A lineage is expected to evolve to maximise its inclusive fitness—the number of copies of its genes passed on globally (rather than by a particular individual). As a result, populations will tend towards an evolutionarily stable strategy. The book also introduces the term meme for a unit of human cultural evolution analogous to the gene, suggesting that such "selfish" replication may also model human culture, in a different sense. Memetics has become the subject of many studies since the publication of the book. 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.
Dawkins uses the term "selfish gene" as a way of expressing the gene-centered view of evolution. As such, the book is not about a particular gene that causes selfish behaviour; in fact, much of the book's content is devoted to explaining the evolution of altruism. In the foreword to the book's 30th-anniversary edition, Dawkins said he "can readily see that [the book's title] might give an inadequate impression of its contents" and in retrospect wishes he had taken Tom Maschler's advice and titled it The Immortal Gene. He laments that “Too many people read it by title only.” In response, he expanded on the evolution of altruism in the BBC documentary Nice Guys Finish First.
Book
Contents
:It is finally time to return to the problem with which we started, to the tension between individual organism and gene as rival candidates for the central role in natural selection...One way of sorting this whole matter out is to use the terms 'replicator' and 'vehicle'. The fundamental units of natural selection, the basic things that survive or fail to survive, that form lineages of identical copies with occasional random mutations, are called replicators. DNA molecules are replicators. They generally, for reasons that we shall come to, gang together into large communal survival machines or 'vehicles'.
::— Richard Dawkins, The Selfish Gene, p. 253 (Anniversary Edition)
The original replicator (Dawkins Replicator) was the initial molecule which first managed to reproduce itself and thus gained an advantage over other molecules within the primordial soup. As replicating molecules became more complex, Dawkins postulates, the replicators became the genes within organisms, with each organism's body serving the purpose of a 'survival machine' for its genes.
Dawkins writes that gene combinations which help an organism to survive and reproduce tend to also improve the gene's own chances of being replicated, and, as a result, "successful" genes frequently provide a benefit to the organism. An example of this might be a gene that protects the organism against a disease. This helps the gene spread, and also helps the organism.
Genes vs organisms
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. Another example is the existence of segregation distorter genes that are detrimental to their host, but nonetheless propagate themselves at its expense. 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. These unselected for but transmitted DNA variations connect the individual genetically to its parents but confer no survival benefit.
These examples might suggest that there is a power struggle between genes and their interactor. In fact, the claim is that there isn't much of a struggle because the genes usually win without a fight. However, the claim is made that if the organism becomes intelligent enough to understand its own interests, as distinct from those of its genes, there can be true conflict.
An example of such a conflict might be a person using birth control to prevent fertilisation, thereby inhibiting the replication of his or her genes. But this action might not be a conflict of the 'self-interest' of the organism with his or her genes, since a person using birth control might also be enhancing the survival chances of their genes by limiting family size to conform with available resources, thus avoiding extinction as predicted under the Malthusian model of population growth.
Altruism
Dawkins says that his "purpose" in writing The Selfish Gene is "to examine the biology of selfishness and altruism." He does this by supporting the claim that "gene selfishness will usually give rise to selfishness in individual behaviour. 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. The claim is made that these "selfish" actions of genes lead to unselfish actions by organisms. 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. In fact, Dawkins has proposed that it is at the level of the extended phenotype:
:We agree [referring to Wilson and Sober's book Unto others: The evolution and psychology of unselfish behavior] that genes are replicators, organisms and groups are not. We agree that the group selection controversy ought to be a controversy about groups as vehicles, and we could easily agree to differ on the answer...I coined the [term] vehicle not to praise it but to bury it...Darwinism can work on replicators whose phenotypic effects (interactors) are too diffuse, too multi-levelled, too incoherent to deserve the accolade of vehicle...Extended phenotypes can include inanimate artifacts like beaver dams...But the vehicle is not something fundamental...Ask rather "Is there a vehicle in this situation and, if so, why?"
::—Richard Dawkins, Burying the Vehicle
Although Dawkins agrees that groups can assist survival, they rank as a "vehicle" for survival only if the group activity is replicated in descendants, recorded in the gene, the gene being the only true replicator. An improvement in the survival lottery for the group must improve that for the gene for sufficient replication to occur. Dawkins argues qualitatively that the lottery for the gene is based upon a very long and broad record of events, and group advantages are usually too specific, too brief, and too fortuitous to change the gene lottery:
::—Richard Dawkins, *The Selfish Gene*, [pp. 254–255](https://books.google.com/books?id=WkHO9HI7koEC&pg=PA254)
Prior to the 1960s, it was common for altruism to be explained in terms of group selection, where the benefits to the organism or even population were supposed to account for the popularity of the genes responsible for the tendency towards that behaviour. Modern versions of "multilevel selection" claim to have overcome the original objections, namely, that at that time no known form of group selection led to an evolutionarily stable strategy. 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.
