This book should be read almost as though it were science fiction. It is designed to appeal to the imagination. But it is not science fiction: it is science. Cliche or not, 'stranger than fiction' expresses exactly how I feel about the truth. We are survival machines - robot vehicles blindly programmed to preserve the selfish molecules known as genes. This is a truth which still fills me with astonishment. Though I have known it for years, I never seem to get fully used to it. One of my hopes is that I may have some success in astonishing others...


Intelligent life on a planet comes of age when it first works out the reason for its own existence. If superior creatures from space ever visit earth, the first question they will ask, in order to assess the level of our civilization, is: 'Have they discovered evolution yet?' 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.

We no longer have to resort to superstition when faced with the deep problems: Is there a meaning to life? What are we for? What is man? After posing the last of these questions, the eminent zoologist G. G. Simpson put it thus: "The point I want to make now is that all attempts to answer that question before 1859 are worthless and that we will be better off if we ignore them completely."

Today the theory of evolution is about as much open to doubt as the theory that the earth goes round the sun, but the full implications of Darwin's revolution have yet to be widely realized. Zoology is still a minority subject in universities, and even those who choose to study it often make their decision without appreciating its profound philosophical significance. Philosophy and the subjects known as 'humanities' are still taught almost as if Darwin had never lived. No doubt this will change in time.

My purpose is to examine the biology of selfishness and altruism. Apart from its academic interest, the human importance of this subject is obvious. It touches every aspect of our social lives, our loving and hating, fighting and cooperating, giving and stealing, our greed and our generosity.

These are claims that could have been made for Lorenz's On Aggression, Ardrey's The Social Contract, and Eibl-Eibesfeldt's Love and Hate. The trouble with these books is that their authors got it totally and utterly wrong. They got it wrong because they misunderstood how evolution works. They made the erroneous assumption that the important thing in evolution is the good of the species (or the group) rather than the good of the individual (or the gene). It is ironic that Ashley Montagu should criticize Lorenz as a 'direct descendant of the "nature red in tooth and claw" thinkers of the nineteenth century . . .'. As I understand Lorenz's view of evolution, he would be very much at one with Montagu in rejecting the implications of Tennyson's famous phrase. Unlike both of them, I think 'nature red in tooth and claw' sums up our modern understanding of natural selection admirably.


Before beginning on my argument itself, I want to explain briefly what sort of an argument it is, and what sort of an argument it is not, If we were told that a man had lived a long and prosperous life in the world of Chicago gangsters, we would be entitled to make some guesses as to the sort of man he was, We might expect that he would have qualities such as toughness, a quick trigger finger, and the ability to attract loyal friends. These would not be infallible deductions, but you can make some inferences about a man's character if you know something about the conditions in which he has survived and prospered. The argument of this book is that we, and all other animals, are machines created by our genes. Like successful Chicago gangsters, our genes have survived, in some cases for millions of years, in a highly competitive world. This entitles us to expect certain qualities in our genes. I shall argue that a predominant quality to be expected in a successful gene is ruthless selfishness. This gene selfishness will usually give rise to selfishness in individual behavior. 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. 'Special' and 'limited' are important words in the last sentence. Much as we might wish to believe otherwise, universal love and the welfare of the species as a whole are concepts that simply do not make evolutionary sense.

This brings me to the first point I want to make about what this book is not. I am not advocating a morality based on evolution. I am saying how things have evolved. I am not saying how we humans morally ought to behave. ...If you wish to extract a moral from it, read it as a warning. Be warned that if you wish, as I do, to build a society in which individuals cooperate generously and unselfishly towards a common good, you can expect little help from biological nature. Let us try to teach generosity and altruism, because we are born selfish. Let us understand what our own selfish genes are up to, because we may then at least have a chance to upset their designs, something that no other species has ever aspired to do.
I stress that, because I know I am in danger of being misunderstood by those people, all too numerous, who cannot distinguish a statement of belief in what is the case from an advocacy of what ought to be the case. Unfortunately, however much we deplore something, it does not stop it being true.

