The Selfish Gene

by Richard Dawkins

Instead of actually having too many children, and then finding out the hard way that it was a mistake, populations use formal contests over status and territory as a means of limiting their size slightly below the level at which starvation itself actually takes its toll.

there are costs as well as benefits in laying a large number of eggs. Increased bearing is bound to be paid for in less efficient caring. Lack’s essential point is that for any given species, in any given environmental situation, there must be an optimal clutch size. Where he differs from Wynne-Edwards is in his answer to the question: ‘Optimal from whose point of view?’. Wynne-Edwards would say the important optimum, to which all individuals should aspire, is the optimum for the group as a whole. Lack would say each selfish individual chooses the clutch size that maximizes the number of children she rears.

They are not practising birth control in order to avoid over-exploiting the group’s resources. They are practising birth control in order to maximize the number of surviving children they actually have, an aim which is the very opposite of that which we normally associate with birth control.

The total amount of food and other resources which an individual female, or a mated pair, can muster is the limiting factor determining the number of children they can rear.

Individuals 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, because 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 and 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.

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 for 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.

Contraception is sometimes attacked as ‘unnatural’. So it is, very unnatural. The trouble is, so is the welfare state.

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.

Our conclusion from this chapter is that individual parents practise family planning, but in the sense that they optimize their birth rates rather than restrict them for public good. They try to maximize the number of surviving children that they have, and this means having neither too many babies nor too few. Genes that make an individual have too many babies tend not to persist in the gene pool, because children containing such genes tend not to survive to adulthood.

Should a mother have favourites, or should she be equally altruistic towards all her children?

what would it mean to say a mother had a favourite child? It would mean she would invest her resources unequally among her children. The resources that a mother has available to invest consist of a variety of things. Food is the obvious one,

Parental investment (P.I.) is defined as ‘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’.

altruism investment measure. Individual A may be said to invest in individual B, when A increases B’s chance of surviving, at the cost of A’s ability to invest in other individuals including herself, all costs being weighted by the appropriate relatedness.

An under-sized runt bears just as many of his mother’s genes as his more thriving litter mates. But his life expectation is less. Another way to put this is that he needs more than his fair share of parental investment, just to end up equal to his brothers. Depending on the circumstances, it may pay a mother to refuse to feed a runt, and allocate all of his share of her parental investment to his brothers and sisters.

if she saves the little brother she will still have to invest some costly resources in him just to get him up to the age of the big brother.

A mother who had some way of knowing that she had had her last child might be expected to continue to invest all her resources in him for the rest of her life, and perhaps suckle him well into adulthood. Nevertheless, she should ‘weigh up’ whether it would not pay her more to invest in grandchildren or nephews and nieces, since although these are half as closely related to her as her own children, their capacity to benefit from her investment may be more than double that of one of her own children.

menopause, the rather abrupt termination of a human female’s reproductive fertility in middle age. This may not have occurred too commonly in our wild ancestors, since not many women would have lived that long anyway. But still, the difference between the abrupt change of life in women and the gradual fading out of fertility in men suggests that there is something genetically ‘deliberate’ about the menopause—

the ‘grandchild altruism’ gene prevails in the gene pool. A woman could not invest fully in her grandchildren if she went on having children of her own. Therefore genes for becoming reproductively infertile in middle age became more numerous, since they were carried in the bodies of grandchildren whose survival was assisted by grandmotherly altruism. This is a possible explanation of the evolution of the menopause in females. The reason why the fertility of males tails off gradually rather than abruptly is probably that males do not invest so much as females in each individual child anyway.

