05/22:CP.8. BATTLE OF THE GENERATIONS

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8

BATTLE OF THE GENERATIONS

Let us begin by tackling the first of the questions posed at the end of the last chapter. Should a mother have favourites, or should she be equally altruistic towards all her children? At the risk of being boring, I must yet again throw in my customary warning. The word ‘favourite’ carries no subjective connotations, and the word ‘should’ no moral ones. I am treating a mother as a machine programmed to do everything in its power to propagate copies of the genes which ride inside it. Since you and I are humans who know what it is like to have conscious purposes, it is convenient for me to use the language of purpose as a metaphor in explaining the behaviour of survival machines.

In practice, 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, together with the effort expended in gathering food, since this in itself costs the mother something. Risk undergone in protecting young from predators is another resource which the mother can ‘spend’ or refuse to spend. Energy and time devoted to nest or home maintenance, protection from the elements, and, in some species, time spent in teaching children, are valuable resources which a parent can allocate to children, equally or unequally as she ‘chooses’.

It is difficult to think of a common currency in which to measure all these resources that a parent can invest. Just as human societies use money as a universally convertible currency which can be translated into food or land or labouring time, so we require a currency in which to measure resources that an individual survival machine may invest in another individual's life, in particular a child's life. A measure of energy such as the calorie is tempting, and some ecologists have devoted themselves to the accounting of energy costs in nature. This is inadequate though, because it is only loosely  {124}  convertible into the currency that really matters, the ‘gold-standard’ of evolution, gene survival. R. L. Trivers, in 1972, neatly solved the problem with his concept of Parental Investment (although, reading between the close-packed lines, one feels that Sir Ronald Fisher, the greatest biologist of the twentieth century, meant much the same thing in 1930 by his ‘parental expenditure’).(1)

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.’ The beauty of Trivers's parental investment is that it is measured in units very close to the units that really matter. When a child uses up some of its mother's milk, the amount of milk consumed is measured not in pints, not in calories, but in units of detriment to other children of the same mother. For instance, if a mother has two babies, X and Y, and X drinks one pint of milk, a major part of the P.I. that this pint represents is measured in units of increased probability that Y will die because he did not drink that pint. P.I. is measured in units of decrease in life expectancy of other children, born or yet to be born.

Parental investment is not quite an ideal measure, because it overemphasizes the importance of parentage, as against other genetic relationships. Ideally we should use a generalized 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. Thus a parent's investment in any one child should ideally be measured in terms of detriment to life expectancy not only of other children, but also of nephews, nieces, herself, etc. In many respects, however, this is just a quibble, and Trivers's measure is well worth using in practice.

Now any particular adult individual has, in her whole lifetime, a certain total quantity of P.I. available to invest in children (and other relatives and in herself, but for simplicity we consider only children). This represents the sum of all the food she can gather or manufacture in a lifetime of work, all the risks she is prepared to take, and all the energy and effort that she is able to put into the welfare of children. How should a young female, setting out on her adult life, invest her life's resources? What would be a wise investment policy for her to follow? We have already seen from the Lack theory that she should not spread her investment too thinly among too many  {125}  children. That way she will lose too many genes: she won't have enough grandchildren. On the other hand, she must not devote all her investment to too few children — spoilt brats. She may virtually guarantee herself some grandchildren, but rivals who invest in the optimum number of children will end up with more grandchildren. So much for even-handed investment policies. Our present interest is in whether it could ever pay a mother to invest unequally among her children, i.e. in whether she should have favourites.

The answer is that there is no genetic reason for a mother to have favourites. Her relatedness to all her children is the same, 1/2 Her optimal strategy is to invest equally in the largest number of children that she can rear to the age when they have children of their own. But, as we have already seen, some individuals are better life insurance risks than others. An undersized 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. Indeed it may pay her to feed him to his brothers and sisters, or to eat him herself, and use him to make milk. Mother pigs do sometimes devour their young, but I do not know whether they pick especially on runts.

Runts constitute a particular example. We can make some more general predictions about how a mother's tendency to invest in a child might be affected by his age. If she has a straight choice between saving the life of one child or saving the life of another, and if the one she does not save is bound to die, she should prefer the older one. This is because she stands to lose a higher proportion of her life's parental investment if he dies than if his little brother dies. Perhaps a better way to put this is that 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.

