INTERACTIONS BETWEEN ABSCISIC-ACID-MEDIATED RESPONSES AND PLANT RESISTANCE TO PATHOGENS AND INSECTS

One goal of phytohormonal ecology is to study the interactions between biotic and abiotic stress at hierarchical levels of biological organization. From an ecological perspective, exposure to one stress may alter the plant's probability of being exposed to another stress. From a mechanistic perspective, hormonal and biochemical signaling in- teractions between responses to each stress may influence the severity or ability to adaptively respond to the subsequent stress. In this article, we consider the relationship between plant water and salt stress and attack by pathogens and herbivores. Empirical data suggest that water stress and the probability of attack by pathogens and herbivores are correlated between habitats. Biochemical interactions between plant responses to water and salt stress and insect and pathogen attack are also interrelated. Initial biochemical models indicated that abscisic acid (ABA), an important hormone in responses to water and salt stress, had a synergistic positive role with jasmonate-induced defenses against herbivores and an an- tagonistic role with salicylate-based resistance to some pathogens. Based on this back- ground, we developed predictions about how water and salt stress would alter plant resis- tance to insects and pathogens and tested the predictions using tomato plants as a model system. We used polyphenol oxidase activity as a marker of the jasmonate response and pathogenesis-related protein P4 as a marker of the salicylate response. First, we examined levels of chemical defense in wild-type and ABA-deficient plants and the ability of these plants to resist insect and pathogen attack. In the second experiment, we exposed plants to short-term salinity stress and tested their subsequent resistance to a chewing insect Spodoptera exigua and the bacterial speck pathogen Pseudomonas syringae pv. tomato. We have two key findings. First, ABA-deficient plants had higher levels of salicylate-mediated responses and were more resistant to bacterial speck disease, consistent with the proposed role of salicylate in defense against pathogens. This suggests linkage between water avail- ability to the plant and salicylate action in pathogenesis through ABA signaling. ABA- deficient plants had reduced resistance to the insect Spodoptera exigua, suggesting a positive correlation between responses to water stress and herbivory. The lack of difference in chemical expression of the jasmonate (JA) response (polyphenol oxidase activity) between wild-type and ABA-deficient plants did not support the proposed mechanism of synergism with the jasmonate response. Second, salt stress reduced the chemical induction (e.g., pathogenesis-related protein P4) of the salicylate response, but this did not affect resistance to the pathogen. Salt stress did not alter resistance to the herbivore Trichoplusia ni, but did alter the negative signal interaction between the jasmonate and salicylate responses. Under control conditions, the jasmonate and salicylate responses are antagonistic to one another, with induction of one response reducing the inducibility of the other. Under salt stress conditions, the negative effect of salicylate on the jasmonate response was reduced. Thus, complex interactions occur between ABA, JA, and SA, hormones that are important regulators of abiotic and biotic stress responses. Phytohormonal ecology is attempting to link ecological and hormonal interactions to develop a predictive framework for how and why plants coordinate responses to the environment.