## Reception
*The Selfish Gene* was extremely popular when first published, causing "a silent and almost immediate revolution in biology", and it continues to be widely read. It has sold over a million copies and has been translated into more than 25 languages. 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.
According to the ethologist Alan Grafen, acceptance of adaptionist theories is hampered by a lack of a mathematical unifying theory and a belief that anything in words alone must be suspect. 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." More generally, critics argue that *The Selfish Gene* oversimplifies the relationship between genes and the organism. (As an example, see Thompson.)
*The Selfish Gene* further popularised sociobiology in Japan after its translation in 1980. With the addition of Dawkins's book to the country's consciousness, the term "meme" entered popular culture. Yuzuru Tanaka of Hokkaido University wrote a book, *Meme Media and Meme Market Architectures*, while the psychologist Susan Blackmore wrote *The Meme Machine* (2000), with a foreword by Dawkins. The information scientist Osamu Sakura has published a book in Japanese and several papers in English on the topic. Nippon Animation produced an educational television program titled *The Many Journeys of Meme*.
In 1976, the ecologist Arthur Cain, one of Dawkins's tutors at Oxford in the 1960s, called it a "young man's book" (which Dawkins points out was a deliberate quote of a commentator on the New College, Oxford 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." 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*. "This hasn't occurred to Dawkins", she says. "He goes on saying the same thing."" However, according to Wilkins and Hull, Dawkins's thinking has developed, although perhaps not defusing this criticism:
: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." 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*
In *Humankind: A Hopeful History*, Rutger Bregman contrasts the portrayal of human nature in *The Selfish Gene* with a more cooperative view.
### Units of selection
As to the unit of selection: "One internally consistent logical picture is that the unit of replication is the gene,...and the organism is one kind of ...entity on which selection acts directly." Dawkins proposed the matter without a distinction between 'unit of replication' and 'unit of selection' that he made elsewhere: "the fundamental unit of selection, and therefore of self-interest, is not the species, nor the group, nor even strictly the individual. It is the gene, the unit of heredity." However, he continues in a later chapter:
:On any sensible view of the matter Darwinian selection does not work on genes directly. ...The important differences between genes emerge only in their *effects*. The technical word *phenotype* is used for the bodily manifestation of a gene, the effect that a gene has on the body...Natural selection favours some genes rather than others not because of the nature of the genes themselves, but because of their consequences—their phenotypic effects...But we shall now see that the phenotypic effects of a gene need to be thought of as *all the effects that it has on the world*. ...The phenotypic effects of a gene are the tools by which it levers itself into the next generation. All I am going to add is that the tools may reach outside the individual body wall...Examples that spring to mind are artefacts like beaver dams, bird nests, and caddis houses.
::— Richard Dawkins, *The Selfish Gene*, Chapter 13, pp. 234, 235, 238
Dawkins's later formulation is in his book *The Extended Phenotype* (1982), where the process of selection is taken to involve every possible phenotypical effect of a gene.
Stephen Jay Gould finds Dawkins's position tries to have it both ways:
: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.
::— Stephen Jay Gould, *The Structure of Evolutionary Theory*, pp. 640-641
The view of *The Selfish Gene* is that selection based upon groups and populations is rare compared to selection on individuals. Although supported by Dawkins and by many others, this claim continues to be disputed. While naïve versions of group selectionism have been disproved, more sophisticated formulations make accurate predictions in some cases while positing selection at higher levels. Both sides agree that very favourable genes are likely to prosper and replicate if they arise and both sides agree that living in groups can be an advantage to the group members. The conflict arises in part over defining concepts:
:Cultural evolutionary theory, however, has suffered from an overemphasis on the experiences and behaviors of individuals at the expense of acknowledging complex group organization...Many important behaviors related to the success and function of human societies are only properly defined at the level of groups.
