Factors influencing the turnover and net isotopic discrimination of hydrogen isotopes in proteinaceous tissue: experimental results using Japanese quail

Stable hydrogen isotopes (δ(2)H) are commonly used in studies of animal movement. Tissue that is metabolically inactive after growth (e.g., feathers) provides spatial or dietary information that reflects only the period of tissue growth, whereas tissues that are metabolically active (e.g., red blood cells) provide a moving window of forensic information. However, using δ(2)H for studies of animal movement relies on the assumption that tissue δ(2)H values reflect dietary δ(2)H values, plus or minus a net diet-tissue discrimination value, and that the turnover rate is known for metabolically active tissue. The metabolic rate of an animal may influence both diet-tissue discrimination values and isotopic tissue turnover rate, but this hypothesis has not been tested experimentally. To examine the metabolic hypothesis, an experimental group of 12 male and 15 female captive Japanese quail (Coturnix japonica) was housed at 8.9°C for 90 d to elevate their metabolic rates (mL CO(2) min(-1)), and a control group of 12 male and 13 female quail was housed at room temperature during the same period. For both experimental and control birds, diet-tissue discrimination values were estimated for red blood cells and feathers. To determine turnover rate, experimental and control birds were switched from a (2)H-enriched diet to a (2)H-depleted diet, with red blood cells sampled before and after diet switch. Metabolic rate did not influence red blood cell hydrogen isotope turnover rate (η(2)(p) = 0.24)) or diet-feather isotope discrimination values (η(2)(p) = 0.86). Diet-feather hydrogen isotopic discrimination had a significant sex plus treatment interaction effect; female feathers were depleted in (2)H relative to food regardless of treatment, whereas male feathers were enriched in (2)H. The effect of sex suggested that experimental studies should examine whether coeval males and females differ in blood δ(2)H levels during certain periods of the annual cycle.     

Effects of elemental composition on the incorporation of dietary nitrogen and carbon isotopic signatures in an omnivorous songbird

The use of stable isotopes to infer diet requires quantifying the relationship between diet and tissues and, in particular, knowing of how quickly isotopes turnover in different tissues and how isotopic concentrations of different food components change (discriminate) when incorporated into consumer tissues. We used feeding trials with wild-caught yellow-rumped warblers (Dendroica coronata) to determine delta15N and delta13C turnover rates for blood, delta15N and delta13C diet-tissue discrimination factors, and diet-tissue relationships for blood and feathers. After 3 weeks on a common diet, 36 warblers were assigned to one of four diets differing in the relative proportion of fruit and insects. Plasma half-life estimates ranged from 0.4 to 0.7 days for delta13C and from 0.5 to 1.7 days for delta15N . Half-life did not differ among diets. Whole blood half-life for delta13C ranged from 3.9 to 6.1 days. Yellow-rumped warbler tissues were enriched relative to diet by 1.7-3.6% for nitrogen isotopes and by -1.2 to 4.3% for carbon isotopes, depending on tissue and diet. Consistent with previous studies, feathers were the most enriched and whole blood and plasma were the least enriched or, in the case of carbon, slightly depleted relative to diet. In general, tissues were more enriched relative to diet for birds on diets with high percentages of insects. For all tissues, carbon and nitrogen isotope discrimination factors increased with carbon and nitrogen concentrations of diets. The isotopic signature of plasma increased linearly with the sum of the isotopic signature of the diet and the discrimination factor. Because the isotopic signature of tissues depends on both elemental concentration and isotopic signature of the diet, attempts to reconstruct diet from stable isotope signatures require use of mixing models that incorporate elemental concentration.     

