Abstract

SummaryThe rice XA21‐mediated immune response is activated on recognition of the RaxX peptide produced by the bacterium Xanthomonas oryzae pv. oryzae (Xoo). The 60‐residue RaxX precursor is post‐translationally modified to form a sulfated tyrosine peptide that shares sequence and functional similarity with the plant sulfated tyrosine (PSY) peptide hormones. The 5‐kb raxX‐raxSTAB gene cluster of Xoo encodes RaxX, the RaxST tyrosylprotein sulfotransferase, and the RaxA and RaxB components of a predicted type I secretion system. To assess raxX‐raxSTAB gene cluster evolution and to determine its phylogenetic distribution, we first identified rax gene homologues in other genomes. We detected the complete raxX‐raxSTAB gene cluster only in Xanthomonas spp., in five distinct lineages in addition to X. oryzae. The phylogenetic distribution of the raxX‐raxSTAB gene cluster is consistent with the occurrence of multiple lateral (horizontal) gene transfer events during Xanthomonas speciation. RaxX natural variants contain a restricted set of missense substitutions, as expected if selection acts to maintain peptide hormone‐like function. Indeed, eight RaxX variants tested all failed to activate the XA21‐mediated immune response, yet retained peptide hormone activity. Together, these observations support the hypothesis that the XA21 receptor evolved specifically to recognize Xoo RaxX.

Highlights

  • Host receptors activate innate immunity pathways upon pathogen recognition (Ronald & Beutler, 2010)

  • These results suggest that RaxX recognition by XA21 is restrained by different sequence and length requirements compared to its recognition by the root growth promoting receptor(s) for plant sulfated tyrosine (PSY) hormone(s)

  • The raxX-raxSTAB gene cluster is present in a subset of Xanthomonas spp

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Summary

Introduction

Host receptors activate innate immunity pathways upon pathogen recognition (Ronald & Beutler, 2010). Oryzae (Xoo) (Wang et al, 1996). Four Xoo genes that are required for activation of XA21-mediated immunity, are located in the raxX-raxSTAB gene cluster (Fig. 1). The 60-residue RaxX predicted precursor protein undergoes sulfation by the RaxST tyrosylprotein sulfotransferase at residue Tyr-41 (Pruitt et al, 2015). Located outside the raxX-raxSTAB gene cluster, the raxC gene, an ortholog of the tolC gene, encodes the predicted outer membrane channel for this secretion complex (da Silva et al, 2004). The raxPQ genes encode enzymes to assimilate sulfate into 3'-phosphoadenosine 5'-phosphosulfate (PAPS) (Shen et al, 2002), the sulfodonor for the RaxST sulfotransferase (Han et al, 2012)

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