The Price of the Induced Defense Against Pests: A Meta-Analysis.

Plants and phytophagous arthropods have co-evolved for millions of years. During this long coexistence, plants have developed defense mechanisms including constitutive and inducible defenses. In an effort to survive upon herbivore attack, plants suffer a resource reallocation to facilitate the prioritization of defense toward growth. These rearrangements usually end up with a penalty in plant growth, development or reproduction directly linked to crop losses. Achieving the balance to maximize crop yield requires a fine tune regulation specific for each host-arthropod combination, which remains to be fully elucidated. The purpose of this work is to evaluate the effects of induced plant defenses produced upon pest feeding on plant fitness and surrogate parameters. The majority of the studies are focused on specific plant-pest interactions based on artificial herbivory damage or simulated defoliation on specific plant hosts. In this meta-analysis, the relevance of the variables mediating plant-pest interactions has been studied. The importance of plant and pest species, the infestation conditions (plant age, length/magnitude of infestation) and the parameters measured to estimate fitness (carbohydrate content, growth, photosynthesis and reproduction) in the final cost have been analyzed through a meta-analysis of 209 effects sizes from 46 different studies. Herbivore infestation reduced growth, photosynthesis and reproduction but not carbohydrate content. When focusing on the analyses of the variables modulating plant-pest interactions, new conclusions arise. Differences on the effect on plant growth and photosynthesis were observed among different feeding guilds or plant hosts, suggesting that these variables are key players in the final effects. Regarding the ontogenetic stage of a plant, negative effects were reported only in infestations during the vegetative stage of the plant, while no effect was observed during the reproductive stage. In addition, a direct relation was found between the durability and magnitude of the infestation, and the final negative effect on plant fitness. Among the parameters used to estimate the cost, growth and photosynthesis revealed more differences among subgroups than reproduction parameters. Altogether, this information on defense-growth trade-offs should be of great help for the scientific community to design pest management strategies reducing costs.