On the other hand, if the choice is not such a stark life or death choice, her best bet might be to prefer the younger one. For instance, suppose her dilemma is whether to give a particular morsel of food to a little child or a big one. The big one is likely to be more capable of finding his own food unaided. Therefore if she stopped feeding him he would not necessarily die. On the other hand, the little one who is too young to find food for himself would be more likely to die if his  {126}  mother gave the food to his big brother. Now, even though the mother would prefer the little brother to die rather than the big brother, she may still give the food to the little one, because the big one is unlikely to die anyway. This is why mammal mothers wean their children, rather than going on feeding them indefinitely throughout their lives. There comes a time in the life of a child when it pays the mother to divert investment from him into future children. When this moment comes, she will want to wean him. 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.

This seems a good moment to mention the puzzling phenomenon known as the 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 — that it is an ‘adaptation’. It is rather difficult to explain. At first sight we might expect that a woman should go on having children until she dropped, even if advancing years made it progressively less likely that any individual child would survive. Surely it would seem always worth trying? But we must remember that she is also related to her grandchildren, though half as closely.

For various reasons, perhaps connected with the Medawar theory of ageing (page 40), women in the natural state became gradually less efficient at bringing up children as they got older. Therefore the life expectancy of a child of an old mother was less than that of a child of a young mother. This means that, if a woman had a child and a grandchild born on the same day, the grandchild could expect to live longer than the child. When a woman reached the age where the average chance of each child reaching adulthood was just less than half the chance of each grandchild of the same age reaching adulthood, any gene for investing in grandchildren in preference to  {127}  children would tend to prosper. Such a gene is carried by only one in four grandchildren, whereas the rival gene is carried by one in two children, but the greater expectation of life of the grandchildren outweighs this, and 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 talis off gradually rather than abruptly is probably that males do not invest so much as females in each individual child anyway. Provided he can sire children by young women, it will always pay even a very old man to invest in children rather than in grandchildren.

So far, in this chapter and in the last, we have seen everything from the parent's point of view, largely the mother's. We have asked whether parents can be expected to have favourites, and in general what is the best investment policy for a parent. But perhaps each child can influence how much his parents invest in him as against his brothers and sisters. Even if parents do not ‘want’ to show favouritism among their children, could it be that children grab favoured treatment for themselves? Would it pay them to do so? More strictly, would genes for selfish grabbing among children become more numerous in the gene pool than rival genes for accepting no more than one's fair share? This matter has been brilliantly analysed by Trivers, in a paper of 1974 called Parent-Offspring Conflict.

A mother is equally related to all her children, born and to be born. On genetic grounds alone she should have no favourites, as we have seen. If she does show favouritism it should be based on differences in expectation of life, depending on age and other things. The mother, like any individual, is twice as closely ‘related’ to herself as she is to any of her children. Other things being equal, this means that she should invest most of her resources selfishly in herself, but other things are not equal. She can do her genes more good by investing a fair proportion of her resources in her children. This is because these are younger and more helpless than she is, and they can therefore benefit more from each unit of investment than she can herself. Genes for investing in more helpless individuals in preference to oneself can prevail in the gene pool, even though the  {128}  beneficiaries may share only a proportion of one's genes. This is why animals show parental altruism, and indeed why they show any kind of kin-selected altruism.

Now look at it from the point of view of a particular child. He is just as closely related to each of is brothers and sisters as his mother is to them. The relatedness is 1/2 in all cases. Therefore he ‘wants’ his mother to invest some of her resources in his brothers and sisters. Genetically speaking, he is just as altruistically disposed to them as his mother is. But again, he is twice as closely related to himself as he is to any brother or sister, and this will dispose him to want his mother to invest in him more than in any particular brother or sister, other things being equal. In this case other things might indeed be equal. If you and your brother are the same age, and both are in a position to benefit equally from a pint of mother's milk, you ‘should’ try to grab more than your fair share, and he should try to grab more than his fair share. Have you ever heard a litter of piglets squealing to be first on the scene when the mother sow lies down to feed them? Or little boys fighting over the last slice of cake? Selfish greed seems to characterize much of child behaviour.

