Fisher, Fitness, and the Fundamentals of Population Genetics
Zachary Hancock is a postdoctoral researcher at the University of Michigan and specializes in evolutionary genetics.
Evolution by natural selection requires that there exists variation in fitness between individuals. This simple truism goes back to Darwin’s Origin of Species, but where Darwin relied on verbal arguments, the statistician and population geneticist R.A. Fisher sought to codify this basic fact mathematically.
In 1930, Fisher published The Genetical Theory of Natural Selection, and within introduced the fundamental theorem of natural selection (FTNS). The FTNS is, in effect, a restatement of Darwin’s idea on the reliance of natural selection on variation. In words, the FTNS states that the rate of change in a population’s average fitness is equal to the additive genetic variance in fitness at that time. An implication of this is that when there’s a ton of variation in fitness, the population’s average fitness can increase rapidly by natural selection. But if there is very little differences in fitness among individuals, any subsequent fitness increase must be slow or absent entirely.
The basic idea behind the FTNS is quite simple, even if the derivation mathematically has caused many biologists to scratch their heads over the many decades since Fisher first introduced it. Some, like Warren Ewens (Ewens 1989), have argued that Fisher’s theorem is useless, while others, such as George Price (Price 1972), Alan Grafen (Grafen 2003), and Sean Rice (Rice 2004) have contended it is, indeed, a general theorem properly understood. Point here is that the FTNS has been a source of much debate for decades.
In 2018, two creationists wandered into this rather esoteric historic dispute. And as is characteristic of creationists, they did so without a clear appreciation of the field they were embarking to critique and with highly suspect motivations. Bill Basener—a mathematician and data scientist at the University of Virginia—and John Sanford—retired Cornell plant geneticist, inventor of the gene gun, and promoter of “genetic entropy”—together (and “BS” hereafter) published a paper in the Journal of Mathematical Biology titled “The fundamental theorem of natural selection with mutations.”