As a corollary to these remarks about teaching, it is a fallacy - incidentally a very common one - to suppose that genetically inherited traits are by definition fixed and unmodifiable. Our genes may instruct us to be selfish, but we are not necessarily compelled to obey them all our lives. It may just be more difficult to learn altruism than it would be if we were genetically programmed to be altruistic. Among animals, man is uniquely dominated by culture, by influences learned and handed down. Some would say that culture is so important that genes, whether selfish or not, are virtually irrelevant to the understanding of human nature. Others would disagree. It all depends where you stand in the debate over 'nature versus nurture' as determinants of human attributes. This brings me to the second thing this book is not: it is not an advocacy of one position or another in the nature/nurture controversy. Naturally I have an opinion on this, but I am not going to express it, except insofar as it is implicit in the view of culture that I shall present in the final chapter. If genes really turn out to be totally irrelevant to the determination of modern human behavior, if we really are unique among animals in this respect, it is, at the very least, still interesting to inquire about the rule to which we have so recently become the exception. And if our species is not so exceptional as we might like to think, it is even more important that we should study the rule.

Before going any further, we need a definition. An entity, such as a baboon, is said to be altruistic if it behaves in such a way as to increase another such entity's welfare at the expense of its own.

It is important to realize that the above definitions of altruism and selfishness are behavioral, not subjective. I am not concerned here with the psychology of motives. I am not going to argue about whether people who behave altruistically are 'really' doing it for secret or subconscious selfish motives. Maybe they are and maybe they aren't, and maybe we can never know, but in any case that is not what this book is about. 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.


Nice guys finish first, because being nice is a mutually beneficial strategy, and will come to predominate in each other's presence.

In many species a mother can be more sure of her young than a father can. the mother lays the visible, tangible egg, or bears the child. She has a good chance of knowing for certain the bearers of her own gene. The poor father is much more vulnerable to deception. It is therefore to be expected that fathers will put less effort than mothers into caring for the young.

There are 2 methods used by females in nature to try and ensure the best chance of survival for her offspring. the first is the 'domestic-bliss' approach, where she will mate with only one mate, and try to ensure that this male remains faithful to her, and stays to help rear the offspring. The female tries to identify signs of faithfullness in the male before she allows him to mate with her. One way for a female to do this is to act coy & play hard to get for a long time. Any male who is not patient enough to wait until the female eventually consents to mate is not likely to be a good bet as a fiathful husband. A long engagement period aims to weed out casual suitors, and obtains a male who has dispalyed his qualities of fidelity & perseverance in advance of copulation. A long engagement period may also benefit males where there is a danger of his being duped into caring for another male's child.

Female's may also force the male to invest so heavily in the well-being of the offspring ( eg building a nest ) before copulation that it would no longer pay the male to desert after copulation.

In the 'He-Man' strategy of harems, females resign themslves to getting no help from the father of their children, and go all-out for good genes instead. They use the weapon of withholding copulation, and refuse to mate with just any male, and go for the best. But what are the best? What qualities should a female look for? If a female can somehow detect good genes in males, using external visual clues, she can benefit her own genes by allying them with good paternal genes. One thing she wants is evidence of ability to survive. Quite a good policy for a female might be to go for old men. Whatever their shortcomings, they have at least proved they can survive, ensuring longevity. Longevity, though, does not ensure virility. A male may have survived for so long because he does not take risks in order to reproduce. What else? Perhaps long-legs for running from predators?

In a society of he-men, though, one of the best things a mother can do is to make a son who will turn out to be a he-man.The result of this is that one of the most desirable qualities a male can have in the eyes of a female is, quite simply, sexual attractiveness itself. Once such qualities become widely accepted as attractive by females, natural selection would continue to favour them simply because they were attractive. The influence of culture over genes in man should be seen in the fact that both the domestic-bliss & he-man approach are found around the world.

