The Death of Greatness
Roger Kitching, Professor of Ecology at
Griffith University in Brisbane, spoke
to Robyn Williams of Australian ABC Radio National's
Ockhams Razor on Sunday 18 June 2000)
This is an obituary and a celebratory talk about the great scientist William Hamilton who died earlier this year.
Nature is red in tooth and claw. The survival of the fittest. Its only human Nature; choose your own cliché. Each one is an alibi for the carnivorous version of destiny, that loving kindness is for wimps and wimps get wiped out by the big boys. Sad, but true.
Well it isn't. True, that is. Today we celebrate the work of someone youve probably never heard of but whose contribution to science and by association to our hope for progress, was simply enormous. He had a full page obituary in both The Economist and the journal, Nature. And today, through Professor Roger Kitching from Griffith University, Ockham's Razor pays tribute to the New Zealander, Dr Bill Hamilton.
Look, evolution works like this: Some new feature, an extra whisker, a different enzyme in the digestive process, or a new behaviour, arises because theres some mistake in the genetic copying when cells replicate before reproduction. In general, the new feature simply disappears from view because whatever change it brings about in the animal or plant is not as good as the 'norm' before the copying mistake. Once in a while though, the new feature is actually an improvement. How do we know? Well, the animals or plants, or bacteria or viruses, that possess the new feature live longer, perform better and somehow, and this is the key, they reproduce more often, producing kids who, in turn, survive better than the other offspring on the block.
Thinking about genetically programmed behaviour, this means that one of the these mis-copied novelties (let's call them mutations) produces some twist in hunting behaviour, competitive behaviour, or what-have-you, that means that that particular individual gets more food, or avoids being eaten somewhat better than its neighbours. This is an individual thing and it is essentially 'selfish'; wittingly or unwittingly, it's about doing better than your neighbour. Yet we see worker ants dying to save not their offspring, because they don't have any, but the overall ant colony; we see elephants and dolphins supporting and protecting injured or weakly neighbours; and we see springboks leaping high into the air to give warning of the predator that they have seen first. These all appear to be altruistic acts; our 'selfish' theory would have the worker ant simply doing no more than feeding itself and acting to enhance its own survival, and if there were possible, reproductive activity. Indeed, how come it gave up reproduction in the first place? The elephants and dolphins would get on with their own feeding, mating and surviving activities, leaving their less fortunate neighbours to cope as best they can. And the springbok that sees the cheetah first would have hared off to escape the predator, leaving its less vigilant herd mates to their own devices.
The evolution of so-called altruism stumped Darwin. The assumption that evolution could not produce something that was altruistic was the basis for Spenserian 'Social Darwinism', not unrelated to the evil underpinnings of Stalinism and Nazism.
And then in 1964 William Hamilton pointed out that the individuals in the so-called 'struggle for existence' to which we had ascribed such selfish behaviour should be regarded not as biological entities in themselves, but as vehicles for the transmission or otherwise of sets of genes. And that the success or otherwise of a mutation should be judged not on the basis of whether or not a particular individual organism survived and reproduced better, but whether the population of genes spread over many individuals actually survived better. To put it another way: if a mutation produces a new gene which actually decreases its owner's chances of survival or reproduction, such as succouring injured mates, but which thereby enhances the survival probabilities of individuals which contain a large proportion of the genes of the apparent altruist, then that gene and the behaviour it codes for, can be selected. To trivialise this, it will be fine to die for more than two siblings, eight cousins, sixteen half-cousins and so on, because by so doing you are doing the whole set of genes you contain a favour, by preserving, on average, rather more copies of them (in your rellies) than you yourself possess. This is the basis for the theory of inclusive fitness and the numbers involved reflect what has become known as Hamilton's Rule.
Now if all this seems familiar and you associate Edward O. Wilson's great tome Sociobiology: the New Synthesis, or, more likely, Richard Dawkins' racy popularisation The Selfish Gene with this idea, you would be one of a large number who have been influenced by these seminal books. Both Wilson and Dawkins brought flair and originality to their writings and added greatly to our understanding of the evolution of social (that is, apparently altruistic) behaviour. But the giant on whose shoulders they stood was Bill Hamilton, according to American scientist Robert Trivers, the only truly original evolutionist since Darwin.
And now Bill is dead!
Bill first shambled into my life in 1964 when I turned up for the first presentation by the new genetics lecturer at Imperial College, London. I was a third-year student at the time. Bill was that lecturer and it is no exaggeration to say, that Bill's lectures were models of everything a lecture should not be. They were disordered, not leavened by anecdote or theatricality, and they juxtaposed the relatively elementary with the very cutting edge of the subject. In fact if we had only known we could have pipped Dawkins on the post by ten years, because as my scrappy notes recall, Bill told us of his discoveries published that very year in his seminal, if somewhat densely written, paper in The Journal of Theoretical Biology. He spoke of male arrhenotoky rather than haplo-diploidy (in fact we knew not what the concept meant under either description) but it was all there.
