Energetics of individual organisms and ecological communities

The biotic regulation concept predicts that natural ecological communities of species should be organized in a manner ensuring their maximum environmental stability. This general principle allows to explain major allometric patterns of distribution of energy fluxes observed at the level individual organisms, biological species and natural ecological communities.

One of the main findings is that, independent of their body size, all living organisms, from bacteria to whales, work to keep their mass-specific metabolic power within universal limits near the metabolic optimum of 1-10 W/kg (, 1981, Makarieva et al., 2008). This result opposes the common mechanistic view on the living matter, according to which biological features of the living organisms are shaped by their external environment, to which they presumably have to continuously adapt. Instead, our studies show that living organisms are able to overcome the physical limitations imposed on them by their environment and their own physical properties, maintaining optimal, preferred biochemical characteristics (in this case, mass-specific metabolic rate). As is demonstrated by other studies, they can also exert a stabilising impact on their immediate environment. In the modern biological paradigm the complexity of living organisms and their environmental abilities are practically ignored.

Another important pattern is that in stable ecosystems large animals are allowed to consume no more than 1% of primary production. In the modern biosphere man has exceeded this ecological quota by ten times (, 1980, 1981, Makarieva et al., 2004).

Fluctuations of population densities of heterotrophs lead to fluctuations of ecosystem energy flows and biomass. Thus, the stability principle demands that animal population densities must be kept within the corridor of ecological sustainability. This can be achieved via genetic encoding the territorial requirements of animals at the species level. (Makarieva et al., 2005). We present evidence in support of the statement that animals are biologically organized to occupy exclusive home range areas where no conspecific intruders are normally tolerated, this being a major mechanism of animal population numbers control in natural ecosystems. Applied to humans, this means that the need/right for a fairly large territory to be controlled by the individual is one of the most essential human needs/rights, as biologically indispensable as are, for example, the need/right for food and water.

Modelling is widely spread in modern natural science. Models differ from theories in that they include immeasurable parameters and unknown dependencies between measurable variables. These dependencies have therefore to be postulated, which is commonly done by fitting the model to the available empirical data. Thus, models are in their essence equivalent to tabulations of relevant data having zero predictive power. The modelling approach where the search for fundamental natural regularities is replaced by formal fitting and computer simulations represents a serious, if not deadly, disease of modern natural science. We illustrate the above statements on the example of a popular ontogenetic growth model, which, among other things, violates the energy conservation law (Makarieva et al., 2004).

Documents are listed in reverse chronological order.

Book chapters (PDF)

V. G. Gorshkov, V. V. Gorshkov, A. M. Makarieva (2000) Biotic Regulation of the Environment: Key Issue of Global Change. Springer-Praxis Series in Environmental Sciences, 367 pp. Praxis: Chichester, Springer: Berlin.
Chapter 3. Ecology of organisms with different body sizes
Chapter 4. Ecology of locomotive animals

V. G. Gorshkov (1995) Physical and Biological Bases of Life Stability. Man. Biota. Environment. Springer: Berlin.
Chapter 4. Stability of the Biosphere's Organization
5.1. Metabolic Power of the Individual; 5.2. Body Size Limits; 5.3. Energetics and Body Size of Photosynthesizing Plants; 5.4. Fluctuations in Synthesis and Destruction of Organic Matter; 5.5. Immotile and Locomotive Organisms; 5.6. Distribution of Consumption by Heterotrophs According to Their Body Size; 5.7. Daily Average Travelling Distance; 5.8. The Maximum Speed of Movement for Animals; 5.9. Maximum Permissible Share of Biomass Consumed by Mobile Animals; 5.10. Settled and Nomadic Lifestyles for Locomotive Animals; 5.11. Carnivores; 5.12. Diffusion of Excreta; 5.13. Detailed Distribution of Destructiveness with Body Size; 5.14. Brief Conclusions

Papers

Makarieva A.M., Gorshkov V.G., Li B.-L. (2009) Comment on "Energy uptake and allocation during ontogeny". Science, 325, 1206-a. Abstract. pdf

Makarieva A.M., Gorshkov V.G., Li B.-L. (2009) Re-calibrating the snake palaeothermometer. Nature, 460, E2-E3. Abstract. pdf Figure 1.

