Effectors, chaperones, and harpins of the Type III secretion system in the fire blight pathogen Erwinia amylovora: a review

Abstract

Erwinia amylovora is a plant pathogenic bacterium that causes the disease fire blight in a wide range of Rosaceous plants including apples and pears. This pathogen utilizes the hypersensitive response and pathogenicity (hrp)-type III secretion system (T3SS), an essential pathogenicity factor of E. amylovora, to deliver proteins from bacteria to plant apoplasts or the cytoplasm to regulate host immune responses and physiology. Proteins secreted by the E. amylovora T3SS include five effectors: DspA/E, Eop1, Eop3, Eop4 (AvrRpt2Ea), and HopPtoCEa, two harpins: HrpN and HrpW, and other proteins such as Eop2, HrpJ, and HrpK. Homologs of these T3SS-secreted proteins have been characterized in plant pathogens like Pseudomonas syringae, Xanthomonas spp., Ralstonia spp., plant symbionts like Rhizobium spp. and animal pathogens such as Yersinia spp., implicating a central role of these effector proteins in manipulating bacterial interactions with diverse host species. Translocation of effectors is crucial for T3SS-mediated disease development in hosts. It is thought to be mediated by cognate chaperones, but the translocation of E. amylovora DspA/E was recently shown to be regulated by several different chaperones as well as a previously reported harpin, HrpN, in addition to its cognate chaperone DspB/F. This suggests that the dynamics of effector translocation are more complicated than previously expected in E. amylovora. In this review, we summarize the current knowledge of E. amylovora T3SS effectors, chaperones, and harpins together with their homologs from other plant-associated bacterial species. We also discuss the roles of T3SS-secreted proteins in pathogenicity and fitness via exploring their localization, activity, and direct or indirect targets in host plants. Finally, we provide future perspectives on studying effector biology and the T3SS in E. amylovora and how this fundamental knowledge provides potential applications in preventing fire blight.