同位素富集-稀释法研究食性转变对鱼类不同组织N同位素转化率的影响

稳定同位素技术广泛地用于描绘生态系统中食物网的食物来源和营养级关系,但是消费者不同组织转化率的研究相对较少。通过锦鲤摄食人工添加15N蓝藻的食性转化实验,研究不同组织N同位素转化率的差异,探讨组织生长和代谢对同位素转化的相对贡献,为不同时间尺度的稳定同位素研究取样奠定基础。结果表明,通过42d的加富蓝藻饲喂,各组织的N稳定同位素发生显著变化。肝的δ15N为(19.3±1.4)‰,显著高于其它组织,其次为鱼鳍((15.6±1.0)‰)和血液((12.6±0.4)‰),肌肉的δ15N‰最低,为(9.9±0.7)‰。在随后的同位素稀释实验中,锦鲤的体重增加,相对生长速率为0.011d-1,鳍肉的转化率最快,达到11.4%/d,半衰期仅为6.1d,其次是血液和肝,肌肉的转化率最低,仅有3.8%/d,半衰期最长,为18.4d。代谢衰减指数c和-1不存在显著差异,表明锦鲤各组织的N同位素转化主要由组织生长引起。结论显示,同位素富集-稀释法可以有效评价鱼类食性转变对不同组织同位素转化的差异,鳍肉和血液同位素分析可以作为锦鲤食性转变快速追踪的手段。     

The maximum oxygen consumption and aerobic scope of birds and mammals: getting to the heart of the matter

Resting or basal metabolic rates, compared across a wide range of organisms, scale with respect to body mass as approximately the 0.75 power. This relationship has recently been linked to the fractal geometry of the appropriate transport system or, in the case of birds and mammals, the blood vascular system. However, the structural features of the blood vascular system should more closely reflect maximal aerobic metabolic rates rather than submaximal function. Thus, the maximal aerobic metabolic rates of birds and mammals should also scale as approximately the 0.75 power. A review of the literature on maximal oxygen consumption and factorial aerobic scope (maximum oxygen consumption divided by basal metabolic rate) suggests that body mass influences the capacity of the cardiovascular system to raise metabolic rates above those at rest. The results show that the maximum sustainable metabolic rates of both birds and mammals are similar and scale as approximately the 0.88 +/- 0.02 power of body mass (and aerobic scope as approximately the 0.15 +/- 0.05 power), when the measurements are standardized with respect to the differences in relative heart mass and haemoglobin concentration between species. The maximum heart beat frequency of birds and mammals is predicted to scale as the -0.12 +/- 0.02 power of body mass, while that at rest should scale as -0.27 +/- 0.04.     

Relative longevity and field metabolic rate in birds

Metabolism is a defining feature of all living organisms, with the metabolic process resulting in the production of free radicals that can cause permanent damage to DNA and other molecules. Surprisingly, birds, bats and other organisms with high metabolic rates have some of the slowest rates of senescence begging the question whether species with high metabolic rates also have evolved mechanisms to cope with damage induced by metabolism. To test whether species with the highest metabolic rates also lived the longest I determined the relationship between relative longevity (maximum lifespan), after adjusting for annual adult survival rate, body mass and sampling effort, and mass-specific field metabolic rate (FMR) in 35 species of birds. There was a strongly positive relationship between relative longevity and FMR, consistent with the hypothesis. This conclusion was robust to statistical control for effects of potentially confounding variables such as age at first reproduction, latitude and migration distance, and similarity in phenotype among species because of common phylogenetic descent. Therefore, species of birds with high metabolic rates senesce more slowly than species with low metabolic rates.     

Metabolic and organ mass responses to selection for high growth rates in the domestic chicken (Gallus domesticus)

Evolutionary hypotheses suggest that higher rates of postembryonic development in birds should either lower the resting metabolic rate (RMR) in a trade-off between the costs of growth and maintenance or increase RMR because of a buildup of metabolic machinery. Furthermore, some suggest that higher rates of postembryonic development in birds should reduce peak metabolic rate (PMR) through delayed tissue maturation and/or an increased energy allocation to organ growth. We studied this by comparing metabolic rates and organ sizes of fast-growing meat-type chickens (broilers) with those of birds from a laying strain, which grow much slower. During the first week of life, despite growing six times faster, the RMR of the broiler chickens was lower than that of birds of the laying strain. The difference between strains in RMR disappeared thereafter, even though broilers continued to grow twice as fast as layers. The differences between strains in growth rate during the first week after hatching were not reflected in similar differences in the relative masses of the heart, liver, and small intestine. However, broilers had heavier intestines once they reached a body mass of 80 g. In contrast, broilers had relatively smaller brains than did layers. There was a positive correlation, over both strains, between RMR and the masses of leg muscles, intestine, and liver. Furthermore, despite delayed maturation of muscle tissue, broilers exhibited significantly higher PMR. We hypothesize that a balance between the larger relative muscle mass but lower muscle maturation level explains this high PMR. Another correlation, between leg muscle mass and PMR, partly explained the positive correlation between RMR and PMR.     

