Fisher’s Fundamental Theorem of Natural Selection Isn’t Fundamental After All

Abstract

Fisher’s Fundamental Theorem of Natural Selection (FTNS) was called “biology’s central theorem” (Fisher, 1930, pgs. 36–37; Brockman, 2011; Royal Society, 2020). FTNS might possibly have been accorded this status for decades because Fisher himself declared his own theorem to be fundamental to biology (Fisher, 1930, pgs. 36–37). However, the idea that Fisher’s theorem is biology’s central theorem is by-and-large a myth promoted by popu- lar science writers like Richard Dawkins (Brockman, 2011). Joseph Felsenstein, when delivering the 2018 Fisher Memorial Lecture declared that FTNS was “alas, not so fundamental” (Felsenstein, 2018; Felsenstein, 2017, pg. 94. One may be hard-pressed to find a biology textbook or biology student who can explain how FTNS helps them understand biology. Even the meaning and proof of the FTNS have re- mained contentious even to this day (Price, 1972; Basener and Sanford, 2018).
 Not only does FTNS do little to nothing to explain biological evolution, but like most population genetic and evolutionary literature, FTNS relies on a definition of fit- ness in terms of population growth rates rather than the biophysical notions of fitness which are more in line with the common-sense intuitions of the medical and engineering communities.
 From the perspective of the biophysical (rather than the population growth) notion of fitness, natural selection might be more accurately described as an agent against the increase of complexity rather than an agent for it. Thus, metaphorically speaking, some sort of anti-Weasel model of natural selection might better describe how selection actu- ally works in nature rather than Dawkins’ Weasel or other man-made genetic algorithms.
 However, the main focus of this communication is to pro- vide some pedagogical insights through simple numerical illustrations of Fisher’s Theorem. The hope is that this will show the general irrelevance of FTNS to the question of the evolution of complexity by means of natural selection, and thus show that Fisher’s Theorem is not so fundamental after all.