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.
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. Now if this is the case, you have been wasting
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 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 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 wel
l. 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 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.
Battle of the sexes
If there is conflict of interest between parents and children, who share 50 per cent of each others' genes, how much more severe must be the conflict between mates, who are not related to each other? All that they have in common is a 50 per cent genetic shareholding in the same children. Since father and mother are both interested in the welfare of different halves of the same children, there may be some advantage for both of them in cooperating with each other in rearing those children. If one parent can get away with investing less than his or her fair share of costly resources in each child, however, he will be better off, since he will have more to spend on other children by other sexual partners, and so propagate more of his genes. Each partner can therefore be thought of as trying to exploit the other, trying to force the other one to invest more. Ideally, what an individual would 'like' (I don't mean physically enjoy, although he might) would be to copulate with as many members of the opposite sex as possible, leaving the partner in each case to bring up the children. As we shall see, this state of affairs is achieved by the males of a number of species, but in other species the males are obliged to share an equal part of the burden of bringing up children. This view of sexual partnership, as a relationship of mutual mistrust and mutual exploitation, has been stressed especially by Trivers. It is a comparatively new one to ethologists. We had usually thought of sexual behaviour, copulation, and the courtship that precedes it, as essentially a cooperative venture undertaken for mutual benefit, or even for the good of the species!
Let us go right back to first principles, and inquire into the fundamental nature of maleness and femaleness. In Chapter 3 we discussed sexuality without stressing its basic asymmetry. We simply accepted that some animals are called male, and others female, without asking what these words really meant. But what is the essence of maleness? What, at bottom, defines a female? We as mammals see the sexes defined by whole syndromes of characteristics-possession of a penis, bearing of the young, suckling by means of special milk glands, certain chromosomal features, and so on. These criteria for judging the sex of an individual are all very well for mammals but, for animals and plants generally, they are no more reliable than is the tendency to wear trousers as a criterion for judging human sex. In frogs, for instance, neither sex has a penis. Perhaps, then, the words male and female have no general meaning. They are, after all, only words, and if we do not find them helpful for describing frogs, we are quite at liberty to abandon them. We could arbitrarily divide frogs into Sex 1 and Sex 2 if we wished. However, 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. The difference is especially pronounced in reptiles and in birds, where a single egg cell is big enough and nutritious enough to feed a developing baby for several weeks. Even in humans, where the egg is microscopic, it is still many times larger than the sperm. As we shall see, it is possible to interpret all the other differences between the sexes as stemming from this one basic difference.
In certain primitive organisms, for instance some fungi, maleness and femaleness do not occur, although sexual reproduction of a kind does. In the system known as isogamy the individuals are not distinguishable into two sexes. Anybody can mate with anybody else. There are not two different sorts of gametes-sperms and eggs-but all sex cells are the same, called isogametes. New individuals are formed by the fusion of two isogametes, each produced by meiotic division. If we have three isogametes, A, B, and C,
A could fuse with B or C,
B could fuse with A or C.
The same is never true of normal sexual systems. If A i
s a sperm and it can fuse with B or C, then B and C must be eggs and B cannot fuse with C.
When two isogametes fuse, both contribute equal numbers of genes to the new individual, and they also contribute equal amounts of food reserves. 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.
Parker and others showed how this asymmetry might have evolved from an originally isogamous state of affairs. In the days when all sex cells were interchangeable and of roughly the same size, there would have been some that just happened to be slightly bigger than others. In some respects a big isogamete would have an advantage over an average-sized one, because it would get its embryo off to a good start by giving it a large initial food supply. There might therefore have been an evolutionary trend towards larger gametes. But there was a catch. The evolution of isogametes that were larger than was strictly necessary would have opened the door to selfish exploitation. Individuals who produced smaller than average gametes could cash in, provided they could ensure that their small gametes fused with extra-big ones. This could be achieved by making the small ones more mobile, and able to seek out large ones actively. The advantage to an individual of producing small, rapidly moving gametes would be that he could afford to make a larger number of gametes, and therefore could potentially have more children. Natural selection favoured the production of sex cells that were small and that actively sought out big ones to fuse with. So we can think of two divergent sexual 'strategies' evolving. There was the large-investment or 'honest' strategy. This automatically opened the way for a small-investment exploitative strategy. Once the divergence between the two strategies had started, it would have continued in runaway fashion. Medium-sized intermediates would have been penalized, because they did not enjoy the advantages of either of the two more extreme strategies. The exploiters would have evolved smaller and smaller size, and faster mobility. The honest ones would have evolved larger and larger size, to compensate for the ever-smaller investment contributed by the exploiters, and they became immobile because they would always be actively chased by the exploiters anyway. Each honest one would 'prefer' to fuse with another honest one. But the selection pressure to lock out exploiters would have been weaker than the pressure on exploiters to duck under the barrier: the exploiters had more to lose, and they therefore won the evolutionary battle. The honest ones became eggs, and the exploiters became sperms.