The origin of altruism remains one of the most enduring puzzles of human behaviour. Indeed, true altruism is often thought either not to exist, or to arise merely as a miscalculation of otherwise selfish behaviour. In this paper, we argue that altruism emerges directly from the way in which distinct human decision-making systems learn about rewards. Using insights provided by neurobiological accounts of human decision-making, we suggest that reinforcement learning in game-theoretic social interactions (habitisation over either individuals or games) and observational learning (either imitative of inference based) lead to altruistic behaviour. This arises not only as a result of computational efficiency in the face of processing complexity, but as a direct consequence of optimal inference in the face of uncertainty. Critically, we argue that the fact that evolutionary pressure acts not over the object of learning (‘what’ is learned), but over the learning systems themselves (‘how’ things are learned), enables the evolution of altruism despite the direct threat posed by free-riders
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