신경 쓰이는 잡다한 이야기

     If you look at what actually changes in evolution it is changes in gene frequencies in gene pools as generations go by. I'm talking now about sexually reproducing organisms such as ourselves, such as mammals, birds, such as all vertebrates, and so on. Now, the reason that gene survival is important is that genes in the form of copies of the DNA are immortal. They are potentially immortal, in the sense that what goes on from generation to generation, is the coded information in the DNA. So potentially a bit of DNA code can persist for tens of millions, hundreds of millions of years. Not all of it does, therefore there's an important difference between those bits of DNA code that do persist for over many generations and those that don't. That is natural selection.

     The reason why I insist on talking about natural selection at the level of the genes is precisely that it really makes a difference at the level of the gene because some genes are going to go on forever for a very long time and others are not. No other units, no other level in the hierarchy of life has that property; individual organisms don't, groups don't, species don't, nothing else has that property of 'potential immortality' which genes have. Now this is nothing new; this is the way that people have looked at natural selection ever since 1930s when the great founders of the population genetics modified Darwinism, Darwin himself didn't know anything about genetics. It, genetics, was imported properly into evolutionary theory in the 1930s, and that time it was understood that natural selection is the differential survival of genes in gene pools.

     Now, as a euphemism, people will sometimes say animals do so and so to perpetuate the species. They realized that survival itself is not enough. Survival, about Darwin realized of course, the survival plus reproduction is what matters. Survival is a means to be end of reproduction. And, a euphemism for reproduction is perpetuating the species. And partly out of the result of that, a sort of myth grew up of what animals are doing, the reason why they do what they do is to perpetuate the species. The real reason is to perpetuate their genes, if you forgive the teleological language, the real reason that is to perpetuate their genes, perpetuating the species is an incidental consequence.

     The idea of perpetuating species of perpetuating the group was semi-formalized into theory of group selection, and some biologists tried to explain the evolution of thins like altruism as a result of the differential survival of groups. Those groups, in which individuals care for each other, survive better than groups in which individuals don't care for each other; that was group selection not aimed to survival, but perhaps bad off other daughter groups. That theory of group selection was correctly rubbished in the 1960s by various people, and perhaps the most significant of them was W. D. Hamilton who invented the phrase 'inclusive fitness,' which the questioner referred to, and I say that inclusive fitness was not the most clear way of explaining the point of view but is the one that Hamilton brilliantly put forward.

     Fitness, in Darwin's time, fitness meant roughly what we in colloquial language mean by fitness it meant being fast running, it meant being strong, it meant being attractive, all the thing that we in all ordinary language might mean by fitness. When the population genetics revolution happened, the word fitness was given a technical meaning which was pretty much 'that which survives,' which meant that survival of the fittest, the phrase that Darwin took over from Herbert Spencer, survival of the fittest became a tautology because the fittest from whole is defined as that which survives.

     The technical meaning of fitness pretty much meant the reproductive success of an individual not only to the next generation but the grandchild generation, the great grandchild generation, and so on. So a 'fit' animal is one who had what it took, who was equipped, to leave a lot of descendants that has obvious connection with the classical meaning of fitness; you can run fast if you got keen eyesight and so on, that helps you, that equips you, to have lots of descendants, but fitness was actually defined mathematically in this precise way.

     When Hamilton came along, he realized that, because what really matters is gene survival, it's not enough to count just children and grandchildren and great-grandchildren direct descendants, because collateral relatives - siblings, nephews and nieces and so on - also share genes. Therefore a gene that makes an individual care for a brother or sister and nephew or niece would survive for the same kind of a reason as a gene for caring for children and grandchildren. Hamilton realized that what matters is the statistical probability of a gene being shared and that statistical probability can be calculated whether or not the descent is lineau or whether we're talking about collateral relatives. And he showed that what matters is the quantity called are the 'coefficient of relationship' which is a half for offspring, a half for full siblings, a quarter for half siblings, a quarter for nephews and nieces, an eighth for first cousins, a sixteenth for second cousins, a quarter for grandchildren, and so on. So natural selection then favors animals that care for those other animals that have a statistical likelihood of sharing the same genes.

