Evolution of wheat blast resistance gene Rmg8 accompanied by differentiation of variants recognizing the powdery mildew fungus.

Wheat blast, a devastating disease having spread recently from South America to Asia and Africa, is caused by Pyricularia oryzae (synonym of Magnaporthe oryzae) pathotype Triticum, which first emerged in Brazil in 1985. Rmg8 and Rmg7, genes for resistance to wheat blast found in common wheat and tetraploid wheat, respectively, recognize the same avirulence gene, AVR-Rmg8. Here we show that an ancestral resistance gene, which had obtained an ability to recognize AVR-Rmg8 before the differentiation of Triticum and Aegilops, has expanded its target pathogens. Molecular cloning revealed that Rmg7 was an allele of Pm4, a gene for resistance to wheat powdery mildew on 2AL, whereas Rmg8 was its homoeologue on 2BL ineffective against wheat powdery mildew. Rmg8 variants with the ability to recognize AVR-Rmg8 were distributed not only in Triticum spp. but also in Aegilops speltoides, Aegilops umbellulata and Aegilops comosa. This result suggests that the origin of resistance gene(s) recognizing AVR-Rmg8 dates back to the time before differentiation of A, B, S, U and M genomes, that is, ~5 Myr before the emergence of its current target, the wheat blast fungus. Phylogenetic analyses suggested that, in the evolutionary process thereafter, some of their variants gained the ability to recognize the wheat powdery mildew fungus and evolved into genes controlling dual resistance to wheat powdery mildew and wheat blast.

Origin and Dynamics of Rwt6, a Wheat Gene for Resistance to Nonadapted Pathotypes of Pyricularia oryzae.

Avirulence of Eleusine isolates of Pyricularia oryzae on common wheat is conditioned by at least five avirulence genes. One is PWT3 corresponding to resistance gene Rwt3 located on chromosome 1D. We identified a resistance gene corresponding to a second avirulence gene, PWT6, and named it Rmg9 (Rwt6). Rwt6 was closely linked to Rwt3. A survey of the population of Aegilops tauschii, the D genome donor to common wheat, revealed that some accessions from the southern coastal region of the Caspian Sea, the birthplace of common wheat, carried both genes. Rwt6 and Rwt3 carriers accounted for 65 and 80%, respectively, of accessions in a common wheat landrace collection. The most likely explanation of our results is that both resistance genes were simultaneously introduced into common wheat at the time of hybridization of Triticum turgidum and A. tauschii. However, a prominent difference was recognized in their geographical distributions in modern wheat; Rwt3 and Rwt6 co-occurred at high frequencies in regions to the east of the Caspian Sea, whereas Rwt6 occurred at a lower frequency than Rwt3 in regions to the west. This difference was considered to be associated with range of pathotypes to which these genes were effective. A. tauschii accessions carrying Rwt3 and Rwt6 also carried Rwt4, another resistance gene involved in the species specificity. We suggest that the gain of the D genome should have given an adaptive advantage to the genus Triticum by conferring disease resistance.