Natural selection. I. Variable environments and uncertain returns on investment

TL;DR

This paper introduces a unified framework for understanding variability in reproductive success and fitness, emphasizing relative success, diminishing returns, and hierarchical population structure, with implications for biology and economics.

Contribution

It provides a concise, integrated theory of variability in populations, clarifying the structure of fitness measures and their hierarchical nature across traits and individuals.

Findings

Relative success measures fitness through comparison to population average.

Diminishing returns reduce fitness as variability increases.

Hierarchical variability affects individual and population-level success.

Abstract

Many studies have analyzed how variability in reproductive success affects fitness. However, each study tends to focus on a particular problem, leaving unclear the overall structure of variability in populations. This fractured conceptual framework often causes particular applications to be incomplete or improperly analyzed. In this paper, I present a concise introduction to the two key aspects of the theory. First, all measures of fitness ultimately arise from the relative comparison of the reproductive success of individuals or genotypes with the average reproductive success in the population. That relative measure creates a diminishing relation between reproductive success and fitness. Diminishing returns reduce fitness in proportion to variability in reproductive success. The relative measurement of success also induces a frequency dependence that favors rare types. Second, variability in populations has a hierarchical structure. Variable success in different traits of an individual affects that individual's variation in reproduction. Correlation between different individuals' reproduction affects variation in the aggregate success of particular alleles across the population. One must consider the hierarchical structure of variability in relation to different consequences of temporal, spatial, and developmental variability. Although a complete analysis of variability has many separate parts, this simple framework allows one to see the structure of the whole and to place particular problems in their proper relation to the general theory. The biological understanding of relative success and the hierarchical structure of variability in populations may also contribute to a deeper economic theory of returns under uncertainty.