The Selfish Gene - Richard Dawkins (2006)

Chapter 7. Family planning

It is easy to see why some people have wanted to separate parental care from the other kinds of kin-selected altruism. Parental care looks like an integral part of reproduction whereas, for example, altruism toward a nephew is not. I think there really is an important distinction hidden here, but that people have mistaken what the distinction is. They have put reproduction and parental care on one side, and other sorts of altruism on the other. But I wish to make a distinction between bringing new individuals into the world, on the one hand, and caring for existing individuals on the other. I shall call these two activities respectively child-bearing and child-caring. An individual survival machine has to make two quite different sorts of decisions, caring decisions and bearing decisions. I use the word decision to mean unconscious strategic move.

The caring decisions are of this form: 'There is a child; its degree of relatedness to me is so and so; its chances of dying if I do not feed it are such and such; shall I feed it?' Bearing decisions, on the other hand, are like this: 'Shall I take whatever steps are necessary in order to bring a new individual into the world; shall I reproduce?' To some extent, caring and bearing are bound to compete with each other for an individual's time and other resources: the individual may have to make a choice:

'Shall I care for this child or shall I bear a new one?'

Depending on the ecological details of the species, various mixes of caring and bearing strategies can be evolutionarily stable. The one thing that cannot be evolutionarily stable is a pure caring strategy. If all individuals devoted themselves to caring for existing children to such an extent that they never brought any new ones into the world, the population would quickly become invaded by mutant individuals who specialized in bearing. Caring can only be evolutionarily stable as part of a mixed strategy-at least some bearing has to go on.

The reason this error has grown up is largely historical. The evolutionary advantage of parental care is so obvious that we did not have to wait for Hamilton to point it out It has been understood ever since Darwin. When Hamilton demonstrated the genetic equivalence of other relationships, and their evolutionary significance, he naturally had to lay stress on these other relationships. In particular, he drew examples from the social insects such as ants and bees, in which the sister/sister relationship is particularly important, as we shall see in a later chapter. I have even heard people say that they thought Hamilton's theory applied only to the social insects!

If anybody does not want to admit that parental care is an example of kin selection in action, then the onus is on him to formulate a general theory of natural selection that predicts parental altruism, but that does not predict altruism between collateral kin. I think he will fail.

The species with which we are most familiar-mammals and birds-tend to be great carers. A decision to bear a new child is usually followed by a decision to care for it. It is because bearing and caring so often go together in practice that people have muddled the two things up. But from the point of view of the selfish genes there is, as we have seen, no distinction in principle between caring for a baby brother and caring for a baby son. Both infants are equally closely related to you. If you have to choose between feeding one or the other, there is no genetic reason why you should choose your own son. But on the other hand you cannot, by definition, bear a baby brother. You can only care for him once somebody else has brought him into the world. In the last chapter we looked at how individual survival machines ideally should decide whether to behave altruistically towards other individuals who already exist. In this chapter we look at how they should decide whether to bring new individuals into the world.

It is over this matter that the controversy about 'group selection', which I mentioned in Chapter 1, has chiefly raged. This is because Wynne-

Edwards, who has been mainly responsible for promulgating the idea of group selection, did so in the context of a theory of 'population regulation'. He suggested that individual animals deliberately and altruistically reduce their birth rates for the good of the group as a whole.

This is a very attractive hypothesis, because it fits so well with what individual humans ought to do. Mankind is having too many children.

Population size depends upon four things: births, deaths, immigrations and emigrations. Taking the world population as a whole, immigrations and emigrations do not occur, and we are left with births and deaths. So long as the average number of children per couple is larger than two surviving to reproduce, the numbers of babies born will tend to increase over the years at an ever-accelerating rate. In each generation the population, instead of going up by a fixed amount, increases by something more like a fixed proportion of the size that it has already reached. Since this size is itself getting bigger, the size of the increment gets bigger. If this kind of growth was allowed to go on unchecked, a population would reach astronomical proportions surprisingly quickly.

