Metabolic Rate In Birds Research Articles

Integrative Physiological Responses to Acute Dehydration in the Rufous-Collared Sparrow: Metabolic, Enzymatic, and Oxidative Traits

Predictions indicate that birds worldwide will be affected by global warming and extreme climatic events which is especially relevant for passerines because the diurnal habits, small body size, and high mass-adjusted metabolic rates of this group make it particularly susceptible to increases in temperature and aridity. Some bird species respond to conditions that stress osmoregulation by increasing their rates of energy expenditure, nevertheless, the effect of dehydration on metabolic rates in birds has produced contrasting results. It also remains unknown whether hydration state may cause shifts in tissue-specific metabolic rates or modify tissue oxidative status. We used the rufous-collared sparrow (Zonotrichia capensis), to experimentally test the effect of dehydration on metabolic enzymes in erythrocytes, tissue oxidative status, basal metabolic rate (BMR), and total evaporative water loss. We found a significant increase in mass-adjusted BMR in water restricted (WR) birds compared to control birds (CT). Activity of cytochrome-c-oxidase (COX) in red blood cells (RBCs) was also significantly higher in the WR group relative to the CT group and this activity was positively correlated with mass-adjusted BMR. We found a moderate effect of water restriction on membrane damage of skeletal muscle. In a second set of individuals subjected to the same experimental conditions, lean mass and total water were tightly correlated and decreased by 10 and 12%, respectively, in birds in the WR group relative to the CT group. Decreases in total water and lean mass leads to an increase in mass-adjusted BMR in WR Z. capensis, suggesting that birds may simultaneously increase protein catabolism and production of metabolic water through oxidation. The significant positive relationship between BMR and COX in RBCs is a finding that requires additional research to determine whether erythrocyte metabolism is affected by dehydration per se and or it more generally reflects rates of energy expenditure in birds.

Seasonal metabolic flexibility is correlated with microclimate variation in horned larks and house sparrows.

Maximum and minimum metabolic rates in birds are flexible traits and such flexibility can be advantageous in variable climates. The climatic variability hypothesis (CVH) posits that more variable climates should result in greater metabolic flexibility for geographically distinct populations. Whether the CVH applies to sympatric species occupying microclimates differing in variability is unknown. Microclimates of open habitats are likely more variable than those of sheltered habitats. If the CVH extends to microclimates, we expect birds from open habitats to show greater flexibility than those from sheltered habitats. To test this extension of the CVH, we compared seasonal variation in microclimates and metabolic rates for sympatric horned larks Eremophila alpestris, which occupy open habitats, and house sparrows Passer domesticus, which occupy sheltered habitats. We measured operative temperature (Te, an integrative measure of the thermal environment), summit metabolic rate (Msum, maximal cold-induced metabolic rate), and basal metabolic rate (BMR, minimal maintenance metabolic rate) in summer and winter. For both winter and summer, daily minimum Te was similar between open and sheltered habitats but maximum Te was higher for open habitats. Winter microclimates, however, were colder for open than for sheltered habitats after accounting for convective differences. Both species increased Msum in winter, but seasonal Msum flexibility was greater for larks (43%) than for sparrows (31%). Winter increases in BMR were 92.5% and 11% for larks and sparrows, respectively, with only the former attaining statistical significance. Moreover, species * season interactions in general linear models for whole-organism metabolic rates were significant for BMR and showed a similar, although not significant, pattern for Msum, with greater seasonal metabolic flexibility in horned larks than in house sparrows. These results suggest that extending the CVH to sympatric bird species occupying different microclimates may be valid.

