Inclusive Fitness

inclusive fitness, theory in evolutionary biology in which an organism’s genetic success is believed to be derived from cooperation and altruistic behaviour. Inclusive fitness theory suggests that altruism among organisms who share a given percentage of genes enables those genes to be passed on to subsequent generations. In this way, an altruistic act that supports the survival of a relative or other individual theoretically enhances the genetic fitness of both the recipient of the act and the altruistic organism. The propagation of shared genes was believed to be an underlying mechanism for the evolution of eusociality (cooperative behaviour characterized by division of labour and group integration that is found in certain species of animals, mainly social insects).

The idea of inclusive fitness was first proposed in 1932 by British geneticist J.B.S. Haldane in The Causes of Evolution. The theory was later named and developed by British evolutionary biologist William Donald Hamilton, who used inclusive fitness to explain direct (reproductive) and indirect (aided by a relative or a colony member) inheritance of genetic traits associated with altruism. Hamilton presented his inclusive fitness theory in 1963; the following year British evolutionary biologist John Maynard Smith coined the term kin selection to describe Hamilton’s theory. Inclusive fitness later came to be understood as forming a general basis for kin selection theory, which attempts to interpret altruistic social behaviour in animals through genetic relatedness and benefits and costs associated with altruistic acts. Thus, in contrast to inclusive fitness, which considers genetic traits in both related and unrelated individuals, kin selection is concerned only with relatives. Hamilton’s inclusive fitness theory, as well as kin selection, seemed to many biologists to reconcile the conflict between natural selection, in which “selfish” genes perpetuate their own fitness through survival of the fittest, and selfless behaviour, in which eusocial genes shared by relatives and colony members influence cooperative behaviours that encourage the propagation of those genes.

Inclusive fitness theory is most commonly applied to eusocial organisms, such as bees and ants, although it has also been invoked to explain cooperative breeding in animals such as birds and the adoption of orphaned young by asocial red squirrels (Tamiasciurus hudsonicus). In certain bird species, such as the Florida scrub jay (Aphelocoma coerulescens) and the groove-billed ani (Crotophaga sulcirostris), some individuals will stay near nesting sites and participate in the rearing of related offspring. Individuals that do not disperse to their own territories have been thought to perceive the inclusive fitness gains of cooperative breeding as being greater than fitness gains offered by dispersal to potentially less-favourable territory. In such instances, inclusive fitness through cooperative breeding is the result of constraints on territory quality and is influenced by factors such as food, mate attraction, and predation. Indeed, in the absence of constraints, staying near relatives is less advantageous, potentially limiting breeding opportunities and thereby making kin selection and inclusive fitness less beneficial to reproductive success. The amount of labour that cooperative breeding individuals contribute to raising relatives is variable. In contrast, eusocial organisms have fixed and stereotyped divisions of labour; castes such as sterile workers presumably accumulate reproductive advantages by helping their relatives in the cooperative raising of young.

Although some researchers still contend that inclusive fitness can be used to describe the evolution of eusociality, the theory’s empirical assumptions and relevance to only very specialized social structures have led others to challenge its validity. American biologists Edward O. Wilson, Martin A. Nowak, and Corina E. Tarnita have provided mathematical explanations for eusociality based on population genetics and natural selection. By analyzing hypothetical populations of organisms in different evolutionary scenarios, the researchers determined that competition between selection for a eusocial allele (one of a pair of genes) and selection for a solitary allele was determined by basic principles guiding natural selection rather than by selection factors that extend beyond standard fitness calculations. The researchers further concluded that genetic relatedness is a consequence of cooperation and eusociality, not a driving force behind the evolution of these characteristics.