Models of large-scale human cooperation take two forms. 'Indirect reciprocity' occurs when individuals help others in order to uphold a reputation and so be included in future cooperation. In 'collective action', individuals engage in costly behaviour that benefits the group as a whole. Although the evolution of indirect reciprocity is theoretically plausible, there is no consensus about how collective action evolves. Evidence suggests that punishing free riders can maintain cooperation, but why individuals should engage in costly punishment is unclear. Solutions to this 'second-order free rider problem' include meta-punishment, mutation, conformism, signalling and group-selection. The threat of exclusion from indirect reciprocity can sustain collective action in the laboratory. Here, we show that such exclusion is evolutionarily stable, providing an incentive to engage in costly cooperation, while avoiding the second-order free rider problem because punishers can withhold help from free riders without damaging their reputations. However, we also show that such a strategy cannot invade a population in which indirect reciprocity is not linked to collective action, thus leaving unexplained how collective action arises.

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Development is typically a constructive process, in which phenotypes incrementally adapt to local ecologies. Here, we present a novel model in which natural selection shapes developmental systems based on the evolutionary ecology, and these systems adaptively guide phenotypic development. We assume that phenotypic construction is incremental and trades off with sampling cues to the environmental state. We computed the optimal developmental programmes across a range of evolutionary ecological conditions. Using these programmes, we simulated distributions of mature phenotypes. Our results show that organisms sample the environment most extensively when cues are moderately, not highly, informative. When the developmental programme relies heavily on sampling, individuals transition from sampling to specialization at different times in ontogeny, depending on the consistency of their sampled cue set; this finding suggests that stochastic sampling may result in individual differences in plasticity itself. In addition, we find that different selection pressures may favour similar developmental mechanisms, and that organisms may incorrectly calibrate development despite stable ontogenetic environments. We hope our model will stimulate adaptationist research on the constructive processes guiding development.

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Sensitive periods, in which experience shapes phenotypic development to a larger extent than other periods, are widespread in nature. Despite a recent focus on neural-physiological explanation, few formal models have examined the evolutionary selection pressures that result in developmental mechanisms that produce sensitive periods. Here, we present such a model. We model development as a specialization process during which individuals incrementally adapt to local environmental conditions, while receiving a constant stream of cost-free, imperfect cues to the environmental state. We compute optimal developmental programmes across a range of ecological conditions and use these programmes to simulate developmental trajectories and obtain distributions of mature phenotypes. We highlight four main results. First, matching the empirical record, sensitive periods often result from experience or from a combination of age and experience, but rarely from age alone. Second, individual differences in sensitive periods emerge as a result of stochasticity in cues: individuals who obtain more consistent cue sets lose their plasticity at faster rates. Third, in some cases, experience shapes phenotypes only at a later life stage (lagged effects). Fourth, individuals might perseverate along developmental trajectories despite accumulating evidence suggesting the alternate trajectory is more likely to match the ecology.. . .we all begin with the natural equipment to live a thousand kinds of life but end in the end having lived only one.-Clifford Geertz, 1973