A woman could not invest fully in her grandchildren if she went on having children of her own. Therefore genes for becoming reproductively infertile in middle age became more numerous, since they were carried in the bodies of grandchildren whose survival was assisted by grandmotherly altruism.

it is of advantage to parent to know when a baby is happy, and it is a good thing for a baby to be able to tell its parents when it is happy. Signals like purring and smiling may have been selected because they enable parents to learn which of their actions are most beneficial to their children. The sight of her child smiling, or the sound of her kitten purring, is rewarding to a mother, in the same sense as food in the stomach is rewarding to a rat in a maze. But once it becomes true that a sweet smile or a loud purr are rewarding, the child is in a position to use the smile or the purr in order to manipulate the parent, and gain more than its fair share of parental investment.

I must emphasize that I am not talking about conscious motives. Nobody is suggesting that children deliberately and consciously deceive their parents because of the selfish genes within them. And I must repeat that when I say something like ‘A child should lose no opportunity of cheating … lying, deceiving, exploiting …’, I am using the word ‘should’ in a special way. I am not advocating this kind of behaviour as moral or desirable. I am simply saying that natural selection will tend to favour children who do act in this way, and that therefore when we look at wild populations we may expect to see cheating and selfishness within families. The phrase ‘the child should cheat’ means that genes that tend to make children cheat have an advantage in the gene pool.

If there is a human moral to be drawn, it is that we must teach our children altruism, for we cannot expect it to be part of their biological nature.

there is one fundamental feature of the sexes which can be used to label males as males, and females as females, throughout animals and plants. This is that the sex cells or ‘gametes’ of males are much smaller and more numerous than the gametes of females. This is true whether we are dealing with animals or plants. One group of individuals has large sex cells, and it is convenient to use the word female for them. The other group, which it is convenient to call male, has small sex cells.

Sperms and eggs too contribute equal numbers of genes, but eggs contribute far more in the way of food reserves: indeed, sperms make no contribution at all and are simply concerned with transporting their genes as fast as possible to an egg. At the moment of conception, therefore, the father has invested less than his fair share (i.e. 50 per cent) of resources in the offspring. Since each sperm is so tiny, a male can afford to make many millions of them every day. This means he is potentially able to beget a very large number of children in a very short period of time, using different females. This is only possible because each new embryo is endowed with adequate food by the mother in each case. This therefore places a limit on the number of children a female can have, but the number of children a male can have is virtually unlimited. Female exploitation begins here.*

What we seek is something equivalent to an evolutionarily stable strategy (ESS), although here, even more than in the chapter on aggression, strategy is just a figure of speech. An individual cannot literally choose the sex of his children. But genes for tending to have children of one sex or the other are possible. If we suppose that such genes, favouring unequal sex ratios, exist, are any of them likely to become more numerous in the gene pool than their rival alleles, which favour an equal sex ratio?

The strategy of producing equal numbers of sons and daughters is an evolutionarily stable strategy, in the sense that any gene for departing from it makes a net loss.

there could be unequal sex ratios that were evolutionarily stable, provided correspondingly unequal amounts of resources were invested in sons and daughters. In the case of the elephant seals, a policy of having three times as many daughters as sons, but of making each son a supermale by investing three times as much food and other resources in him, could be stable.

Each individual wants as many surviving children as possible. The less he or she is obliged to invest in any one of those children, the more children he or she can have. The obvious way to achieve this desirable state of affairs is to induce your sexual partner to invest more than his or her fair share of resources in each child, leaving you free to have other children with other partners. This would be a desirable strategy for either sex, but it is more difficult for the female to achieve. Since she starts by investing more than the male, in the form of her large, food-rich egg, a mother is already at the moment of conception ‘committed’ to each child more deeply than the father is. She stands to lose more if the child dies than the father does. More to the point, she would have to invest more than the father in the future in order to bring a new substitute child up to the same level of development.

the possible courses of action open to a mother who has been deserted by her mate. Best of all for her would be to try to deceive another male into adopting her child, ‘thinking’ it is his own. This might not be too difficult if it is still a foetus, not yet born. Of course, while the child bears half her genes, it bears no genes at all from the gullible step-father. Natural selection would severely penalize such gullibility in males and indeed would favour males who took active steps to kill any potential step-children as soon as they mated with a new wife.

