The hop metabolome important for the brewing industry and for medical purposes is endangered worldwide due to multiple viroid infections affecting hop physiology. Combinatorial biolistic hop inoculation with Citrus bark cracking viroid (CBCVd), Apple fruit crinkle viroid (AFCVd), Hop latent viroid, and Hop stunt viroid (HSVd) showed a low CBCVd compatibility with HSVd, while all other viroid combinations were highly compatible. Unlike to other viroids, single CBCVd propagation showed a significant excess of (-) over (+) strands in hop, tomato, and Nicotiana benthamiana, but not in citruses. Inoculation of hop with all viroids led to multiple infections with unstable viroid levels in individual plants in the pre- and post-dormancy periods, and to high plant mortality and morphological disorders. Hop isolates of CBCVd and AFCVd were highly stable, only minor quasispecies were detected. CBCVd caused a strong suppression of some crucial mRNAs related to the hop prenylflavonoid biosynthesis pathway, while AFCVd-caused effects were moderate. According to mRNA degradome analysis, this suppression was not caused by a direct viroid-specific small RNA-mediated degradation. CBCVd infection led to a strong induction of two hop transcription factors from WRKY family and to a disbalance of WRKY/WDR1 complexes important for activation of lupulin genes.

Biology Centre ASCR v v i Institute of Plant Molecular Biology Branišovská 31 České Budějovice 370 05 Czech Republic

Biology Centre ASCR v v i Institute of Plant Molecular Biology Branišovská 31 České Budějovice 370 05 Czech Republic; University of South Bohemia Faculty of Science Branišovská 31 370 05 České Budějovice Czech Republic

Faculty of Agriculture and Life Science Hirosaki University Bubkyo cho Hirosaki 036 8561 Japan

Institute of Physical Biology Heinrich Heine Universität Düsseldorf D 40204 Düsseldorf Germany

Slovenian Institute of Hop Research and Brewing Cesta Žalskega tabora 2 SI 3310 Žalec Slovenia

University of Ljubljana Biotechnical Faculty Department of Agronomy Jamnikarjeva 101 SI 1000 Ljubljana Slovenia

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