When does it pay off to prime for defense? A modeling analysis.

Plants can prepare for future herbivore attack through a process called priming. Primed plants respond more strongly and/or faster to insect attack succeeding the priming event than nonprimed plants, while the energetic costs of priming are relatively low. To better understand the evolution of priming, we developed a simulation model, partly parameterized for Brassica nigra plants, to explore how the fitness benefits of priming change when plants are grown in different biotic environments. Model simulations showed that herbivore dynamics (arrival probability, arrival time, and feeding rate) affect the optimal duration, the optimal investment and the fitness benefits of priming. Competition for light increases the indirect costs of priming, but may also result in a larger payoff when the nonprimed plant experiences substantial leaf losses. This modeling approach identified some important knowledge gaps: herbivore arrival rates on individual plants are rarely reported but they shape the optimal duration of priming, and it would pay off if the likelihood, severity and timing of the attack could be discerned from the priming cue, but it is unknown if plants can do so. In addition, the model generated some testable predictions, for example that the sensitivity to the priming cue decreases with plant age.