The dynamics of a quantitative trait in an age-structured population living in a variable environment

Time series of rapid phenotypic change have been documented in age-structured populations living in the wild. Researchers are often interested in identifying the processes responsible for such change. We derive an equation to exactly decompose change in the mean value of a phenotypic trait into contributions from fluctuations in the demographic structure and age-specific viability selection, fertility selection, phenotypic plasticity, and differences between offspring and parental trait values. We treat fitness as a sum of its components rather than as a scalar and explicitly consider age structure by focusing on short time steps, which are appropriate for describing phenotypic change in species with overlapping generations. We apply the method to examine stasis in birth weight in a well-characterized population of red deer. Stasis is achieved because positive viability selection for an increase in birth weight is countered by parents producing offspring that are, on average, smaller than they were at birth. This is one of many ways in which equilibria in the mean value of a phenotypic trait can be maintained. The age-structured Price equation we derive has the potential to provide considerable insight into the processes generating now frequently reported cases of rapid phenotypic change.