But there is more to it than this. If I am competing with my brother for a morsel of food, and if he is much younger than me so that he could benefit from the food more than I could, it might pay my genes to let him have it. An elder brother may have exactly the same grounds for altruism as a parent: in both cases, as we have seen, the relatedness is 1/2, and in both cases the younger individual can make better use of the resource than the elder. If I possess a gene for giving up food, there is a 50 per cent chance that my baby brother contains the same gene. Although the gene has double the chance of being in my own body — 100 per cent, it is in my body — my need of the food may be less than half as urgent. In general, a child ‘should’ grab more than his share of parental investment, but only up to a point. Up to what point? Up to the point where the resulting net cost to his brothers and sisters, born and potentially to be born, is just double the benefit of the grabbing to himself.

Consider the question of when weaning should take place. A mother wants to stop suckling her present child so that she can prepare for the next one. The present child, on the other hand, does not want to be weaned yet, because milk is a convenient, trouble-free source of food, and he does not want to have to go out and work for his living. To be more exact, he does want eventually to go out and  {129}  work for his living, but only when he can do his genes more good by leaving his mother free to rear his little brothers and sisters, than by staying behind himself. The older a child is, the less relative benefit does he derive from each pint of milk. This is because he is bigger, and a pint of milk is therefore a smaller proportion of his requirement, and also he is becoming more capable offending for himself if he is forced to. Therefore when an old child drinks a pint that could have been invested in a younger child, he is taking relatively more parental investment for himself than when a young child drinks a pint. As a child grows older, there will come a moment when it would pay his mother to stop feeding him, and invest in a new child instead. Somewhat later there will come a time when the old child too would benefit his genes most by weaning himself. This is the moment when a pint of milk can do more good to the copies of his genes that may be present in his brothers and sisters than it can to the genes that are present in himself.

The disagreement between mother and child is not an absolute one, but a quantitative one, in this case a disagreement over timing. The mother wants to go on suckling her present child up to the moment when investment in him reaches his ‘fair’ share, taking into account his expectation of life and how much she has already invested in him. Up to this point there is no disagreement. Similarly, both mother and child agree in not wanting him to go on sucking after the point when the cost to future children is more than double the benefit to him. But there is disagreement between mother and child during the intermediate period, the period when the child is getting more than his share as the mother sees it, but when the cost to other children is still less than double the benefit to him.

Weaning time is just one example of a matter of dispute between mother and child. It could also be regarded as a dispute between one individual and all his future unborn brothers and sisters, with the mother taking the part of her future unborn children. More directly there may be competition between contemporary rivals for her investment, between litter mates or nest mates. Here, once again, the mother will normally be anxious to see fair play.

Many baby birds are fed in the nest by their parents. They all gape and scream, and the parent drops a worm or other morsel in the open mouth of one of them. The loudness with which each baby screams is, ideally, proportional to how hungry he is. Therefore, if the parent always gives the food to the loudest screamer, they should all tend to  {130}  get their fair share, since when one has had enough he will not scream so loudly. At least that is what would happen in the best of all possible worlds, if individuals did not cheat. But in the light of our selfish gene concept we must expect that individuals will cheat, will tell lies about how hungry they are. This will escalate, apparently rather pointlessly because it might seem that if they are all lying by screaming too loudly, this level of loudness will become the norm, and will cease, in effect, to be a lie. However, it cannot deescalate, because any individual who takes the first step in decreasing the loudness of his scream will be penalized by being fed less, and is more likely to starve. Baby bird screams do not become infinitely loud, because of other considerations. For example, loud screams tend to attract predators, and they use up energy.

Sometimes, as we have seen, one member of a litter is a runt, much smaller than the rest. He is unable to fight for food as strongly as the rest, and runts often die. We have considered the conditions under which it would actually pay a mother to let a runt die. We might suppose intuitively that the runt himself should go on struggling to the last, but the theory does not necessarily predict this. As soon as a runt becomes so small and weak that his expectation of life is reduced to the point where benefit to him due to parental investment is less than half the benefit that the same investment could potentially confer on the other babies, the runt should die gracefully and willingly. He can benefit his genes most by doing so. That is to say, a gene that gives the instruction ‘Body, if you are very much smaller than your litter-mates, give up the struggle and die’, could be successful in the gene pool, because it has a 50 per cent chance of being in the body of each brother and sister saved, and its chances of surviving in the body of the runt are very small anyway. There should be a point of no return in the career of a runt. Before he reaches this point he should go on struggling. As soon as he reaches it he should give up and preferably let himself be eaten by his litter-mates or his parents.