Incidentally, a thing that is sometimes not realized even by people who worry about population problems is that population growth depends on when people have children, as well as on how many they have. Since populations tend to increase by a certain proportion per generation, it follows that if you space the generations out more, the population will grow at a slower rate per year. Banners that read 'Stop at Two' could equally well be changed to 'Start at Thirty'! But in any case, accelerating population growth spells serious trouble.

We have probably all seen examples of the startling calculations that can be used to bring this home. For instance, the present population of Latin America is around 300 million, and already many of them are under-nourished. But if the population continued to increase at the present rate, it would take less than 500 years to reach the point where the people, packed in a standing position, formed a solid human carpet over the whole area of the continent. This is so, even if we assume them to be very skinny-a not unrealistic assumption. In 1,000 years from now they would be standing on each other's shoulders more than a million deep.

By 2,000 years, the mountain of people, travelling outwards at the speed of light, would have reached the edge of the known universe.

It will not have escaped you that this is a hypothetical calculation! It will not really happen like that for some very good practical reasons. The names of some of these reasons are famine, plague, and war; or, if we are lucky, birth control. It is no use appealing to advances in agricultural science-'green revolutions' and the like. Increases in food production may temporarily alleviate the problem, but it is mathematically certain that they cannot be a long-term solution; indeed, like the medical advances that have precipitated the crisis, they may well make the problem worse, by speeding up the rate of the population expansion. It is a simple logical truth that, short of mass emigration into space, with rockets taking off at the rate of several million per second, uncontrolled birth-rates are bound to lead to horribly increased death-rates. It is hard to believe that this simple truth is not understood by those leaders who forbid their followers to use effective contraceptive methods. 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.

But of course the unease that such long-term calculations arouse is based on concern for the future welfare of our species as a whole.

Humans (some of them) have the conscious foresight to see ahead to the disastrous consequences of over-population. It is the basic assumption of this book that survival machines in general are guided by selfish genes, who most certainly cannot be expected to see into the future, nor to have the welfare of the whole species at heart. This is where Wynne-Edwards parts company with orthodox evolutionary theorists. He thinks there is a way in which genuine altruistic birth-control can evolve.

A point that is not emphasized in the writings of Wynne-Edwards, or in Ardrey's popularization of his views, is that there is a large body of agreed facts that are not in dispute. It is an obvious fact that wild animal populations do not grow at the astronomical rates of which they are theoretically capable. Sometimes wild animal populations remain rather stable, with birth-rates and death-rates roughly keeping pace with each other. In many cases, lemmings being a famous example, the population fluctuates widely, with violent explosions alternating with crashes and near extinction. Occasionally the result is outright extinction, at least of the population in a local area. Sometimes, as in the case of the Canadian lynx-where estimates are obtained from the numbers of pelts sold by the Hudson's Bay Company in successive years-the population seems to oscillate rhythmically. The one thing animal populations do not do is go on increasing indefinitely.

Wild animals almost never die of old age: starvation, disease, or predators catch up with them long before they become really senile. Until recently this was true of man too. Most animals die in childhood, many never get beyond the egg stage. Starvation and other causes of death are the ultimate reasons why populations cannot increase indefinitely. But as we have seen for our own species, there is no necessary reason why it ever has to come to that. If only animals would regulate their birth-rates, starvation need never happen. It is Wynne-Edwards's thesis that that is exactly what they do. But even here there is less disagreement than you might think from reading his book. Adherents of the selfish gene theory would readily agree that animals do regulate their birth-rates. Any given species tends to have a rather fixed clutch-size or litter-size: no animal has an infinite number of children. The disagreement comes not over whether birth-rates are regulated. The disagreement is over why they are regulated: by what process of natural selection has family-planning evolved? In a nutshell, the disagreement is over whether animal birth-control is altruistic, practised for the good of the group as a whole; or selfish, practised for the good of the individual doing the reproducing. I will deal with the two theories in order.