     Hamilton could've left it of that, and that was indeed the way he phrased it in his first paper published in 1963 in the American Naturalist. However, he wanted to take over the concept of fitness and modify it, in the way that it needs to be modified in order to take account of collateral relatives, because his audience, professional biologists, were already accustomed to talking about fitness, and Hamilton, what he said 'look, you can keep your fitness in your equations but you must modify it to take account of collateral relatives.' And the modified former fitness that he invented was 'inclusive fitness' and he did a lot of rather difficult mathematics in order to show how to calculate inclusive fitness. Informally you can define inclusive fitness as that quantity which an individual will appear to be maximizing, when what is really being maximized is gene survival.

     The great triumph of Hamilton's theory was the social insects; it works for all animals, it applies to all animals in the form of a simple equation called Hamilton's rule which states that an altruistic act will spread through the population if RB is greater than C, where R is the coefficient of relationship, that fraction I told you about a half a quarter whatever it is, B is the benefit to the recipient, and C is the cost to the donor - the cost to the altruist. But as I say he wanted to express, he wanted to find a modification of fitness that you need in order to the others were rescued the concept of fitness.

     And I said that the great triumph of his theory was with social insects but I also said that R be greater than C works for all animals and all plants even if R is so small, or B and CR so large, that it you can more or less, forget about collateral relatives, and that is true for many animals. In the case of the social insects, R is high, social insects are, the workers are sterile, they're closely related to their fertile relatives, so worker ant doesn't reproduce in most cases it doesn't reproduce itself, but is closely related to the young queens and males, which is going to be churned out by the nest. And that's why the ants work, that's why worker ants do work they do, because they share genes with the queens and males which are going to propagate copies of those very same genes into the next generation.

     Now the question I was asked why such eminent biologists as E. O. Wilson the great entomologist, the great ant specialist, the world's leading expert on ants indeed, why he has turned against inclusive fitness, it's not entirely clear to me, I mean, it's as though he thinks, and this actually goes back to his great book, 'sociobiology,' which already in 1975 betrays a certain misunderstanding of the idea of inclusive fitness. Wilson clearly thought that caring for relatives is a kind of unparsimonious addition to the ordinary theory of natural selection which was what we were quite happy with for the 1930s onwards. So for Wilson, you don't move on to inclusive fitness unless you absolutely have to, because classical fitness, which applies only to direct offspring and descendants, grandchildren and so on, will do the job.

     My reply to that would be, you can't escape from inclusive fitness, you can't escape from looking at collateral relatives whether you like it or not, RB greater than C is the correct formulation of the way of natural selection works, and if you've got at species in which there is no altruism toward collateral relatives that's absolutely fine, it just means that the B's and C's and R's work out that way. But you cannot get away from inclusive fitness; it follows deductively from the neo-Darwinian synthesis from the 1930s; it's not an add-on, it's not an additional thing that has to be justified by empirical evidence. It's there deductively in the mathematics. To go around looking for species of animal and testing Hamilton theory is a bit like testing Pythagoras' theorem by going out with a ruler and measuring right angled triangles - you don't do that, you prove Pythagoras' theorem by mathematical deduction and that's what inclusive fitness is, like.

     Now I think, is a way of interpreting why Wilson and his colleagues, I mean nothing else the weakened, watered-down version of their objection which is that inclusive fitness itself is difficult to calculate in practice, and that's true. It is probably a difficult thing to do in practice unless you could measure very, very precisely lot of difficult economic quantities. So, it's a reasonable objection to say that inclusive fitness is not a practical measure for a field biologist. But you cannot say that 'therefore altruistic behavior is not explained by Hamilton's rule, RB greater than C, and my own preference would be actually to forget about inclusive fitness and go straight to the level of the gene, and think in terms of genes manipulating individual organisms, rather than think in terms of the individual organisms trying to maximize their inclusive fitness.