An early precursor to kin selection was the theory of inclusive fitness. This was proposed by J.B.S. Haldane in 1932 but developed and named by William Donald Hamilton in 1964. Hamilton’s theory is the basis of Richard Dawkins famous book, “The Selfish Gene” and argues that it is the survival and reproduction of genes, rather than organisms, that is the principal driver behind evolution. As a result, an organism can display altruism if this leads to a greater propagation of the genes it holds than would be the case if it acted solely out of personal self-interest. This relies on the individual organism being able to identify those genes in others. There are two main ways of doing so. Firstly, by knowing its kin or related family members and, secondly, by recognising external characteristics displayed by others with the relevant gene. However, there are several difficulties with the latter, for example whether the gene does in fact express itself in the form of recognisable traits and whether the organism can see those traits. Because such traits are often only skin deep, there is the potential for imposters to display them to benefit from altruistic behaviour.
The more specific theory of kin selection developed from Hamilton’s work. This theory states that an organism can behave in a way which maximises the propagation of its genes by behaving in an altruistic manner towards close relatives likely to hold the same genes.
Individuals in a species have approximately 99% of their genes in common. The remaining 1% constitutes their variable genome which accounts for physical variation within the species. The fitness of the 99% is well established and, thus, only genes in the variable genome, including any mutations, compete to propagate themselves. 50% of the variable genome is inherited from each parent. On average, therefore, an individual will share 50% with each parent, child, and sibling and, on average, 25% with each grandparent, uncle, aunt, nephew, niece, or grandchild. The theory of kin selection proposes, therefore, that it is advantageous in terms of the propagation of the variable genome to favour the survival and reproduction of three siblings over that of the self. Thus, genetically driven behaviour which facilitates this will propagate within the species.
Kin selection behaviour relies on the ability of an individual to recognise its kin. Nurture kinship, i.e., having raised, been raised by, or having been raised with another nuclear family member, is clearly an important factor, and can be observed in other species. However, the recognition of more remotely related kin, e.g., aunts, uncles, and other members of the extended family, requires considerable cognitive skill and, so, is probably limited to the more intelligent species.
As individuals become more remotely related, it only becomes possible to recognise kinship through physical appearance and, in the case of humans, cues such as language, dress, beliefs, etc. Thus, kin selection suggests that an individual is more likely to behave altruistically towards others of similar appearance and culture because these factors also suggest a similar variable genome.
Intuitively, kin selection operates within humanity. There is also a great deal of objective evidence for its presence. For example, research has shown that non-reciprocal help is far more likely to occur in kin relationships than non-kin relationships. It has also been shown that, when wills are written, there is a close correlation between kinship and the proportion of wealth passed on.
A small number of species can be described as eusocial. These species co-operatively rear their young across multiple generations. They also divide labour through the surrender, by some members, of all or part of their personal reproductive success to increase the reproductive success of others. In this way they benefit the overall reproductive success of the group. Eusociality arose late in the history of life and is extremely rare. Only nineteen species are known to display this characteristic: two species of mole rat, some species of brine shrimp, insects such as wasps, bees, and ants and, of course, mankind. In eusocial species, group level natural selection takes place due to competition between groups. In the case of the eusocial insects, the group is the nest or hive. Individual workers will lose their lives in the interest of the hive as a whole. It can be argued that this form of behaviour in insects is entirely altruistic and an inherited form of kin selection. However, in the case of humanity, this argument does not hold true because human groups display both kin altruism and non-kin co-operation.
However, there remain doubts whether individual and kin selection fully explain natural selection and human social behaviour since natural selection may also occur at higher biological levels. This will be explored further in subsequent posts.