Reversible plasticity in phenotypic traits allows organisms to cope with environmental variation within lifetimes, but costs of plasticity may limit just how well the phenotype matches the environmental optimum. An additional adaptive advantage of plasticity might be to reduce fitness variance, in other words: bet-hedging to maximize geometric (rather than simply arithmetic) mean fitness. Here, we model the evolution of plasticity in the form of reaction norm slopes, with increasing costs as the slope or degree of plasticity increases. We find that greater investment in plasticity (i.e. a steeper reaction norm slope) is favoured in scenarios promoting bet-hedging as a response to multiplicative fitness accumulation (i.e. coarser environmental grains and fewer time steps prior to reproduction), because plasticity lowers fitness variance across environmental conditions. In contrast, in scenarios with finer environmental grain and many time steps prior to reproduction, bet-hedging plays less of a role and individual-level optimization favours evolution of shallower reaction norm slopes. However, the opposite pattern holds if plasticity costs themselves result in increased fitness variation, as might be the case for production costs of plasticity that depend on how much change is made to the phenotype each time step. We discuss these contrasting predictions from this partitioning of adaptive plasticity into short-term individual benefits versus long-term genotypic (bet-hedging) benefits, and how this approach enhances our understanding of the evolution of optimum levels of plasticity in examples from thermal physiology to advances in avian lay dates.
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