Does death drive the scaling of life?

Abstract

The magnitude of many kinds of biological structures and processes scale with organismal size, often in regular ways that can be described by power functions. Traditionally, many of these "biological scaling" relationships have been explained based on internal geometric, physical, and energetic constraints according to universal natural laws, such as the "surface law" and "3/4-power law". However, during the last three decades it has become increasingly apparent that biological scaling relationships vary greatly in response to various external (environmental) factors. In this review, I propose and provide several lines of evidence supporting a new ecological perspective that I call the "mortality theory of ecology" (MorTE). According to this viewpoint, mortality imposes time limits on the growth, development, and reproduction of organisms. Accordingly, small, vulnerable organisms subject to high mortality due to predation and other environmental hazards have evolved faster, shorter lives than larger, more protected organisms. A MorTE also includes various corollary, size-related internal and external causative factors (e.g. intraspecific resource competition, geometric surface area to volume effects on resource supply/transport and the protection of internal tissues from environmental hazards, internal homeostatic regulatory systems, incidence of pathogens and parasites, etc.) that impact the scaling of life. A mortality-centred approach successfully predicts the ranges of body-mass scaling slopes observed for many kinds of biological and ecological traits. Furthermore, I argue that mortality rate should be considered the ultimate (evolutionary) driver of the scaling of life, that is expressed in the context of other proximate (functional) drivers such as information-based biological regulation and spatial (geometric) and energetic (metabolic) constraints.