Assessing temporal and spatial trends in estuarine nutrient dynamics using a multi-species stable isotope approach

Coastal urbanisation can alter estuarine nutrient dynamics through the input of point-source and diffuse pollutants, and nutrient concentrations can be highly influenced by seasonal and episodic rainfall and river flow. Understanding of both the spatial and temporal variability of nutrient dynamics is therefore critical to managing these estuaries. This can be achieved by periodically analysing the stable isotopes a range of aquatic taxa with variable nutrient turnover rates, mobility and distribution within the estuary. In two subtropical urban estuaries with different land use patterns, we analysed the carbon and nitrogen stable isotopes of phytoplankton, shrimp, prawns and fish at various proximities to pollution sources in dry and wet seasons. The fast nutrient turnover rates and ubiquity of phytoplankton in the estuary resulted in stable isotopes varying over fine-scale spatial scales, particularly in relation to proximity to point-source pollution. The slower nutrient turnover rates and localised habitat use of prawns, resulted in stable isotopes varying over larger spatial (between pollution sources) and temporal (seasonal) scales. The much slower nutrient turnover rates and high mobility throughout the estuary of fish resulted in stable isotopes varying over very large-scale spatial scales (between estuaries). These results illustrate a wide range of spatial and temporal changes to estuarine nutrient dynamics in subtropical urban estuaries in relation to rainfall conditions and nutrient inputs. This research also highlights the application of stable isotopes in assessing estuarine trophodynamics, and provides direction on the types of organisms that should be used to assess different spatial and temporal trends.     

The intraspecies relationship between tissue turnover and metabolic rate in rats

Ecologists interested in studying fluctuating relationships between consumers and nutrient sources are increasingly involved in modeling the rate at which consumers incorporate dietary components. In mammals a correlation between resting metabolic rate (RMR) and tissue turnover may exist across a range of species. Less is known about the variation of tissue turnover rate within a species, and how that correlates with RMR. Here we examine two strains of rats (Rattus norvegicus) with different RMR to test whether variation in RMR is positively correlated with tissue turnover rate within a species. If RMR, a relatively simple measurement, can be correlated with tissue turnover, then this relationship could be used to better interpret ecological functions, including impact of migratory or seasonally available nutrient sources. Here, the changing isotope signature in rat whole blood was modeled using a modified exponential decay equation and a reaction progress variable model. The modeled rate of turnover, metabolic rate (O₂ consumed), and mass were then compared between strains of rats. The mass and RMRs (conditions during which RMRs were determined modified from the ideal, as outlined in the Methods) were significantly different between strains, but half-life and the metabolic tissue replacement component of turnover (as opposed to turnover from mass gain) were not. No significant correlation was found between RMR and metabolic tissue replacement between the strains. Results suggest that within a species showing a range of RMRs, blood tissue turnover should not vary significantly.     

Development of maximum metabolic rate and pulmonary diffusing capacity in the superprecocial Australian Brush Turkey Alectura lathami: an allometric and morphometric study.