Local Perception of Climate Change, COVID-19 and their Impact on Birds in Jammu and Kashmir

Weather is of major significance for the population dynamics of birds, but the implications of climate change have only recently begun to be addressed. There is already compelling support that birds have been affected by recent climate changes. The metabolic rate of birds is not only affected by weather (e.g. cold weather requiring increased energy expenditure for body maintenance)but also exerts other direct and indirect effects on bird behaviour. The extreme weather events, like prolonged frozen spells and droughts, can have catastrophic effects on bird populations, including long-term effects on whole cohorts. Birds are generally admired for their beauty, songs, and the beauty of their near miraculous ability to fly without recognizing their role direct or indirect to the Ecosystem Services. Birds are marvellous indicators of environmental health which offer humans pleasure, joy and spiritual motivation. The main reason to study climate change, COVID-19 and their impact on birds is to promote understanding of the ecosystems that support all life on earth, including human beings. In this study, a well designed and validated questionnaire was used to collect the information from a sample of 400 local people from Kashmir valley. The main aim of writing this paper was to find out the perception and approach of local people towards climate change, COVID-19 and their impact on birds. The data collected from the survey was analyzed using standard statistical tools. The result of our study revealed from people perception that increase in temperature, unpredictable rainfall, increase in frequency of disaster and decrease in food production has direct impact on birds. Further, the study revealed that local people showed interest towards birds, nature and wildlife preservation. The respondents in majority reported the sudden increase in bird sightings during COVID-19 lockdown. Finally, to cope with climate change impact, it was suggested that education programmers be introduced among the people to increase awareness about the role played by bird in our lives, significance of wildlife conservation and management amidst changing climate. Further, to cope with COVID-19 threat, the research advised to follow WHO and health ministry guidelines.

Metabolic Scaling of Stress Hormones in Vertebrates.

Glucocorticoids (GCs) are stress hormones that can strongly influence physiology, behavior, and an organism's ability to cope with environmental change. Despite their importance, and the wealth of studies that have sought to understand how and why GC concentrations vary within species, we do not have a clear understanding of how circulating GC levels vary within and across the major vertebrate clades. New research has proposed that much interspecific variation in GC concentrations can be explained by variation in metabolism and body mass. Specifically, GC concentrations should vary proportionally with mass-specific metabolic rates and, given known scaling relationships between body mass and metabolic rate, GC concentrations should scale to the -1/4 power of body mass and to the power of 1 with mass-specific metabolic rate. Here, we use HormoneBase, the newly compiled database that includes plasma GC concentrations from free-living and unmanipulated vertebrates, to evaluate this hypothesis. Specifically, we explored the relationships between body mass or mass-specific metabolic rate and either baseline or stress-induced GC (cortisol or corticosterone) concentrations in tetrapods. Our phylogenetically-informed models suggest that, whereas the relationship between GC concentrations and body mass across tetrapods and among mammals is close to -1/4 power, this relationship does not exist in amphibians, reptiles, and birds. Moreover, with the exception of a positive association between stress-induced GC concentrations and mass-specific metabolic rate in birds, we found little evidence that GC concentrations are linked to metabolic rate, although the number of species sampled was quite limited for amphibians and somewhat so for reptiles and mammals. Nevertheless, these results stand in contrast to the generally accepted association between the two and suggest that our observed positive association between body mass and GC concentrations may not be due to the well-established link between mass and metabolism. Large-scale comparative approaches can come with drawbacks, such as pooling and pairing observations from separate sources. However, these broad analyses provide an important counterbalance to the majority of studies examining variation in GC concentrations at the population or species level, and can be a powerful approach to testing both long-standing and new questions in biology.

Diurnal rhythms of standard metabolic rate and respiratory quotient in the coal tit (Periparus Ater, Aves) during the autumn-winter period

Standard metabolic rate of birds has been measured by flow-through respirometry based on their standard O2 consumption and CO2 release during the day- and nighttimes in a darkened chamber at a constant temperature of 25°C. The research was carried out in October–March 2009–2013, in the western part of Moscow region. The standard metabolic rate and respiratory quotient of coal tits have well-pronounced diurnal rhythms reaching their minimum in the nighttime (from 2:00 a.m. to 5:00 a.m.) and maximum in the daytime (from 11:00 a.m. to 2:00 p.m.). A significant linear dependence has been found between an increase in the standard metabolic rate and that in the respiratory quotient. The maximum difference between values of the standard metabolic rate during the day- and nighttimes is 55%. It may be concluded that the main elements oxidized at night are fat reserves, whereas proteins, as well as a mixture of proteins, fats, and carbohydrates, are oxidized during the daytime.