I did not mention it when we were discussing Lack's theory of clutch size, but the following is a reasonable strategy for a parent who is undecided as to what is her optimum clutch size for the current year. She might lay one more egg than she actually ‘thinks’ is likely to be the true optimum. Then, if the year's food crop should turn out to be a better one than expected, she will rear the extra child. If not, she can cut her losses. By being careful always to feed the young in the  {131}  same order, say in order of size, she sees to it that one, perhaps a runt, quickly dies, and not too much food is wasted on him, beyond the initial investment of egg yolk or equivalent. From the mother's point of view, this may be the explanation of the runt phenomenon. He represents the hedging of the mother's bets. This has been observed in many birds.

Using our metaphor of the individual animal as a survival machine behaving as if it had the ‘purpose’ of preserving its genes, we can talk about a conflict between parents and young, a battle of the generations. The battle is a subtle one, and no holds are barred on either side. A child will lose no opportunity of cheating. It will pretend to be hungrier than it is, perhaps younger than it is, more in danger than it really is. It is too small and weak to bully its parents physically, but it uses every psychological weapon at its disposal: lying, cheating, deceiving, exploiting, right up to the point where it starts to penalize its relatives more than its genetic relatedness to them should allow. Parents, on the other hand, must be alert to cheating and deceiving, and must try not to be fooled by it. This might seem an easy task. If the parent knows that its child is likely to lie about how hungry it is, it might employ the tactic of feeding it a fixed amount and no more, even though the child goes on screaming. One trouble with this is that the child may not have been lying, and if it dies as a result of not being fed the parent would have lost some of its precious genes. Wild birds can die after being starved for only a few hours.

A. Zahavi has suggested a particularly diabolical form of child blackmail: the child screams in such a way as to attract predators deliberately to the nest. The child is ‘saying’ ‘Fox, fox, come and get me.’ The only way the parent can stop it screaming is to feed it. So the child gains more than its fair share of food, but at a cost of some risk to itself. The principle of this ruthless tactic is the same as that of the hijacker threatening to blow up an aeroplane, with himself on board, unless he is given a ransom. I am sceptical about whether it could ever be favoured in evolution, not because it is too ruthless, but because I doubt if it could ever pay the blackmailing baby. He has too much to lose if a predator really came. This is clear for an only child, which is the case Zahavi himself considers. No matter how much his mother may already have invested in him, he should still value his own life more than his mother values it, since she has only half of his genes. Moreover, the tactic would not pay even if the blackmailer was one of a clutch of vulnerable babies, all in the nest together, since  {132}  the blackmailer has a 50 per cent genetic ‘stake’ in each of his endangered brothers and sisters, as well as a 100 per cent stake in himself. I suppose the theory might conceivably work if the predominant predator had the habit of only taking the largest nestling from a nest. Then it might pay a smaller one to use the threat of summoning a predator, since it would not be greatly endangering itself. This is analogous to holding a pistol to your brother's head rather than threatening to blow yourself up.

More plausibly, the blackmail tactic might pay a baby cuckoo. As is well known, cuckoo females lay one egg in each of several ‘foster’ nests, and then leave the unwitting foster-parents, of a quite different species, to rear the cuckoo young. Therefore a baby cuckoo has no genetic stake in his foster brothers and sisters. (Some species of baby cuckoo will not have any foster brothers and sisters, for a sinister reason which we shall come to. For the moment I assume we are dealing with one of those species in which foster brothers and sisters co-exist alongside the baby cuckoo.) If a baby cuckoo screamed loudly enough to attract predators, it would have a lot to lose — its life — but the foster mother would have even more to lose, perhaps four of her young. It could therefore pay her to feed it more than its share, and the advantage of this to the cuckoo might outweigh the risk.

This is one of those occasions when it would be wise to translate back into respectable gene language, just to reassure ourselves that we have not become too carried away with subjective metaphors. What does it really mean to set up the hypothesis that baby cuckoos ‘blackmail’ their foster parents by screaming ‘Predator, predator, come and get me and all my little brothers and sisters’? In gene terms it means the following.

