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The wheat pathogen  Stagonospora nodorum  produces multiple necrotrophic effectors (also called host-selective toxins) that promote disease by interacting with corresponding host sensitivity gene products. SnTox1 was the first necrotrophic effector identified in  S. nodorum , and was shown to induce necrosis on wheat lines carrying  Snn1 . Here, we report the molecular cloning and validation of  SnTox1  as well as the preliminary characterization of the mechanism underlying the SnTox1- Snn1  interaction which leads to susceptibility.  SnTox1  was identified using bioinformatics tools and verified by heterologous expression in  Pichia pastoris .  SnTox1  encodes a 117 amino acid protein with the first 17 amino acids predicted as a signal peptide, and strikingly, the mature protein contains 16 cysteine residues, a common feature for some avirulence effectors. The transformation of  SnTox1  into an avirulent  S. nodorum  isolate was sufficient to make the strain pathogenic. Additionally, the deletion of  SnTox1  in virulent isolates rendered the  SnTox1  mutated strains avirulent on the  Snn1  differential wheat line.  SnTox1  was present in 85% of a global collection of  S .  nodorum  isolates. We identified a total of 11 protein isoforms and found evidence for strong diversifying selection operating on  SnTox1 . The SnTox1- Snn1  interaction results in an oxidative burst, DNA laddering, and pathogenesis related (PR) gene expression, all hallmarks of a defense response. In the absence of light, the development of SnTox1-induced necrosis and disease symptoms were completely blocked. By comparing the infection processes of a GFP-tagged avirulent isolate and the same isolate transformed with  SnTox1 , we conclude that SnTox1 may play a critical role during fungal penetration. This research further demonstrates that necrotrophic fungal pathogens utilize small effector proteins to exploit plant resistance pathways for their colonization, which provides important insights into the molecular basis of the wheat- S. nodorum  interaction, an emerging model for necrotrophic pathosystems.

The Cysteine Rich Necrotrophic Effector SnTox1 Produced by Stagonospora nodorum Triggers Susceptibility of Wheat Lines Harboring Snn1