Wynne-Edwards supposed that individuals have fewer children than they are capable of, for the benefit of the group as a whole. He recognized that normal natural selection cannot possibly give rise to the evolution of such altruism: the natural selection of lower-than-average reproductive rates is, on the face of it, a contradiction in terms. He therefore invoked group selection, as we saw in Chapter 1. According to him, groups whose individual members restrain their own birth-rates are less likely to go extinct than rival groups whose individual members reproduce so fast that they endanger the food supply. Therefore the world becomes populated by groups of restrained breeders. The individual restraint that Wynne-Edwards is suggesting amounts in a general sense to birth-control, but he is more specific than this, and indeed comes up with a grand conception in which the whole of social life is seen as a mechanism of population regulation. For instance, two major features of social life in many species of animals are territoriality and dominance hierarchies, already mentioned in Chapter 5.

Many animals devote a great deal of time and energy to apparently defending an area of ground which naturalists call a territory. The phenomenon is very widespread in the animal kingdom, not only in birds, mammals, and fish, but in insects and even sea-anemones. The territory may be a large area of woodland which is the principal foraging ground of a breeding pair, as in the case of robins. Or, in herring gulls for instance, it may be a small area containing no food, but with a nest at its centre.

Wynne-Edwards believes that animals who fight over territory are fighting over a token prize, rather than an actual prize like a bit of food.

In many cases females refuse to mate with males who do not possess a territory. Indeed it often happens that a female whose mate is defeated and his territory conquered promptly attaches herself to the victor. Even in apparently faithful monogamous species, the female may be wedded to a male's territory rather than to him personally.

If the population gets too big, some individuals will not get territories, and therefore will not breed. Winning a territory is therefore, to Wynne-Edwards, like winning a ticket or licence to breed. Since there is a finite number of territories available, it is as if a finite number of breeding licences is issued. Individuals may fight over who gets these licences, but the total number of babies that the population can have as a whole is limited by the number of territories available. In some cases, for instance in red grouse, individuals do, at first sight, seem to show restraint, because those who cannot win territories not only do not breed; they also appear to give up the struggle to win a territory. It is as though they all accepted the rules of the game: that if, by the end of the competition season, you have not secured one of the official tickets to breed, you voluntarily refrain from breeding and leave the lucky ones unmolested during the breeding season, so that they can get on with propagating the species.

Wynne-Edwards interprets dominance hierarchies in a similar way. In many groups of animals, especially in captivity, but also in some cases in the wild, individuals learn each other's identity, and they learn whom they can beat in a fight, and who usually beats them. As we saw in Chapter 5, they tend to submit without a struggle to individuals who they 'know' are likely to beat them anyway. As a result a naturalist is able to describe a dominance hierarchy or 'peck order' (so called because it was first described for hens)-a rank-ordering of society in which everybody knows his place, and does not get ideas above his station. Of course sometimes real earnest fights do take place, and sometimes individuals can win promotion over their former immediate bosses. But we saw in Chapter 5, the overall effect of the automatic submission by lower-ranking individuals is that few prolonged fights actually take place, and serious injuries seldom occur.

Many people think of this as a 'good thing' in some vaguely group-selectionist way. Wynne-Edwards has an altogether more daring interpretation. High-ranking individuals are more likely to breed than low-ranking individuals, either because they are preferred by females, or because they physically prevent low-ranking males from getting near females. Wynne-Edwards sees high social rank as another ticket of entitlement to reproduce. Instead of fighting directly over females themselves, individuals fight over social status, and then accept that if they do not end up high on the social scale they are not entitled to breed.

They restrain themselves where females are directly concerned, though they may try even now and then to win higher status, and therefore could be said to compete indirectly over females. But, as in the case of territorial behaviour, the result of this Voluntary acceptance' of the rule that only high-status males should breed is, according to Wynne-Edwards, that populations do not grow too fast. 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.

Perhaps the most startling of Wynne-Edwards's ideas is that of epideictic behaviour, a word that he coined himself. Many animals spend a great deal of time in large flocks, herds, or shoals. Various more or less common-sense reasons why such aggregating behaviour should have been favoured by natural selection have been suggested, and I will talk about some of them in Chapter 10. Wynne-Edwards's idea is quite different. He proposes that when huge flocks of starlings mass at evening, or crowds of midges dance over a gatepost, they are performing a census of their population. Since he is supposing that individuals restrain their birth-rates in the interests of the group as a whole, and have fewer babies when the population density is high, it is reasonable that they should have some way of measuring the population density. Just so; a thermostat needs a thermometer as an integral part of its mechanism.

