Inclusive fitness
Inclusive fitness
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Inclusive fitness

Inclusive fitness is a conceptual framework in evolutionary biology first defined by W. D. Hamilton in 1964. It is primarily used to aid the understanding of how social traits are expected to evolve in structured populations. It involves partitioning an individual's expected fitness returns into two distinct components: direct fitness returns - the component of a focal individual’s fitness that is independent of who it interacts with socially; and indirect fitness returns - the component that is dependent on who it interacts with socially. The direct component of an individual's fitness is often called its personal fitness, while an individual’s direct and indirect fitness components taken together are often called its inclusive fitness.

Under an inclusive fitness framework, direct fitness returns are realised through the offspring a focal individual produces independent of who it interacts with, while indirect fitness returns are realised by adding up all the effects our focal individual has on the (number of) offspring produced by those it interacts with weighted by the relatedness of our focal individual to those it interacts with. This can be visualised in a sexually reproducing system (assuming identity by descent) by saying that an individual's own child, who carries one half of that individual's genes, represents one offspring equivalent. A sibling's child, who will carry one-quarter of the individual's genes, will then represent 1/2 offspring equivalent, and so on (see coefficient of relationship).

Neighbour-modulated fitness is the conceptual inverse of inclusive fitness. Where inclusive fitness calculates an individual’s indirect fitness component by summing the fitness that focal individual receives through modifying the productivities of those it interacts with (its neighbours), neighbour-modulated fitness instead calculates it by summing the effects an individual’s neighbours have on that focal individual’s productivity. When taken over an entire population, these two frameworks give functionally equivalent results. Hamilton’s rule is a particularly important result in the fields of evolutionary ecology and behavioral ecology that follows naturally from the partitioning of fitness into direct and indirect components, as given by inclusive and neighbour-modulated fitness, revealing how the average trait value of a population is expected to evolve under the assumption of small mutational steps.

Kin selection is a well known case whereby inclusive fitness effects can influence the evolution of social behaviours. Kin selection relies on positive relatedness (driven by identity by descent) to enable individuals who positively influence the fitness of those they interact with at a cost to their own personal fitness, to outcompete individuals employing more selfish strategies. It is thought to be one of the primary mechanisms underlying the evolution of altruistic behaviour, alongside the less prevalent reciprocity (see also reciprocal altruism), and to be of particular importance in enabling the evolution of eusociality among other forms of group living. Inclusive fitness has also been used to explain the existence of spiteful behaviour, where individuals negatively influence the fitness of those they interact with at a cost to their own personal fitness.

Inclusive fitness and neighbour-modulated fitness are both frameworks that leverage the individual as the unit of selection. It is from this that the gene-centered view of evolution emerged: a perspective that has facilitated much of the work done into the evolution of conflict (examples include parent-offspring conflict, interlocus sexual conflict, and intragenomic conflict).

The British evolutionary biologist W. D. Hamilton showed mathematically that, because other members of a population may share one's genes, a gene can also increase its evolutionary success by indirectly promoting the reproduction and survival of other individuals who also carry that gene. This is variously called "kin theory", "kin selection theory" or "inclusive fitness theory". The most obvious category of such individuals is close genetic relatives, and where these are concerned, the application of inclusive fitness theory is often more straightforwardly treated via the narrower kin selection theory. Hamilton's theory, alongside reciprocal altruism, is considered one of the two primary mechanisms for the evolution of social behaviors in natural species and a major contribution to the field of sociobiology, which holds that some behaviors can be dictated by genes, and therefore can be passed to future generations and may be selected for as the organism evolves.

Belding's ground squirrel provides an example; it gives an alarm call to warn its local group of the presence of a predator. By emitting the alarm, it gives its own location away, putting itself in more danger. In the process, however, the squirrel may protect its relatives within the local group (along with the rest of the group). Therefore, if the effect of the trait influencing the alarm call typically protects the other squirrels in the immediate area, it will lead to the passing on of more copies of the alarm call trait in the next generation than the squirrel could leave by reproducing on its own. In such a case natural selection will increase the trait that influences giving the alarm call, provided that a sufficient fraction of the shared genes include the gene(s) predisposing to the alarm call.

Synalpheus regalis, a eusocial shrimp, is an organism whose social traits meet the inclusive fitness criterion. The larger defenders protect the young juveniles in the colony from outsiders. By ensuring the young's survival, the genes will continue to be passed on to future generations.

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