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The wheat  Pm3  resistance gene against the powdery mildew pathogen occurs as an allelic series encoding functionally different immune receptors which induce resistance upon recognition of isolate-specific avirulence (AVR) effectors from the pathogen. Here, we describe the identification of five effector proteins from the mildew pathogens of wheat, rye, and the wild grass  Dactylis glomerata , specifically recognized by the PM3B, PM3C and PM3D receptors. Together with the earlier identified AVRPM3 A2/F2 , the recognized AVRs of PM3B/C, (AVRPM3 B2/C2 ), and PM3D (AVRPM3 D3 ) belong to a large group of proteins with low sequence homology but predicted structural similarities.  AvrPm3   b2/c2   and  AvrPm3   d3   are conserved in all tested isolates of wheat and rye mildew, and non-host infection assays demonstrate that  Pm3b ,  Pm3c , and  Pm3d  are also restricting the growth of rye mildew on wheat. Furthermore, divergent AVR homologues from non-adapted rye and  Dactylis  mildews are recognized by PM3B, PM3C, or PM3D, demonstrating their involvement in host specificity.

The AvrPm3-Pm3 effector-NLR interactions control both race-specific resistance and host-specificity of cereal mildews on wheat