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Hamilton and Haldane's ideas of 'inclusive fitness'


The first idea of population regulation by group selection
John Maynard Smith Scientist
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In '62, I think it was, Wynne-Edwards published his book in which he argued that the regulation of animal population numbers was brought about because animals limited their own population numbers behaviourally by displays revealing the density of the population. The idea being that if they could regulate their own numbers, it would be better for the survival of the species than if they all bred as fast as they could and there was kind of population disaster and starvation. So the notion was that in order to avoid starving you regulate your own population by display. And Wynne-Edward's great merit, of course, was that he realised that this is a Group Selection argument, it can only be true if Natural Selection operates by favouring some species at the expense of others rather than some individuals. And this book was really enormously influential. My friends and colleagues all swallowed it whole except the population of geneticists and a few people like David Lack at Oxford who were very hostile to it. And I tried to model mathematically Wynne-Edwards' argument and showed that, in fact, you can have selection operating between groups and it can be effective, but in order for that to be true, the groups have to be genetically very homogenous within the groups and different between groups, otherwise there's nothing selection can get hold of. And in order for that to be true, it has to be true of the groups founded by, in the model I made, one pregnant female, so that all the offspring are closely related. And I wrote this up and then, at about the same time, I can't remember the exact order, I'd come across this paper of Hamilton's. And also at about that time, David Lack rang me up and said that he and Tinbergen had been discussing this idea of Wynne-Edwards and would I like to come to Oxford and talk to them about it, not as a seminar, but just to sit around and talk about it one evening. And, in fact, four of us had a long talk in Oxford: Lack, Tinbergen and Arthur Cain and myself. And we all agreed that Wynne-Edwards' ideas, although interesting, were basically false, had to be, because some group structure was wrong. But I do remember saying at that time, 'Look, there's this paper by this chap, Hamilton, which isn't wrong, is not the same idea, it's an important idea.' And I remember we agreed that it was important to distinguish between what Hamilton was saying, which was right and interesting, and what Wynne-Edwards was saying, which was wrong, interesting but false, if you like. So when I wrote my paper about my Group Selection Model, the paper is actually published in Nature and it's actually Group Selection and Kin Selection. And we invented the term kin selection, and it emerged that evening, and I can't remember who first suggested it, I think it wasn't me, though I first used it in print. And the idea of the term kin selection was to distinguish it from group selection, so that the two idea... two notions would have different names that people... people can't keep things distinct in their minds unless they have different names, in my experience, you have to give a name to something. But I do think there were problems with Bill that he had the feeling that somehow or other I'd been trying to hold up his publication until I'd got in first. I can understand his feelings on this. He was a young man, he had no name, his ideas were not taken seriously. I think it's one of these horrid misunderstandings.

[Q] I don't think he thinks that anymore.

I don't think he does. No, I'll talk later if I may about how that... no I think Bill and I are fine now, I mean, I think we understand one another very well and admire one another. I admire him enormously and respect him. But no doubt, that from 1964 to the late 1970s, it was a problem. I think it is to our credit that we did not encourage our students to quarrel with one another, and we passed students from one to the other, and we didn't have public rows about it.

[Q] Unlike Fisher and Haldane.

Unlike Fisher and Haldane. I was very conscious of the fact that Fisher and Haldane had done a lot of damage by public rows, you know, and I wasn't going to do the same. And in any case, there was nothing intellectual to argue about. I didn't disagree with Bill intellectually, it was just that Bill felt, I think with some justification, that I had not been as helpful as I should have been, the justification being that I could have helped him when I was a student, and I didn't. And maybe I wasn't as generous to his ideas as I should have been, I don't know, it's hard to say. But I don't think, as far as the notion of inclusive fitness, kin selection, and so on, I have to say, I don't think I contributed anything significant to that idea. I think that idea is essentially Bill Hamilton's. I contributed to popularising it but not to having it. I mean, the idea is essentially his.

The late British biologist John Maynard Smith (1920-2004) is famous for applying game theory to the study of natural selection. At Eton College, inspired by the work of old Etonian JBS Haldane, Maynard Smith developed an interest in Darwinian evolutionary theory and mathematics. Then he entered University College London (UCL) to study fruit fly genetics under Haldane. In 1973 Maynard Smith formalised a central concept in game theory called the evolutionarily stable strategy (ESS). His ideas, presented in books such as 'Evolution and the Theory of Games', were enormously influential and led to a more rigorous scientific analysis and understanding of interactions between living things.

Listeners: Richard Dawkins

Richard Dawkins was educated at Oxford University and has taught zoology at the universities of California and Oxford. He is a fellow of New College, Oxford and the Charles Simonyi Professor of the Public Understanding of Science at Oxford University. Dawkins is one of the leading thinkers in modern evolutionary biology. He is also one of the best read and most popular writers on the subject: his books about evolution and science include "The Selfish Gene", "The Extended Phenotype", "The Blind Watchmaker", "River Out of Eden", "Climbing Mount Improbable", and most recently, "Unweaving the Rainbow".

Tags: 1962, Animal Dispersion in Relation to Social Behavior, group selection, natural selection, Oxford University, Group Selection and Kin Selection, Nature, VC Wynne-Edwards, David Lack, WD Hamilton, Nikolaas Tinbergen, AJ Cain, Robert Fisher, JBS Haldane

Duration: 5 minutes, 13 seconds

Date story recorded: April 1997

Date story went live: 24 January 2008