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absence of the Rcr3 pathogenicity target. The characterization of a Cf0/rcr3 mutant will reveal whether Rcr3 is necessary for full pathogenicity of C. fulvum.

The Guard hypothesis predicts that for each R protein there is both a corresponding pathogen Avr product and a host target. Such a model would explain the dual recognition capacity of some NB-LRR proteins such as RPM1 and Mi-1 if they “guard” the same host component targeted by unrelated Avr products. Evolutionary mechanisms sustaining R gene diversity are essential for the plant to be able to detect distinct pathogen (a)virulence products that target R protein-“guarded ” host components.

All members of the Jones lab are thanked for useful discussion. We thank Sara Perkins, Margaret Shailer, and Justine Campling for their excellent horticultural service. This work was supported in part by the United Kingdom Biotechnology and Biological Sciences Research Council (M.S.D.) and by a Gatsby Ph.D. studentship (C.G.). The Sainsbury Laboratory is supported by the Gatsby Charitable Foundation.

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