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The gene-for-gene hypothesis states that for each gene controlling resistance in the host, there is a corresponding, specific gene controlling avirulence in the pathogen. Allelic series of the cereal mildew resistance genes  Pm3  and  Mla  provide an excellent system for genetic and molecular analysis of resistance specificity. Despite this opportunity for molecular research, avirulence genes in mildews remain underexplored. Earlier work in barley powdery mildew ( B.g. hordei ) has shown that the reaction to some  Mla  resistance alleles is controlled by multiple genes. Similarly, several genes are involved in the specific interaction of wheat mildew ( B.g. tritici ) with the  Pm3  allelic series. We found that two mildew genes control avirulence on  Pm3f : one gene is involved in recognition by the resistance protein as demonstrated by functional studies in wheat and the heterologous host  Nicotiana benthamiana . A second gene is a suppressor, and resistance is only observed in mildew genotypes combining the inactive suppressor and the recognized  Avr . We propose that such suppressor/avirulence gene combinations provide the basis of specificity in mildews. Depending on the particular gene combinations in a mildew race, different genes will be genetically identified as the “avirulence” gene. Additionally, the observation of two LINE retrotransposon-encoded avirulence genes in  B.g. hordei  further suggests that the control of avirulence in mildew is more complex than a canonical gene-for-gene interaction. To fully understand the mildew–cereal interactions, more knowledge on avirulence determinants is needed and we propose ways how this can be achieved based on recent advances in the field.

Avirulence Genes in Cereal Powdery Mildews: The Gene-for-Gene Hypothesis 2.0