Among- and within-individual correlations between basal and maximal metabolic rates in birds.

The aerobic capacity model proposes that endothermy is a by-product of selection favouring high maximal metabolic rates (MMR) and its mechanistic coupling with basal metabolic rate (BMR). Attempts to validate this model in birds are equivocal and restricted to phenotypic correlations (rP), thus failing to distinguish among- and within-individual correlations (rind and re). We examined 300 paired measurements of BMR and MMR from 60 house sparrows before and after two levels of experimental manipulation - testosterone implants and immune challenge. Overall, repeatability was significant in both BMR (R=0.25±0.06) and MMR (R=0.52±0.06). Only the testosterone treatment altered the rP between BMR and MMR, which resulted from contrasting effects on rind and re. While rind was high and significant (0.62±0.22) in sham-implanted birds, re was negative and marginally non-significant (-0.15±0.09) in testosterone-treated birds. Thus, the expected mechanistic link between BMR and MMR was apparent, but only in birds with low testosterone levels.

Intraspecific Correlations of Basal and Maximal Metabolic Rates in Birds and the Aerobic Capacity Model for the Evolution of Endothermy

The underlying assumption of the aerobic capacity model for the evolution of endothermy is that basal (BMR) and maximal aerobic metabolic rates are phenotypically linked. However, because BMR is largely a function of central organs whereas maximal metabolic output is largely a function of skeletal muscles, the mechanistic underpinnings for their linkage are not obvious. Interspecific studies in birds generally support a phenotypic correlation between BMR and maximal metabolic output. If the aerobic capacity model is valid, these phenotypic correlations should also extend to intraspecific comparisons. We measured BMR, Msum (maximum thermoregulatory metabolic rate) and MMR (maximum exercise metabolic rate in a hop-flutter chamber) in winter for dark-eyed juncos (Junco hyemalis), American goldfinches (Carduelis tristis; Msum and MMR only), and black-capped chickadees (Poecile atricapillus; BMR and Msum only) and examined correlations among these variables. We also measured BMR and Msum in individual house sparrows (Passer domesticus) in both summer, winter and spring. For both raw metabolic rates and residuals from allometric regressions, BMR was not significantly correlated with either Msum or MMR in juncos. Moreover, no significant correlation between Msum and MMR or their mass-independent residuals occurred for juncos or goldfinches. Raw BMR and Msum were significantly positively correlated for black-capped chickadees and house sparrows, but mass-independent residuals of BMR and Msum were not. These data suggest that central organ and exercise organ metabolic levels are not inextricably linked and that muscular capacities for exercise and shivering do not necessarily vary in tandem in individual birds. Why intraspecific and interspecific avian studies show differing results and the significance of these differences to the aerobic capacity model are unknown, and resolution of these questions will require additional studies of potential mechanistic links between minimal and maximal metabolic output.

