Optimizing the Effectiveness of Induced Resistance in Tomato for Bacterial Disease Management
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Optimizing the Effectiveness of Induced Resistance in Tomato for Bacterial Disease Management by Cheryl Trueman A Thesis presented to The University of Guelph In partial fulfilment of requirements for the degree of Doctor of Philosophy in Environmental Biology Guelph, Ontario, Canada © Cheryl Trueman, December, 2017 1 i ABSTRACT OPTIMIZING THE EFFECTIVENESS OF INDUCED RESISTANCE IN TOMATO FOR BACTERIAL DISEASE MANAGEMENT Cheryl Lynn Trueman Advisor: University of Guelph, 2017 Professor P. H. Goodwin As alternatives for managing bacterial speck (Pseudomonas syringae pv. tomato (Pst)) and bacterial spot (Xanthomonas gardneri (Xg)) of tomato (Solanum lycopersicum), the synthetic chemical defense activator, acibenzolar-s-methyl (ASM) combined with the synthetic plant growth regulator (PGR) uniconazole (UNI), the natural chemical defense activator, para-aminobenzoic acid (PABA), and the putative biological defense activator, B. mycoides/weihenstephanensis R17, were examined. No consistent benefits for bacterial spot and speck control or tomato growth were observed in field experiments from 2011-2013 with the combination of ASM and UNI, whether applied to seedlings (<6 weeks old) or post-transplanting. However, greenhouse applied ASM or ASM+UNI reduced late season disease severity 18 to 24% compared to nontreated and CuOH-treated controls for cv. TSH4 in 2012, indicating that ASM applied to tomato seedlings can have long-term benefits under certain conditions. Fitness costs in terms of reduced growth are a concern with ASM, so UNI was added to ASM to determine if that could be ameliorated, but this approach was ineffective. Greenhouse ASM applications to cv. H9909 in 2012 reduced total yield by 20% compared to the nontreated control, indicating a fitness cost, and ASM+UNI treated plants showed a similar loss. Fitness costs may have occurred in 2012 due to stress from dry conditions after transplanting. For PABA, effectiveness was affected by application method, concentration and host genotype. Despite optimizing PABA efficacy under controlled conditions, PABA was ineffective in the field. A bacterial endophyte, R17, was isolated from Solanum arcanum and its ability to putatively induce resistance against Pst under controlled conditions was affected by its concentration, application method, and host genotype. PABA and R17 reduced bacterial speck lesion incidence up to 43 and 51%, respectively, but the lesions that developed were larger in treated than nontreated plants resulting in no reduction in Pst population or total symptomatic leaf area. This suggests that certain defense activators can reduce the ability of Pst to infect but then allow for greater Pst population growth post-infection. While defense activators have potential, they need to be more effective and consistent before they are integrated into bacterial disease management strategies of tomato. 2 ii DEDICATION For Justin, you believed in me, and Alexis, you helped me believe in myself. Also for Dad, who told me to find a summer job in Ontario, and Mom for all those kilometres you drove with me and for me. iii 3 ACKNOWLEDGEMENTS Thank you to my advisor, Dr. Paul Goodwin, and the members of my advisory committee, Dr. Kari Dunfield, Dr. Tom Hsiang, Dr. Annette Nassuth, and Dr. Istvan Rajcan for their guidance and advice during this journey. Your time and patience is truly appreciated. This research was completed at the Ridgetown Campus, University of Guelph, and there are many people who contributed time and effort toward its completion. These people include the Field Vegetable Pest Management Technician, Phyllis May, former co-op students Sherri Tedford and Tina Simonton, and many summer students and research assistants. Thank you for your dedication to the program. I would also like to thank the many colleagues at the Ridgetown Campus who provided advice and support during this journey including Janice LeBoeuf, Steve Loewen, Ken McEwan, Elaine Roddy, Dr. Darren Robinson, Dr. Art Schaafsma, and Dr. Laura Van Eerd. I would also like to thank Dr. Diane Cuppels, with whom I had a valuable conversation about bacterial disease in tomato after her retirement from Agriculture and Agri-Food Canada in London. To my partner in life, Justin Kritikos, thank you for being my soft place to fall. Your patience and encouragement did not go unnoticed. Thank you for embracing what it means to be an equal partner in life. Funding for this research was provided in part by the Ontario Tomato Research Institute, Syngenta Canada, Valent Canada, Ridgetown Campus – University of Guelph, and the Ontario Ministry and Food and Rural Affairs. iv 4 TABLE OF CONTENTS ABSTRACT .................................................................................................................................................. ii DEDICATION ............................................................................................................................................. iii ACKNOWLEDGEMENTS ......................................................................................................................... iv TABLE OF CONTENTS .............................................................................................................................. v LIST OF TABLES ........................................................................................................................................ x LIST OF FIGURES .................................................................................................................................... xv LIST OF ABBREVIATIONS .................................................................................................................... xxi Chapter 1: Literature Review ........................................................................................................................ 1 1.1 Introduction ................................................................................................................................... 1 1.2 Processing tomatoes in Ontario ..................................................................................................... 1 1.3 Tomato genetics ............................................................................................................................ 3 1.4 Bacterial endophytes of tomato ..................................................................................................... 3 1.5 Bacterial speck pathogen of tomato .............................................................................................. 4 1.6 Bacterial spot pathogen of tomato ................................................................................................ 9 1.7 Current bacterial speck and spot management practices in tomato ............................................ 13 1.8 SIR and its use in bacterial disease management ........................................................................ 15 1.8.1 SAR ..................................................................................................................................... 15 1.8.1.1 Mechanisms of SAR ................................................................................................... 15 1.8.1.2 Biological activators of SAR ...................................................................................... 17 1.8.1.3 PAMP activators of SAR ............................................................................................ 18 1.8.1.4 Plant activators of SAR ............................................................................................... 18 1.8.1.5 Synthetic activators of SAR ........................................................................................ 19 1.8.1.6 Plant fitness costs associated with SAR ...................................................................... 20 1.8.1.7 Variation in SAR response among cultivars ............................................................... 22 1.8.1.8 SAR in disease management ....................................................................................... 23 1.8.1.9 SAR in management of bacterial speck and spot of tomato ....................................... 24 1.8.2 ISR ...................................................................................................................................... 26 1.8.2.1 Mechanisms of ISR ..................................................................................................... 26 1.8.2.2 Biological activators of ISR ........................................................................................ 28 1.8.2.3 PAMP activators of ISR .............................................................................................. 30 v 1.8.2.4 Synthetic activators of ISR ......................................................................................... 31 1.8.2.5 Plant growth effects associated with ISR .................................................................... 31 1.8.2.6 Factors affecting ISR .................................................................................................. 33 1.8.2.7 ISR as a disease management tool .............................................................................. 35 1.8.2.8 ISR as a disease management tool in tomato .............................................................. 36 1.8.3 Interactions and relationships between SAR and ISR pathways ......................................... 38 1.9 GA-related PGRs .......................................................................................................................