Abstract
Endoplasmic reticulum (ER)-mediated protein secretion and quality control have been shown to play an important role in immune responses in both animals and plants. In mammals, the ER membrane-located IRE1 kinase/endoribonuclease, a key regulator of unfolded protein response (UPR), is required for plasma cell development to accommodate massive secretion of immunoglobulins. Plant cells can secrete the so-called pathogenesis-related (PR) proteins with antimicrobial activities upon pathogen challenge. However, whether IRE1 plays any role in plant immunity is not known. Arabidopsis thaliana has two copies of IRE1, IRE1a and IRE1b. Here, we show that both IRE1a and IRE1b are transcriptionally induced during chemically-induced ER stress, bacterial pathogen infection and treatment with the immune signal salicylic acid (SA). However, we found that IRE1a plays a predominant role in the secretion of PR proteins upon SA treatment. Consequently, the ire1a mutant plants show enhanced susceptibility to a bacterial pathogen and are deficient in establishing systemic acquired resistance (SAR), whereas ire1b is unaffected in these responses. We further demonstrate that the immune deficiency in ire1a is due to a defect in SA- and pathogen-triggered, IRE1-mediated cytoplasmic splicing of the bZIP60 mRNA, which encodes a transcription factor involved in the expression of UPR-responsive genes. Consistently, IRE1a is preferentially required for bZIP60 splicing upon pathogen infection, while IRE1b plays a major role in bZIP60 processing upon Tunicamycin (Tm)-induced stress. We also show that SA-dependent induction of UPR-responsive genes is altered in the bzip60 mutant resulting in a moderate susceptibility to a bacterial pathogen. These results indicate that the IRE1/bZIP60 branch of UPR is a part of the plant response to pathogens for which the two Arabidopsis IRE1 isoforms play only partially overlapping roles and that IRE1 has both bZIP60-dependent and bZIP60-independent functions in plant immunity.
Highlights
Plants and their pathogens are engaged in a constant, coevolutionary battle for dominance
An experiment conducted in the ire1 mutants showed that IRE1a and IRE1b are induced independently as the IRE1a expression in ire1b-4 and the IRE1b transcript in ire1a-2, ire1a-3 and ire1a-4 mutants are comparable to Col-0
Given that salicylic acid (SA) can induce unfolded protein response (UPR) and IRE1 is required for efficient pathogenesis-related 1 (PR1) secretion as well as mounting effective systemic acquired resistance (SAR), we further investigated whether SA is a signal capable of activating the IRE1/bZIP60 amplicon smaller in size (bZIP60s) signaling pathway
Summary
Plants and their pathogens are engaged in a constant, coevolutionary battle for dominance. Plants lack mobile phagocytic cells or somatic adaptive immune systems. They have evolved highly sophisticated innate immune systems to initiate effective defense responses [1,2]. Plants trigger a robust disease resistance at the site of infection [3]. Stimulation of defense responses occurs locally and in distal areas of the plant where the state of resistance is heightened, a phenomenon known as systemic acquired resistance (SAR) [4]. Plants lacking functional NPR1 are impaired in their abilities to express PR genes and are almost completely defective in mounting SAR in response to pathogen infection [5,6]
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