Dichotomy of eutherian reproduction and metabolism

How anatomical, physiological and ecological (life history) features scale with body mass is a fundamental question in biology. There is an ongoing debate in the scientific literature whether allometric scaling follows a universal pattern that can be described in a single model, or differs between groups. However, recently some analyses were published demonstrating a change in scaling across the body mass range: brain‐size allometry of mammals indicates that scaling follows a curvilinear pattern in double‐logarithmic space, and a quadratic pattern in double‐logarithmic space was found in one of the largest physiological datasets, on basal metabolic rate (MR) in mammals. Here, we analysed a variety of independent datasets on anatomical, physiological and ecological characteristics in mammals, birds and reptiles to answer the question whether the quadratic scaling is a universal biological law, or a pattern unique to mammals. The pattern was present in mammalian basal and field MR, brain size, and reproduction parameters, but neither in other organ allometries in mammals, nor in the scaling of MR in birds and reptiles. However, the curvature was better explained by separate allometric scaling of three different mammalian reproduction strategies: marsupials, and eutherian mammals with one and with many offspring. The two latter strategies are distributed unequally over the body mass range in eutherian mammals. Our findings show that a quadratic model, as well as a traditional allometric model with a universal scaling exponent (such as 0.67 or 0.75), may be inappropriate in mammals as they are a result of different scalings within these three reproductive groups. We propose that the observed distribution pattern is the result of the eutherian mammal clade's uniquely pronounced dichotomy of reproductive strategies.

Estimation of heritability and repeatability of resting metabolic rate in birds by the example of free-living pied flycatchers Ficedula hypoleuca (Aves: Passeriformes)

Repeatability of a trait estimated over long time intervals and its heritability make it possible to assess whether it can serve as an individual characteristic and be subject to selection. Heritability and repeatability of energetic traits are still poorly studied in birds. The most important physiological characteristic of the homoiotherms is the minimal level of energy expenditure, referred to as the resting metabolic rate (RMR), which, in the absence of the expenditure for biomass growth, does not exceed the basal metabolic rate (BMR). We have estimated the BMR repeatability in adult free-living pied flycatchers in Moscow-region (55°44′ N, 36°51′ E; 1992–2008) and Tomsk (56°20′ N, 84°56′ E; 2008–2009) populations over intervals of 40 days to 3 years. In the Moscow-region pied flycatcher population, the BMR repeatability recorded during the same period of the reproductive season was τ = 0.34 ± 0.10 for a 1-year interval (n = 80), τ = 0.60 ± 0.15 for a 2-year interval (n = 19), and τ = 0.85 ± 0.13 for a 3-year interval (n = 6). In the Tomsk population, the BMR repeatability for the 1-year interval was τ = 0.49 ± 0.11 (n = 50). The repeatability of a trait is a measure of its constancy in time and specifies the upper limit of its heritability. RMR heritability was assessed in cross-fostering experiments in the Moscow-region free-living pied flycatcher population in 2003–2005. The RMRs of chicks and BMRs of their biological fathers displayed a positive correlation, whereas no correlation was found between the RMRs of chicks and their foster fathers. The RMR heritability value was h2 = 0.43 ± 0.17 (n = 210). The obtained estimates for repeatability and heritability of fundamental energetic traits in pied fly-catcher are rather high for a physiological trait and suggest the existence of evolutionary stable diversity of the avian population in the BMR.

The use of body mass loss to estimate metabolic rate in birds

During starvation, energy production occurs at the expense of body reserve utilisation which results in body mass loss. Knowing the role of the fuels involved in this body mass loss, along with their energy density, can allow an energy equivalent of mass loss to be calculated. Therefore, it is possible to determine daily energy expenditure (DEE) if two body mass loss measurements at an interval of a few days are obtained. The technique can be cheap, minimally stressful for the animals involved, and the data relatively simple to gather. Here we review the use of body mass loss to estimate DEE in birds through critiquing the strengths and weaknesses of the technique, and detail the methodology and considerations that must be adhered to for accurate measures of DEE to be obtained. Owing to the biology of the species, the use of the technique has been used predominantly in Antarctic seabirds, particularly penguins and albatrosses. We demonstrate how reliable the technique can be in predicting DEE in a non-Antarctic species, common eiders ( Somateria mollissima), the female of which undergoes a fasting period during incubation. We conclude that using daily body mass loss to estimate DEE can be a useful and effective approach provided that (1) the substrate being consumed during mass loss is known, (2) the kinetics of body mass loss are understood for the species in question and (3) only species that enter a full phase II of a fast (where substrate catabolism reaches a steady state) and are not feeding for a period of time are appropriate for this method.

