Investigating the initial signalling mechanisms underpinning gene-for-gene mediated Systemic Acquired Resistance in Arabidopsis thaliana

Abstract

Plants deploy two key active defensive strategies to combat microbial pathogens; (i) Recognition of Pathogen-Associated Molecular Patterns (PAMPs) by extracellular surface receptors leading to the activation of PAMP-Triggered Immunity (PTI); (ii) Recognition of pathogen effector activity, usually intracellularly, by host Resistance (R) proteins leading to Effector-Triggered Immunity (ETI). ETI is characterised by a rapid localised Hypersensitive Response (HR). HR induces Systemic Acquired Resistance (SAR) through the production of an inducible immune signal(s), leading to broad spectrum systemic resistance. I investigated the earliest events associated with SAR signalling using plant electrophysiology, SAR mutants and a unique promoter-luciferase fusion that captures early systemic transcriptional events associated with ETI. We describe the transcriptional dynamics of A70 (At5g56980), a gene of unknown function (Truman et al. 2007), in local and systemic tissue following challenge with different elicitors and virulent or avirulent pathogen challenges. We provide evidence that A70 responds to a jasmonate (JA) related signal that is rapidly generated following ETI recognition. We further evaluate A70::LUC reporter activity in response to JA stimulus and correlate activity with histological expression of a JA repressor reporter (JAZ10::GUS) and A70::GFP reporter in systemically responding leaves following avirulent pathogen challenges. Finally, we examine changes in electrophysiological signals following ETI in local and systemic leaves. Focussing on events underpinning initiation, propagation and perception of SAR-inducing signals within the first 6-8 h of pathogen challenge we provide new insight into the integrated signalling mechanisms, dynamics and connectivity underpinning systemic immune responses. We conclude that there are multicomponent signals that link ETI induced transcriptional and electrical signals, with a COI1 receptor dependent propagative transcriptional wave the leads to rapid temporal spatial activation of jasmonate responsive genes in systemic responding leaves.