The Selfish Gene

Title

Dawkins recalls showing The Selfish Gene to Tom Maschler, who “liked the book but not the title". He suggested The Immortal Gene. Dawkins writes that "Maschler may have been right. Many critics, especially vociferous ones learned in philosophy as I have discovered, prefer to read a book by title only. No doubt this works well enough for The Tale of Benjamin Bunny or The Decline and Fall of the Roman Empire, but I can readily see that ‘The Selfish Gene’ on its own, without the large footnote of the book itself, might give an inadequate impression of its contents.”^[10]: x

Contents

1. Why Are People?

Dawkins writes that “Intelligent life on a planet comes of age when it first works out the reason for its own existence. ... Living organisms had existed on earth, without ever knowing why, for over three thousand million years before the truth finally dawned on one of them. His name was Charles Darwin.”^[10]: 1 Darwin (and Alfred Russel Wallace, working independently) discovered the mechanism of evolution: natural selection.

2. The Replicators

Dawkins introduces the term replicator to describe self-replicating molecules like DNA and RNA.^[10]: 19 He considers the origin of life with the emergence of replicators. The original replicator was the first molecule which managed to reproduce itself and thus gained an advantage over other molecules within the primordial soup. As replicating molecules became more complex, Dawkins postulates, they evolved cells serving as survival machines. Cells joined to form bodies.

3. Immortal Coils

Dawkins expands on DNA, its helical structure and its organization into chromosomes. Genes are DNA segments which are translated into proteins. Darwin’s coeval Gregor Mendel worked out the laws of inheritance and found that traits are inherited as discrete units. In meiosis, the production of gametes, genes are recombined during crossing over.

4. The Gene Machine

Dawkins discusses the evolution of behavior. Genes encoding behaviors that cause those genes to be passed on will naturally be selected for. He provides various examples.

5. Aggression

Dawkins discusses John Maynard Smith’s evolutionarily stable strategy, "a strategy which, if most members of a population adopt it, cannot be bettered by an alternative strategy … once an ESS is achieved it will stay: selection will penalize any deviation from it."^[10]: 90 A 50:50 ratio of 'hawks' (aggressors) and 'doves' (nonaggressors) is evolutionarily stable.

6. Genesmanship

Dawkins discusses kin selection: "Close relatives – kin – have a greater than average chance of sharing genes. It has long been clear that this must be why altruism by parents toward their young is so common. What R. A. Fisher, J. B. S. Haldane and especially W. D. Hamilton realized was that the same applies to other close relations—brothers and sisters, nephews and nieces, close cousins. If an individual dies in order to save ten close relatives, one copy of the kin-altruism gene may be lost, but a larger number of copies of the gene is saved.”^[10]: 116

7. Family Planning

Dawkins discusses David Lack’s principle. Natural selection, according to Lack, adjusts initial clutch size so as to take maximum advantage of these limited resources.

8. Battle of the Generations

Dawkins discusses R. L. Trivers’s concept of parental investment, ‘any investment by the parent in an individual offspring that increases the offspring’s chance of surviving (and hence reproductive success) at the cost of the parent’s ability to invest in other offspring’.^[10]: 160

9. Battle of the Sexes

Dawkins discusses Darwin’s theory of sexual selection. Females can afford to be choosy in their mates, and select for attractive traits in males. Fisher's principle explains why a 50:50 ratio of males to females is evolutionarily stable. This is true even in an extreme case like the harem-keeping elephant seal, where 4% of the males get 88% of copulations. In that case, the strategy of having a female offspring is safe, as she'll have a pup, but the strategy of having a male can bring a large return (dozens of pups), even though many males live out their lives as bachelors.

10. You Scratch My back, I’ll Ride on Yours

Dawkins discusses reciprocal altruism. Amotz Zahavi's theory of honest signaling explains stotting as a selfish act that improves the springbok's chances of escaping from a predator by indicating how difficult the chase would be. Dawkins discusses why many species live in groups, achieving mutual benefits through mechanisms such as Hamilton's selfish herd model: each individual behaves selfishly but the result is herd behaviour. Altruism can evolve, as in the social insects such as ants and bees, where workers give up the right to reproduce in favour of a sister, the queen; in their case, the unusual (haplodiploid) system of sex determination may have helped to bring this about, as females in a nest are exceptionally closely related.