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

Abstract 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), summit metabolic rate (Msum; maximal thermoregulatory metabolic rate), and maximal metabolic rate (MMR; maximal exercise metabolic rate)], the factors associated with this variation, the evidence for functional links between these metabolic traits, and the ecological and evolutionary significance of avian metabolic diversity. Both lower and upper limits to metabolic power production are phenotypically flexible traits, and vary in association with numerous ecological and evolutionary factors. For both inter- and intraspecific comparisons, lower and upper limits to metabolic power production are generally upregulated in response to energetically demanding conditions and downregulated when energetic demands are relaxed, or under conditions of energetic scarcity. Positive correlations have been documented between BMR, Msum and MMR in some, but not all studies on birds, providing partial support for the idea of a functional link between lower and upper limits to metabolic power production, but more intraspecific studies are needed to determine the robustness of this conclusion. Correlations between BMR and field metabolic rate (or daily energy expenditure) in birds are variable, suggesting that the linkage between these traits is subject to behavioral adjustment, and studies of the relationship between field and maximal metabolic rates are lacking. Our understanding of avian metabolic diversity would benefit from future studies of: (1) the functional and mechanistic links between lower and upper limits of metabolic power output; (2) the environmental and ecological cues driving phenotypically flexible metabolic responses, and how responses to such cues might impact population responses to climate change; (3) the shapes of metabolic reaction norms and their association with environmental variability; and (4) the relationship of metabolic variation to fitness, including studies of repeatability and heritability of minimum and maximum metabolic power output.

Heterothermy in small, migrating passerine birds during stopover: use of hypothermia at rest accelerates fuel accumulation

For small endothermic animals, heterothermy serves as an energy-saving mechanism for survival in challenging environments, but it may also accelerate fat accumulation in individuals preparing for fuel-demanding activities. This is the first study to demonstrate adaptive hypothermic responses in migrating passerines. While monitoring body temperature (T(b)) of eight blackcaps (Sylvia atricapilla) by radiotelemetry, we found that during daytime T(b)=42.5+/-0.4 degrees C (mean +/- s.d.); at night T(b) decreased to a minimum between 33 and 40 degrees C. We determined the lower limit for normothermy at 37.4 degrees C and found that on 12 out of 34 bird-nights of observations under semi-natural conditions blackcaps reduced their T(b) below normothermic resting levels with minimum values of 33 and 34.5 degrees C compared with rest-phase normothermic T(b) of 38.8+/-0.8 degrees C. In birds of body mass (m(b)) <16.3 g, minimum T(b) at night correlated with the individual's m(b) (r=0.67, P<0.01, N=17), but this was not the case in birds with m(b)>16.3 g. Minimum nocturnal T(b) did not correlate with night-time air temperature (T(a)). Measurements of metabolic rate in birds subjected to a T(a) of 15 degrees C showed that hypothermia of this magnitude can lead to a reduction of some 30% in energy expenditure compared with birds remaining normothermic. Our data suggest that by reducing the T(b)-T(a) gradient, blackcaps accelerate their rate of fuel accumulation at a stopover. When body energy reserves are low blackcaps may achieve this reduction by entering hypothermia. Since hypothermia, as seen in blackcaps, may lead to significant energy savings and facilitate body mass gain, we predict that it is common among small migrating passerines.