Cuckoo genes for screaming loudly became more numerous in the cuckoo gene pool because the loud screams increased the probability that the foster parents would feed the baby cuckoos. The reason the foster parents responded to the screams in this way was that genes for responding to the screams had spread through the gene pool of the foster-species. The reason these genes spread was that individual foster parents who did not feed the cuckoos extra food, reared fewer of their own children — fewer than rival parents who did feed their cuckoos extra. This was because predators were attracted to the nest by the cuckoo cries. Although cuckoo genes for not screaming were less likely to end up in the bellies of predators than  {133}  screaming genes, the non-screaming cuckoos paid the greater penalty of not being fed extra rations. Therefore the screaming genes spread through the cuckoo gene pool.

A similar chain of genetic reasoning, following the more subjective argument given above, would show that although such a blackmailing gene could conceivably spread through a cuckoo gene pool, it is unlikely to spread through the gene pool of an ordinary species, at least not for the specific reason that it attracted predators. Of course, in an ordinary species there could be other reasons for screaming genes to spread, as we have already seen, and these would incidentally have the effect of occasionally attracting predators. But here the selective influence of predation would be, if anything, in the direction of making the cries quieter. In the hypothetical case of the cuckoos, the net influence of predators, paradoxical as it sounds at first, could be to make the cries louder.

There is no evidence, one way or the other, on whether cuckoos, and other birds of similar ‘brood-parasitic’ habit, actually employ the blackmail tactic. But they certainly do not lack ruthlessness. For instance, there are honeyguides who, like cuckoos, lay their eggs in the nests of other species. The baby honeyguide is equipped with a sharp, hooked beak. As soon as he hatches out, while he is still blind, naked, and otherwise helpless, he scythes and slashes his foster brothers and sisters to death: dead brothers do not compete for food! The familiar British cuckoo achieves the same result in a slightly different way. It has a short incubation-time, and so the baby cuckoo manages to hatch out before its foster brothers and sisters. As soon as it hatches, blindly and mechanically, but with devastating effectiveness, it throws the other eggs out of the nest. It gets underneath an egg, fitting it into a hollow in its back. Then it slowly backs up the side of the nest, balancing the egg between its wing-stubs, and topples the egg out on to the ground. It does the same with all the other eggs, until it has the nest, and therefore the attention of its foster parents, entirely to itself.

One of the most remarkable facts I have learned in the past year was reported from Spain by F. Alvarez, L. Arias de Reyna, and H. Segura. They were investigating the ability of potential foster parents — potential victims of cuckoos — to detect intruders, cuckoo eggs or chicks. In the course of their experiments they had occasion to introduce into magpie nests the eggs and chicks of cuckoos, and, for comparison, eggs and chicks of other species such as swallows.  {134}  On one occasion they introduced a baby swallow into a magpie's nest. The next day they noticed one of the magpie eggs lying on the ground under the nest. It had not broken, so they picked it up, replaced it, and watched. What they saw is utterly remarkable. The baby swallow, behaving exactly as if it was a baby cuckoo, threw the egg out. They replaced the egg again, and exactly the same thing happened. The baby swallow used the cuckoo method of balancing the egg on its back between its wing-stubs, and walking backwards up the side of the nest until the egg toppled out.

Perhaps wisely, Alvarez and his colleagues made no attempt to explain their astonishing observation. How could such behaviour evolve in the swallow gene pool? It must correspond to something in the normal life of a swallow. Baby swallows are not accustomed to finding themselves in magpie nests. They are never normally found in any nest except their own. Could the behaviour represent an evolved anti-cuckoo adaptation? Has the natural selection been favouring a policy of counterattack in the swallow gene pool, genes for hitting the cuckoo with his own weapons? It seems to be a fact that swallows’ nests are not normally parasitized by cuckoos. Perhaps this is why. According to this theory, the magpie eggs of the experiment would be incidentally getting the same treatment, perhaps because, like cuckoo eggs, they are bigger than swallow eggs. But if baby swallows can tell the difference between a large egg and a normal swallow egg, surely the mother should be able to as well. In this case why is it not the mother who ejects the cuckoo egg, since it would be so much easier for her to do so than the baby? The same objection applies to the theory that the baby swallow's behaviour normally functions to remove addled eggs or other debris from the nest. Once again, this task could be — and is — performed better by the parent. The fact that the difficult and skilled egg-rejecting operation was seen to be performed by a weak and helpless baby swallow, whereas an adult swallow could surely do it much more easily, compels me to the conclusion that, from the parent's point of view, the baby is up to no good.

