Optimal Control and Cold War Dynamics between Plant and Herbivore

Herbivores eat the leaves that a plant needs for photosynthesis. However, the degree of antagonism between plant and herbivore may depend critically on the timing of their interactions and the intrinsic value of a leaf. We present a model that investigates whether and when the timing of plant defense and herbivore feeding activity can be optimized by evolution so that their interactions can move from antagonistic to neutral. We assume that temporal changes in environmental conditions will affect intrinsic leaf value, measured as potential carbon gain. Using optimal-control theory, we model herbivore evolution, first in response to fixed plant strategies and then under coevolutionary dynamics in which the plant also evolves in response to the herbivore. In the latter case, we solve for the evolutionarily stable strategies of plant defense induction and herbivore hatching rate under different ecological conditions. Our results suggest that the optimal strategies for both plant and herbivore are to avoid direct conflict. As long as the plant has the capability for moderately lethal defense, the herbivore will modify its hatching rate to avoid plant defenses, and the plant will never have to use them. Insights from this model offer a possible solution to the paradox of sublethal defenses and provide a mechanism for stable plant-herbivore interactions without the need for natural enenmy control.