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THE PLANT CELL, Vol 9, Issue 3 367-379, Copyright © 1997 by American Society of Plant Biologists
The Biotrophic Fungus Cladosporium fulvum Circumvents Cf-4[mdash]Mediated Resistance by Producing Unstable AVR4 Elicitors
MHAJ. Joosten, R. Vogelsang, T. J. Cozijnsen, M. C. Verberne and PJGM. De Wit
Department of Phytopathology, Wageningen Agricultural University, Binnenhaven 9, 6709 PD Wageningen, The Netherlands
The avirulence gene Avr4 conditions avirulence of the biotrophic fungus
Cladosporium fulvum on tomato genotypes carrying resistance gene Cf-4
(MM-Cf4). Strains of the fungus that circumvent Cf-4[mdash]specific
resistance show various single point mutations in the coding region of the
Avr4 gene. Similar to expression of the Avr4 gene, expression of the
various virulent avr4 alleles is specifically induced during pathogenesis.
Polyclonal antibodies raised against the AVR4 elicitor, however, did not
detect AVR4 isoforms in MM-Cf4 plants infected by the different virulent
strains, indicating that these isoforms are unstable. To analyze whether
the AVR4 isoforms still possess specific elicitor activity, the avr4
alleles were expressed in MM-Cf4 plants by using the potato virus X
(PVX)-based expression system. Inoculation with PVX::Avr4 resulted in the
development of spreading lesions, eventually leading to plant death,
whereas the various PVX::avr4 derivatives induced symptoms ranging from
severe necrosis to no lesions at all. We conclude that instability of the
AVR4 isoforms that are produced by virulent strains is a crucial factor in
circumvention of Cf-4[mdash]mediated resistance.
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