First, altruism must be defined so we can see how it defies the conventional theory of natural selection. Perhaps more importantly we must then ask why, when inexplicable using Darwin 's original model for selection, we still frequently observe altruism in nature. Three answers to this question have been put forward: kin selection, group selection and reciprocation. These will be examined in order.
When an actor 's behaviour increases the fitness of the behaviour 's receiver at the expense of the actor, then the behaviour is said to be altruistic. This definition is simplistic but provides a good starting point; throughout the discussion it will be sharpened. In contrast to so-called selfish behaviour increases the actor 's fitness while reducing the fitness of the receiver. If both the actor and the receiver gain in fitness following a behaviour, then the behaviour is cooperative. We must keep in mind that these costs and benefits are measures of reproductive success, the number of offspring surviving to reproductive age and thus the number of genes passed to the next generation.
It is worth mentioning that much altruistic behaviour may directly benefit those performing the behaviour in a way that may not be immediately obvious to the observer.
References: Dawkins. 1989. The selfish gene. 2nd Ed, OUP. Dawkins. 1995. Unravelling animal behaviour. Trivers. 1985. Social evolution. Benjamin/Cummings. Hamilton. 1964. The genetical evolution of social behaviour. J. Theor. Biol. 7, 1-52. Maynard-Smith. 1964. Group selection and kin selection. Nature, 201, 1145-1147. Duffy. 1996. Eusociality in a coral reef shrimp. Nature, 381, 512-514. Axelrod and Hamilton. 1981. The evolution of cooperation. Science, 211, 1390-1396.