In *The Social Conquest of Earth* (2012), the entomologist E. O. Wilson contends that although the selfish-gene approach was accepted "until 2010 [when] Martin Nowak, Corina Tarnita, and I demonstrated that inclusive fitness theory, often called kin selection theory, is both mathematically and biologically incorrect." Chapter 18 of *The Social Conquest of Earth* describes the deficiencies of kin selection and outlines group selection, which Wilson argues is a more realistic model of social evolution. He criticises earlier approaches to social evolution, saying: "unwarranted faith in the central role of kinship in social evolution has led to the reversal of the usual order in which biological research is conducted. 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." According to Wilson: "People must have a tribe...Experiments conducted over many years by social psychologists have revealed how swiftly and decisively people divide into groups, and then discriminate in favor of the one to which they belong." (pp. 57, 59) According to Wilson: "Different parts of the brain have evolved by group selection to create groupishness." (p. 61)
Some authors consider facets of this debate between Dawkins and his critics about the level of selection to be blather:
:"The particularly frustrating aspects of these constantly renewed debates is that, even though they seemed to be sparked by rival theories about how evolution works, in fact they often involve only rival metaphors for the very same evolutionary logic and [the debates over these aspects] are thus empirically empty."
::— Laurent Keller, *Levels of Selection in Evolution*, p.4
Other authors say Dawkins has failed to make some critical distinctions, in particular, the difference between group selection for group advantage and group selection conveying individual advantage.
### 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. Summarizing the Dawkins-Gould difference of view, Sterelny says:
:Gould thinks gene differences do not cause evolutionary changes in populations, they register those changes.
::—Kim Sterelny: *Dawkins *vs.* Gould*, p. 83
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.
Like Gould, Gabriel Dover in criticizing *The Selfish Gene* says:
: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.
::— Gabriel Dover: *Dear Mr. Darwin*, p. 56
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. The term "replicator" includes genes as the most fundamental replicators but possibly other agents, and *interactor* includes organisms but maybe other agents, much as do Dawkins's 'vehicles'. The distinction is as follows:
:*replicator*: an entity that passes on its structure largely intact in successive replications.
:*interactor*: an entity that interacts as a cohesive whole with its environment in such a way that this interaction *causes* replication to be differential.
:*selection*: a process in which the differential extinction or proliferation of interactors causes the differential perpetuation of the replicators that produced them.
Hull suggests that, despite some similarities, Dawkins takes too narrow a view of these terms, engendering some of the objections to his views. According to Godfrey-Smith, this more careful vocabulary has cleared up "misunderstandings in the "units of selection" debates."
### Enactive arguments
Behavioural genetics entertains the view:
:that genes are dynamic contributors to behavioral organization and are sensitive to feedback systems from the internal and external environments. Technically behavior is not inherited; only DNA molecules are inherited. From that point on behavioral formation is a problem of constant interplay between genetic potential and environmental shaping.
::—D.D. Thiessen, *Mechanism specific approaches in behavior genetics*, p. 91
This view from 1970 is still espoused today, and conflicts with Dawkins's view of "the gene as a form of "information [that] passes through bodies and affects them, but is not affected by them on its way through"".
The philosophical/biological field of enactivism stresses the interaction of the living agent with its environment and the relation of probing the environment to cognition and adaptation. Gene activation depends upon the cellular milieu. An extended discussion of the contrasts between enactivism and Dawkins's views, and with their support by Dennett, is provided by Thompson.
In *Mind in Life*, the philosopher Evan Thompson has assembled a multi-sourced objection to the "selfish gene" idea. Thompson takes issue with Dawkin's reduction of "life" to "genes" and "information":
:Life is just bytes and bytes and bytes of digital information.
::— Richard Dawkins: *River out of Eden: A Darwinian View of Life*, p. 19
:On the bank of the Oxford canal...is a large willow tree, and it is pumping downy seeds into the air...It is raining instructions out there; it's raining programs; it's raining tree-growing, fluff-spreading algorithms. That is not a metaphor, it is the plain truth.
::— Richard Dawkins: *The Blind Watchmaker*, p. 111
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.
::— Sahotra Sarkar *Biological information: a skeptical look at some central dogmas of molecular biology*, p. 187
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" 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
Another criticism of the book is its treatment of morality, and more particularly altruism, as existing only as a form of selfishness:
:It is important to realize that the above definitions of altruism and selfishness are *behavioural*, not subjective. I am not concerned here with the psychology of motives...My definition is concerned only with whether the effect of an act is to lower or raise the survival prospects of the presumed altruist and the survival prospects of the presumed beneficiary.
::— Richard Dawkins, *The Selfish Gene*, p. 12
: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.
::— Richard Dawkins, *The Selfish Gene*, p. 179
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. Part of Mary Midgley's concern is that Richard Dawkins's account of *The Selfish Gene* serves as a moral and ideological justification for selfishness to be adopted by modern human societies as simply following "nature", providing an excuse for behavior with bad consequences for future human society.
