Does life consistently maximise energy intensity?
Article first published online: 4 MAR 2010
© 2010 The Authors. Biological Reviews © 2010 Cambridge Philosophical Society
Volume 85, Issue 4, pages 859–879, November 2010
How to Cite
Milewski, A. V. and Mills, A. J. (2010), Does life consistently maximise energy intensity?. Biological Reviews, 85: 859–879. doi: 10.1111/j.1469-185X.2010.00131.x
- Issue published online: 4 MAR 2010
- Article first published online: 4 MAR 2010
- (Received 26 September 2009; revised 9 February 2010; accepted 10 February 2010)
We propose that a basic biological imperative of all organisms is to maximise energy (E) intensity, defined as the average rate of energy use per unit area of the Earth's surface. The dominant organism in any given environment is predicted to be that exerting the greatest E intensity regardless of evolutionary origin. Our ‘theory of biological E intensity’ thus explains variation in life form in terms of adaptations as opposed to accidents of biological history. It defines the competitive criterion in all metabolic pathways and industrial processes as the average rate of kinetic energy use, excluding heating but including all directed biological kinesis at scales up to the whole organism. A suggested unit for E intensity is joules per square meter per year. Because catalysts are crucial to extremely rapid use of energy (and therefore maximisation of E intensity), catalytic nutrient elements can be viewed as the ultimate means of life. It follows that a common denominator of all dominant organisms would be the acquisition of an optimal catalytic formula as determined by concentrations and ratios of C, H, O, N, S, Na, Mg, P, K, Ca, V, Cr, Mn, Fe, Co, Ni, Cu, Zn, Se, Mo, Cd, I, W, and Hg. The likely local shortages of various of these elements can theoretically be alleviated by various changes in the size, shape, and/or behaviour of organisms, depending on the environment. Thus, the availability, and potential for supplementation, of catalytic elements would be the ultimate basis for adaptation, largely determining which life form dominates in any particular location. The theory predicts the following. (1) In nutrient-rich environments offering the optimal catalytic formula, dominant organisms will be microbes. This is because microbes, and prokaryotes in particular, excel in E intensity through rapid biomolecular turnover, enabling them to usurp resources despite minimising biomass, complexity, and information. (2) Where the environment is catabolically dystrophic (i.e. scarce in certain nutrients required for catabolism), macrobes (e.g. humans and trees) will be superior competitors because they can collect and supplement nutrients and thereby approach the optimal catalytic formula. This enables macrobes, despite having considerably slower metabolism per unit body mass, to enhance E intensity relative to competing microbes constrained by catabolic dystrophy. Finally, (3) where the environment is anabolically dystrophic (i.e. scarce in certain nutrients required for anabolism) microbes will again dominate because biomolecular turnover can be relatively free from constraint given the limited fuel available. We suggest that an important and overlooked way to achieve power is to reuse energy, and that all organisms maximise E intensity by converting chemical potential energy (i.e. in fuel) into circuits of electromagnetic energy comprising electric charge, photons, and excited electrons. Because space and time merge subatomically, these electromagnetic circuits represent a concentration in spacetime of energy that (1) is concurrently kinetic and static, hence available for immediate use yet also conserved with minimal dissipation, and (2) ultimately promotes catalysis, which we assert is the primary biological tactic for maximising E intensity and thus fitness.