For Wynne-Edwards, epideictic behaviour is deliberate massing in crowds to facilitate population estimation. He is not suggesting conscious population estimation, but an automatic nervous or hormonal mechanism linking the individuals' sensory perception of the density of their population with their reproductive systems.

I have tried to do justice to Wynne-Edwards's theory, even if rather briefly. If I have succeeded, you should now be feeling persuaded that it is, on the face of it, rather plausible. But the earlier chapters of this book should have prepared you to be sceptical to the point of saying that, plausible as it may sound, the evidence for Wynne-Edwards's theory had better be good, or else. ... And unfortunately the evidence is not good. It consists of a large number of examples which could be interpreted in his way, but which could equally well be interpreted on more orthodox

'selfish gene' lines.

Although he would never have used that name, the chief architect of the selfish gene theory of family planning was the great ecologist David Lack.

He worked especially on clutch-size in wild birds, but his theories and conclusions have the merit of being generally applicable. Each bird species tends to have a typical clutch size. For instance, gannets and guillemots incubate one egg at a time, swifts three, great tits half a dozen or more. There is variation in this: some swifts lay only two at a time, great tits may lay twelve. It is reasonable to suppose that the number of eggs a female lays and incubates is at least partly under genetic control, like any other characteristic. That is say there may be a gene for laying two eggs, a rival allele for laying three, another allele for laying four, and so on, although in practice it is unlikely to be quite as simple as this.

Now the selfish gene theory requires us to ask which of these genes will become more numerous in the gene pool. Superficially it might seem that the gene for laying four eggs is bound to have an advantage over the genes for laying three or two. A moment's reflection shows that this simple 'more means better' argument cannot be true, however. It leads to the expectation that five eggs should be better than four, ten better still, 100 even better, and infinity best of all. In other words it leads logically to an absurdity. Obviously 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.

If three is the optimum clutch size for swifts, what this means, for Lack, is that any individual who tries to rear four will probably end up with fewer children than rival, more cautious individuals who only try to rear three. The obvious reason for this would be that the food is so thinly spread between the four babies that few of them survive to adulthood.

This would be true both of the original allocation of yolk to the four eggs, and of the food given to the babies after hatching. According to Lack, therefore, individuals regulate their clutch size for reasons that are anything but altruistic. 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.

Rearing baby birds is a costly business. The mother has to invest a large quantity of food and energy in manufacturing eggs. Possibly with her mate's help, she invests a large effort in building a nest to hold her eggs and protect them. Parents spend weeks patiently sitting on the eggs.

Then, when the babies hatch out, the parents work themselves nearly to death fetching food for them, more or less non-stop without resting. As we have already seen, a parent great tit brings an average of one item of food to the nest every 30 seconds of daylight. Mammals such as ourselves do it in a slightly different way, but the basic idea of reproduction being a costly affair, especially for the mother, is no less true. It is obvious that if a parent tries to spread her limited resources of food and effort among too many children, she will end up rearing fewer than if she had set out with more modest ambitions. She has to strike a balance between bearing and caring. 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. Natural selection, according to the Lack theory, adjusts initial clutch size (litter size etc.) so as to take maximum advantage of these limited resources.

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. 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 for altruistic restraint in the birth-rate, because there is no welfare state in nature. Any gene for overindulgence is promptly punished: the children containing that gene starve. Since we humans do not want to return to the old selfish 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 substituted 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 the welfare state is highly desirable. But you cannot have an unnatural welfare state, unless you also have unnatural birth-control, otherwise the end result will be misery even greater than that which obtains in nature. 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 so seem to me less free from suspicion.