The Australian Brush Turkey Alectura lathami is a member of the Megapodiidae, the mound-building birds that produce totally independent, "superprecocial" hatchlings. This study examined the post-hatching development of resting and maximal metabolic rates, and the morphometrically determined changes in pulmonary gas exchange anatomy, in chicks during 3.7 months of growth from hatchlings (122 g) to subadults (1.1 kg). Allometric equations of the form y=aM(b) related gas exchange variables (y) to body mass (M, g). Metabolic rates were measured with open-flow respirometry (mL O2 min(-1)) of chicks resting in the dark and running above the aerobic limit on a treadmill. Resting metabolic rate (RMR=0.02 M(0.99)) and maximal metabolic rate (MMR=0.05 M(1.07)) scaled with exponents significantly above those of interspecific allometries of adult birds. However MMR was below that expected for other species of adult birds in flapping flight, consistent with the Brush Turkey's ground-dwelling habits. Total lung volumes (mL) increased faster than isometrically (V(L)=0.0075 M(1.19)), as did the surface area (cm(2)) of the blood-gas barrier (S(t)=7.80 M(1.23)), but the data overlapped those of adult species. Harmonic mean thickness of the blood-gas barrier was independent of body size (mean tau(ht),=0.39 microm) and was about twice that expected for flying birds. Diffusing capacity (mL O2 min(-1) kPa(-1)) of the blood-gas tissue barrier increased faster than isometrically (Dto2=0.049 M(1.23)); in hatchling Brush Turkeys, it was about 30% expected for adult birds, but this difference disappeared when they became subadults. When compared to altricial Australian pelicans that hatch at similar body masses, superprecocial Brush Turkeys had higher MMR and higher Dto2 at the same body size. A parallel allometry between MMR and Dto2 in Brush Turkeys and pelicans is consistent with the concept of symmorphosis during development.     

Development of maximum metabolic rate and pulmonary diffusing capacity in the superprecocial Australian Brush Turkey Alectura lathami: an allometric and morphometric study

The Australian Brush Turkey Alectura lathami is a member of the Megapodiidae, the mound-building birds that produce totally independent, "superprecocial" hatchlings. This study examined the post-hatching development of resting and maximal metabolic rates, and the morphometrically determined changes in pulmonary gas exchange anatomy, in chicks during 3.7 months of growth from hatchlings (122 g) to subadults (1.1 kg). Allometric equations of the form y=aM(b) related gas exchange variables (y) to body mass (M, g). Metabolic rates were measured with open-flow respirometry (mL O2 min(-1)) of chicks resting in the dark and running above the aerobic limit on a treadmill. Resting metabolic rate (RMR=0.02 M(0.99)) and maximal metabolic rate (MMR=0.05 M(1.07)) scaled with exponents significantly above those of interspecific allometries of adult birds. However MMR was below that expected for other species of adult birds in flapping flight, consistent with the Brush Turkey's ground-dwelling habits. Total lung volumes (mL) increased faster than isometrically (V(L)=0.0075 M(1.19)), as did the surface area (cm(2)) of the blood-gas barrier (S(t)=7.80 M(1.23)), but the data overlapped those of adult species. Harmonic mean thickness of the blood-gas barrier was independent of body size (mean tau(ht),=0.39 microm) and was about twice that expected for flying birds. Diffusing capacity (mL O2 min(-1) kPa(-1)) of the blood-gas tissue barrier increased faster than isometrically (Dto2=0.049 M(1.23)); in hatchling Brush Turkeys, it was about 30% expected for adult birds, but this difference disappeared when they became subadults. When compared to altricial Australian pelicans that hatch at similar body masses, superprecocial Brush Turkeys had higher MMR and higher Dto2 at the same body size. A parallel allometry between MMR and Dto2 in Brush Turkeys and pelicans is consistent with the concept of symmorphosis during development.     

Availability of water affects renewal of tissues in migratory blackcaps during stopover

Migrating blackcaps (Sylvia atricapilla) were used to test the predictions that (1) the rebuilding of the digestive tract, as reflected by mass-specific consumption of food on the first 2-3 days of a stopover, is faster in birds with access to drinking water than in birds without, and (2) that adipose tissue and pectoral muscles grow faster and to a greater extent in birds with unlimited access to water. We simulated migratory stopover in two experiments. In Experiment I, each of 31 birds was randomly assigned to one of three experimental groups for 6 days. Along with mealworms (～64% water) ad libitum, Group 1 received drinking water ad libitum; Group 2 had 0.5 h/day access to water; and Group 3 had no access to water. In Experiment II, 30 birds were offered a mixed diet for insectivorous birds (～33% water) ad libitum for 6 days, while randomly assigned to two groups: (1) Water ad libitum-control; and (2) 30 min access to water twice a day. We measured lean mass and fat mass using dual energy X-ray absorptiometry, as well as body mass (m(b)), pectoral muscle index (PMI), and daily intake of food and water. Mean daily water intake was significantly different among the groups in both experiments. However, the availability of drinking water positively affected the rates of gain of lean and fat mass only in birds fed with the mixed, relatively dry diet. Furthermore, mass-specific daily food intake was affected by the availability of drinking water only in the mixed diet experiment, in which birds with unlimited access to drinking water reached an asymptote, 1 day earlier than birds in the water-restricted group. We suggest that in birds consuming diets with low water content, the lack of sufficient drinking water may result in slower rebuilding of the digestive tract, or may influence biochemical processes in the gut that result in slower growth of tissue. Although blackcaps obtained sufficient water from preformed and metabolic water to renew lost tissues when eating mealworms, given access to water, the birds drank prodigiously. Our results also suggest that if drinking water is unavailable to migrating blackcaps, their choices are restricted to water-rich foods, which may constrain their rate of feeding and thus the rate at which they deposit fat. Consequently, drinking water may have an important influence on birds' migratory strategies with respect to habitat selection, use of energy, and the saving of time.     

