Transcriptomic changes under combined drought and nonhost bacteria reveal novel and robust defenses in Arabidopsis thaliana

Plants in the natural conditions are often challenged by a combination of two or more stressors. A combination of drought and pathogen is one of the most pressing threats to the plant's growth and survival in the field, and thus warrants a mechanistic understanding. Susceptible plant-pathogen interaction, owing to effector-mediated suppression of plant defense responses, limits its scope for combined stress studies. In the present study, we have investigated the morpho-physiological responses of Arabidopsis thaliana to simultaneous drought and nonhost bacterial pathogen Pseudomonas syringae pv. tabaci. Combined stress treatment provoked an early and more pronounced hypersensitive response in the plant as compared to the non-host pathogen treatment. We have further deciphered the molecular basis for the robust defense response observed under combined stress by transcriptomic profiling carried out using whole-genome microarray. We found that the enhanced resistance to the combined stress is accompanied by a massive transcriptional reprogramming involving several transcripts specifically responding to the stress combination. A prominent over-representation of genes involved in basal defense-related machinery was observed under the combined stress. Genes involved in various defense signaling cascades, accumulation of secondary metabolites and those encoding for receptor-like kinases were highly up regulated under the combined stress. Up-regulated genes related to redox homeostasis and hypersensitive response (HR)-mediated cell death were also found to be markedly enriched under combined stress. We also compared the global gene expression profile of A. thaliana subjected to combined drought-nonhost bacteria to those treated with a combination of drought-host bacteria Pseudomonas syringae pv. tomato DC3000. A significant induction of genes responding to drought as well as bacteria was observed during both the interactions. However, the amplitude of induction was more pronounced under the combination of drought and nonhost bacteria. Our results also indicate that plant activates multiple defense pathways upon exposure to combined stress which strengthens the overall basal immunity of the plant, characterized by a stronger HR response.