动物长期能量收支理论及研究进展

阐述了持续代谢率和持续代谢范围的概念,并介绍有关制约因子的４种假说：食物可利用性假说、外周制约假说、中心制约假说以及同形态椹制约假说。此外,还就持续代谢率的进化原因、实验验证的方法和结论,以及与最小维持消耗（ＢＭＲ）的关系和机制进行了分析。持续代谢率对动物生活具有影响,具有生态学意义。最后,分析了本领域未来的４个发展方向。     

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。     

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。     

长爪沙鼠的代谢率与器官的关系

我们测定了野生长爪沙鼠（Meriones unguiculatus)的基础代谢率和冷诱导的最大代谢率,分析了动物体内11种器官或组织的大小与代谢率的关系。长爪沙鼠的基础代谢率为118.10mlO2/h,最大代谢率为659.83mlO2/h。经过残差分析表明,基础代谢率并不与任何一种器官或组织相关,而最大代谢率与小肠湿重（n=20,r=-0.478,P=0.033)和消化道全长（n=20,r=-0.487,P=0.030)显著相关,表明体内器官重量的差别并不是造成种内基础代谢率差别的原因;体内存在着与最大代谢率相关的“代谢机器”,消化系统（特别是小肠）是这一代谢机器的重要组成部分,但代谢机器的大小并不能通过基础代谢率反映出来。基础代谢率与最大代谢率不相关,因此不支持“较高的基础代谢率能够产生较高的非基础代谢率（最大代谢率等）”的假设。     

高原鼠兔和根田鼠的最大代谢率

采用He-O2混合诱导方法测定了高原鼠兔（Ochotona curzoniae）和根田鼠（Microtus oeconomus）的最大代谢率（MMR）。高原鼠兔的最大代谢率在夏季和冬季分别为5．93（体重为118g）和6．33（体重为115．8g）mlO2/g.h,而根田鼠在夏季和冬季分别为12．70（体重为27．8g）和18．29（体重为17．5g）mlO2/g.h,并且根田鼠的最大代谢率的变化幅度大于高原鼠兔,存在种间差异。MMR季节变化的不显著性来源于动物环境温度在夏季较低,而在冬季为越冬亦采用行为调节等其它机制。栖于青藏高原的高原鼠兔和根田鼠同时受到低温和低氧的腔迫,而两者对最大代谢率作用相反,导致两种动物的最大代谢率与各自期望值相比差异不大。     

温度和光周期对高原鼠兔和根田鼠最大代谢率的影响

研究了光周期和温度对高原鼠兔（Ochotona curzoniae)和根田鼠（Microtus oeconomus)最大代谢率（MMR）的影响。结果表明在高原鼠兔中,光周期对MMR的影响较小,而低温可显著地导致MMR的增加（P＜0．01）,由此提出温度是高原鼠兔MMR季节性变化的主要生理信号;在根田鼠中,光周期和温度对MMR的影响皆达到显著水平（P＜0．01）,短光照和低温都可导致MMR的增加。     

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。     

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。     

当年生高山姬鼠最大代谢率

用开放式呼吸仪测定了当年生和往年生高山姬鼠的冷诱导和运动诱导最大代谢率。当年生高山姬鼠的冷诱导最大代谢率为:(7.25±0.25)mLO2/(g.h),运动诱导最大代谢率为(9.22±0.27)mLO2/(g.h);当年生鼠冷诱导最大代谢率雌雄间无显著差异,运动诱导最大代谢率雌雄间差异显著。当年生高山姬鼠的冷诱导最大代谢率低于往年个体,运动诱导最大代谢率高于往年生个体。结果表明:当年生高山姬鼠忍受寒冷的能力低于往年个体,可能是对横断山地区年平均气温较低的自然环境的胁迫产生的适应;而雄性运动诱导最大代谢率大于雌性,可能是为了适应避免近亲繁殖和激烈竞争而在性成熟后从出生地向外扩散。     

Encephalization and the evolution of longevity in mammals

Allometric principles account for most of the observed variation in maximum life span among mammals. When body-size effects are controlled for, most of the residual variance in mammalian life span can be explained by variations in brain size, metabolic rate and body temperature. It is shown that species with large brains for a given body size and metabolic rate, such as anthropoid primates, also have long maximum life spans. Conversely, mammals with relatively high metabolic rates and low levels of encephalization, as in most insectivores and rodents, tend to have short life spans. The hypothesis is put forward that encephalization and metabolic rate, which may govern other life history traits, such as growth and reproduction, are the primary determinants directing the evolution of mammalian longevity.     

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.     

Do the high energy lifestyles of shorebirds result in high maximal metabolic rates? Basal and maximal metabolic rates in least and pectoral sandpipers during migration

Shorebirds have high resting and field metabolic rates relative to many other bird groups, and this is posited to be related to their high‐energy lifestyle. Maximum metabolic outputs for cold or exercise are also often high for bird groups with energetically demanding lifestyles. Moreover, shorebirds demonstrate flexible basal and maximal metabolic rates, which vary with changing energy demands throughout the annual cycle. Consequently, shorebirds might be expected to have high maximum metabolic rates, especially during migration periods. We captured least Calidris minutilla and pectoral C. melanotos sandpipers during spring and fall migration in southeastern South Dakota and measured maximal exercise metabolic rate (MMR; least sandpipers only), summit metabolic rate (M_sum, maximal cold‐induced metabolic rate) and basal metabolic rate (BMR, minimum maintenance metabolic rate) with open‐circuit respirometry. BMR for both least and pectoral sandpipers exceeded allometric predictions by 3–14%, similar to other shorebirds, but M_sum and MMR for both species were either similar to or lower than allometric predictions, suggesting that the elevated BMR in shorebirds does not extend to maximal metabolic capacities. Old World shorebirds show the highest BMR during the annual cycle on the Arctic breeding grounds. Similarly, least sandpiper BMR during migration was lower than on the Arctic breeding grounds, but this was not the case for pectoral sandpipers, so our data only partially support the idea of similar seasonal patterns of BMR variation in New World and Old World shorebirds. We found no correlations of BMR with either M_sum or MMR for either raw or mass‐independent data, suggesting that basal and maximum aerobic metabolic rates are modulated independently in these species.     