Extent of phenotypic flexibility during long‐distance flight is determined by tissue‐specific turnover rates: a new hypothesis

Phenotypic flexibility in organ size of migratory birds is typically explained in functional terms in accordance with the principal of economic design. However, proposed functional hypotheses do not adequately explain differences in phenotypic flexibility between organs during fasting and in‐flight starvation. We show that the extent of phenotypic flexibility in organ mass in five species of migratory birds during actual migration or simulated in‐flight starvation consistently ranked as follows from highest to lowest mass change: small intestine, liver, kidney, gizzard, heart, flight and leg muscle. This pattern of phenotypic flexibility in organ mass was not consistent with proposed functional hypotheses, and was almost completely explained by differences in tissue‐specific turnover rate measured in vivo using nutrients differing in their isotopic values. Thus, the fundamental process of tissue‐specific protein turnover determines extent of organ mass changes for birds during migration, this likely applies to other organisms during fasting, and no further functional explanation(s) for differences in the magnitude of phenotypic flexibility between organs is required.     

Reply to Hussey et al.: The requirement for accurate diet-tissue discrimination factors for interpreting stable isotopes in sharks

Carbon and nitrogen stable isotope analyses have improved our understanding of food webs and movement patterns of aquatic organisms. These techniques have recently been applied to diet studies of elasmobranch fishes, but isotope turnover rates and isotope diet–tissue discrimination are still poorly understood for this group. We performed a diet switch experiment on captive sandbar sharks (Carcharhinus plumbeus) as a model shark species to determine tissue turnover rates for liver, whole blood, and white muscle. In a second experiment, we subjected captive coastal skates (Leucoraja spp.) to serial salinity reductions to measure possible impacts of tissue urea content on nitrogen stable isotope values. We extracted urea from spiny dogfish (Squalus acanthias) white muscle to test for effects on nitrogen stable isotopes. Isotope turnover was slow for shark tissues and similar to previously published estimates for stingrays and teleost fishes with low growth rates. Muscle isotope data would likely fail to capture seasonal migrations or diet switches in sharks, while liver and whole blood would more closely reflect shorter term movement or shifts in diet. Nitrogen stable isotope values of skate blood and skate and dogfish white muscle were not affected by tissue urea content, suggesting that available diet–tissue discrimination estimates for teleost fishes with similar physiologies would provide accurate estimates for elasmobranchs.     

The effect of cold-induced increased metabolic rate on the rate of <Superscript>13</Superscript>C and <Superscript>15</Superscript>N incorporation in house sparrows (<Emphasis Type="Italic">Passer domesticus</Emphasis>)

Animals with high metabolic rates are believed to have high rates of carbon and nitrogen isotopic incorporation. We hypothesized that (1) chronic exposure to cold, and hence an increase in metabolic rate, would increase the rate of isotopic incorporation of both ¹³C and ¹⁵N into red blood cells; and (2) that the rate of isotopic incorporation into red blood cells would be allometrically related to body mass. Two groups of sparrows were chronically exposed to either 5 or 22°C and switched from a ¹³C-depleted C₃-plant diet to a more ¹³C-enriched C₄-plant one. We used respirometry to estimate the resting metabolic rate of birds exposed chronically to our two experimental temperatures. The allometric relationship between the rate of ¹³C incorporation into blood and body mass was determined from published data. The of birds at 5°C was 1.9 times higher than that of birds at 22°C. Chronic exposure to a low temperature did not have an effect on the rate of isotopic incorporation of ¹⁵N save for a very small effect on the incorporation of ¹³C. The isotopic incorporation rate of ¹³C was 1.5 times faster than that of ¹⁵N. The fractional rate of ¹³C incorporation into avian blood was allometrically related to body mass with an exponent similar to −1/4. We conclude that the relationship between metabolic rate and the rate of isotopic incorporation into an animal’s tissues is indirect. It is probably mediated by protein turnover and thus more complex than previous studies have assumed.     

