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  • A Grafen
  • 1990
An ESS model of Zahavi's handicap principle is constructed. This allows a formal exposition of how the handicap principle works, and shows that its essential elements are strategic. The handicap model is about signalling, and it is proved under fairly general conditions that if the handicap principle's conditions are met, then an evolutionarily stable(More)
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)
The formal Darwinism project aims to provide a mathematically rigorous basis for optimisation thinking in relation to natural selection. This paper deals with the situation in which individuals in a population belong to classes, such as sexes, or size and/or age classes. Fisher introduced the concept of reproductive value into biology to help analyse(More)
  • A Grafen
  • 1990
A population genetic model of sexual selection is constructed in which, at equilibrium, males signal their quality by developing costly ornaments, and females pay costs to use the ornaments in mate choice. It is shown that the form of the equilibrium is uninfluenced by the Fisher process, that is, by self-reinforcement of female preferences. This is a(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)
Genetic recombination has important consequences, including the familiar rules of Mendelian genetics. Here we present a new argument for the evolutionary function of recombination based on the hypothesis that meiotic drive systems continually arise to threaten the fairness of meiosis. These drive systems act at the expense of the fitness of the organism as(More)