Condition-dependence resolves the paradox of missing plasticity costs

Phenotypic plasticity plays a key role in adaptation to changing environments. However, plasticity is neither perfect nor ubiquitous, implying that fitness costs must limit the evolution of phenotypic plasticity in nature. The measurement of such costs of plasticity has proved elusive; decades of experiments show that fitness costs of plasticity are often weak or nonexistent. Here, we show that this paradox can be at least partially explained by condition-dependence. We develop two models differing in their assumptions about how condition-dependence arises; both models show that variation in condition can readily mask costs of plasticity even when such costs are substantial. This can be shown simply in a model where costly plasticity itself evolves condition-dependence. Yet similar effects emerge from an alternative model where trait expression is condition-dependent. In this more complex model, average condition in each environment and genetic covariance in condition across environments both determine when costs of plasticity can be revealed. Analogous to the paradox of missing trade-offs between life history traits, our models show that variation in condition masks costs of plasticity even when costs exist, and suggests this conclusion may be robust to the details of how condition affects trait expression. Our models demonstrate that condition dependence can also account for the often-observed pattern of elevated plasticity costs inferred in stressful environments, the maintenance of genetic variance in plasticity, and provides insight into experimental and biological scenarios ideal for revealing a cost of phenotypic plasticity.