Population‐wide body mass increment at stopover sites is an unreliable indicator of refuelling rates in migrating waders

Estimates of refuelling rates in migrating waders are best based on intra‐seasonal recaptures of individually marked birds. This method, however, has methodological problems associated with capture effects and difficulties in attaining sufficient sample sizes. An alternative method had been proposed whereby refuelling rates are approximated by the body mass increment from the slope of the regression between body masses of all birds caught at a site and date. We investigated the appropriateness of this indicator with a simulation study in non‐synchronized migratory species (i.e. arrivals and departures of individuals at the stopover site are not synchronized). Simulation results indicated that the mass increment across the population may be used as an approximation of refuelling rate only in populations with low turnover rates (percentage of birds arriving at/departing from stopover site per day <2%) and ideally with constant numbers of staging birds. The mass increment of non‐synchronized populations with moderate or high turnover rates (higher than 5%) depends mainly on body masses of arriving birds and is not indicative of the individual rate of refuelling. The results of the simulation study were confirmed with empirical data gathered from Wood Sandpiper Tringa glareola and Common Snipe Gallinago gallinago during their autumn migration at a stopover site in Poland. The population mass increment methods considerably underestimated refuelling rate obtained from the recapture‐based approach of individual birds in both species. As a consequence, we suggest that population mass increment should not be used as an indicator of refuelling rates in non‐synchronized stopover populations of migrating waders.     

Reduced norepinephrine turnover in brown adipose tissue of pre-obese mice treated with monosodium-L-glutamate

Norepinephrine (NE) turnover, an index of sympathetic nervous system (SNS) activity, was measured in interscapular brown adipose tissue (IBAT), heart and pancreas of 3-weeks-old pre-obese monosodium-L-glutamate (MSG) mice and at 6-weeks-old mildly obese MSG mice. In IBAT, rates of NE turnover were slower not only in 3-weeks-old MSG mice but also in older obese MSG mice than in their saline controls. In heart, rates of NE turnover were slower in 6-weeks-old mildly obese MSG mice, but not in pre-obese MSG mice. No significant difference in NE turnover in pancreas was observed at either age. The low NE turnover in IBAT of MSG-treated mice prior to the onset of gross obesity suggests that low SNS activity may be an initial contributor to their high energy efficiency and resultant obesity.     

Metabolic rate is negatively linked to adult survival but does not explain latitudinal differences in songbirds

Survival rates vary dramatically among species and predictably across latitudes, but causes of this variation are unclear. The rate‐of‐living hypothesis posits that physiological damage from metabolism causes species with faster metabolic rates to exhibit lower survival rates. However, whether increased survival commonly observed in tropical and south temperate latitudes is associated with slower metabolic rate remains unclear. We compared metabolic rates and annual survival rates that we measured across 46 species, and from literature data across 147 species of birds in northern, southern and tropical latitudes. High metabolic rates were associated with lower survival but survival varied substantially among latitudinal regions independent of metabolism. The inability of metabolic rate to explain latitudinal variation in survival suggests (1) species may evolve physiological mechanisms that mitigate physiological damage from cellular metabolism and (2) extrinsic rather than intrinsic sources of mortality are the primary causes of latitudinal differences in survival.     