Assuming then that a deserted female cannot fool a new male into adopting her child, what else can she do? Much may depend on how old the child is. If it is only just conceived, it is true that she has invested the whole of one egg in it and perhaps more, but it may still pay her to abort it and find a new mate as quickly as possible. In these circumstances it would be to the mutual advantage both of her and of the potential new husband that she should abort—since we are assuming she has no hope of fooling him into adopting the child.

Another option open to a deserted female is to stick it out, and try and rear the child on her own. This will especially pay her if the child is already quite old. The older he is the more has already been invested in him, and the less it will take out of her to finish the job of rearing him. Even if he is still quite young, it might yet pay her to try to salvage something from her initial investment, even if she has to work twice as hard to feed the child,

Is there anything a female can do to reduce the extent to which her mate exploits her in the first place? She has a strong card in her hand. She can refuse to copulate. She is in demand, in a seller’s market.

The female looks the males over, and tries to spot signs of fidelity and domesticity in advance.

One way for a female to do this is to play hard to get for a long time, to be coy. Any male who is not patient enough to wait until the female eventually consents to copulate is not likely to be a good bet as a faithful husband.

Could females force males to invest so heavily in their offspring before they allow copulation that it would no longer pay the males to desert after copulation? The idea is appealing. A male who waits for a coy female eventually to copulate with him is paying a cost: he is forgoing the chance to copulate with other females, and he is spending a lot of time and energy in courting her. By the time he is finally allowed to copulate with a particular female, he will inevitably be heavily ‘committed’ to her. There will be little temptation for him to desert her, if he knows that any future female he approaches will also procrastinate in the same manner before she will get down to business.

the word ‘strategy’ refers to a blind unconscious behaviour program. Our two female strategies will be called coy and fast, and the two male strategies will be called faithful and philanderer. The behavioural rules of the four types are as follows. Coy females will not copulate with a male until he has gone through a long and expensive courtship period lasting several weeks. Fast females will copulate immediately with anybody. Faithful males are prepared to go on courting for a long time, and after copulation they stay with the female and help her to rear the young. Philanderer males lose patience quickly if a female will not copulate with them straight away: they go off and look for another female; after copulation too they do not stay and act as good fathers, but go off in search of fresh females.

Imagine we have a population in which all the females are coy, and all the males are faithful. It is an ideal monogamous society. In each couple, the male and the female both get the same average pay-off. They get +15 for each child reared; they share the cost of rearing it (−20) equally between the two of them, an average of −10 each. They both pay the −3 point penalty for wasting time in prolonged courtship. The average pay-off for each is therefore + 15 − 10 − 3 = + 2. Now suppose a single fast female enters the population. She does very well. She does not pay the cost of delay, because she does not indulge in prolonged courtship. Since all the males in the population are faithful, she can reckon on finding a good father for her children whomever she mates with. Her average pay-off per child is + 15 − 10 = + 5. She is 3 units better off than her coy rivals. Therefore fast genes will start to spread. If the success of fast females is so great that they come to predominate in the population, things will start to change in the male camp too. So far, faithful males have had a monopoly. But now if a philanderer male arises in the population, he starts to do better than his faithful rivals. In a population where all the females are fast, the pickings for a philanderer male are rich indeed. He gets the +15 points if a child is successfully reared, and he pays neither of the two costs. What this lack of cost mainly means to him is that he is free to go off and mate with new fem...

The system would converge to a stable state.* If you do the sums, it turns out that a population in which of the females are coy, and of the males are faithful, is evolutionarily stable.

Demanding that a prospective mate should build a nest is one effective way for a female to trap him.