Environment, Migratory Tendency, Phylogeny and Basal Metabolic Rate in Birds

Basal metabolic rate (BMR) represents the minimum maintenance energy requirement of an endotherm and has far-reaching consequences for interactions between animals and their environments. Avian BMR exhibits considerable variation that is independent of body mass. Some long-distance migrants have been found to exhibit particularly high BMR, traditionally interpreted as being related to the energetic demands of long-distance migration. Here we use a global dataset to evaluate differences in BMR between migrants and non-migrants, and to examine the effects of environmental variables. The BMR of migrant species is significantly higher than that of non-migrants. Intriguingly, while the elevated BMR of migrants on their breeding grounds may reflect the metabolic machinery required for long-distance movements, an alternative (and statistically stronger) explanation is their occupation of predominantly cold high-latitude breeding areas. Among several environmental predictors, average annual temperature has the strongest effect on BMR, with a 50% reduction associated with a 20°C gradient. The negative effects of temperature variables on BMR hold separately for migrants and non-migrants and are not due their different climatic associations. BMR in migrants shows a much lower degree of phylogenetic inertia. Our findings indicate that migratory tendency need not necessarily be invoked to explain the higher BMR of migrants. A weaker phylogenetic signal observed in migrants supports the notion of strong phenotypic flexibility in this group which facilitates migration-related BMR adjustments that occur above and beyond environmental conditions. In contrast to the findings of previous analyses of mammalian BMR, primary productivity, aridity or precipitation variability do not appear to be important environmental correlates of avian BMR. The strong effects of temperature-related variables and varying phylogenetic effects reiterate the importance of addressing both broad-scale and individual-scale variation for understanding the determinants of BMR.

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.

Effects of metabolic level on the body size scaling of metabolic rate in birds and mammals

Metabolic rate is traditionally assumed to scale with body mass to the 3/4-power, but significant deviations from the '3/4-power law' have been observed for several different taxa of animals and plants, and for different physiological states. The recently proposed 'metabolic-level boundaries hypothesis' represents one of the attempts to explain this variation. It predicts that the power (log-log slope) of metabolic scaling relationships should vary between 2/3 and 1, in a systematic way with metabolic level. Here, this hypothesis is tested using data from birds and mammals. As predicted, in both of these independently evolved endothermic taxa, the scaling slope approaches 1 at the lowest and highest metabolic levels (as observed during torpor and strenuous exercise, respectively), whereas it is near 2/3 at intermediate resting and cold-induced metabolic levels. Remarkably, both taxa show similar, approximately U-shaped relationships between the scaling slope and the metabolic (activity) level. These predictable patterns strongly support the view that variation of the scaling slope is not merely noise obscuring the signal of a universal scaling law, but rather is the result of multiple physical constraints whose relative influence depends on the metabolic state of the organisms being analysed.

Basal metabolic rate of birds is associated with habitat temperature and precipitation, not primary productivity

A classic example of ecophysiological adaptation is the observation that animals from hot arid environments have lower basal metabolic rates (BMRs, ml O2min-1) than those from non-arid (luxuriant) ones. However, the term 'arid' conceals within it a multitude of characteristics including extreme ambient temperatures (Ta, degrees C) and low annual net primary productivities (NPPs, gCm-2), both of which have been shown to correlate with BMR. To assess the relationship between environmental characteristics and metabolic rate in birds, we collated BMR measurements for 92 populations representing 90 wild-caught species and examined the relationships between BMR and NPP, Ta, annual temperature range (Tr), precipitation and intra-annual coefficient of variation of precipitation (PCV). Using conventional non-phylogenetic and phylogenetic generalized least-squares approaches, we found no support for a relationship between BMR and NPP, despite including species captured throughout the world in environments spanning a 35-fold range in NPP. Instead, BMR was negatively associated with Ta and Tr, and positively associated with PCV.

The Evolution of Concepts on the Evolution of Endothermy in Birds and Mammals

Warm-blooded animals, mammals and birds, are unique not because they are endothermic in the strict sense of the term but because they use an extravagant economy: they have high energy budgets and spend a large part of their energy resources on basic maintenance. Although several advantages of endothermy are easy to indicate, mechanisms behind evolution of such a wasteful life strategy remain unclear and have been subject to intensive debate. For two decades, the aerobic capacity model has been widely recognized as a promising hypothesis and has catalyzed a new direction in ecological and evolutionary physiology--the study of correlated evolution of behavioral and morphophysiological traits. Recently, two alternative models have been proposed, both of which see evolution of high metabolic rates in birds and mammals as an element in evolution of intensive parental care. Unlike previous models, which treated individuals as static objects of fixed properties, the parental care models explicitly incorporate life histories into a evolutionary-physiology research program. The aim of this article was to outline the process of evolution of major concepts in the field, which reflects development of the paradigm of modern evolutionary physiology.