So, like generations of students before and since, we taught ourselves such elements of the topic as we decided we needed to know. But, different from modern students perhaps, we did not resent the fact. In fact we pursued Bill to the pubs of South Kensington and tried to fathom why this particular young scientist had been chosen by the omniscient Professor O.W. Richards to join his select staff. This immediately became obvious. Bill lived life on an intellectual plane that few of us could see, let alone share. He was the one who brought into my life the notion that behavioural, ecological and evolutionary ecology could and should have a mathematical base. Of course this idea preceded him in the writings of Fisher, Sewall Wright, Maynard Smith and others within theoretical genetics but my eleven student colleagues and I were perhaps the last generation of classically trained biologists who just had to be field naturalists before we became zoologists, and mathematicians we were not. Bill, like Darwin before him, was also an avid field naturalist with a deep knowledge of the natural world. So when he spoke of the importance of formulae and equations it rang true; it set me on a road which led to mathematical ecology, simulation modelling and systems ecology, old hat now, but almost unheard of in the mid 60s.
In 1977 Bill moved on to the University of Michigan. Ultimately he returned to the UK as Royal Society Research Professor at Oxford. He tackled other research topics and always the big, intractable questions: the origin of insect diversity under bark, the evolution of sex ratios, evolutionarily stable strategies, and, for much of the last 20 years, the relationship between parasite challenge and the evolution of sexual reproduction. This last challenge was ultimately to take his life, but I'll come to that. The problem with sexual reproduction from an evolutionary point of view is that if you reproduce asexually, as all life on earth did for billions of years, then each time you produce offspring you pass on 100% of your genes. Why then suddenly evolve a system, sexual reproduction, by which an individual automatically gives up 50% of its genome at each reproductive event?
Like many evolutionary innovations it is possible to justify the feature after the fact, but how could the actual switch occur? Bill picked up and developed an old idea that sexual reproduction was a means by which a host organism could throw co-evolving parasites (viruses, bacteria, spirochaetes) off the track, by suddenly jumbling its genes and starting afresh with at least a 50% new set. Micro-organisms reproducing within cells of animals or plants many times a day can rapidly evolve more and more effective ways of diverting the host's productivity into themselves. The multi-celled hosts reproduce much less often and when they do, they get a once-only chance to throw the parasites back many generations. This is the selective pressure and sexual reproduction is the mechanism.
It was in pursuit of this idea that Bill travelled to the Congo to study the chimpanzees that, one theory has it, were the agents from which the HIV virus 'jumped' to the chimp's closest relatives, humans. And it was another parasite, the malaria organism, that infected Bill, and the haemorrhagic complications of this attack that killed him.
Bill's written products were relatively few, but each paper was so influential that almost without exception each engendered a whole school of derivative work. He was a Fellow of the Royal Society and won the Darwin Medal, Kyoto and Crafoord prizes, the highest accolades available in evolutionary biology. As a lifelong academic I spend my life among smart people, a lot of whom are clever and a few of whom are very sharp indeed. But it is a quantum leap from these day-to-day intellectuals to the few who you know have the qualities of genius. I have known just three or four of these: Bill was one, and, incidentally, the young Australian expatriate who will probably fill his shoes, David Haig, is another, significantly, employed not in Australia, but at Harvard. It is these few that put the rest of us scientists into context. The over-dramatisation of prosaic and even somewhat exceptional scientific results by publicity-conscious universities and research organisations, have devalued words like 'breakthrough', 'brilliance' and 'genius'. We of the scientific community have in general gone along with this as part of survival in the black-and-white, or should that perhaps be grey? world of science as driven by economic rationalism. But we know in our hearts that we're kidding ourselves and it is in the white light of genius that we are reminded of the polychrome fascination of the world that drew us to science and research in the first place. There are few such lights: Bill Hamilton was one of them and his untimely passing is a matter for global mourning.
I cannot close this obituary and celebratory piece without noting one sobering contrast: Bill Hamilton produced few papers, far less than one for each year of his working life; he gained few grants, excluding his prize windfalls, and he was, by his own acknowledgment, an indifferent lecturer. In my university currently, he would not get tenure or promotion, he would be annually admonished because of adverse personal comments by his juvenile student audiences, and he would not receive the paltry $2,000 a year university research allowance as he did not meet the minimum requirements of two refereed papers each year. In fact, in the current Australian system he would probably have long ago given up on his scientific career and moved on to writing novels, (there is an unpublished manuscript) selling insurance or steak knives, or bolstering the Social Darwinists in the banking profession. Of course few academics are as brilliant or influential as Bill was able to be, but it has been the great strength of the western university tradition that the freedom to think 'outside the square' allows such revolutionaries to flourish perhaps once in a generation in each field. That is all it takes. It justifies the existence of the entire system, warts and all, and it is on this that our cultural, intellectual and even economic wellbeing depends.
Of course this is threatening to small minds.
Vale, Bill, vale.
Another site which pays homage to Bill Hamilton can be found here