thumb|upright=1.3|The co-operative behaviour of [[social insects like the honey bee can be explained by kin selection.]]
Kin selection is a process whereby natural selection favours a trait due to its positive effects on the reproductive success of an organism's relatives, even when at a cost to the organism's own survival and reproduction. Kin selection can lead to the evolution of altruistic behaviour. It is related to inclusive fitness, which combines the number of offspring produced with the number an individual can ensure the production of by supporting others (weighted by the relatedness between individuals). A broader definition of kin selection includes selection acting on interactions between individuals who share a gene of interest even if the gene is not shared due to common ancestry. Hamilton proposed two mechanisms for kin selection. First, kin recognition allows individuals to be able to identify their relatives. Second, in viscous populations, populations in which the movement of organisms from their place of birth is relatively slow, local interactions tend to be among relatives by default. The viscous population mechanism makes kin selection and social cooperation possible in the absence of kin recognition. In this case, nurture kinship, the interaction between related individuals, simply as a result of living in each other's proximity, is sufficient for kin selection, given reasonable assumptions about population dispersal rates. Kin selection is not the same thing as group selection, where natural selection is believed to act on the group as a whole.
In humans, altruism is both more likely and on a larger scale with kin than with unrelated individuals; for example, humans give presents according to how closely related they are to the recipient. In other species, vervet monkeys use allomothering, where related females such as older sisters or grandmothers often care for young, according to their relatedness. The social shrimp Synalpheus regalis protects juveniles within highly related colonies.
Historical overview
thumb|upright|[[Charles Darwin wrote that selection could be applied to the family as well as to the individual.
In this passage "the family" and "stock" stand for a kin group. These passages and others by Darwin about kin selection are highlighted in D.J. Futuyma's textbook of reference Evolutionary Biology and in E. O. Wilson's Sociobiology.
Kin selection was briefly referred to by R.A. Fisher in 1930 and J.B.S. Haldane in 1932 and 1955. J.B.S. Haldane grasped the basic quantities in kin selection, famously writing "I would lay down my life for two brothers or eight cousins". Haldane's remark alluded to the fact that if an individual loses its life to save two siblings, four nephews, or eight cousins, it is a "fair deal" in evolutionary terms, as siblings are on average 50% identical by descent, nephews 25%, and cousins 12.5% (in a diploid population that is randomly mating and previously outbred). But Haldane also joked that he would truly die only to save more than a single identical twin of his or more than two full siblings. In 1955 he clarified:
W. D. Hamilton, in 1963 and especially in 1964
thumb|upright|The evolutionary biologist [[John Maynard Smith used the term "kin selection" in 1964.]]
John Maynard Smith may have coined the actual term "kin selection" in 1964:
Kin selection causes changes in gene frequency across generations, driven by interactions between related individuals. This dynamic forms the conceptual basis of the theory of sociobiology. Some cases of evolution by natural selection can only be understood by considering how biological relatives influence each other's fitness. Under natural selection, a gene encoding a trait that enhances the fitness of each individual carrying it should increase in frequency within the population; and conversely, a gene that lowers the individual fitness of its carriers should be eliminated. However, a hypothetical gene that prompts behaviour which enhances the fitness of relatives but lowers that of the individual displaying the behaviour, may nonetheless increase in frequency, because relatives often carry the same gene. According to this principle, the enhanced fitness of relatives can at times more than compensate for the fitness loss incurred by the individuals displaying the behaviour, making kin selection possible. This is a special case of a more general model, "inclusive fitness". This analysis has been challenged, Wilson writing that "the foundations of the general theory of inclusive fitness based on the theory of kin selection have crumbled" and that he now relies instead on the theory of eusociality and "gene-culture co-evolution" for the underlying mechanics of sociobiology. Inclusive fitness theory is still generally accepted however, as demonstrated by the publication of a rebuttal to Wilson's claims in Nature from over a hundred researchers.
Hamilton's rule
Formally, genes for a particular behavior should increase in frequency when
:<math>rB > C</math>
where
:r = the genetic relatedness of the recipient to the actor, often defined as the probability that a gene picked randomly from each at the same locus is identical by descent.
:B = the additional reproductive benefit gained by the recipient of the altruistic act,
:C = the reproductive cost to the individual performing the act.
This inequality is known as Hamilton's rule after W. D. Hamilton who in 1964 published the first formal quantitative treatment of kin selection. Modern formulations of the rule use Alan Grafen's definition of relatedness based on the theory of linear regression.
A 2014 review of many lines of evidence for Hamilton's rule found that its predictions were confirmed in a wide variety of social behaviours across a broad phylogenetic range of birds, mammals and insects, in each case comparing social and non-social taxa. Among the experimental findings, a 2010 study used a wild population of red squirrels in Yukon, Canada. Surrogate mothers adopted related orphaned squirrel pups but not unrelated orphans. The cost of adoption was calculated by measuring a decrease in the survival probability of the entire litter after increasing the litter by one pup, while benefit was measured as the increased chance of survival of the orphan. The degree of relatedness of the orphan and surrogate mother for adoption to occur depended on the number of pups the surrogate mother already had in her nest, as this affected the cost of adoption. Females always adopted orphans when rB was greater than C, but never adopted when rB was less than C, supporting Hamilton's rule.
Mechanisms
Altruism occurs where the instigating individual suffers a fitness loss while the receiving individual experiences a fitness gain. The sacrifice of one individual to help another is an example.
Hamilton outlined two ways in which kin selection altruism could be favoured: