Papers are in English unless otherwise stated. See also an extended list of publications arranged by date and books on biotic regulation. For extended texts of presentations of V.G. Gorshkov and A.M. Makarieva at the Open Science Conference "Challenges of a Changing Earth", 10-13 July, 2001, Amsterdam, see abstracts and publications under Sections IV and V.

 

The biotic regulation concept Biotic regulation of the global carbon cycle Genetics in the biotic regulation concept The climate instability problem Information and orderliness of living systems Energetics of individual organisms and ecological communities WATER

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I. The biotic regulation concept

The following publications give a general outline of the biotic regulation concept and its implications for the global change science.
 

Gorshkov V.G., Makar'eva A.M., Losev K.S. (2006) A strategy for the survival of the humanity is on agenda. Herald of the Russian Academy of Sciences, 76(2), 139-143, doi:10.1134/S1019331606020055. PDF (500 Kb, Russian). The authors discuss the most important fallacies in modern climatology and ecology and priorities in solving the problems of environment and nature protection.

Gorshkov V.G., Makarieva A.M., Gorshkov V.V. (2004) Revising the fundamentals of ecological knowledge: the biota-environment interaction. Ecological Complexity, 1(1), 17-36, doi:10.1016/j.ecocom.2003.09.002. Abstract, PDF (320 Kb). Copyright 2003 Elsevier B. V. Further reproduction or electronic distribution is not permitted.
     We would recommend this paper as an up-to-date introduction to the biotic regulation concept.

Gorshkov V.V., Gorshkov V.G., Danilov-Danil'yan V.I., Losev K.S., Makar'eva A.M. (1999) Biotic control of the environment. Russian Journal of Ecology, 30(2), 87-96. Abstract, PDF (240 Kb).

Gorshkov V.G., Makarieva A.M., Mackey B., Gorshkov V.V. (2001) Biological theory and global change science. Global Change NewsLetter, 48, 11-14. PDF (0.3 Mb).

Gorshkov V.G., Makarieva A.M., Mackey B., Gorshkov V.V. (2002) How valid are the biological and ecological principles underpinning Global Change science? Energy & Environment, 13(3), 299-310. Abstract, PDF (500 Kb).

 

II. Biotic regulation of the global carbon cycle

Gorshkov V.G., Makar'eva A.M. (2002) Changes in the global carbon cycle: Evidence from the measurements of O₂/N₂ in the atmosphere and CO₂ partial pressure at the ocean-atmosphere boundary. Geochemistry International, 40(5), 472-480. Abstract, PDF (200 Kb).

Gorshkov V.G., Makarieva A.M. (1998) Impact of terrestrial and oceanic biota on the modern carbon and oxygen cycles. Ecological Chemistry, 7(2), 129-137. Abstract, PDF (electronic version, 240 Kb), PDF (offprint, 1.22 Mb).

Gorshkov V.G., Grassl H., Kondratyev K.Ya., Losev K.S. (1995) On the natural biological regulation of the environment. Environmental Conservation, 22(2), 170-174. Full text (PDF, 1.0 Mb).

 

III. Genetics in the biotic regulation concept

The following publications deal with the genetic aspect of the biotic regulation concept. Information needed for the highly-organized process of biotic regulation is written in the genomes of natural biological species. The evidence presented suggests that the observed genetic polymorphism of natural species is a consequence of a finite sensitivity of the process of natural selection, which allows a certain number of genetic defects to remain unnoticed in the course of competitive interaction of individuals and, consequently, to be preserved in the population.

Genetic polymorphism is thus interpreted not as an adaptive potential of a species (see here on the conflict between the biotic regulation and genetic adaptation concepts), but as a permissible level of erosion of the meaningful genetic information of species. Instead of being continuously changing in a certain direction, the genetic information of a species fluctuates randomly around the normal genome during the whole period of the species' existence, the magnitude of fluctuations being determined by the limit of sensitivity of natural selection.
 

Makarieva A.M., Gorshkov V.G. (2004) On the dependence of speciation rates on species abundance and characteristic population size. Journal of Biosciences, 29(1), 119-128. Abstract, PDF (110 Kb). Copyright Indian Academy of Sciences.

One of the widely spread postulates of modern biology is the statement that the driving force of evolution is the adaptation of biological species to changing environmental conditions. This statement contradicts the biotic regulation theory. If it were true, one could expect that species consisting of many individuals (e.g. bacteria with characteristic population density of a million per cubic millilitre) would evolve faster than species with a small number of individuals (e.g. mammals, 1 ind. per square km). In simple words, it seems easier to find an organism well fitted to a new environment if you choose among billions, rather than among a few, individuals. It is easy to calculate that the rate of speciation would differ by ten and more orders of magnitude among small numerous and large non-numerous organisms.
In this paper we analyze the available empirical data to show that in reality all organisms speciate at approximately the same rate, the mean time of species duration being of the order of several million years, be that a unicell or vertebrate species. This confirms the biotic regulation statement that evolution cannot be caused by changes of the environment, the latter being itself under control of the biota.
To learn more about the significance of this paper to the biotic regulation concept read Chapter 11 of Gorshkov et al. (2000) Biotic regulation of the environment.

Makarieva A.M. (2001) Variance of protein heterozygosity in different species of mammals with respect to the number of loci studied. Heredity, 87(1), 41-51. Abstract, PDF (0.4 Mb), dataset (HTML, 0.1 Mb).

Gorshkov V.G., Makar'eva A.M. (1999) Haldane's Rule and somatic mutations. Russian Journal of Genetics, 35(6), 611-617. Abstract, PDF (184 Kb).

Gorshkov V.G., Makar'eva A.M. (1997) Dependence of heterozygosity on body weight in mammals. Doklady Biological Sciences, 355, 384-386. PDF (127 Kb).

 

IV. The climate instability problem

It is shown, based on the analysis of the physical behaviour of the terrestrial greenhouse effect, that the climate with an extensive liquid hydrosphere like that of Earth is physically unstable with respect to spontaneous transitions to either of the two stable but life-incompatible states, that of complete evaporation of the hydrosphere with global mean surface temperature higher than +400°C and that of complete glaciation of the planet with temperature lower than -80° C.

The available paleodata testify for a stable maintenance of the values of global mean surface temperatures within the interval 5-25°C during the last seven hundred million years. In the absence of a physical mechanism that could ensure such a stability, the observed sustainability of the life-compatible values of surface temperature can be only explained by accepting the existence of biotic regulation of the global cycle of water (the major greenhouse gas on Earth) and surface temperature.
 

Gorshkov V.G., Makarieva A.M. (2002) Greenhouse effect dependence on atmospheric concentrations of greenhouse substances and the nature of climate stability on Earth. Atmospheric Chemistry and Physics Discussions, 2, 289-337. Abstract, PDF and discussion.

Makarieva A.M., Gorshkov V.G., Pujol T. (2003) Height of convective layer in planetary atmospheres with condensable and non-condensable greenhouse substances. Atmospheric Chemistry and Physics Discussions, 3, 6701-6720. Abstract, PDF and discussion.

Makar'eva A.M., Gorshkov V.G. (2001) The greenhouse effect and the stability of the global mean surface temperature. Doklady Earth Sciences, 377(2), 210-214. PDF (173 Kb).

Gorshkov V.G., Makarieva A.M. (2001) Diffusion of thermal photons in the atmosphere, pp. 141-142 in: Dakhno L.G. (Ed.) PNPI XXX (Petersburg Nuclear Physics Institute XXX-th Anniversary). Scientific Highlights. Theoretical Physics Division. PNPI, Gatchina, 145 pp., ISBN 5-86763-043-9. PDF (0.1 Mb).

Gorshkov V.G., Makarieva A.M. (2000) Environmental safety, climate stability and the non-perturbed biota. Global Change NewsLetter, 43, 24-25. PDF (0.1 Mb). recommended as introductory reading

Makarieva A.M. (2000) Biotically maintained stability of the Earth's mean global surface temperature. Petersburg Nuclear Physics Institute, Preprint No. 2384, 42 pp. PDF (0.4 Mb).

 

V. Information and orderliness of living systems

We point out that the quantitative gap between the level of orderliness inherent to living systems and that of open dynamic systems of physical nature 'self-organized' at the expense of external energy fluxes reaches twenty four orders of magnitude. The environmental fluxes of free energy being virtually disordered as compared to living systems, the orderliness of life cannot be maintained by any physical process. To counteract the spontaneous decay of the highly ordered state of life, the living matter must be divided into a sufficiently large number of equivalent objects (individuals), among which a competitive interaction is switched on. As soon as the level of orderliness of any given object diminishes below the sensitivity of competitive interaction, such an object loses competitiveness and is replaced by a copy of a normal object retaining the initial high level of order. Such a mechanism of maintenance of order is unique to living matter and differentiates it from the inanimate world.

We also show that the biotic regulation mechanism cannot be replaced by technology, as far as the information processing capacity of the civilisation is twenty orders of magnitude lower than that used by the natural biota for environmental control.
 

Gorshkov V.V., Gorshkov V.G., Danilov-Danil'yan V.I., Losev K.S., Makar'eva A.M. (2002) Information in the animate and inanimate worlds. Russian Journal of Ecology, 33(3), 149-155. Abstract, PDF (145 Kb).

Gorshkov V.G., Makar'eva A.M. (2001) On the possibility of physical self-organization of biological and ecological systems. Doklady Biological Sciences, 378, 258-261. PDF (123 Kb).

Makarieva A.M., Gorshkov V.G. (2000) Order in physical and living systems: Principal differences in quantitative characteristics and mechanisms of maintenance do not allow a similar description. Petersburg Nuclear Physics Institute, Preprint No. 2388, 17 pp. Abstract, PDF (1.3 Mb).

