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Journal of Bacteriology, April 2008, p. 2880-2891, Vol. 190, No. 8
0021-9193/08/$08.00+0     doi:10.1128/JB.01702-07
Copyright © 2008, American Society for Microbiology. All Rights Reserved.

The HopZ Family of Pseudomonas syringae Type III Effectors Require Myristoylation for Virulence and Avirulence Functions in Arabidopsis thaliana{triangledown} ,{dagger}

Jennifer D. Lewis,1 Wasan Abada,1 Wenbo Ma,2 David S. Guttman,1,3,{ddagger} and Darrell Desveaux1,3*,{ddagger}

Department of Cell & Systems Biology, University of Toronto, 25 Willcocks St., Toronto, Ontario M5S 3B2, Canada,1 Department of Plant Pathology and Microbiology, University of California, Riverside, 900 University Ave., Riverside, California 92521,2 Centre for the Analysis of Genome Evolution & Function, University of Toronto, Toronto, Ontario, Canada3

Received 23 October 2007/ Accepted 28 January 2008

Pseudomonas syringae utilizes the type III secretion system to translocate effector proteins into plant cells, where they can contribute to the pathogen's ability to infect and cause disease. Recognition of these effectors by resistance proteins induces defense responses that typically include a programmed cell death reaction called the hypersensitive response. The YopJ/HopZ family of type III effector proteins is a common family of effector proteins found in animal- and plant-pathogenic bacteria. The HopZ family in P. syringae includes HopZ1aPsyA2, HopZ1bPgyUnB647, HopZ1cPmaE54326, HopZ2Ppi895A and HopZ3PsyB728a. HopZ1a is predicted to be most similar to the ancestral hopZ allele and causes a hypersensitive response in multiple plant species, including Arabidopsis thaliana. Therefore, it has been proposed that host defense responses have driven the diversification of this effector family. In this study, we further characterized the hypersensitive response induced by HopZ1a and demonstrated that it is not dependent on known resistance genes. Further, we identified a novel virulence function for HopZ2 that requires the catalytic cysteine demonstrated to be required for protease activity. Sequence analysis of the HopZ family revealed the presence of a predicted myristoylation sequence in all members except HopZ3. We demonstrated that the myristoylation site is required for membrane localization of this effector family and contributes to the virulence and avirulence activities of HopZ2 and HopZ1a, respectively. This paper provides insight into the selective pressures driving virulence protein evolution by describing a detailed functional characterization of the diverse HopZ family of type III effectors with the model plant Arabidopsis.

* Corresponding author. Mailing address: Department of Cell & Systems Biology, University of Toronto, 25 Willcocks St., Toronto, Ontario M5S 3B2, Canada. Phone: (416) 978-7153. Fax: (416) 978-5878. E-mail: desveaux{at}csb.utoronto.ca

{triangledown} Published ahead of print on 8 February 2008.

{dagger} Supplemental material for this article may be found at http://jb.asm.org/.

{ddagger} D.D. and D.S.G. made equal contributions to this paper.

Journal of Bacteriology, April 2008, p. 2880-2891, Vol. 190, No. 8
0021-9193/08/$08.00+0     doi:10.1128/JB.01702-07
Copyright © 2008, American Society for Microbiology. All Rights Reserved.



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