In raising awareness of Hamilton's ideas, as well as making its own valuable contributions to the field, the book has also stimulated research on human inclusive fitness.

[8] Dawkins begins by discussing the altruism that people display, indicating that he will argue it is explained by gene selfishness, and attacking group selection as an explanation.

He describes organisms as apparently purposive but fundamentally simple survival machines, which use negative feedback to achieve control.

He attacks the idea of group selection for the good of the species as proposed by V. C. Wynne-Edwards, arguing instead that each parent necessarily behaves selfishly.

Similarly, Dawkins argues, there are conflicts of interest between males and females, but he notes that R. A. Fisher showed that the optimal sex ratio is 50:50.

Amotz Zahavi's theory of honest signalling explains stotting as a selfish act, he argues, improving the springbok's chances of escaping from a predator by indicating how difficult the chase would be.

Altruism too 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.

The final chapter of the first edition introduced the idea of the meme, a culturally-transmitted entity such as a hummable tune, by analogy to genetic transmission.

His contention is that the genes that are passed on are the ones whose evolutionary consequences serve their own implicit interest (to continue the anthropomorphism) in being replicated, not necessarily those of the organism.

For example, Andrew Brown has written:[12] Donald Symons also finds it inappropriate to use anthropomorphism in conveying scientific meaning in general, and particularly in this instance.

There are other times when the implicit interests of the vehicle and replicator are in conflict, such as the genes behind certain male spiders' instinctive mating behaviour, which increase the organism's inclusive fitness by allowing it to reproduce but shorten its life by exposing it to the risk of being eaten by the cannibalistic female.

[16] Likewise, the persistence of junk DNA that [Dawkins believed at that time] provides no benefit to its host can be explained on the basis that it is not subject to selection.

However, as we shall see, there are special circumstances in which a gene can achieve its own selfish goals best by fostering a limited form of altruism at the level of individual animals."

Gene selection provides one explanation for kin selection and eusociality, where organisms act altruistically, against their individual interests (in the sense of health, safety or personal reproduction), namely the argument that by helping related organisms reproduce, a gene succeeds in "helping" copies of themselves (or sequences with the same phenotypic effect) in other bodies to replicate.

A requirement upon this claim, supported by Dawkins in Chapter 10: "You scratch my back, I'll ride on yours" by examples from nature, is the need to explain how genes achieve kin recognition, or manage to orchestrate mutualism and coevolution.

Although Dawkins (and biologists in general) recognize these phenomena result in more copies of a gene, evidence is inconclusive whether this success is selected for at a group or individual level.

The claim still is made by some that it would take only a single individual with a tendency towards more selfish behaviour to undermine a population otherwise filled only with the gene for altruism towards non-kin.

[22] Proponents argue that the central point, that replicating the gene is the object of selection, usefully completes and extends the explanation of evolution given by Charles Darwin before the basic mechanisms of genetics were understood.

[23] According to Grafen, these difficulties along with an initial conflict with population genetics models at the time of its introduction "explains why within biology the considerable scientific contributions it [The Selfish Gene] makes are seriously underestimated, and why it is viewed mainly as a work of exposition.

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] This point also was made by the philosopher Mary Midgley: "The same thing happened to AJ Ayer, she says, but he spent the rest of his career taking back what he'd written in Language, Truth and Logic.

[35][36] While naïve versions of group selectionism have been disproved, more sophisticated formulations make accurate predictions in some cases while positing selection at higher levels.

The proven best way in evolutionary biology, as in most of science, is to define a problem arising during empirical research, then select or devise the theory that is needed to solve it.

Almost all research in inclusive-fitness theory has been the opposite: hypothesize the key roles of kinship and kin selection, then look for evidence to test that hypothesis."

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).

[57][58] On BBC-2 TV, Dawkins pointed to evidence for a "Tit-for-Tat" strategy (shown to be successful in game theory[59]) as the most common, simple, and profitable choice.

[60] 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.

[61] The Selfish Gene was first published by Oxford University Press in 1976 in eleven chapters with a preface by the author and a foreword by Robert Trivers.