An altruistic act is one in which an individual incurs a cost that results in a benefit to others. Giving money or time to those less fortunate than ourselves is one example, as is giving up one’s seat on a bus. At first, one might consider such behaviour hopelessly naive in a world in which natural selection seemingly rewards selfishness in the competitive struggle for existence. As the saying goes, ‘nice guys finish last’. Yet examples of apparent altruism are commonplace. Meerkats will spend hours in the baking sun keeping lookout for predators that might attack their colony mates. Vampire bats will regurgitate blood to feed their starving roost fellows, while baboons will take the time and effort to groom other baboons. Some individuals, such as honeybee workers, forego their own reproduction to help their queen and will even die in her defence. The common gut bacterium Escherichia coli commits suicide when it is infected by a bacteriophage, thereby protecting its clones from being infected. If helping incurs a cost, then surely an individual that accepts a cooperative act yet gives nothing in return would do better than cooperators? What, then, allows these cases of apparent altruism to persist? In his last presidential address to the Royal Society of London in November 2005, Robert M. May argued, ‘The most important unanswered question in evolutionary biology, and more generally in the social sciences, is how cooperative behaviour evolved and can be maintained’. In this chapter, we document a number of examples of cooperation in the natural world and ask how it is maintained despite the obvious evolutionary pressure to ‘cheat’. We will see that, while it is tempting to see societies as some form of higher organism, to fully understand cooperation, it helps to take a more reductionist view of the world, frequently a gene-centred perspective. Indeed, thinking about altruism has led to one of the greatest triumphs of the ‘selfish gene’ approach, namely the theory of kin selection. Ultimately, as the quote from Mandeville indicates, we will see that cooperation frequently arises simply out of pure self-interest—it just so happens that individuals (or, more precisely, genes) in the business of helping themselves sometimes help others.
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