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).
Dawkins claims he merely describes how things are under evolution, and makes no moral arguments. On BBC-2 TV, Dawkins pointed to evidence for a "Tit-for-Tat" strategy (shown to be successful in game theory
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.
## Publication
*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. A second edition was published in 1989. This edition added two extra chapters, and substantial endnotes to the preceding chapters, reflecting new findings and thoughts. It also added a second preface by the author, but the original foreword by Trivers was dropped. The book contains no illustrations.
The book has been translated into at least 23 languages including Arabic, Thai and Turkish.
In 2006, a 30th-anniversary edition was published with the Trivers foreword and a new introduction by the author in which he states, "This edition does, however---and it is a source of particular joy to me---restore the original Foreword by Robert Trivers." This edition was accompanied by a *festschrift* entitled *Richard Dawkins: How a Scientist Changed the Way We Think* (2006). In March 2006, a special event entitled *The Selfish Gene*: Thirty Years On was held at the London School of Economics. In March 2011, Audible Inc published an audiobook edition narrated by Richard Dawkins and Lalla Ward.
In 2016, Oxford University Press published *The Extended Selfish Gene*, a 40th anniversary edition with a new epilogue, in which Dawkins describes the continued relevance of the gene's eye view of evolution and states that it, along with coalescence analysis "illuminates the deep past in ways of which I had no inkling when I first wrote *The Selfish Gene*..." It contains two chapters from his later *The Extended Phenotype*. He credits Yan Wong, "my co-author of *The Ancestor's Tale*, from whom I learned everything I know about coalescence theory and much else besides."
### Editions
::data[format=table]
| Year | Title | ISBN | Format |
|---|---|---|---|
| 1976 | The Selfish Gene (1st ed.) | | Hardback |
| 1978 | The Selfish Gene (Scientific Book Club ed.) | | Hardback |
| 1978 | The Selfish Gene (1st ed. Reprint) | | Paperback |
| 1989 | The Selfish Gene (2nd ed.) | | Paperback |
| 2006 | The Selfish Gene: 30th Anniversary Edition (3rd ed.) | | Hardback |
| | Paperback | | |
| 2011 | The Selfish Gene (MP3 CD) | | Audiobook |
| 2016 | The Selfish Gene: 40th Anniversary Edition (4th ed.) | | Paperback |
| 2016 | The 'Extended' Selfish Gene (4th ed.) | | Hardback |
::
## Awards and recognition
In April 2016, *The Selfish Gene* was listed in ''The Guardian'''s list of the 100 best nonfiction books, by Robert McCrum.
Ian McEwan writes that There has never been a science book quite like it. Drawing on the work of a handful of scientists, it bound together genetics and Darwinian natural selection in a creative synthesis that amazed even those few who were already familiar with the concepts. It hastened a sea change in evolutionary theory, it affected profoundly the teaching of biology, it enticed an enthusiastic younger generation into the subject, and spawned a huge literature, and eventually a new discipline - memetics. At the same time, and this is the measure of its achievement, it addressed itself without condescension to the layman. It did so provocatively, and with style.
In the years since then, Dawkins' work might be seen as one extended invitation addressed to us non-scientists to enjoy science, to indulge ourselves at a feast of human ingenuity. Just as we can sit around the kitchen table and discuss operas, movies or novels without being composers, directors or novelists, so we can engage with this subject, one more sublime achievement of accumulated creativity. We can make it "ours" just as we might the music of Bach or Bill Evans.
*The Selfish Gene* stood at the beginning of a golden age of science writing. With a fine sense of literary tradition, the physicist Steven Weinberg, in his book *Dreams of a Final Theory*, revisited Huxley's lecture on chalk in order to make the case for reductionism. Steven Pinker's application of Darwinian thought to Chomskyan linguistics in *The Language Instinct* is one of the finest celebrations of language I know. Among many other indispensable 'classics', I would propose EO Wilson's *The Diversity of Life* on the ecological wonders of the Amazon rain forest, and on the teeming micro-organisms in a handful of soil; David Deutsch's masterly account of the Many Worlds theory in *The Fabric of Reality*; Jared Diamond's melding of history with biological thought in *Guns, Germs and Steel*...
Weinberg included it on his list of the 13 best science books for the general reader.
In July 2017, a poll to celebrate the 30th anniversary of the Royal Society Science Book Prize listed *The Selfish Gene* as the most influential science book of all time.
## Notes
## References
## Bibliography
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|editor1-first = Alan
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