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The first fully explicit argument is given that broadly supports a widespread belief among whole-organism biologists that natural selection tends to lead to organisms acting as if maximizing their inclusive fitness. The use of optimization programs permits a clear statement of what this belief should be understood to mean, in contradistinction to the common(More)
Adaptation is conventionally regarded as occurring at the level of the individual organism. However, in recent years there has been a revival of interest in the possibility for group adaptations and superorganisms. Here, we provide the first formal theory of group adaptation. In particular: (1) we clarify the distinction between group selection and group(More)
  • A Grafen
  • 2007
For 8 years I have been pursuing in print an ambitious and at times highly technical programme of work, the 'Formal Darwinism Project', whose essence is to underpin and formalize the fitness optimization ideas used by behavioural ecologists, using a new kind of argument linking the mathematics of motion and the mathematics of optimization. The value of the(More)
  • A Grafen
  • 2007
A recent model studies the evolution of cooperation on a network, and concludes with a result connecting the benefits and costs of interactions and the number of neighbours. Here, an inclusive fitness analysis is conducted of the only case solved analytically, of a cycle, and the identical result is obtained. This brings the result within a biologically(More)
The Darwin unification project is pursued. A meta-model encompassing an important class of population genetic models is formed by adding an abstract model of the number of successful gametes to the Price equation under uncertainty. A class of optimization programs are defined to represent the "individual-as-maximizing-agent analogy" in a general way. It is(More)
A recent model shows that altruism can evolve with limited migration and variable group sizes, and the authors claim that kin selection cannot provide a sufficient explanation of their results. It is demonstrated, using a recent reformulation of Hamilton's original arguments, that the model falls squarely within the scope of inclusive fitness theory, which(More)
Hamilton's rule explains when natural selection will favor altruism between conspecifics, given their degree of relatedness. In practice, indicators of relatedness (such as scent) coevolve with strategies based on these indicators, a fact not included in previous theories of kin recognition. Using a combination of simulation modeling and mathematical(More)