Returning to wild animals, the Lack clutch-size argument can be generalized to all the other examples Wynne-Edwards uses: territorial behaviour, dominance hierarchies, and so on. Take, for instance, the red grouse that he and his colleagues have worked on. These birds eat heather, and they parcel out the moors in territories containing apparently more food than the territory owners actually need. Early in the season they fight over territories, but after a while the losers seem to accept that they have failed, and do not fight any more. They become outcasts who never get territories, and by the end of the season they have mostly starved to death. Only territory owners breed. That non-territory owners are physically capable of breeding is shown by the fact that if a territory owner is shot his place is promptly filled by one of the former outcasts, who then breeds. Wynne-Edwards's interpretation of this extreme territorial behaviour is, as we have seen, that the outcasts

'accept' that they have failed to gain a ticket or licence to breed; they do not try to breed.

On the face of it, this seems an awkward example for the selfish gene theory to explain. Why don't the outcasts try, try, and try again to oust a territory holder, until they drop from exhaustion? They would seem to have nothing to lose. But wait, perhaps they do have something to lose.

We have already seen that if a territory-holder should happen to die, an outcast has a chance of taking his place, and therefore of breeding. If the odds of an outcast's succeeding to a territory in this way are greater than the odds of his gaining one by fighting, then it may pay him, as a selfish individual, to wait in the hope that somebody will die, rather than squander what little energy he has in futile fighting. For Wynne-Edwards, the role of the outcasts in the welfare of the group is to wait in the wings as understudies, ready to step into the shoes of any territory holder who dies or. the main stage of group reproduction. We can now see that this may also be their best strategy purely as selfish individuals. As we saw in Chapter 4, we can regard animals as gamblers. The best strategy for a gambler may sometimes be a wait-and-hope strategy, rather than a bull-at-a-gate strategy.

Similarly, the many other examples where animals appear to 'accept'

non-reproductive status passively can be explained quite easily by the selfish gene theory. The general form of the explanation is always the same: the individual's best bet is to restrain himself for the moment, in the hope of better chances in the future. A seal who leaves the harem-holders unmolested is not doing it for the good of the group. He is biding his time, waiting for a more propitious moment. Even if the moment never comes and he ends up without descendants, the gamble might have paid off, though, with hindsight we can see that for him it did not.

And when lemmings flood in their millions away from the centre of a population explosion, they are not doing it in order to reduce the density of the area they leave behind! They are seeking, every selfish one of them, a less crowded place in which to live. The fact that any particular one may fail to find it, and dies, is something we can see with hindsight. It does not alter the likelihood that to stay behind would have been an even worse gamble.

It is a well-documented fact that overcrowding sometimes reduces birth-rates. This is sometimes taken to be evidence for Wynne-Edwards's theory. It is nothing of the kind. It is compatible with his theory, and it is also just as compatible with the selfish gene theory. For example, in one experiment mice were put in an outdoor enclosure with plenty of food, and allowed to breed freely. The population grew up to a point, then levelled off. The reason for the levelling-off turned out to be that the females became less fertile as a consequence of over-crowding: they had fewer babies. This kind of effect has often been reported. Its immediate cause is often called 'stress', although giving it a name like that does not of itself help to explain it. In any case, whatever its immediate cause may be, we still have to ask about its ultimate, or evolutionary explanation.

Why does natural selection favour females who reduce their birth-rate when their population is over-crowded?

Wynne-Edwards's answer is clear. Group selection favours groups in which the females measure the population and adjust their birth-rates so that food supplies are not over-exploited. In the condition of the experiment, it so happened that food was never going to be scarce, but the mice could not be expected to realize that. They are programmed for life in the wild, and it is likely that in natural conditions over-crowding is a reliable indicator of future famine.

What does the selfish gene theory say? Almost exactly the same thing, but with one crucial difference. You will remember that, according to Lack, animals will tend to have the optimum number of children from their own selfish point of view. If they bear too few or too many, they will end up rearing fewer than they would have if they had hit on just the right number. Now, 'just the right number' is likely to be a smaller number in a year when the population is overcrowded than in a year when the population is sparse. We have already agreed that overcrowding is likely to foreshadow famine. Obviously, if a female is presented with reliable evidence that a famine is to be expected, it is in her own selfish interests to reduce her own birth-rate. Rivals who do not respond to the warning signs in this way will end up rearing fewer babies, even if they actually bear more. We therefore end up with almost exactly the same conclusion as Wynne-Edwards, but we get there by an entirely different type of evolutionary reasoning.

