Detecting small environmental differences: risk-response curves for predator-induced behavior and morphology.

Most organisms possess traits that are sensitive to changes in the environment (i.e., plastic traits) which results in the expression of environmentally induced polymorphisms. While most phenotypically plastic traits have traditionally been treated as threshold switches between induced and uninduced states, there is growing evidence that many traits can respond in a continuous fashion. In this experiment we exposed larval anurans (wood frog tadpoles, Rana sylvatica) to an increasing gradient of predation risk to determine how organisms respond to small environmental changes. We manipulated predation risk in two ways: by altering the amount of prey consumed by a constant number of predators (Dytiscus sp.) and by altering the number of predators that consume a constant amount of prey. We then quantified the expression of predator-induced behavior, morphology, and mass to determine the level of risk that induced each trait, the level of risk that induced the maximal phenotypic response for each trait, whether the different traits exhibited a plateauing response, and whether increasing risk via increasing predator number or via increasing prey consumption induced similar phenotypic changes. We found that all of the traits exhibited fine-tuned, graded responses and most of them exhibited a plateauing response with increased predation risk, suggesting either a limit to plasticity or the reflection of high costs of the defensive phenotype. For many traits, a large proportion of the maximum induction occurred at low levels of risk, suggesting that the chemical cues of predation are effective at extremely low concentrations. In contrast to earlier work, we found that behavioral and morphological responses to increased predator number were simply a response to increased total prey consumption. These results have important implications for models of plasticity evolution, models of optimal phenotypic design, expectations for how organisms respond to fine-grained changes (i.e., within generation) in their environment, and impacts on ecological communities via trait-mediated indirect effects.