Stable Isotope Turnover and Half-Life in Animal Tissues: A Literature Synthesis

Stable isotopes of carbon, nitrogen, and sulfur are used as ecological tracers for a variety of applications, such as studies of animal migrations, energy sources, and food web pathways. Yet uncertainty relating to the time period integrated by isotopic measurement of animal tissues can confound the interpretation of isotopic data. There have been a large number of experimental isotopic diet shift studies aimed at quantifying animal tissue isotopic turnover rate λ (%·day^-1, often expressed as isotopic half-life, ln(2)/λ, days). Yet no studies have evaluated or summarized the many individual half-life estimates in an effort to both seek broad-scale patterns and characterize the degree of variability. Here, we collect previously published half-life estimates, examine how half-life is related to body size, and test for tissue- and taxa-varying allometric relationships. Half-life generally increases with animal body mass, and is longer in muscle and blood compared to plasma and internal organs. Half-life was longest in ecotherms, followed by mammals, and finally birds. For ectotherms, different taxa-tissue combinations had similar allometric slopes that generally matched predictions of metabolic theory. Half-life for ectotherms can be approximated as: ln (half-life) = 0.22*ln (body mass) + group-specific intercept; n = 261, p<0.0001, r² = 0.63. For endothermic groups, relationships with body mass were weak and model slopes and intercepts were heterogeneous. While isotopic half-life can be approximated using simple allometric relationships for some taxa and tissue types, there is also a high degree of unexplained variation in our models. Our study highlights several strong and general patterns, though accurate prediction of isotopic half-life from readily available variables such as animal body mass remains elusive.     

Metabolism during flight in two species of bats, Phyllostomus hastatus and Pteropus gouldii.

The energetic cost of flight in a wind-tunnel was measured at various combinations of speed and flight angle from two species of bats whose body masses differ by almost an order of magnitude. The highest mean metabolic rate per unit body mass measured from P. hastatus (mean body mass, 0.093 kg) was 130.4 Wkg-1, and that for P. gouldii (mean body mass, 0.78 kg) was 69.6 Wkg-1. These highest metabolic rates, recorded from flying bats, are essentially the same as those predicted for flying birds of the same body masses, but are from 2.5 to 3.0 times greater than the highest metabolic rates of which similar-size exercising terrestrial mammals appear capable. The lowest mean rate of energy utilization per unit body mass P. hastatus required to sustain level flight was 94.2 Wkg-1 and that for P. gouldii was 53.4 Wkg-1. These data from flying bats together with comparable data for flying birds all fall along a straight line when plotted on double logarithmic coordinates as a function of body mass. Such data show that even the lowest metabolic requirements of bats and birds during level flight are about twice the highest metabolic capabilities of similar-size terrestrial mammals. Flying bats share with flying birds the ability to move substantially greater distance per unit energy consumed than walking or running mammals. Calculations show that P. hastatus requires only one-sixth the energy to cover a given distance as does the same-size terrestrial mammal, while P. gouldii requires one-fourth the energy of the same-size terrestrial mammal. An empirically derived equation is presented which enables one to make estimates of the metabolic rates of bats and birds during level flight in nature from body mass data alone. Metabolic data obtained in this study are compared with predictions calculated from an avian flight theory.     

Slow pace of life in tropical sedentary birds: a common-garden experiment on four stonechat populations from different latitudes

It has been hypothesized that organisms living at different latitudes or in different environments adjust their metabolic activity to the prevailing conditions. However, do differences in energy turnover simply represent a phenotypic adaptation to the local environment, or are they genetically based? To test this, we obtained nestling stonechats (Saxicola torquata) from equatorial Kenya (0 degrees N), Ireland (51.5 degrees N), Austria (47.5 degrees N) and Kazakhstan (51.5 degrees N). Birds were hand-raised and kept in Andechs, Germany. We measured their resting metabolic rates (RMR) and locomotor activity at an age of ca. 14 months (July) and 20 months (January), when birds went through postnuptial moult (July), and neither moulted nor exhibited enlarged gonads or migratory activity (January). RMR was generally higher during moult, but differed among populations: RMR was lowest in the resident Kenyan birds, higher in mostly sedentary Irish birds, and highest in migratory Austrian and Kazakhstan birds. Thus our data demonstrate that even in birds kept from early life under common-garden conditions, the ''pace of life'', as indicated by metabolic turnover, is lower in sedentary tropical than in north-temperate migratory individuals of the same species. Such intrinsically low energy expenditure in sedentary tropical birds may have important implications for slow development, delayed senescence and high longevity in many tropical organisms.