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Summary  One of the most devastating fungal diseases of soybean in the southern USA is Cercospora leaf blight (CLB), which is caused mainly by  Cercospora  cf.  flagellaris . Recent studies found that the fungal effector AVR4, originally identified in  Cladosporium fulvum  as a chitin‐binding protein, is highly conserved among other  Cercospora  species. We wanted to determine whether it is present in  C.  cf.  flagellaris  and, if so, whether it plays a role in the pathogen infection of soybean. We cloned the  Avr4  gene and created  C.  cf.  flagellaris  ∆ avr4  mutants, which produced little cercosporin and significantly reduced expression of cercosporin biosynthesis genes. The ∆ avr4  mutants were also more sensitive to chitinase and showed reduced virulence on soybean compared to the wild‐type. The observed reduced virulence of  C.  cf.  flagellaris  ∆ avr4  mutants on detached soybean leaves is likely due to reduced cercosporin biosynthesis. The phenotypes of reduced cercosporin production and cercosporin pathway gene expression, similar to those of the ∆ avr4  mutants, were reproduced when wild‐type  C.  cf.  flagellaris  was treated with double‐stranded RNA targeting  Avr4   in vitro . These two independent approaches demonstrated for the first time the direct involvement of AVR4 in the biosynthesis of cercosporin.

The AVR4 effector is involved in cercosporin biosynthesis and likely affects the virulence of Cercospora cf. flagellaris on soybean