energy availability or both. A variety of mechanisms may be employed to attain and conserve energy for such periods. Wild grey seals demonstrate seasonal patterns of energy storage and foraging behaviour that appear to maximize the allocation of energy to reproduction—a period characterized by both high energy demand and low food availability. We examined seasonal patterns in resting rates of oxygen consumption as a proxy for metabolic rate (RMR) and body composition in female grey seals (four adults and six juveniles), testing the hypothesis that adults would show seasonal changes in RMR related to the reproductive cycle but that juveniles would not. There was significant seasonal variation in rates of resting oxygen consumption of adult females, with rates being highest in the spring and declining through the summer months into autumn. This variation was not related to changes in water temperature. Adults increased in total body mass and in fat content during the same spring to autumn period that RMR declined. RMR of juveniles showed no clear seasonal patterns, but did increase with increasing mass. These data support the hypothesis that seasonal variation in RMR in female grey seals is related to the high costs of breeding.     

Genetic correlation between resting metabolic rate and exploratory behaviour in deer mice (Peromyscus maniculatus)

According to the 'pace-of-life' syndrome hypothesis, differences in resting metabolic rate (RMR) should be genetically associated with exploratory behaviour. A large number of studies reported significant heritability for both RMR and exploratory behaviour, but the genetic correlation between the two has yet to be documented. We used a quantitative genetic approach to decompose the phenotypic (co)variance of several metabolic and behavioural measures into components of additive genetic, common environment and permanent environment variance in captive deer mice. We found significant additive genetic variance for two mass-independent metabolic measures (RMR and the average metabolic rate throughout the respirometry run) and two behavioural measures (time spent in centre and distance moved in a novel environment). We also detected positive additive genetic correlation between mass-independent RMR and distance moved (r(A) = 0.78 ± 0.23). Our results suggest that RMR and exploratory behaviour are functionally integrated traits in deer mice, providing empirical support for one of the connections within the pace-of-life syndrome hypothesis.     

Improved Equations for Estimating the Resting Metabolic Rate

New models were developed for predicting the resting metabolic rate (RMR) with sufficient accuracy for use in epidemiologic studies and for weight control of individuals. For this purpose, the RMR of 213 women and 76 men was measured, and physical measurements were taken. The RMR was regressed on correlates of RMR, avoiding harmful degrees of collinearity by rejecting interregressor correlations exceeding r=0.5. For women, the best model (R₂=0.71) included the regressors age, race, weight, pulse rate, smoking, and body temperature. The best model for males (R2 = 0.81) included age, race, weight, blood pressure, smoking, time (of day the RMR was measured), and whether subjects had a meal prior to calorimetry. The models were cross validated internally and also validated using an external database. In both cases, the mean estimated RMR did not differ significantly from the measured RMR. The accuracy of the models was compared with four models reported in the literature, three of which overestimated the RMR by up to 17%. In conclusion, improved RMR prediction models have been developed, more accurate than existing models, rendering them suitable for application to epidemiological databases and for individual weight control programs.     

Moult and basal metabolic costs in males of two subspecies of stonechats: the European Saxicola torquata rubicula and the East African S. t. axillaris

The circannual patterns in resting metabolic rate (RMR) of males of two subspecies of stonechats, the European Saxicola torquata rubicula and the East African S. t. axillaris, are compared. As the birds from the two subspecies were raised and kept under comparable laboratory conditions, differences in metabolic rate between the two subspecies had to be genetically determined. RMR peaked during moult in both subspecies. During the rest of the year RMR was fairly constant in both subspecies and assumed to reflect basal metabolic rate (BMR). African stonechats had a 22% lower mass specific BMR than European stonechats, which is thought to reflect a genetical physiological adaptation to the differences in environmental circumstances they experience in the field. A low BMR makes an animal more susceptible to cold. Hence, the relatively high plumage mass in the African compared to the European stonechat may be functionally linked to its relatively low BMR. Moult costs, calculated from the plumage masses and the differences in RMR inside and outside the moult period, tended to be higher in the European compared to the African stonechats. These data and an interspecific comparison of moult costs over various species of birds support the earlier notion by Lindström et al. (1993) that moult costs are more closely linked with BMR than with body mass or rate of moult. The relation between moult costs and BMR and the fact that the efficiency of moult is extremely low (3.8 and 6.4% for European and African stonechats, respectively) suggest that the maintenance of specific tissues necessary for moult is a large cost factor. Alternatively, impaired insulation during moult may necessitate an increased metabolic capacity which may be associated with an increased RMR.