This figure illustrates how the proposed optimality model that links phenotypes to vital rates, and captures verbal models of the DTA, fits into the hierarchy illustrated in Fig 1 and how it yields age-specific directional selection. The optimality functions (lower 2 plots in the center) are assumed to be identical at both ages. However, because selection emphasizes improvements to vital rates more in early life than at late life (upper 2 plots in the center), the sensitivity of fitness to changes in the early-age phenotypes (left) is greater than the fitness sensitivity to changes in the late-life phenotype (right). These outer functions define age-specific directional selection, which is the slope at the population-mean phenotype under our model assumptions. The blue circles represent these means, and the blue lines are the slopes at those points. Note that the same amounts of directional selection can exist at both ages, provided that the late-age phenotype is further removed from its optimum. DTA, developmental theory of aging.