Constitutive and Induced Defenses in Long-lived Pines Do Not Trade Off but Are Influenced by Climate

Plants resist herbivores and pathogens by using constitutive (baseline) and inducible (change in defense after an attack) defenses. Inducibility has long been predicted to trade off with constitutive defense, reflecting the economic use of resources. However, empirical evidence for such tradeoffs is variable, and we still lack understanding about when and where defense trade-offs occur. We tested for tradeoffs between constitutive and induced defenses in natural populations of three species of long-lived pines (Pinus balfouriana, P. flexilis, P. longaeva) that differ greatly in constitutive defense and resistance to mountain pine beetle (MPB, Dendroctonus ponderosae). We also assessed how climate influenced constitutive and inducible defenses. At seven high-elevation sites in the western U.S., we simulated MPB attack to induce defenses and measured concentrations of terpene-based phloem defenses on days 0, 15, and 30. Constitutive and induced defenses did not trade off among or within species. Simulated MPB attack induced large increases in defense concentrations in all species independent of constitutive levels. MPB and its symbiotic fungi typically kill trees and thus could be selective forces maintaining strong inducibility within and among species. The contrasting constitutive concentrations in these species could be driven by the adaptation for specializing in harsh, high-elevation environments (e.g., P. balfouriana and P. longaeva) or by competition (e.g., P. flexilis), though these hypotheses have not been empirically examined. Climate influenced defenses, with the greatest concentrations of constitutive and induced defenses occurring at the coldest and driest sites. The interactions between climate and defenses have implications for these species under climate change.