11. Memes

Dawkins discusses cultural evolution, which is in some ways analogous to biological evolution. Dawkins proposes that units of information can propagate themselves like genes. “The new soup is the soup of human culture. We need a name for the new replicator or a unit of imitation. ‘Mimeme’ comes from a suitable Greek root, but I want a monosyllable that sounds a bit like ‘gene’. I hope my classicist friends will forgive me if I abbreviate mimeme to meme. If it is any consolation, it could alternatively be thought of as being related to ’memory’, or to the French word même. It should be pronounced to rhyme with ‘cream’.”^[10]: 249

Critiques

According to comparative psychologist Nicky Hayes, "Dawkins presented a version of sociobiology that rested heavily on metaphors drawn from animal behavior, and extrapolated these...One of the weaknesses of the sociological approach is that it tends only to seek confirmatory examples from among the huge diversity of animal behavior. Dawkins did not deviate from this tradition."^[16] More generally, critics argue that The Selfish Gene oversimplifies the relationship between genes and the organism.

In 1976, the ecologist Arthur Cain, one of Dawkins's tutors at Oxford in the 1960s, called it a "young man's book", a quote of a critique of the New College, Oxford^[a] philosopher A. J. Ayer's Language, Truth, and Logic (1936). Dawkins noted that he had been "flattered by the comparison, [but] knew that Ayer had recanted much of his first book and [he] could hardly miss Cain's pointed implication that [he] should, in the fullness of time, do the same."^[2] The philosopher Mary Midgley mused that "This hasn't occurred to Dawkins. He goes on saying the same thing."^[17] However, according to Wilkins and Hull,^[18] Dawkins's thinking has developed:

In Dawkins's early writings, replicators and vehicles played different but complementary and equally important roles in selection, but as Dawkins honed his view of the evolutionary process, vehicles became less and less fundamental...In later writings Dawkins goes even further and argues that phenotypic traits are what really matter in selection and that they can be treated independently of their being organized into vehicles...Thus, it comes as no surprise when Dawkins proclaims that he "coined the term 'vehicle' not to praise it but to bury it."^[19] As prevalent as organisms might be, as determinate as the causal roles that they play in selection are, reference to them can and must be omitted from any perspicuous characterization of selection in the evolutionary process. Dawkins is far from a genetic determinist, but he is certainly a genetic reductionist.

— John S Wilkins, David Hull, Dawkins on Replicators and Vehicles, The Stanford Encyclopedia of Philosophy

As to the unit of selection, Stephen Jay Gould, in The Structure of Evolutionary Theory, finds Dawkins's position tries to have it both ways:^[20]

Dawkins claims to prefer genes and to find greater insight in this formulation. But he allows that you or I might prefer organisms—and it really doesn't matter.

Choice of words

A good deal of objection to The Selfish Gene stemmed from its failure to be always clear about "selection" and "replication". Dawkins says the gene is the fundamental unit of selection, and then points out that selection does not act directly upon the gene, but upon "vehicles" or '"extended phenotypes". Stephen Jay Gould took exception to calling the gene a 'unit of selection' because selection acted only upon phenotypes.^[21] Summarizing the Dawkins-Gould difference of view, Sterelny says:^[22]

Gould thinks gene differences do not cause evolutionary changes in populations, they register those changes.

The word "cause" here is somewhat tricky: does a change in lottery rules (for example, inheriting a defective gene "responsible" for a disorder) "cause" differences in outcome that might or might not occur? It certainly alters the likelihood of events, but a concatenation of contingencies decides what actually occurs. Dawkins thinks the use of "cause" as a statistical weighting is acceptable in common usage.^[23] Like Gould, Gabriel Dover in criticizing The Selfish Gene says:^[24]

It is illegitimate to give 'powers' to genes, as Dawkins would have it, to control the outcome of selection...There are no genes for interactions, as such: rather, each unique set of inherited genes contributes interactively to one unique phenotype...the true determinants of selection.