An unattractive or cautious male may live as long as a female, but he has few children, and his genes are not passed on. What shall it profit a man if he shall gain the whole world, and lose his immortal genes?

What interests us at the moment is that the optimal compromise for a male seems to be different from that of a female. This is compatible with our view of males as high-risk, high-reward gamblers. Because a male produces many millions of sperms to every egg produced by a female, sperms heavily outnumber eggs in the population. Any given egg is therefore much more likely to enter into sexual fusion than any given sperm is. Eggs are a relatively scare resource, and therefore, a female does not need to be so sexually attractive as a male does to ensure that her eggs are fertilized. A male is perfectly capable of siring all the children born to a large population of females.

This leads to females being more fussy than males about whom they mate with. For males, the word excess has no meaning wrt copulations. Where incest taboos exist, we should expect females to be more rigid in their adherence to the taboos than males, since their investment in any particular child tends to be greater, and incestous children are more likely to have lethal or damaging double-recessive genes.

Males may have to 'advertise' to females the fact that they are healthy. Males can be forced by females ( eg Peacock's tail ) to evolve clinical thermometers permanently sticking out of their mouths, clearly displayed for females to read.

The Peacock's Tail, it should be noted, is actually a handicap for the male, in that it actually makes it less likely to survive and is costly to produce. However, evolution favours those males that advertise their handicap, as their handicap will reveal true information about their overall quality as a mate. Only a well-fed & healthy peacock will be able to grow the handicap of a long tail. Even 'bad' possible mates have to advertise their quality by displaying a short tail, as females will not mate with males of unknown quality. If we observe an animal doing something really silly, like standing on its head instead of running away from a lion, it may simply be trying to impress a possible mate. Natural selection, of course, will stop before we reach infinte madness, as it will remove 'suicidal' genes.

Curiously, in modern civilization, it is the females that display the equivalent of the peacock's tail - they paint their faces with make-up, they are more interested in their personal appearance. Faced with these facts, a biologist would be forced to suspect that he was looking at a society in which females compete for males, rather than vice versa. In nature, males are bright & ostentatious beacuse females are in demand and can afford to be choosy. Has the human male really become the sought after sex, the one that is in demand and can afford to be choosy? If so, why?


Most of what is unusual about man can be summed up in one word: Culture.

A MEME is the name I give to a replicator which acts, not on genes, but on cultural ideas like songs, religions, sports, fashions, art, methods of construction, ideas like evolution, gravity, faith etc. The survival value for a meme depends not on whether an individual which carries it survives, but its likelyhood of being accepted by a new generation of humans, its catchiness or appeal.They can become self-perpetuating, because of their deep psychological impact.

When we die, there are two things we can leave behind us: genes and memes. But as each generation passes, the contribution of your genes is halved. Socrates may or may not have a gene or two alive in the world today, as G.C Williams remarked, but who cares? The meme-complexes of Socrates, Leonardo, Copernicus and Marconi are still going strong.

Blind faith is powerful enough to immunize people against all appeals to pity, to forgiveness, to decent human feelings. It even immunizes them against fear, if they honestly believe that a martyr's death will send them straight to heaven. What a weapon! Religious faith deserves a chapter to itself in the annals of war technology, on an even footing with the longbow, the warhorse, the tank, and the hydrogen bomb.
Blind faith can justify anything. If a man believes in a different god, or even if he uses a different ritual for worshipping the same god, blind faith can decree that he should die - on the cross, at the stake, skewered on a Crusader's sword, shot in a Beirut street, or blown up in a bar in Belfast. Memes for blind faith have their own ruthless ways of propagating themselves. This is true of patriotic and political as well as religious blind faith.

We, alone on Earth, can rebel against the tyranny of the selfish replicators ( genes or memes ). We have the power to defy the selfish genes of our birth, and, if necessary, the selfish memes of our indoctrination.


