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Powdery mildew is an important disease of cereals. It is caused by one species,  Blumeria graminis , which is divided into  formae speciales  each of which is highly specialized to one host. Recently, a new form capable of growing on triticale ( B.g. triticale ) has emerged through hybridization between wheat and rye mildews ( B.g. tritici  and  B.g. secalis , respectively). In this work, we used RNA sequencing to study the molecular basis of host adaptation in  B.g. triticale . We analyzed gene expression in three  B.g. tritici  isolates, two  B.g. secalis  isolates and two  B.g. triticale  isolates and identified a core set of putative effector genes that are highly expressed in all  formae speciales . We also found that the genes differentially expressed between isolates of the same form as well as between different  formae speciales  were enriched in putative effectors. Their coding genes belong to several families including some which contain known members of mildew avirulence ( Avr ) and suppressor ( Svr ) genes. Based on these findings we propose that effectors play an important role in host adaptation that is mechanistically based on  Avr-Resistance gene-Svr  interactions. We also found that gene expression in the  B.g. triticale  hybrid is mostly conserved with the parent-of-origin, but some genes inherited from  B.g. tritici  showed a  B.g. secalis -like expression. Finally, we identified 11 unambiguous cases of putative effector genes with hybrid-specific, non-parent of origin gene expression, and we propose that they are possible determinants of host specialization in triticale mildew. These data suggest that altered expression of multiple effector genes, in particular  Avr  and  Svr  related factors, might play a role in mildew host adaptation based on hybridization.

frontiers in plant science

Non-parent of Origin Expression of Numerous Effector Genes Indicates a Role of Gene Regulation in Host Adaption of the Hybrid Triticale Powdery Mildew Pathogen