It seems to me just conceivable that the true explanation has nothing to do with cuckoos at all. The blood may chill at the thought, but could this be what baby swallows do to each other? Since the firstborn is going to compete with his yet unhatched brothers and sisters for parental investment, it could be to his advantage to begin his life by throwing out one of the other eggs.  {135} 

The Lack theory of clutch size considered the optimum from the parent's point of view. If I am a mother swallow, the optimum clutch-size from my point of view is, say five. But if I am a baby swallow, the optimum clutch size as I see it may well be a smaller number, provided I am one of them! The parent has a certain amount of parental investment, which she ‘wishes’ to distribute even-handedly among five young. But each baby wants more than his allotted one fifth share. Unlike a cuckoo, he does not want all of it, because he is related to the other babies. But he does want more than one fifth. He can acquire a 1/4 share simply by tipping out one egg; a 1/3 share by tipping out another. Translating into gene language, a gene for fratricide could conceivably spread through the gene pool, because it has 100 per cent chance of being in the body of the fratricidal individual, and only a 50 per cent chance of being in the body of his victim.

The chief objection to this theory is that it is very difficult to believe that nobody would have seen this diabolical behaviour if it really occurred. I have no convincing explanation for this. There are different races of swallow in different parts of the world. It is known that the Spanish race differs from, for example, the British one, in certain respects. The Spanish race has not been subjected to the same degree of intensive observation as the British one, and I suppose it is just conceivable that fratricide occurs but has been overlooked.

My reason for suggesting such an improbable idea as the fratricide hypothesis here is that I want to make a general point. This is that the ruthless behaviour of a baby cuckoo is only an extreme case of what must go on in any family. Full brothers are more closely related to each other than a baby cuckoo is to its foster brothers, but the difference is only a matter of degree. Even if we cannot believe that outright fratricide could evolve, there must be numerous lesser examples of selfishness where the cost to the child, in the form of losses to his brothers and sisters, is outweighed, more than two to one, by the benefit to himself. In such cases, as in the example of weaning time, there is a real conflict of interest between parent and child.

Who is most likely to win this battle of the generations? R. D. Alexander has written an interesting paper in which he suggests that there is a general answer to this question. According to him the parent will always win.(2) Now if this is the case, you have been wasting  {136}  your time reading this chapter. If Alexander is right, much that is of interest follows. For instance, altruistic behaviour could evolve, not because of benefit to the genes of the individual himself, but solely because of benefit to his parents’ genes. Parental manipulation, to use Alexander's term, becomes an alternative evolutionary cause of altruistic behaviour, independent of straightforward kin selection. It is therefore important that we examine Alexander's reasoning, and convince ourselves that we understand why he is wrong. This should really be done mathematically, but we are avoiding explicit use of mathematics in this book, and it is possible to give an intuitive idea of what is wrong with Alexander's thesis.

His fundamental genetic point is contained in the following abridged quotation. ‘Suppose that a juvenile. . . cause(s) an uneven distribution of parental benefits in its own favor, thereby reducing the mother's own overall reproduction. A gene which in this fashion improves an individual's fitness when it is a juvenile cannot fail to lower its fitness more when it is an adult, for such mutant genes will be present in an increased proportion of the mutant individual's offspring.’ The fact that Alexander is considering a newly mutated gene is not fundamental to the argument. It is better to think of a rare gene inherited from one of the parents. ‘Fitness’ has the special technical meaning of reproductive success. What Alexander is basically saying is this. A gene that made a child grab more than his fair share when he was a child, at the expense of his parent's total reproductive output, might indeed increase his chances of surviving. But he would pay the penalty when he came to be a parent himself, because his own children would tend to inherit the same selfish gene, and this would reduce his overall reproductive success. He would be hoist with his own petard. Therefore the gene cannot succeed, and parents must always win the conflict.