This leads to what I have called the Central Theorem of the Extended Phenotype: An animal's behaviour tends to maximize the survival of the genes 'for' that behaviour, whether or not those genes happen to be in the body of the particular animal performing it. I was writing in the context of animal behaviour, but the theorem could apply, of course, to colour, size, shape - to anything.

A selfish gene is trying to become more numerous in the gene pool. It does this by helping to program the bodies in which it finds itself to survive and to reproduce. But now we are emphasising it as a distributed agency, existing in many different individuals at once. The key point of this chapter is that a gene might be able to assist *replicas* of itself that are sitting in other bodies. If so, this would appear as individual altruism but would be brought about by gene selfishness. For example, if the gene for being an albino just happened to to cause its bodies to behave altruistically towards other albinos, it would tend to become more numerous in the gene pool as a result.

If animals had a tendency to behave altruistically to individuals who physically resembled them ( to identify as being related ), they might indirectly be doing their kin a bit of good. Much would depend on details of the species concerned. Such a rule would, in any case, only lead to 'right' decisions in a statistical sense. If conditions changed, for example if a species started living in much larger groups, it could lead to wrong decision. Conceivably, racial prejudice could be interpreted as an irrational generalization of a kin-selected tendency to identify with individuals resembling oneself, and to be nasty to those different in appearance.

Now we are in a position to talk about genes for kin-altruism much more precisely. A gene for suicidally saving 5 cousins would not become more numerous in the population, but a gene for saving 5 brothers or 10 first cousins would. Such a gene, on average, tends to live on in the bodies of enough individuals saved by the altruist to compensate for the death of the altruist itself.


Why do we die of old age (senile decay)? One theory is the accumulation of deletrius copying erros and other kinds of gene damage which occur during a person's lifetime. Another theory is advanced by Sir Peter Medawar. A general quality that a successful gene must have is the tendency to postpone the death of their survival machines ( ie us ) at least until after reproduction. Not a single one of your ancestors died young! A gene that makes its possessors die is called a lethal gene, and evolution will tend to remove these from the gene pool. However, a late-acting lethal gene may still be successful in the gene pool, as its effects will not harm the individual until he has had a chance to have some offspring. Therefore, death from old age is a by-product of the accumulation in the gene pool of late-acting lethal genes, which have been allowed to slip through the net of natural selection simply beacuse they are late-acting.

An offshoot of this *theory* is that if we want to increase the human life span we could ban reproduction before a certain age, say 40. After some centuries of this, raise it to 50, and so on. Conceivably, human longevity could be pushed up to several centuries by this method. I cannot imagine that anyone would seriously consider implementing such a policy. Another offshoot would be trying to identify chemicals in the body which are indicative of old-age, and attempting to fool these lethal-genes into thinking that the body is younger than it actually is.


Curiously, peace time appeals for individuals to make some small sacrifice in the rate at which they increase their standard of living seem to be less effective than war time appeals for individuals to lay down their lives.

We tend to regard erratic copying as a bad thing, and in the case of human documents it is hard to think of examples where errors can be described as as improvements. I suppose the scholars of the Septuagint could at least be said to have started something big when they mistranslated the Hebrew word for 'young woman' into the Greek word for virgin', coming up with the prophecy : "Behold, a virgin shall conceive and bear a son..."

Perhaps consciousness arises when the brain's simulation of the world becomes so complete that it must include a model of itself.

There are some species of ant that take slaves. True warfare in which large rival armies fight to the death is known only in man and in social insects ( ie those with Queens ). Slaving raids are a particular kind of war effort where the slavers mount an attack on a nest of ants belonging to a different specis, attempt to kill the defending workers or soldiers, and carry off the unhatched young. When these young hatch, they will act as normal workers in the way they would have in their own nest.


