In addition to weak altruism, several other processes that increase the personal reproduction of the actor (direct
Most recently, explanations for altruistic-like behaviors have focused upon a somewhat sinister mechanism: enforcement. This idea can be traced not only to the 1970s and Richard Alexander who proposed parental manipulation to explain insect workers (Figure 3d), but also to Darwin, whose writings suggest something similar. While policing and punishment can explain apparent acts of altruism, however, one still needs an explanation for how policing, which carries a personal cost, can evolve the so-called 'second-order problem'. For this, one must appeal again to some or all of the above theories: inclusive fitness, group selection, and direct benefits.
If a helping behavior has arisen completely through enforcement, the primary evolutionary adaptation is in the enforcer, rather than the helping individual. The helping behavior, therefore, should probably not be considered an altruistic adaptation. This objection can be overturned, however, when an altruistic action evolves through a combination of enforcement and inclusive-fitness effects, as occurs in the social insects (below).
Enforcement, punishment, and policing are central to cementing the altruism in many social groups. This includes queen and worker policing in many species ofsocial insects, whereby the queen and workers suppress the reproduction of other workers. The suppression means that natural selection favors workers that invest more in the indirect fitness from helping than direct fitness from their own reproduction, which increases altruistic self-restraint (Figure 4). In addition, dominant males in macaque societies police and punish noncooperative individuals, and dominance hierarchies help to resolve breeding conflicts in many insect and vertebrate groups.
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