Our suspicions should be immediately aroused by this argument, because it rests on the assumption of a genetic asymmetry which is not really there. Alexander is using the words ‘parent’ and ‘offspring’ as though there was a fundamental genetic difference between them. As we have seen, although there are practical differences between parent and child, for instance parents are older than children, and children come out of parents’ bodies, there is really no fundamental genetic asymmetry. The relatedness is 50 per cent, whichever way round you look at it. To illustrate what I mean, I am going to repeat Alexander's words, but with ‘parent’, ‘juvenile’ and other appropriate  {137}  words reversed. ‘Suppose that a parent has a gene that tends to cause an even distribution of parental benefits. A gene which in this fashion improves an individual's fitness when it is a parent could not fail to have lowered its fitness more when it was a juvenile.’ We therefore reach the opposite conclusion to Alexander, namely that in any parent/offspring conflict, the child must win!

Obviously something is wrong here. Both arguments have been put too simply. The purpose of my reverse quotation is not to prove the opposite point to Alexander, but simply to show that you cannot argue in that kind of artificially asymmetrical way. Both Alexander's argument, and my reversal of it, erred through looking at things from the point of view of an individual — in Alexander's case, the parent, in my case, the child. I believe this kind of error is all too easy to make when we use the technical term ‘fitness’. This is why I have avoided using the word in this book. There is really only one entity whose point of view matters in evolution, and that entity is the selfish gene. Genes in juvenile bodies will be selected for their ability to outsmart parental bodies; genes in parental bodies will be selected for their ability to outsmart the young. There is no paradox in the fact that the very same genes successively occupy a juvenile body and a parental body. Genes are selected for their ability to make the best use of the levers of power at their disposal: they will exploit their practical opportunities. When a gene is sitting in a juvenile body its practical opportunities will be different from when it is sitting in a parental body. Therefore its optimum policy will be different in the two stages in its body's life history. There is no reason to suppose, as Alexander does, that the later optimum policy should necessarily overrule the earlier.

There is another way of putting the argument against Alexander. He is tacitly assuming a false asymmetry between the parent/child relationship on the one hand, and the brother/sister relationship on the other. You will remember that, according to Trivers, the cost to a selfish child of grabbing more than his share, the reason why he only grabs up to a point, is the danger of loss of his brothers and sisters who each bear half his genes. But brothers and sisters are only a special case of relatives with a 50 per cent relatedness. The selfish child's own future children are no more and no less ‘valuable’ to him than his brothers and sisters. Therefore the total net cost of grabbing more than your fair share of resources should really be measured, not only in lost brothers and sisters, but also in lost future offspring  {138}  due to their selfishness among themselves. Alexander's point about the disadvantage of juvenile selfishness spreading to your own children, thereby reducing your own long-term reproductive output, is well taken, but it simply means we must add this in to the cost side of the equation. An individual child will still do well to be selfish so long as the net benefit to him is at least half the net cost to close relatives. But ‘close relatives’ should be read as including, not just brothers and sisters, but future children of one's own as well. An individual should reckon his own welfare as twice as valuable as that of his brothers, which is the basic assumption Trivers makes. But he should also value himself twice as highly as one of his own future children. Alexander's conclusion that there is a built-in advantage on the parent's side in the conflict of interests is not correct.

In addition to his fundamental genetic point, Alexander also has more practical arguments, stemming from undeniable asymmetries in the parent/child relationship. The parent is the active partner, the one who actually does the work to get the food, etc., and is therefore in a position to call the tune. If the parent decides to withdraw its labour, there is not much that the child can do about it, since it is smaller, and cannot hit back. Therefore the parent is in a position to impose its will, regardless of what the child may want. This argument is not obviously wrong, since in this case the asymmetry that it postulates is a real one. Parents really are bigger, stronger and more worldly-wise than children. They seem to hold all the good cards. But the young have a few aces up their sleeves too. For example, it is important for a parent to know how hungry each of its children is, so that it can most efficiently dole out the food. It could of course ration the food exactly equally between them all, but in the best of all possible worlds this would be less efficient than a system of giving a little bit more to those that could genuinely use it best. A system whereby each child told the parent how hungry he was would be ideal for the parent, and, as we have seen, such a system seems to have evolved. But the young are in a strong position to lie, because they know exactly how hungry they are, while the parent can only guess whether they are telling the truth or not. It is almost impossible for a parent to detect a small lie, although it might see through a big one.

Then again, it is of advantage to a 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  {139}  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.

