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Università Degli Studi Di Palermo UNIVERSITÀ DEGLI STUDI DI PALERMO PhD course in Agricultural, Forestry and Environmental Sciences Curriculum: Agro-Mediterranean Ecosystems Plant Pathology AGR 12 STUDY OF THE INTERACTIONS BETWEEN BROAD BEAN WILT VIRUS 1 AND ITS HOST PLANTS COORDINATOR PhD STUDENT PROF. VINCENZO BAGARELLO CATERINA CARPINO CO-TUTORS TUTOR DOTT. LUIS GALIPIENSO TORREGROSA PROF. SALVATORE DAVINO PROF. EZIO PERI CICLO XXX 2018 Table of contents Lists of abbreviations ........................................................................................................... 5 Abstract ................................................................................................................................. 7 Chapter 1 Introduction .......................................................................................................................... 9 1.1 Plant viruses ................................................................................................................. 9 1.1.1 Evolution of plant viruses ................................................................................... 11 1.1.2 Biology of RNA plant viruses ............................................................................. 12 1.1.3 Transmission of RNA vector-borne plant viruses ............................................... 13 1.2 Host defensive responses and pathological process induction .................................. 14 1.2.1 Hypersensitive response and systemic acquired resistance ................................ 15 1.2.2 RNA silencing ..................................................................................................... 15 1.2.3 Pathogenicity determinants ................................................................................. 17 1.2.4 Viral suppressors of gene silencing ..................................................................... 18 1.3 Control of plant viruses .............................................................................................. 19 1.4 RNA viral infectious clones ....................................................................................... 19 1.5 Plant virus used in this study: Broad bean wilt virus 1 ............................................. 21 1.5.1 Distribution, host range and transmission of fabaviruses ................................... 22 1.5.2 Virion and genome organization of Broad bean wilt virus 1 .............................. 23 Chapter 2 Interest and objectives ....................................................................................................... 25 Chapter 3 Biological characterisation and viral accumulation ........................................................ 29 of BBWV-1 isolates ............................................................................................................. 29 3.1 Introduction ................................................................................................................ 31 3.2 Materials and methods ............................................................................................... 32 3.2.1 BBWV-1 isolates and mechanical inoculation in herbaceous hosts ................... 32 3.2.2 RNA extraction ................................................................................................... 33 3.2.3 Virus quantitation by real-time RT-PCR ............................................................ 33 3.3 Results ........................................................................................................................ 34 3.3.1 Infectivity of different BBWV-1 isolates ............................................................ 34 3.3.2 Symptomatology of different BBWV-1 isolates ................................................. 35 3.3.3 Accumulation of different BBWV-1 viral isolates ............................................. 37 3.4 Discussion .................................................................................................................. 40 1 Chapter 4 Role of BBWV-1 RNA2 encoded VP37 protein in the induction of plants symptoms and suppression of post-transcriptional gene silencing ................................................. 43 4.1 Introduction ............................................................................................................... 45 4.2 Materials and methods ............................................................................................... 46 4.2.1 Plant and virus materials ..................................................................................... 46 4.2.2 Construction of full-length genome cDNA clones of BBWV-1 ......................... 47 4.2.3 Agroinfiltration of cDNA BBWV-1 constructions ............................................. 48 4.2.4 Real-time quantitative RT-PCR .......................................................................... 49 4.2.5 Virion purification and analysis .......................................................................... 49 4.2.6 PVX-based vector expression assay ................................................................... 50 4.1.7 NBT and Trypan blue staining ............................................................................ 50 4.2.8 Transient expression assay in N. benthamiana 16c ............................................ 51 4.2.9 pTCV-GFP complementation assay ................................................................... 52 4.2.10 RNA extraction and northern blot hybridization .............................................. 52 4.2.11 RT-PCR and sequencing ................................................................................... 53 4.3 Results ....................................................................................................................... 54 4.3.1 BBWV-1 VP37 protein is involved in plant symptom induction and host specificity ..................................................................................................................... 54 4.3.2 BBWV-1 VP37 protein enhances the PVX-induced symptoms and activates the systemic necrosis associated with programed cell death in N. benthamiana plants .... 61 4.3.3 BBWV-1 VP37 protein is an RNA silencing suppressor ................................... 64 4.4 Discussion .................................................................................................................. 66 Chapter 5 Role of BBWV-1 RNA2-encoded MP, SCP and LCP proteins in the suppression of PTGS and induction of plant symptoms .......................................................................... 71 5.1 Introduction ............................................................................................................... 73 5.2 Materials and methods ............................................................................................... 74 5.2.1 Plasmid construction ........................................................................................... 74 5.2.2 N. benthamiana 16c transient expression assay .................................................. 75 5.2.3 RNA extraction, northern blot and RT-PCR ....................................................... 75 5.2.4 NBT and TB staining .......................................................................................... 76 5.2.5 pTCV-GFP complementation assay ................................................................... 76 5.3 Results ....................................................................................................................... 76 5.3.1 BBWV-1 encoded MP and SCP proteins are suppressors of post-transcriptional gene silencing in N. benthamiana plants ..................................................................... 76 2 5.3.2 BBWV-1 SCP protein enhances the PVX-induced symptoms and activates the systemic necrosis associated with programed cell death in N. benthamiana plants .... 80 5.4 Discussion .................................................................................................................. 84 Chapter 6 General conclusions ............................................................................................................ 87 References............................................................................................................................ 91 Acknowledgements ........................................................................................................... 105 3 4 List of abbreviations Ago Argonaute protein Avr Pathogen avirulence gene product BBTV Banana bunchy top virus BBVV Broad bean V virus BBWV-1 Broad bean wilt virus 1 BBWV-2 Broad bean wilt virus 2 BCMNV Bean common necrotic virus BCMV Bean common mosaic virus BPMV Bean pod mottle virus BYMV Bean yellow mosaic virus BYV Beet yellows virus CaMV Cauliflower mosaic virus ChLCB Chili leaf curl betasatellite CMV Cucumber mosaic virus CP Capsid protein CPMV Cowpea mosaic virus CTV Citrus tristeza virus CuMMV Cucurbit mild mosaic virus DCLs Dicer-like ribonuclease proteins dsRNA Double-stranded RNA FHV Flock house virus FLUAV Human influenza virus GeMV Gentian mosaic virus GFLV Grapevine fanleaf virus HDV Hepatitis delta virus HEL Helicase HR Hypersensitive response LMMV Lamium mosaic virus MCDV Maize chlorotic dwarf virus MP Movement protein NBT Nitro blue tetrazolium 5 PCD Programmed cell death PDs Plasmodesmata PepMV Pepino mosaic virus PNRS Prunus necrotic ringspot virus POL Polymerase PPV Plum pox virus PRO-CO Protease cofactor PRs Pathogenesis-related genes PTGS Post-transcriptional gene silencing PVX Potato virus X R Dominant
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