We analyze weak fluctuating selection on a quantitative character in an age-structured population not subject to density regulation.We assume that early in the first year of life before selection, during a critical state of development, environments exert a plastic effect on the phenotype, which remains constant throughout the life of an individual. Age-specific selection on the character affects survival and fecundity, which have intermediate optima subject to temporal environmental fluctuations with directional selection in some age classes as special cases. Weighting individuals by their reproductive value, as suggested by Fisher, we show that the expected response per year in the weighted mean character has the same form as for models with no age structure. Environmental stochasticity generates stochastic fluctuations in the weighted mean character following a first-order autoregressive model with a temporally autocorrelated noise term and stationary variance depending on the amount of phenotypic plasticity. The parameters of the process are simple weighted averages of parameters used to describe age-specific survival and fecundity. The "age-specific selective weights" are related to the stable distribution of reproductive values among age classes. This allows partitioning of the change in the weighted mean character into age-specific components.
K E Y W O R D S :Age structure, evolutionary dynamics, fluctuating selection, life history, plasticity, reproductive value.During the past few decades a large number of empirical estimates of phenotypic selection have appeared, facilitated by Lande and Arnold's (1983) realization that multiple regression of individual fitness (or a component of fitness) on individual characters provides estimates of selection gradients on quantitative characters. This approach has produced considerable general insight into operation of directional and stabilizing selection (Endler 1986;Kingsolver et al. 2001; Siepielski et al. 2009). In combination with estimates of additive genetic variances and covariances based, for example, on the animal model (Kruuk 2004), a mechanistic understanding of factors causing variation in the rate of evolutionary response to selection in fluctuating environments has now started to emerge (Ellegren and Sheldon 2008;Kruuk et al. 2008; Siepielski et al. 2009; Walsh and Blows 2009). However, we still have little knowledge about how variable environments affect selection pressures and rates of evolutionary responses to environmental changes.The method of Lande and Arnold (1983) is still based on several simplifying assumptions, including no age-structure. The step from an evolutionary theory for nonoverlapping generations to populations with age or stage structure is a large one, introducing heterogeneity among individuals of different ages, temporal interactions between age-specific survival and fecundity, and transient fluctuations in age structure (Caswell 2001). Based on earlier ideas of Hamilton (1966), Medawar (1946, 1952), and Fisher (1930, 1958, de...