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The vascular plant pathogen  Verticillium nonalfalfae  causes  Verticillium  wilt in several important crops.  Vna SSP4.2 was recently discovered as a  V .  nonalfalfae  virulence effector protein in the xylem sap of infected hop. Here, we expanded our search for candidate secreted effector proteins (CSEPs) in the  V .  nonalfalfae  predicted secretome using a bioinformatic pipeline built on  V .  nonalfalfae  genome data, RNA-Seq and proteomic studies of the interaction with hop. The secretome, rich in carbohydrate active enzymes, proteases, redox proteins and proteins involved in secondary metabolism, cellular processing and signaling, includes 263 CSEPs. Several homologs of known fungal effectors (LysM, NLPs, Hce2, Cerato-platanins, Cyanovirin-N lectins, hydrophobins and CFEM domain containing proteins) and avirulence determinants in the PHI database (Avr-Pita1 and MgSM1) were found. The majority of CSEPs were non-annotated and were narrowed down to 44 top priority candidates based on their likelihood of being effectors. These were examined by spatio-temporal gene expression profiling of infected hop. Among the highest  in planta  expressed CSEPs, five deletion mutants were tested in pathogenicity assays. A deletion mutant of  VnaUn . 279 , a lethal pathotype specific gene with sequence similarity to SAM-dependent methyltransferase (LaeA), had lower infectivity and showed highly reduced virulence, but no changes in morphology, fungal growth or conidiation were observed. Several putative secreted effector proteins that probably contribute to  V .  nonalfalfae  colonization of hop were identified in this study. Among them,  LaeA  gene homolog was found to act as a potential novel virulence effector of  V .  nonalfalfae . The combined results will serve for future characterization of  V .  nonalfalfae  effectors, which will advance our understanding of  Verticillium  wilt disease.

Comprehensive analysis of Verticillium nonalfalfae in silico secretome uncovers putative effector proteins expressed during hop invasion