A fundamental but unanswered biological question asks how much energy, on average, Earth's different life forms spend per unit mass per unit time to remain alive. Here, using the largest database to date, for 3,006 species that includes most of the range of biological diversity on the planet-from bacteria to elephants, and algae to sapling trees-we show that metabolism displays a striking degree of homeostasis across all of life. We demonstrate that, despite the enormous biochemical, physiological, and ecological differences between the surveyed species that vary over 10 20 -fold in body mass, mean metabolic rates of major taxonomic groups displayed at physiological rest converge on a narrow range from 0.3 to 9 W kg ؊1 . This 30-fold variation among life's disparate forms represents a remarkably small range compared with the 4,000-to 65,000-fold difference between the mean metabolic rates of the smallest and largest organisms that would be observed if life as a whole conformed to universal quarterpower or third-power allometric scaling laws. The observed broad convergence on a narrow range of basal metabolic rates suggests that organismal designs that fit in this physiological window have been favored by natural selection across all of life's major kingdoms, and that this range might therefore be considered as optimal for living matter as a whole.allometry ͉ body size ͉ breathing ͉ scaling ͉ energy consumption T he process of life is critically dependent on consumption of energy from the environment. The amount of energy-per unit time per unit mass-required to sustain life can rightfully be considered one of the fundamental questions in biology. Yet a general quantitative answer to this question is lacking, despite the long history and the considerable number of studies devoted to various aspects of organismal energetics in all fields of bioscience. One reason for this persistent knowledge gap is that this fundamental question is typically approached in markedly different ways depending on the organisms being investigated. We show herein how differences in types, protocols, and units of measurements of metabolism have presented a challenge to the development of quantitative generalizations regarding the metabolic rates of organisms. We then use a comprehensive dataset to reconcile such differences and to characterize the remarkable similarity that emerges from comparisons of mass-specific metabolic rates across all of life. Problem SettingStudies of animal energetics have frequently focused on the allometric relationship between the whole-body metabolic rate Q and body mass M, Q ϭ Q 0 (M/M 0 ) b , where Q 0 is metabolic rate of an organism with body mass M 0 . Either M 0 or Q 0 can be chosen arbitrarily, whereas the second of these parameters is unambiguously defined by the choice of the first one. Usually, M 0 is chosen to be one mass unit-e.g., M 0 ϭ 1 g. For the mass-specific metabolic rate q ' Q/M, we have q ϭ q 0 (M/M 0 ) ␤ , ␤ ϭ b Ϫ 1, q 0 ϭ Q 0 /M 0 . Much of the current debate concerns the value of b, an...

Power laws describing the dependence of metabolic rate on body mass have been established for many taxa, but not for prokaryotes, despite the ecological dominance of the smallest living beings. Our analysis of 80 prokaryote species with cell volumes ranging more than 1 000 000-fold revealed no significant relationship between mass-specific metabolic rate q and cell mass. By absolute values, mean endogenous mass-specific metabolic rates of non-growing bacteria are similar to basal rates of eukaryote unicells, terrestrial arthropods and mammals. Maximum mass-specific metabolic rates displayed by growing bacteria are close to the record tissue-specific metabolic rates of insects, amphibia, birds and mammals. Minimum mass-specific metabolic rates of prokaryotes coincide with those of larger organisms in various energy-saving regimes: sit-and-wait strategists in arthropods, poikilotherms surviving anoxia, hibernating mammals. These observations suggest a size-independent value around which the mass-specific metabolic rates vary bounded by universal upper and lower limits in all body size intervals.

Abstract. In this paper the basic geophysical and ecological principles are jointly analyzed that allow the landmasses of Earth to remain moistened sufficiently for terrestrial life to be possible. 1. Under gravity, land inevitably loses water to the ocean. To keep land moistened, the gravitational water runoff must be continuously compensated by the atmospheric ocean-to-land moisture transport. Using data for five terrestrial transects of the International Geosphere Biosphere Program we show that the mean distance to which the passive geophysical air fluxes can transport moisture over non-forested areas, does not exceed several hundred kilometers; precipitation decreases exponentially with distance from the ocean. 2. In contrast, precipitation over extensive natural forests does not depend on the distance from the ocean along several thousand kilometers, as illustrated for the Amazon and Yenisey river basins and Equatorial Africa. This points to the existence of an active biotic pump transporting atmospheric moisture inland from the ocean. 3. Physical principles of the biotic moisture pump are investigated based on the previously unstudied properties of atmospheric water vapor, which can be either in or out of hydrostatic equilibrium depending on the lapse rate of air temperature. A novel physical principle is formulated according to which the low-level air moves from areas with weak evaporation to areas with more intensive evaporation. Due to the high leaf area index, natural forests maintain high transpiration fluxes, which support the ascending air motion over the forest and "suck in" moist air from the ocean, which is the essence of the biotic pump of atmospheric moisture. In the result, the gravitational runoff water losses from the optimally moistened forest soil can be fully compensated by the biotically enhanced precipitation at any distance from the ocean. 4. It is discussed how a continent-scale biotic water pump mechanism could be produced by natural selection acting on individual trees. 5. Replacement of the natural forest cover by a low leaf index vegetation leads to an up to tenfold reduction in mean continental precipitation and runoff, in contrast to the previously available estimates made without accounting for the biotic moisture pump. The analyzed body of evidence testifies that the long-term stability of an intense terrestrial water cycle is unachievable without the recovery of natural, self-sustaining forests on continent-wide areas.