However, from a comparison with Dawkins's discussion of this very same point, it would seem both Gould's and Dover's comments are more a critique of his sloppy usage than a difference of views.^[25] Hull suggested a resolution based upon a distinction between replicators and interactors.^{[26][26][27][27]}

Andrew Brown has written:^[28]

"Selfish", when applied to genes, doesn't mean "selfish" at all. It means, instead, an extremely important quality for which there is no good word in the English language: "the quality of being copied by a Darwinian selection process." This is a complicated mouthful. There ought to be a better, shorter word—but "selfish" isn't it.

Donald Symons also finds it inappropriate to use anthropomorphism in conveying scientific meaning in general, and particularly in this instance. He writes in The Evolution of Human Sexuality (1979):^[29]

In summary, the rhetoric of The Selfish Gene exactly reverses the real situation: through [the use of] metaphor genes are endowed with properties only sentient beings can possess, such as selfishness, while sentient beings are stripped of these properties and called machines...The anthropomorphism of genes...obscures the deepest mystery in the life sciences: the origin and nature of mind.

Enactive arguments

In Mind in Life, the philosopher Evan Thompson has assembled a multi-sourced objection to the "selfish gene".^[30] Thompson objects that the gene cannot operate by itself, since it requires an environment such as a cell, and life is "the creative outcome of highly structured contingencies". Thompson quotes Sarkar: "there is no clear technical notion of "information" in molecular biology. It is little more than a metaphor that masquerades as a theoretical concept and ...leads to a misleading picture of the nature of possible explanations in molecular biology."^[31]

Thompson follows with a detailed examination of the concept of DNA as a look-up-table and the role of the cell in orchestrating the DNA-to-RNA transcription, indicating that by anyone's account the DNA is hardly the whole story. Thompson goes on to suggest that the cell-environment interrelationship has much to do with reproduction and inheritance, and a focus on the gene as a form of "information [that] passes through bodies and affects them but is not affected by them on its way through"^[32]: 4 is tantamount to adoption of a form of material-informational dualism that has no explanatory value and no scientific basis. (Thomson, p. 187) The enactivist view, however, is that information results from the probing and experimentation of the agent with the agent's environment subject to the limitations of the agent's abilities to probe and process the result of probing, and DNA is simply one mechanism the agent brings to bear upon its activity.

Moral arguments

Dawkins's critics accuse him of promoting genetic determinism. However, he emphasizes that we are not at the mercy of our genes. Dawkins writes: "Let us understand what our own selfish genes are up to, because we may then at least have the chance to upset their design, something no other species has ever aspired to do."^[10]: 4

Nor does he promote selfishness: “We can even discuss ways of cultivating and nurturing pure, disinterested altruism, something that has no place in nature, something that has never existed before in the whole history of the world.”^[10]: 260 The philosopher Mary Midgley has suggested this position is a variant of Hobbes's explanation of altruism as enlightened self-interest, and that Dawkins goes a step further to suggest that our genetic programming can be overcome by what amounts to an extreme version of free will.^[33] Dawkins's major concluding theme, that humanity is finally gaining power over the "selfish replicators" by virtue of their intelligence, is criticized also by primatologist Frans de Waal, who refers to it as an example of a "veneer theory" (the idea that morality is not fundamental, but is laid over a brutal foundation).^[34]

Dawkins expanded on the evolution of cooperation in the documentary Nice Guys Finish First. He pointed to evidence for a "Tit-for-Tat" strategy (shown to be successful in game theory^[35]) as the most common, simple, and profitable choice.^[36] More generally, the objection has been made that The Selfish Gene discusses philosophical and moral questions that go beyond biological arguments, relying upon anthropomorphisms and careless analogies.^[37][3]

Memes

The psychologist Susan Blackmore wrote The Meme Machine (2000), with a foreword by Dawkins.^[38] James Gleick describes Dawkins's meme as "his most famous and memorable invention".^[39] The Selfish Gene popularized sociobiology in Japan after its translation in 1980.^[40] The "meme" entered the country's consciousness. Yuzuru Tanaka of Hokkaido University wrote Meme Media and Meme Market Architectures. The information scientist Osamu Sakura has published a book in Japanese and several papers in English on the topic.^[41] Nippon Animation produced the educational series The Many Dream Journeys of Meme.^[42]

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