The selfish gene theory has no trouble even with 'epideictic displays'. You will remember that Wynne-Edwards hypothesized that animals deliberately display together in large crowds in order to make it easy for all the individuals to conduct a census, and regulate their birth-rates accordingly. There is no direct evidence that any aggregations are in fact epideictic, but just suppose some such evidence were found. Would the selfish gene theory be embarrassed? Not a bit.

Starlings roost together in huge numbers. Suppose it were shown, not only that over-crowding in winter reduced fertility in the following spring, but that this was directly due to the birds' listening to each other's calls.

It might be demonstrated experimentally that individuals exposed to a tape-recording of a dense and very loud starling roost laid fewer eggs than individuals exposed to a recording of a quieter, less dense, roost. By definition, this would indicate that the calls of starlings constituted an epideictic display. The selfish gene theory would explain it in much the same way as it handled the case of the mice.

Again, we start from the assumption that genes for having a larger family than you can support are automatically penalized, and become less numerous in the gene pool. The task of an efficient egg-layer is one of predicting what is going to be the optimum clutch size for her, as a selfish individual, in the coming breeding season. You will remember from Chapter 4 the special sense in which we are using the word prediction. Now how can a female bird predict her optimum clutch size?

What variables should influence her prediction? It may be that many species make a fixed prediction, which does not change from year to year.

Thus on average the optimum clutch size for a gannet is one. It is possible that in particular bumper years for fish the true optimum for an individual might temporarily rise to two eggs. If there is no way for gannets to know in advance whether a particular year is going to be a bumper one, we cannot expect individual females to take the risk of wasting their resources on two eggs, when this would damage their reproductive success in an average year.

But there may be other species, perhaps starlings, in which it is in principle possible to predict in winter whether the following spring is going to yield a good crop of some particular food resource. Country people have numerous old sayings suggesting that such clues as the abundance of holly berries may be good predictors of the weather in the coming spring. Whether any particular old wives' tale is accurate or not, it remains logically possible that there are such clues, and that a good prophet could in theory adjust her clutch size from year to year to her own advantage. Holly berries may be reliable predictors or they may not but, as in the case of the mice, it does seem quite likely that population density would be a good predictor. A female starling can in principle know that, when she comes to feed her babies in the coming spring, she will be competing for food with rivals of the same species. If she can somehow estimate the local density of her own species in winter, this could provide her with a powerful means of predicting how difficult it is going to be to get food for babies next spring. If she found the winter population to be particularly high, her prudent policy, from her own selfish point of view, might well be to lay relatively few eggs: her estimate of her own optimum clutch size would have been reduced.

Now the moment it becomes true that individuals are reducing their clutch size on the basis of their estimate of population density, it will immediately be to the advantage of each selfish individual to pretend to rivals that the population is large, whether it really is or not. If starlings are estimating population size by the volume of noise in a winter roost, it would pay each individual to shout as loudly as possible, in order to sound more like two starlings than one. This idea of animals pretending to be several animals at once has been suggested in another context by J.

R. Krebs, and is named the Beau Geste Effect after the novel in which a similar tactic was used by a unit of the French Foreign Legion. The idea in our case is to try to induce neighbouring starlings to reduce their clutch size to a level lower than the true optimum. If you are a starling who succeeds in doing this, it is to your selfish advantage, since you are reducing the numbers of individuals who do not bear your genes. I therefore conclude that Wynne-Edwards's idea of epideictic displays may actually be a good idea: he may have been right all along, but for the wrong reasons. More generally, the Lack type of hypothesis is powerful enough to account, in selfish gene terms, for all evidence that might seem to support the group-selection theory, should any such evidence turn up.

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.

So much, then, for quantitative considerations of family size. We now come on to conflicts of interest within families. Will it always pay a mother to treat all her children equally, or might she have favourites?

Should the family function as a single cooperating whole, or are we to expect selfishness and deception even within the family? Will all members of a family be working towards the same optimum, or will they

'disagree' about what the optimum is? These are the questions we try to answer in the next chapter. The related question of whether there may be conflict of interest between mates, we postpone until Chapter 9.