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Tuesday, October 27, 2026

The Selfish Gene: a critic and a defender are both wrong!

Richard Dawkins published The Selfish Gene in 1976 and in 2017 the Royal Society conducted a popular poll that voted it the most inspiring science book of all time.

I don't think that book is a good representation of modern evolutionary thinking and I've expressed that opinion many times on this blog [The selfish gene vs the lucky allele] ["The Selfish Gene" turns 40] [The "selfish gene" is not a good metaphor to describe evolution] [Die, selfish gene, die!].

The issues are complicated—here's how Dawkins described his book in the 1989 edition.

The selfish gene theory is Darwin's theory, expressed in a way that Darwin did not choose but whose aptness, I should like to think, he would instantly have recognized and delighted in. It is in fact a logical outgrowth of orthodox neo-Darwinism, but expressed as a novel image. Rather than focus on the individual organism, it takes a gene's-eye view of nature. It is a different way of seeing, not a different theory.

My objections are that the book is far too adaptationist and too "Darwinian." It fails to convey the complications of evolution to the general public, including the dominance of chance, accident, and contingency. It misrepresents some views of evolution, especially punctuated equilibria. It focuses too much on genes (alleles) as the unit of selection (evolution?) at the expense of individuals, groups, and species. (Species sorting may not be important but it deserved more than passing dismissal.) It focuses too much on the evolution of animals, especially animal behavior and that feeds into the biases of the general public who know very little about the vast majority of species outside of the animal kingdom.

Nathaniel Comfort has just published an article in Nature expressing his own disappointment about The Selfish Gene [The Selfish Gene, 50 years on — it’s time for an update]. He also doesn't like the book but for entirely different reasons.

Both components of the modern synthesis focus on the gene as the sole determinant of heredity. Both halves reached essentially universal acceptance well before Dawkins came along — the spit and swagger of the field’s proponents keeping challenges and alternatives at bay.

To a historian, this universal acceptance sounds close to ideology. Fortunately, since The Selfish Gene was published, scientists’ understanding of the basic mechanisms that drive evolution has changed drastically.

For instance, neo-Darwinism teaches that evolution is a random walk through changing environments, with selection favouring whatever traits help an organism to survive. Today, many scholars argue that there has to be direction to evolution. In How Life Works (2023), for instance, journalist Philip Ball points out that in a single cell, there is an unfathomably large number of possible interactions between different kinds of molecule — and there are about 37 trillion cells in one human body. “Such a system,” Ball writes, “can only hope to be stable if, out of all this complexity, only a limited number of collective ways of being may emerge”. Living organisms are forced to evolve in those few directions.

Firstly, "scientists' understanding of the basic mechanisms" changed BEFORE The Selfish Gene was written. That change occurred in the late 1960s with the development of neutral theory and the recognition that random genetic drift was the dominant mechanism of evolution. Secondly, Philip Ball is dead wrong about all kinds of things, including agency. [Philip Ball doesn't understand sloppy genomes]

Comfort keeps digging deeper with an appeal to the extended evolutionary synthesis, the epigenetics myth, and the microbiome. (What percentage of all evolution over 3.5 billion years has anything to do with probiotics?)

For years, a group of biologists, philosophers and historians has been calling for an extended evolutionary synthesis that accounts for phenomena beyond this isolated window of transmission. Heredity operates at many levels at once. For example, there’s some evidence that epigenetic changes — molecular modifications to the genome that can alter gene expression without changing DNA — are heritable. Ecosystems involve many life forms that adapt to one another, creating stability and developing a kind of homeostasis — the self-regulating process through which balance is maintained — that confers resilience against catastrophic change. Your body consists of more microbial cells than cells derived from your parents’ genes. The microbiome doesn’t participate directly in your genetic heredity, but you can’t live without it.

Jerry Coyne didn't like the Nathaniel Comfort article [Nathaniel Comfort’s misguided take on The Selfish Gene at 50]. Coyne argues that there's nothing new in the so-called extended synthesis and he quotes Douglas Futuyma as support.

That's good but unfortunately Coyne gets so caught up in defending the Dawkins' view that he doesn't recognize the more legitimate criticisms that many people have raised. Let me quote from another article by Douglas Futuyma (Futuyma, 2023).

There is no question that by the 1960s, selection was thought to be by far the most important of the factors that influence allele frequency change, owing to diverse studies in ecological change. ... The neutral theory of molecular evolution, proposed by Kimura contrasted strikingly with the current thinking, and resulted in the neutralist-selectionist controversy that engendered a large, diverse range of research programs in evolutionary biology for the next decade or more. ... The outcome, as we now know, is that sequence variation within and among species includes both selected and neutral (or nearly neutral) sites with immense consequences for understanding genome variation and evolution. ... Of all the events in evolutionary biology that I have witnessed, this is the single one that has most changed our understanding revolutionary.

Futuyma goes on to say, "The only controversy that might be called a substantial 'paradigm shift' was the neutral theory of molecular evolution."

Futuyma, like Coyne, dismisses the so-called extended evolutionary synthesis as trivial or unimportant but, unlike Coyne or Dawkins he proclaims the development of neutral theory a revolutionary change that might even be called a paradigm shift. Coyne is right to question the silliness of Nathaniel Comfort's article (Philip Ball - really?) but Coyne is wrong to ignore the fact that Dawkins said nothing about the revolutionary change in evolutionary theory that occurred years before he published The Selfish Gene.


Futuyma, D.J. (2023) The Evolving Evolutionary Synthesis. Evolutionary Biology: Contemporary and Historical Reflections Upon Core Theory. T. E. Dickens and B. J. A. Dickens, Springer: 517-541.

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