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“Although a version of Hamilton's rule is indeed a fully general evolutionary truth, nothing about individual maximization can be deduced directly from this generalized form of the rule, contrary to what is sometimes implied. In particular, although the generalized Hamilton's rule yields the rb > c condition for the social trait to spread, one cannot conclude that that individuals behave as if trying to maximize the quantity (rb - c); they cannot do this, as this quantity is frequency-dependent.” — Samir Okasha

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Although a version of Hamilton's rule is indeed a fully general evolutionary truth, nothing about individual maximization can be deduced directly from this generalized form of the rule, contrary to what is sometimes implied. In particular, although the generalized Hamilton's rule yields the rb > c condition for the social trait to spread, one cannot conclude that that individuals behave as if trying to maximize the quantity (rb - c); they cannot do this, as this quantity is frequency-dependent.
— Samir Okasha