We show that the largest tropical species of terrestrial poikilotherms from 25 taxa exceed, in linear body size, the largest representatives of the same taxa from the temperate (e.g. Great Britain) and polar (e.g. Wrangel Island) zones by 3.2 and 5.7 times, respectively. Here we develop a theoretical approach which quantitatively explains the observed body size patterns and relates them to ambient temperature under the assumption that there exists a temperature‐independent critical minimum value of mass‐specific metabolic rate bmin, a fall below which is incompatible with successful biological performance. This value sets an upper limit to linear body size within a taxon. Mass‐specific metabolic rate decreases with increasing body size but, in poikilotherms, grows exponentially with ambient temperature. Such compensation of the size‐related drop in mass‐specific metabolic rate b by higher ambient temperature extends the permitted range of body sizes for which b≥bmin. As a result, the maximum linear body size grows approximately twofold per each ten degrees of increase in ambient temperature. We also discuss why this prediction does not apply to interspecific comparisons of aquatic poikilotherms, for which an opposite trend is to be expected. We quantify in theory the maximum body size patterns recently reported for benthic gammaridean amphipods.

Abstract. In this paper the basic geophysical and ecological principles are jointly analyzed that allow the landmasses of Earth to remain moistened sufficiently for terrestrial life to be possible. 1. Under gravity, land inevitably loses water to the ocean. To keep land moistened, the gravitational water runoff must be continuously compensated by the atmospheric ocean-to-land moisture transport. Using data for five terrestrial transects of the International Geosphere Biosphere Program we show that the mean distance to which air fluxes can transport moisture over non-forested areas, does not exceed several hundred kilometers; precipitation decreases exponentially with distance from the ocean. 2. In contrast, precipitation over extensive natural forests does not depend on the distance from the ocean along several thousand kilometers, as illustrated for the Amazon and Yenisey river basins and Equatorial Africa. This points to the existence of an active biotic pump transporting atmospheric moisture inland from the ocean. 3. Physical principles of the biotic moisture pump are investigated based on the previously unstudied properties of atmospheric water vapor, which can be either in or out of aerostatic equilibrium depending on the lapse rate of air temperature. A novel physical principle is formulated according to which the low-level air moves from areas with weak evaporation to areas with more intensive evaporation. Due to the high leaf area index, natural forests maintain high evaporation fluxes, which support the ascending air motion over the forest and "suck in" moist air from the ocean, which is the essence of the biotic pump of atmospheric moisture. In the result, the gravitational runoff water losses from the optimally moistened forest soil can be fully compensated by the biotically enhanced precipitation at any distance from the ocean. 4. It is discussed how a continent-scale biotic water pump mechanism could be produced by natural selection acting on individual trees. 5. Replacement of the natural forest Correspondence to: A. M. Makarieva (elba@infopro.spb.su) cover by a low leaf index vegetation leads to an up to tenfold reduction in the mean continental precipitation and runoff, in contrast to the previously available estimates made without accounting for the biotic moisture pump. The analyzed body of evidence testifies that the long-term stability of an intense terrestrial water cycle is unachievable without the recovery of natural, self-sustaining forests on continent-wide areas.

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The influence of forest loss on rainfall remains poorly understood. Addressing this challenge Spracklen et al. recently presented a pan-tropical study of rainfall and landcover that showed that satellite-derived rainfall measures were positively correlated with the degree to which model-derived air trajectories had been exposed to forest cover. This result confirms the influence of vegetation on regional rainfall patterns suggested in previous studies. However, we find that the conclusion of Spracklen et al. -that differences in rainfall reflect air moisture content resulting from evapotranspiration while the circulation pattern remains unchanged -appears undermined by methodological inconsistencies. We identify methodological problems with the underlying analyses and the quantitative estimates for rainfall change predicted if forest cover is lost in the Amazon. We discuss some alternative explanations that include the distinct role of forest evapotranspiration in creating low pressure systems that draw moisture from the oceans to the continental hinterland. Our analysis of meteorological data from three regions in Brazil, including the central Amazon forest, reveal a tendency for rainy days during the wet season with column water vapor (CWV) exceeding 50 mm to have higher pressure than rainless days; while at lower CWV rainy days tend to have lower pressure than rainless days. The coupling between atmospheric moisture content and circulation dynamics underlines that the danger posed by forest loss is greater than suggested by focusing only on moisture recycling alone.

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