Explorative behaviour is not associated with metabolism in the European Pied Flycatcher Ficedula hypoleuca

The pace‐of‐life syndrome hypothesis (POLS) represents an attractive theoretical framework suggesting that physiological and behavioural traits have evolved together with environmental conditions and life‐history strategies. POLS predicts that metabolic differences covary with behavioural variation such that high metabolic rate is associated with risk‐prone behaviour and a faster pace‐of‐life, whereas a low metabolic rate is associated with risk‐averse behaviour and a slower pace‐of‐life. We tested the POLS hypothesis in captive European Pied Flycatchers during their first year by examining the relationship between explorative behaviour and basal metabolic rate. Our results are inconsistent with POLS. The positive association of explorative behaviour with basal metabolic rate was not recovered for either sex, possibly due to foraging conditions in the aviaries where control and trial groups were fed twice a day, the birds' young age, developmental plasticity, or a non‐existent syndrome.     

Sources and significance of variation in basal,summit and maximal metabolic rates in birds

The rates at which birds use energy may have profound effects on fitness, thereby influencing physiology, behavior, ecology and evolution. Comparisons of standardized metabolic rates (e.g., lower and upper limits of metabolic power output) present a method for elucidating the effects of ecological and evolutionary factors on the interface between physiology and life history in birds. In this paper we review variation in avian metabolic rates [basal metabolic rate (BMR; minimum normothermic metabolic rate), ...     

Gluttony in migratory waders – unprecedented energy assimilation rates in vertebrates

Maximum energy assimilation rate has been implicated as a constraint on maximal sustained energy expenditure, on biomass production, and in various behavioural and life history models. Data on the upper limit to energy assimilation rate are scarce, and the factors that set the limit remain poorly known. We studied migratory waders in captivity, given unlimited food supply around the clock. Many of these waders assimilated energy at rates of seven to ten times basal metabolism, exceeding maximum rates reported for vertebrates during periods of high energy demand, for example during reproduction and in extreme cold. One factor allowing the high energy assimilation rates may be that much of the assimilated energy is stored and not concomitantly expended by muscles or other organs. The remarkable digestive capacity in waders is probably an adaptation to long and rapid migrations, putting a premium on high energy deposition rates. The upper limit to daily energy assimilation in vertebrates is clearly higher than hitherto believed, and food availability, total daily feeding time and, possibly, the fate of assimilated energy may be important factors to take into account when estimating limits to energy budgets in animals.     

Individual variation and repeatability of the aerobic performance in Brandt's voles (Lasiopodomys brandtii)

Metabolism is thought to play an important role in shaping behaviour, ecology and physiology in animals. To study the changes of metabolism among different ages or generations as well as the repeatability during the ontogeny, we carried out the research in Brandt's voles (Lasiopodomys brandtii), which covered two generations’ life. Meanwhile, we estimated the among-family variations to facilitate the heritability evaluation. Resting metabolic rate within the thermoneutral zone, resting metabolic rate at 5 °C, non-shivering thermogenesis and maximal metabolic rate during thermogenesis in both juveniles and adults were simultaneously measured. Population-average values of aerobic traits were generally consistent among different ages or generations; however, there was no repeatability at the level of individual variation during the ontogeny, which indicated that the aerobic traits of the young were not good indicators for that of later life. At the same time, the coefficient of intraclass correlation for full sibs failed to reach statistical significance, suggesting that heritability of aerobic traits in Brandt's voles was not high.     

Fluctuating selection on basal metabolic rate

BMR (Basal metabolic rate) is an important trait in animal life history as it represents a significant part of animal energy budgets. BMR has also been shown to be positively related to sustainable work rate and maximal thermoregulatory capacity. To this date, most of the studies have focused on the causes of interspecific and intraspecific variation in BMR, and fairly little is known about the fitness consequences of different metabolic strategies. In this study, we show that winter BMR affects local survival in a population of wild blue tits (Cyanistes caeruleus), but that the selection direction differs between years. We argue that this fluctuating selection is probably a consequence of varying winter climate with a positive relation between survival and BMR during cold and harsh conditions, but a negative relation during mild winters. This fluctuating selection can not only explain the pronounced variation in BMR in wild populations, but will also give us new insights into how energy turnover rates can shape the life‐history strategies of animals. Furthermore, the study shows that the process of global warming may cause directional selection for a general reduction in BMR, affecting the general life‐history strategy on the population level.     

The evolution of gigantism in active marine predators

A novel hypothesis to better understand the evolution of gigantism in active marine predators and the diversity of body sizes, feeding strategies and thermophysiologies of extinct and living aquatic vertebrates is proposed. Recent works suggest that some aspects of animal energetics can act as constraining factors for body size. Given that mass-specific metabolic rate decreases with body mass, the body size of active predators should be limited by the high metabolic demand of this feeding strategy. In this context, we propose that shifts towards higher metabolic levels can enable the same activity and feeding strategy to be maintained at bigger body sizes, offering a satisfactory explanation for the evolution of gigantism in active predators, including a vast quantity of fossil taxa. Therefore, assessing the metabolic ceilings of living aquatic vertebrates and the thermoregulatory strategies of certain key extinct groups is now crucial to define the energetic limits of predation and provide quantitative support for this model.