 

VI. 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 in natural ecological communities, in particular, the decline of the share of consumption of primary productivity with growing body size of the community's heterotrophs. So far most part of the original publications on this topic are in Russian. For compilations of the results obtained, and to see how these studies are related to the biotic regulation concept, the interested reader is referred to Chapters 1 (HTML, 100 Kb), 3 (PDF, 2.5 Mb) and 4 (PDF, 2.1 Mb) of Gorshkov et al. (2000) Biotic regulation of the environment. Key issue of global change. Springer, London, see also the book's contents.
 

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, doi:10.1111/j.1365-2435.2006.01070.x. Abstract, PDF (proofs, 350 Kb), supplementary data (WinWord). Copyright 2006 The Authors Journal Compilation; Copyright 2006 British Ecological 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, doi:10.1098/rspb.2005.3225. Abstract, PDF (submitted manuscript, 220 Kb). Copyright 2005 The Royal Society. 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, doi:10.1111/j.1365-2435.2005.01005.x. Abstract, PDF (200 Kb, proofs). 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, doi:10.1016/j.jtbi.2005.04.016. Abstract, PDF (270 Kb). Copyright 2005 Elsevier Ltd. 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, doi:10.1098/rspb.2005.3223. Abstract, PDF (submitted manuscript, 150 Kb). Copyright 2005 The Royal Society. Further reproduction or electronic distribution is not permitted.

Makarieva A.M., Gorshkov V.G., Li B.-L. (2005) Temperature-associated upper limits to body size in terrestrial poikilotherms. OIKOS, 111(3), 425-436. Abstract, PDF (330 Kb, proofs). Copyright 2005 OIKOS. Further reproduction or electronic distribution is not permitted.

The main result of the above studies consists in the demonstration of the fact 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 1-10 W/kg (Gorshkov, 1981). 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. For the lay person, this means that the growing environmental and ecological problems currrently faced by the humanity are attempted to be solved on a predominantly technological basis (which is a dead-end, according to the biotic regulation theory), without involving the huge regulatory potential of natural ecological communities of living organisms. These and other biotic regulation studies aim to change this outdated paradigm by emphasizing the active role of life in building both internal and external environments and keeping the Earth habitable for the humans.

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, doi:10.1016/j.ecocom.2005.04.006. Abstract, PDF (210 Kb). Copyright 2005 Elsevier B. V. Further reproduction or electronic distribution is not permitted.

According to the biotic regulation concept, natural ecological communities are organized such as to ensure maximum possible stability of their environment. 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. In the present paper we show that this can be achieved via encoding the territorial requirements of animals at the species level. 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. The necessity to satisfy this need make people move across large territories even when they cannot individually control them, like in the modern overpopulated cities. Human passion for tourism can be similarly explained. Deprivation of the freedom of movement (with simultaneous satisfaction of all other basic human needs like feeding, entertainment etc.) is the main punishment to which modern humans are exposed (imprisonment). Imprisonment in small cells could not become a punishment for our species if the biological organization of humans were compatible with existence on tiny areas equal to the inverse density of modern human population, about 100 square meters per individual, like in modern cities.

 

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(2), 139-175, doi:10.1016/j.ecocom.2004.02.003. Abstract, PDF (380 Kb). Copyright 2004 Elsevier B. V. Further reproduction or electronic distribution is not permitted.

According to the biotic regulation theory, natural ecological communities are organized such as to ensure maximum possible stability of their environment. In this paper it is shown that this principle allows to quantitatively predict patterns of energy partitioning among differently-sized organisms in stable ecosystems. It is shown 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.

 

Makarieva A.M., Gorshkov V.G., Li B.-L. (2004) Ontogenetic growth: models and theory. Ecological Modelling, 176, 15-26, doi:10.1016/j.ecolmodel.2003.09.037. Abstract, PDF (270 Kb).

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. In this paper we illustrate the above statements on the example of a popular ontogenetic growth model, which, among other things, violates the energy conservation law.

 

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(2), 301-307, doi:10.1006/jtbi.2003.3185. Abstract, PDF (140 Kb). Copyright 2002 Elsevier Science Ltd. Further reproduction or electronic distribution is not permitted.

Gorshkov V.G. (1985) Natural selection of communities and the stability of biogeochemical cycles. In: J. Mlikovsky and V.J.A. Novak (Eds.) Evolution and Morphogenesis, pp. 787-794. Academia, Praha. Abstract, PDF (240 Kb).

The following publications are in Russian with English abstracts:

Gorshkov V.G. (1984) Energetical efficiency of flight and swimming. Journal of General Biology, 45(6), 779-795 (in Russian). Abstract (engl), PDF (740 Kb).

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 (engl), PDF (1.0 Mb).

Gorshkov V.G. (1980) The structure of the biospheric energy flows. Botanical Journal, 65(11), 1579-1590 (in Russian). Abstract (engl), PDF (1.5 Mb).

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 (engl), PDF (700 Kb).