Mammalian metabolism: insights from arid zone reptiles.

Mammals, being endotherms have very high metabolic rates compared to ectothermic reptiles. Similarly, small mammals have high rates of mass-specific metabolism compared to larger mammals. This review examines the mechanistic basis of why particular mammal species have a specific metabolic rate. Initial studies compared mammals with arid zone reptile species of the same size and Tb. Mammals have larger internal organs, with more mitochondrial membrane surface area than the reptiles. The cells of mammals are leakier to Na+ ions and their mitochondrial membranes are leakier to H+ ions than in reptile cells. These leakier membranes have membrane lipids that are polyunsaturated and less monounsaturated than their less leaky counterparts. Examination of the cellular basis of allometric variation in metabolism in mammals reveals very similar findings with polyunsaturated membranes associated with the high mass-specific metabolic rates of small mammal species and monounsaturated membranes with low rates of metabolism of large mammals. These findings have resulted in the development of the ?membrane pacemaker? theory of metabolism, which proposes that membrane bilayer composition is regulated in animals and that highly polyunsaturated membranes result in enhanced molecular activity of membrane proteins and in turn this results in an elevated metabolic rate of cells, tissues and consequently whole animals. This theory is also supported by the recent examination of the basis of body-size variation in the metabolic rates of birds. The ?membrane pacemaker? theory of metabolism is currently the only explanation of the mechanisms determining the metabolic rate and thus the cost of living of animals. It has implications for the effect of food habits on metabolism and the relationship between metabolism and lifespan.

Low mitochondrial free radical production per unit O2 consumption can explain the simultaneous presence of high longevity and high aerobic metabolic rate in birds.

Birds are unique since they can combine a high rate of oxygen consumption at rest with a high maximum life span (MLSP). The reasons for this capacity are unknown. A similar situation is present in primates including humans which show MLSPs higher than predicted from their rates of O2 consumption. In this work rates of oxygen radical production and O2 consumption by mitochondria were compared between adult male rats (MLSP = 4 years) and adult pigeons (MLSP = 35 years), animals of similar body size. Both the O2 consumption of the whole animal at rest and the O2 consumption of brain, lung and liver mitochondria were higher in the pigeon than in the rat. Nevertheless, mitochondrial free radical production was 2-4 times lower in pigeon than in rat tissues. This is possible because pigeon mitochondria show a rate of free radical production per unit O2 consumed one order of magnitude lower than rat mitochondria: bird mitochondria show a lower free radical leak at the respiratory chain. This result, described here for the first time, can possibly explain the capacity of birds to simultaneously increase maximum longevity and basal metabolic rate. It also suggests that the main factor relating oxidative stress to aging and longevity is not the rate of oxygen consumption but the rate of oxygen radical production. Previous inconsistencies of the rate of living theory of aging can be explained by a free radical theory of aging which focuses on the rate of oxygen radical production and on local damage to targets relevant for aging situated near the places where free radicals are continuously generated.

On the Relation Between Basal and Field Metabolic Rates in Birds and Mammals

Basal metabolic rate (BMR) is widely assumed to be an index of energy expenditure of free-living animals. However, few attempts have been made to check if the assumption is valid. In this study field metabolic rates (FMR) of 58 species of birds and mammals have been compared with their BMRs. FMR tends to increase with increasing body mass slower than BMR, but the difference was significant only in marsupials. A correlation between mass-independent FMR and BMR (residuals from regressions) is very high in eutherians (mainly rodents), but it is weak in birds and approaches zero in marsupials. The assumption of a proportionality between BMR and energy expenditure of animals does not have a sound experi-