Individuals in the animal world who have too many children are penalized, not because the whole population goes extinct, but simply because fewer of their children survive. Genes for having too many children are just not passed on to the next generation in large numbers, beacuse few of the children bearing these genes reach adulthood. What has happened in modern civilized man is that family sizes are no longer limited by the finite resources that the individual parents can provide. If a husband & wife have more children than they can feed, the state, which means the rest of the population, simply steps in and keeps the surplus children alive and healthy. There is, in fact, nothing to stop a couple with no material resources at all having and rearing precisely as many children as the woman can physically bear. But the welfare state is a very unnatural thing. In nature, parents who have more children than they can support do not have many grandchildren, and their genes are not passed on to future generations. there is no need ofr altruistic restraint in the birth-rate, because there is no welfare state in nature. Any gene for over-indulgence is promptly punished : the children containing that gene starve. Since we humans do not want a return to the selfish old ways where we let the children of too-large families starve to death, we have abolished the family as a unit of economic self-sufficiency, and substitued the state. But the privilege of guaranteed support for children should not be abused.

Contraception is sometimes attacked as 'unnatural'.So it is, very unnatural. The trouble is, so is the welfare state. I think that most of us believe that the welfare state is highly desirable. The welfare state is perhaps the greatest altruistic system the animal kingdom has ever known. But any altruistic system is inherently unstable, because it is open to abuse by selfish individuals, ready to exploit it. Individual humans who have more children than they are capable of rearing are probably too ignorant in most cases to be accused of conscious malevolent exploitation. Powerful institutions and leaders who deliberately encourage them to do seem to me less free from suspicion.

They express a preference for 'natural' methods of population limitation, and a natural method is exactly what they are going to get. It is called starvation.


The replicators which survived (from the dawn of Earth) were the ones that built survival machines for themselves to live in... Now they swarm in huge colonies, safe inside gigantic lumbering robots... They are in you and me; they created us, body and mind; and their preservation is the ultimate rationale for our existence. They have come a long way those replicators. Now they go by the name of genes, and we are their survival machines.


Evoloution is only as much a theory, in reality, as Newton’s theory of gravity. If we are not to sink into the intellectual dark ages, there is no other bulwark than the truth, which The Selfish Gene overwhelmingly embodies.Truth may not always seem a sufficient defence against determined, wicked ignorance; but it is some consolation to have it so easily and convincingly available as in this great book.
        - Phillip Hensher, reviewing the updated "Selfish Gene" in "The Spectator" (2006)

In 1975, I showed the partially completed book to Tom Maschler, doyen of London publishers, and we discussed it in his room at Jonathan Cape. He liked the book but not the title. “Selfish”, he said, was a “down” word. Why not call it The Immortal Gene? Immortal was an “up” word, the immortality of genetic information was a central theme of the book, and “immortal gene” had almost the same intriguing ring as “selfish gene” (neither of us, I think, noticed the resonance with Oscar Wilde’s The Selfish Giant). I now think 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, without the large footnote of the book itself, might give an inadequate impression of its contents. Nowadays, an American publisher would in any case have insisted on a subtitle.
Unwriting a book is one thing. Unreading it is something else. What are we to make of the following verdict, from a reader in Australia? “Fascinating, but at times I wish I could unread it . . . On one level, I can share in the sense of wonder Dawkins so evidently sees in the workings-out of such complex processes . . . But at the same time, I largely blame The Selfish Gene for a series of bouts of depression I suffered from for more than a decade . . . Never sure of my spiritual outlook on life, but trying to find something deeper — trying to believe, but not quite being able to — I found that this book just about blew away any vague ideas I had along these lines, and prevented them from coalescing any further. This created quite a strong personal crisis for me some years ago.”
        - Richard Dawkins, assessing its impact after 30 years in "The Times"

"Nice means never being the first to defect."
        - Richard Dawkins, from "Nice Guys Finish First" BBC Horizon documentary

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