Richard Dawkins

Richard Dawkins, The Selfish Gene, New York, Oxford University Press, 1976[1989].

The more you think about it, the more you realize that life is riddled with Iterated Prisoner's Dilemma games, not just human life but animal and plant life too. Plant life? Yes, why not? Remember that we are not talking about conscious strategies (though at times we might be), but about strategies in the 'Maynard Smithian' sense, strategies of the kind that genes might preprogram. Later we shall meet plants, various animals and even bacteria, all playing the game of Iterated Prisoner's Dilemma. [208]

Tit for Tat turned out to be a robust strategy. But the set of strategies that people happen to have submitted is an arbitrary set. This was the point that worried us above. It just so happened that in Axelrod's original tournament about half the entries were nice. Tit for Tat won in this climate, and Tit for Two Tats would have won in this climate if it had been submitted. But suppose that nearly all the entries had just happened to be nasty. This could very easily have occurred. After all, 6 out of the 14 strategies submitted were nasty. If 13 of them had been nasty, Tit for Tat wouldn't have won. The 'climate' would have been wrong for it. [215]

Suppose we start with a population already setting on the Always Defect side. The few Tit for Tat individuals don't meet each other often enough to be of mutual benefit. So natural selection pushes the population even further towards the Always Defect extreme. If only the population could just manage, by random drift, to get itself over the knife-edge, it could coast down the slope to the Tit for Tat side, and everyone would do much better at the banker's (or 'nature's') expense. But of course populations have no group will, no group intention or purpose. They cannot strive to leap the knife-edge. They will cross it only if the undirected forces of nature happen to lead them across.
How could this happen? One way to express the answer is that it might happen by 'chance'. But 'chance' is just a word expressing ignorance. It means 'determined by some as yet unknown, or unspecified, means'. We can do a little better than 'chance'. We can try to think of practical ways in which a minority of Tit for Tat individuals might happen to increase to the critical mass. This amounts to a quest for possible ways in which Tit for Tat individuals might happen to cluster together in sufficient numbers that they can all benefit at the banker's expense.
This line of thought seems to be promising, but it is rather vague. How exactly might mutually resembling individuals find themselves clustered together, in local aggregations? In nature, the obvious way is through genetic relatedness - kinship. Animals of most species are likely to find themselves living close to their sisters, brothers and cousins, rather than to random members of the population. This is not necessarily through choice. It follows automatically from 'viscosity' in the population. Viscosity means any tendency for individuals to continue living close to the place where they were born. [219]

Coming back to our knife-edge, then, Tit for Tat could surmount it. All that is required is a little local clustering, of a sort that will naturally tend to arise in natural populations. Tit for Tat has a built-in gift, even when rare, for crossing the knife-edge over to its own side. It is as though there were a secret passage underneath the knife-edge. But that secret passage contains a one-way valve: there is an assymetry. Unlike Tit for Tat, Always Defect, though a true ESS, cannot use local clustering to cross the knife-edge. On the contrary. Local clusters of Always Defect individuals, far from prospering by each other's presence, do especially badly in each other's presence. [219]
[But ALWAYS COOPERATEs don't!]
[Tit for Tat>Always Defect>Always Cooperate>Tit for Tat]

But we shall now see that the phenotypic effects of a gene need to be thought of as all the effects that it has on the world. [238]