If all females of a population forced males to do some difficult and costly deed, like slaying a dragon or climbing a mountain, before they would consent to copulate with them, they could in theory be reducing the temptation for the males to desert after copulation. Any male tempted to desert his mate and try to spread more of his genes by another female would be put off by the thought that he would have to kill another dragon. In practice, however, it is unlikely that females would impose such arbitrary tasks as dragon-killing, or Holy-Grail-seeking on their suitors. The reason is that a rival female who imposed a task no less arduous, but more useful to her and her children, would have an advantage over more romantically minded females who demanded a pointless labour of love.

A female, playing the domestic-bliss strategy, who simply looks the males over and tries to recognize qualities of fidelity in advance, lays herself open to deception. Any male who can pass himself off as a good loyal domestic type, but who in reality is concealing a strong tendency towards desertion and unfaithfulness, could have a great advantage. As long as his deserted former wives have any chance of bringing up some of the children, the philanderer stands to pass on more genes than a rival male who is an honest husband and father. Genes for effective deception by males will tend to be favoured in the gene pool. Conversely, natural selection will tend to favour females who become good at seeing through such deception. One way they can do this is to play especially hard to get when they are courted by a new male, but in successive breeding seasons to be increasingly ready to accept quickly the advances of last year’s mate. This will automatically penalize young males embarking on their first breeding season, whether they are deceivers or not.

a female fish can afford to take the ‘risk’ of spawning early. The male dare not take this risk, since if he spawns too early his sperms will have diffused away before the female is ready, and she will then not spawn herself, because it will not be worth her while to do so. Because of the diffusion problem, the male must wait until the female spawns, and then he must shed his sperms over the eggs. But she has had a precious few seconds in which to disappear, leaving the male in possession, and forcing him on to the horns of Trivers’s dilemma. So this theory neatly explains why paternal care is common in water but rare on dry land.

the other main female strategy, the he-man strategy. In species where this policy is adopted the females, in effect, resign themselves to getting no help from the father of their children, and go all-out for good genes instead. Once again they use their weapon of withholding copulation. They refuse to mate with just any male, but exercise the utmost care and discrimination before they will allow a male to copulate with them.

From the point of view of a female trying to pick good genes with which to ally her own, what is she looking for? One thing she wants is evidence of ability to survive.

What other evidence? There are many possibilities. Perhaps strong muscles as evidence of ability to catch food, perhaps long legs as evidence of ability to run away from predators. A female might benefit her genes by allying them with such traits, since they might be useful qualities in both her sons and her daughters.

In a society where males compete with each other to be chosen as he-men by females, one of the best things a mother can do for her genes is to make a son who will turn out in his turn to be an attractive he-man. If she can ensure that her son is one of the fortunate few males who wins most of the copulations in the society when he grows up, she will have an enormous number of grandchildren. 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. A female who mates with a super-attractive he-man is more likely to have sons who are attractive to females of the next generation, and who will make lots of grandchildren for her.

He suggests that the tails of birds of paradise and peacocks, the huge antlers of deer, and the other sexually-selected features which have always seemed paradoxical because they appear to be handicaps to their possessors evolve precisely because they are handicaps. A male bird with a long and cumbersome tail is showing off to females that he is such a strong he-man that he can survive in spite of his tail.

the various different kinds of breeding system that we find among animals—monogamy, promiscuity, harems, and so on—can be understood in terms of conflicting interests between males and females.

Notions of females withholding copulation until a male shows some evidence of long-term fidelity may strike a familiar chord. This might suggest that human females play the domestic-bliss rather than the he-man strategy. Many human societies are indeed monogamous. In our own society, parental investment by both parents is large and not obviously unbalanced. Mothers certainly do more direct work for children than fathers do, but fathers often work hard in a more indirect sense to provide the material resources that are poured into the children. On the other hand, some human societies are promiscuous, and many are harem-based. What this astonishing variety suggests is that man’s way of life is largely determined by culture rather than by genes. However, it is still possible that human males in general have a tendency towards promiscuity, and females a tendency towards monogamy, as we would predict on evolutionary grounds.