There is, then, no general answer to the question of who is more likely to win the battle of the generations. What will finally emerge is a compromise between the ideal situation desired by the child and that desired by the parent. It is a battle comparable to that between cuckoo and foster parent, not such a fierce battle to be sure, for the enemies do have some genetic interests in common — they are only enemies up to a point, or during certain sensitive times. However, many of the tactics used by cuckoos, tactics of deception and exploitation, may be employed by a parent's own young, although the parent's own young will stop short of the total selfishness that is to be expected of a cuckoo.

This chapter, and the next in which we discuss conflict between mates, could seem horribly cynical, and might even be distressing to human parents, devoted as they are to their children, and to each other. Once again 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.

CHAPTER 8
Battle of the generations

p. 124 R. L. Trivers, in 1972, neatly solved the problem. . .

Robert Trivers, whose papers of the early 1970s were among the most important inspirations for me in writing the first edition of this book, and whose ideas especially dominated Chapter 8, has finally produced his own book, Social Evolution. I recommend it, not only for its content but for its style: clear-thinking, academically correct but with just enough anthropomorphic irresponsibility to tease the pompous, and spiced with personal autobiographical asides. I cannot resist quoting one of these: it is so characteristic. Trivers is describing his excitement on observing the relationship between two rival male baboons in Kenya: ‘There was another reason for my excitement and this was an unconscious identification with Arthur. Arthur was a superb young male in his prime . . .’ Trivers's new chapter on parent-offspring conflict brings the subject up to date. There is indeed rather little to add to his paper of 1974, apart from some new factual examples. The theory has stood the test of time. More detailed mathematical and genetic models have confirmed that Trivers's largely verbal arguments do indeed follow from currently accepted Darwinian theory.

p. 135 According to him the parent will always win.

Alexander has generously conceded, in his 1980 book Darwinism and Human Affairs (p. 39), that he was wrong to argue that parental victory in parent-offspring conflict follows inevitably from fundamental Darwinian assumptions. It now seems to me that his thesis, that parents enjoy an asymmetrical advantage over their offspring in the battle of the generations, could be bolstered by a different kind of argument, which I learnt from Eric Charnov.

Charnov was writing about social insects and the origins of sterile castes, but his argument applies more generally and I shall put it in general terms. Consider a young female of a monogamous species, not necessarily an insect, on the threshold of adult life. Her dilemma is whether to leave and try to reproduce on her own, or to stay in the parental nest and help rear her younger brothers and sisters. Because of the breeding habits of her species, she can be confident that her mother will go on giving her full brothers and sisters for a long time to come. By Hamilton's logic, these sibs are just as  {299}  genetically ‘valuable’ to her as her own offspring would be. As far as genetic relatedness is concerned, the young female will be indifferent between the two courses of action; she doesn't ‘care’ whether she goes or stays. But her parents will be far from indifferent to which she does. Looked at from the point of view of her old mother, the choice is between grandchildren or children. New children are twice as valuable, genetically speaking, as new grandchildren. If we speak of conflict between parents and offspring over whether the offspring leaves or stays and helps at the nest, Charnov's point is that the conflict is a walk-over for the parents for the very good reason that only the parents see it as a conflict at all!

It is a bit like a race between two athletes, where one has been offered £1,000 only if he wins, while his opponent has been promised £1,000 whether he wins or loses. We should expect that the first runner will try harder and that, if the two are otherwise evenly matched, he will probably win. Actually Charnov's point is stronger than this analogy suggests, because the costs of running flat out are not so great as to deter many people, whether they are financially rewarded or not. Such Olympic ideals are too much of a luxury for the Darwinian games: effort in one direction is always paid for as lost effort in another direction. It is as if the more effort you put into any one race, the less likely you are to win future races because of exhaustion.

Conditions will vary from species to species, so we can't always forecast the results of Darwinian games. Nevertheless, if we consider only closeness of genetic relatedness and assume a monogamous mating system (so that the daughter can be sure that her sibs are full sibs), we can expect an old mother to succeed in manipulating her young adult daughter into staying and helping. The mother has everything to gain, while the daughter herself will have no inducement to resist her mother's manipulation because she is genetically indifferent between the available choices.

Once again, it is important to stress that this has been an ‘other things being equal’ kind of argument. Even though other things will usually not be equal, Charnov's reasoning could still be useful to Alexander or anyone else advocating a parental manipulation theory. In any case, Alexander's practical arguments for expecting parental victory — parents being bigger, stronger and so on — are well taken.  {300} 

[veronica]

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