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First published online August 11, 2006; 10.1105/tpc.106.043307

The Plant Cell 18:2402-2414 (2006)
© 2006 American Society of Plant Biologists

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Multiple Avirulence Paralogues in Cereal Powdery Mildew Fungi May Contribute to Parasite Fitness and Defeat of Plant Resistance

Christopher J. Ridouta,b,1, Pari Skamniotia,b,2, Oliver Porritta, Soledad Sacristana, Jonathan D.G. Jonesb and James K.M. Browna

a Department of Disease and Stress Biology, John Innes Centre, Colney, Norwich, NR4 7UH, United Kingdom
b Sainsbury Laboratory, John Innes Centre, Colney, Norwich, NR4 7UH, United Kingdom

1 To whom correspondence should be addressed. E-mail christopher.ridout{at}bbsrc.ac.uk; fax 44-01603-450000.

Powdery mildews, obligate biotrophic fungal parasites on a wide range of important crops, can be controlled by plant resistance (R) genes, but these are rapidly overcome by parasite mutants evading recognition. It is unknown how this rapid evolution occurs without apparent loss of parasite fitness. R proteins recognize avirulence (AVR) molecules from parasites in a gene-for-gene manner and trigger defense responses. We identify AVRa10 and AVRk1 of barley powdery mildew fungus, Blumeria graminis f sp hordei (Bgh), and show that they induce both cell death and inaccessibility when transiently expressed in Mla10 and Mlk1 barley (Hordeum vulgare) varieties, respectively. In contrast with other reported fungal AVR genes, AVRa10 and AVRk1 encode proteins that lack secretion signal peptides and enhance infection success on susceptible host plant cells. AVRa10 and AVRk1 belong to a large family with >30 paralogues in the genome of Bgh, and homologous sequences are present in other formae speciales of the fungus infecting other grasses. Our findings imply that the mildew fungus has a repertoire of AVR genes, which may function as effectors and contribute to parasite virulence. Multiple copies of related but distinct AVR effector paralogues might enable populations of Bgh to rapidly overcome host R genes while maintaining virulence.




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