The Pace of Life: Metabolic Energy, Biological Time, and Life History

New biophysical theory and electronic databases raise the prospect of deriving fundamental rules of life, a conceptual framework for how the structures and functions of molecules, cells and individual organisms give rise to emergent patterns and processes of ecology, evolution and biodiversity. This framework is very general, applying across taxa of animals from 10-10 g protists to 108 g whales, and across environments from deserts and abyssal depths to rain forests and coral reefs. It has several hallmarks: 1) Energy is the ultimate limiting resource for organisms and the currency of biological fitness. 2) Most organisms are nearly equally fit, because in each generation at steady state they transfer an equal quantity of energy (22.4 kJ/g) and biomass (1 g/g) to surviving offspring. This is the equal fitness paradigm (EFP) of Brown et al. (2018). 3) The enormous diversity of life histories is due largely to variation in metabolic rates (e.g., energy uptake and expenditure via assimilation, respiration and production) and biological times (e.g., generation time). As in standard allometric and metabolic theory, most physiological and life history traits scale approximately as quarter-power functions of body mass, m (rates as ∼m-1/4 and times as ∼m1/4), and as exponential functions of temperature. 4) Time is the fourth dimension of life. Generation time is the pace of life. 5) There is, however, considerable variation not accounted for by the above scalings and existing theories. Much of this "unexplained" variation is due to natural selection on life history traits to adapt the biological times of generations to the clock times of geochronological environmental cycles. 7) Most work on biological scaling and metabolic ecology has focused on respiration rate. The emerging synthesis applies conceptual foundations of energetics and the EFP to shift the focus to production rate and generation time.