Makarieva A.M., Gorshkov V.G., Li B.-L., Chown S.L., Reich P.B., Gavrilov V.M. (2008) Mean mass-specific metabolic rates are strikingly similar across life's major domains: Evidence for life's metabolic optimum. Proceedings of the National Academy of Sciences U.S.A., 105, 16994-16999. Abstract. pdf http://www.pnas.org/content/105/44/16994, Supplementary Information Complete File, 212 pp. (PDF, 1.2 Mb)

Makarieva A.M., Gorshkov V.G., Li B.-L. (2006) Distributive network model of Banavar, Damuth, Maritan and Rinaldo (2002): Critique and perspective. Journal of Theoretical Biology, 239, 394-397. No abstract. pdf doi:10.1016/j.jtbi.2005.08.018.

Makarieva A.M., Gorshkov V.G., Li B.-L., Chown S.L. (2006) Size- and temperature-independence of minimum life-supporting metabolic rates. Functional Ecology, 20, 83-96. Abstract. pdf Supplementary data.

Makarieva A.M., Gorshkov V.G., Li B.-L. (2005) Why do population density and inverse home range scale differently with body size? Implications for ecosystem stability. Ecological Complexity, 2, 259-271. Abstract. pdf doi:10.1016/j.ecocom.2005.04.006. Copyright 2005 Elsevier B. V. Further reproduction or electronic distribution is not permitted.

Makarieva A.M., Gorshkov V.G., Li B.-L. (2005) Biochemical universality of living matter and its metabolic implications. Functional Ecology, 19, 547-557. Abstract. pdf doi:10.1111/j.1365-2435.2005.01005.x. Copyright 2005 British Ecological Society. Further reproduction or electronic distribution is not permitted.

Makarieva A.M., Gorshkov V.G., Li B.-L. (2005) Revising the distributive networks models of West, Brown and Enquist (1997) and Banavar, Maritan and Rinaldo (1999): Metabolic inequity of living tissues provides clues for the observed allometric scaling rules. Journal of Theoretical Biology, 237, 291-301. Abstract. pdf doi:10.1016/j.jtbi.2005.04.016.

Makarieva A.M., Gorshkov V.G., Li B.-L. (2005) Temperature-associated upper limits to body size in terrestrial poikilotherms. OIKOS, 111, 425-436. Abstract. pdf Copyright 2005 OIKOS. Further reproduction or electronic distribution is not permitted.

Makarieva A.M., Gorshkov V.G., Li B.-L. (2005) Gigantism, temperature and metabolic rate in terrestrial poikilotherms. Proceedings of the Royal Society of London, Biological Series, 272, 2325-2328. Abstract. pdf doi:10.1098/rspb.2005.3223. Copyright 2005 The Royal Society. Further reproduction or electronic distribution is not permitted.

Makarieva A.M., Gorshkov V.G., Li B.-L. (2005) Energetics of the smallest: Do bacteria breathe at the same rate as whales? Proceedings of the Royal Society of London, Biological Series, 272, 2219-2224. Abstract. pdf doi:10.1098/rspb.2005.3225. Copyright 2005 The Royal Society. Further reproduction or electronic distribution is not permitted.

Makarieva A.M., Gorshkov V.G., Li B.-L. (2004) Ontogenetic growth: models and theory. Ecological Modelling, 176, 15-26. Abstract. pdf

Makarieva A.M., Gorshkov V.G., Li B.-L., Losev K.S. (2004) The upper and lower ecological limits of specific metabolic power of different organisms. Russian Journal of Ecology, 35, 10-15. (Translated from Ekologiya, No. 1, 2004, 13-20.) Abstract. pdf

Makarieva A.M., Gorshkov V.G., Li B.-L. (2004) Body size, energy consumption and allometric scaling: a new dimension in the diversity-stability debate. Ecological Complexity, 1, 139-175. Abstract. pdf doi:10.1016/j.ecocom.2004.02.003. Copyright 2004 Elsevier B. V. Further reproduction or electronic distribution is not permitted.

Makarieva A.M., Gorshkov V.G., Li B.-L. (2003) A note on metabolic rate dependence on body size in plants and animals. Journal of Theoretical Biology, 221, 301-307. Abstract. pdf Copyright 2002 Elsevier Science Ltd. Further reproduction or electronic distribution is not permitted. Abstract provided by the authors.

Gorshkov V.G. (1984) Energetical efficiency of flight and swimming. Journal of General Biology, 45, 779-795. [in Russian] Abstract. pdf

Gorshkov V.G. (1983) Power and rate of locomotion in animals of different sizes. Journal of General Biology, 44(5), 661-678. [in Russian] Abstract. pdf

Gorshkov V.G. (1981) The distribution of energy flow among the organisms of different dimensions. Journal of General Biology, 42(3), 417-429. [in Russian] Abstract. pdf

Gorshkov V.G. (1980) The structure of the biospheric energy flows. Botanical Journal, 65(11), 1579-1590. [in Russian] Abstract. pdf