It is especially easy to appreciate this balance when it is struck between two rather separate aspects of life: peacock survival versus beauty in the eyes of peahens, for instance. Darwinian theory tells us that all survival is just a means to the end of gene propagation, but this does not stop us partitioning the body into those components, like legs, that are primarily concerned with individual survival and those, like penises, that are concerned with reproduction. Or, those, like antlers, that are devoted to competing with rival individuals versus those, like legs and penises, whose importance does not depend upon the existence of rival individuals. Many insects impose a rigid separation between radically different stages in their life history. {125} Caterpillars are devoted to gathering food and growing. Butterflies are like the flowers they visit, devoted to reproducing. They do not grow, and they suck nectar only to burn it immediately as aviation fuel. When a butterfly reproduces successfully, it spreads the genes not just for being an efficient flying and mating butterfly but for being the efficient feeding caterpillar that it was, as well. Mayflies feed and grow as underwater nymphs for up to three years. They then emerge as flying adults that live only a matter of hours. Many of them are eaten by fish, but even if they were not they would soon die anyway, because they cannot feed and they do not even possess guts (Henry Ford would have loved them). Their job is to fly until they find a mate. Then, having passed on their genes – including the genes for being an efficient nymph capable of feeding underwater for three years – they die. A mayfly is like a tree that takes years to grow, then flowers for a single glorious day and dies. The adult mayfly is the flower that briefly blooms at the end of life and the beginning of new life.
A young salmon migrates down the stream of its birth and spends the bulk of its life feeding and growing in the sea. When it reaches maturity it again seeks out, probably by smell, the mouth of its native stream. In an epic and much-celebrated journey the salmon swims upstream, leaping falls and rapids, home to the headwaters from which it sprang a lifetime ago. There it spawns and the cycle renews. At this point there is typically a difference between Atlantic and Pacific salmon. The Atlantic salmon, having spawned, may return to the sea with some chance of repeating the cycle a second time. Pacific salmon die, spent, within days of spawning. {126}
A typical Pacific salmon is like a mayfly but without the anatomically clear-cut separation between nymph and adult phases in the life history. The effort of swimming upstream is so great that it cannot pay to do it twice. Therefore natural selection favors individuals that put every ounce of their resources into one “big bang” reproductive effort. Any resources left after breeding would be wasted – the equivalent of Henry Ford's overdesigned kingpins. The Pacific salmon have evolved toward whittling down their postreproductive survival until it approaches zero, the resources saved being diverted into eggs or milt. The Atlantic salmon were drawn toward the other route. Perhaps because the rivers they have to mount tend to be shorter and spring from less formidable hills, individuals that keep some resources back for a second reproductive cycle can sometimes do well by it. The price these Atlantic salmon pay is that they cannot commit so much to their spawn. There is a trade-off between longevity and reproduction, and different kinds of salmon have opted for different equilibria. The special feature of the salmon life cycle is that the grueling odyssey of their migration imposes a discontinuity. There is no easy continuum between one breeding season and two. Commitment to a second breeding season drastically cuts into efficiency in the first. Pacific salmon have evolved toward an unequivocal commitment to the first breeding season, with the result that a typical individual unequivocally dies immediately after its single titanic spawning effort.
The same kind of trade-off marks every Ufe, but it is usually less dramatic. Our own death is probably programmed in something like the same sense as that of the salmon but in a {127} less downright and clear-cut fashion. Doubtless a eugenicist could breed a race of superlatively long-lived humans. You would choose for breeding those individuals who put most of their resources into their own bodies at the expense of their children: individuals, for example, whose bones are massively reinforced and hard to break but who have little calcium left over to make milk. It is easy enough to live a bit longer, if you are cosseted at the expense of the next generation. The eugenicist could do the cosseting and exploit the trade-offs in the desired direction of longevity. Nature will not cosset in this way, because genes for scrimping the next generation will not penetrate the future.
Nature's Utility Function never values longevity for its own sake but only for the sake of future reproduction. Any animal that, like us but unlike a Pacific salmon, breeds more than once faces trade-offs between the current child (or litter) and future children. A rabbit that devoted all her energy and resources to her first litter would probably have a superior first litter. But she would have no resources left to carry her on to a second litter. Genes for keeping something in reserve will tend to spread through the rabbit population, carried in the bodies of second- and third-litter babies. It is genes of this kind that so conspicuously did not spread through the population of Pacific salmon, because the practical discontinuity between one breeding season and two is so formidable.
As we grow older our chances of dying within the next year, after initially decreasing and then plateauing for a while, settle down to a long climb. What is happening in this long increase in mortality? It is basically the same principle as {128} for the Pacific salmon, but spread out over an extended period instead of being concentrated in a brief precipitous orgy of death after the orgy of spawning. The principle of how senescence evolved was originally worked out by the Nobel laureate and medical scientist Sir Peter Medawar in the early 1950s, with various modifications to the basic idea added by the distinguished Darwinians G.C. Williams and W.D. Hamilton.
The essential argument is as follows: First, as we saw in chapter 1, any genetic effect will normally be switched on at a particular time during the life of the organism. Many genes are switched on in the early embryo, but others – like the gene for Huntington's chorea, the disease that tragically killed the folk poet and singer Woody Guthrie – are not switched on until middle age. Second, the details of a genetic effect, including the time at which it is switched on, may be modified by other genes. A man possessing the Huntington's chorea gene can expect to die from the disease, but whether it kills him when he is forty or when he is fifty-five (as Woody Guthrie was) may be influenced by other genes. It follows that by selection of “modifier” genes the time of action of a particular gene can either be postponed or brought forward in evolutionary time.
A gene like the Huntington's chorea gene, which switches on between the ages of thirty-five and fifty-five, has plenty of opportunity to be passed on to the next generation before it kills its possessor. If, however, it were switched on at the age of twenty, it would be passed on only by people who reproduce rather young, and therefore it would be strongly selected against. If it were switched on at the age of ten, it would {129} essentially never be passed on. Natural selection would favor any modifier genes that had the effect of postponing the age of switching on of the Huntington's chorea gene. According to the Medawar/Williams theory, this would be exactly why it normally does not switch on until middle age. Once upon a time it may well have been an early maturing gene, but natural selection has favored a postponing of its lethal effect until middle age. No doubt there is still slight selection pressure to push it on into old age, but this pressure is weak because so few victims die before reproducing and passing the gene on.