Abstract
The advantage of mutual help is threatened by defectors, who exploit the benefits provided by others without providing benefits in return. Cooperation can only be sustained if it is preferentially channeled toward cooperators and away from defectors. But how? A deceptively simple idea is to distinguish cooperators from defectors by tagging them. It clearly is in the interest of cooperators to use some distinctive cue to assort with their like. Such an assortment, however, conflicts with the interests of the cheaters, who have every incentive to also acquire that tag. This makes for an inherently unstable situation. The history of evolutionary thinking on this issue is long. An article in this issue of PNAS by Antal et al. (1) opens new ground by providing an in-depth analysis of a selection-mutation model. The first to investigate a tag for altruism was W. D. Hamilton (2). He conceived what he called a supergene, able to produce (i) a distinctive phenotypic trait, (ii) the faculty to recognize the trait in others, and (iii) the propensity to direct benefits toward bearers of that trait, even though this entails a fitness cost. Soon afterward, Richard Dawkins described Hamilton's thought experiment by using as phenotypic trait the fanciful example of a green beard. The supergene was now termed “green beard gene,” in part to acknowledge its inherent unlikelihood. “Too good to be true,” were Dawkins' words (3): for the gene would have to be able to program for 3 effects, namely the feature, its recognition, and the altruistic propensity. The green-beard concept relates to both major approaches to cooperation in evolutionary biology, namely kin selection (2) and reciprocal altruism (4). It helps in promoting assortment between cooperators; as a result, cooperators can get more than they give, so that altruism becomes a thriving business. Because wearers …