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Management of Damping-Off Caused by Pythium Spp. in Organic MANAGEMENT OF DAMPING-OFF CAUSED BY PYTHIUM SPP. IN ORGANIC VEGETABLE PRODUCTION IN THE PACIFIC NORTHWEST By ANA VIDA CRISOSTOMO ALCALA A thesis submitted in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY WASHINGTON STATE UNIVERSITY Department of Plant Pathology JULY 2013 To the Faculty of Washington State University: The members of the Committee appointed to examine the dissertation of ANA VIDA CRISOSTOMO ALCALA find it satisfactory and recommend that it be accepted. __________________________________ Lindsey du Toit, Ph.D., Chair __________________________________ Carol Miles, Ph.D. __________________________________ Tim Paulitz, Ph.D. __________________________________ Lyndon Porter, Ph.D. ii ACKNOWLEDGEMENTS Professional I am greatly indebted to four great scientists who mentored me in my PhD. My deepest gratitude is to Dr. Lindsey du Toit, my major advisor, for opening the door of opportunity for me to pursue my Ph.D. in the US, and for providing me with every chance to develop a variety of skills essential for this profession. Her enthusiasm, passion, and dedication to research have been inspirational to me. To my committee members, Dr. Carol Miles, Dr. Tim Paulitz, and Dr. Lyndon Porter, who generously took time to mentor me, by distance or whenever I got a chance to work on campus, as well as for generously sharing their expertise. To the very hardworking VSP crew: Mike, Barb, Sarah, John, and Anita, for all their help in one or many aspects of this project. Particularly to Mike, for his expertise and tremendous help in my field, greenhouse, and lab trials; Barb, for always being ready to help even beyond the call of duty, and for always making what seemed an impossible task possible. To Dr. Tim Murray, for accommodating me in his lab during my first year in Pullman and providing me with everything I needed to be able to carry out the isolation and baiting aspects of this project. To Dr. Lyndon Porter and the Legume Pathology crew of ARS USDA Prosser: Ginny, Tyler, and Nick. I am truly grateful to Dr. Porter for graciously sharing his expertise in many aspects of this project, and making all the field trials possible. Ginny, for all your hard work in all the pea field trials, and for always providing a home for us in Prosser. Tyler and Nick, for the long hours of work without complaining. To the ARS USDA Wheat Pathology Lab in Pullman, especially to Dr. Tim Paulitz and Dr. Kurt Schroeder, for accommodating me in their lab to carry out several aspects of this project iii and their willingness to impart their expertise. I am truly grateful to Dr. Schroeder for patiently teaching and assisting me with the molecular aspects of this project. I would like to extend my gratitude to all the faculty and staff in WSU Mount Vernon NWREC for all their support and encouragement, as well as to the faculty and staff of the Plant Pathology department in Pullman for their untiring assistance. Personal I am richly blessed with family, friends, and colleagues who supported and encouraged me throughout my journey in PhD. I’d like to thank my friends with whom I shared the joys and pains of grad school, in the Plant Pathology department: Renuka, Muditha, Christie, Jeremiah, and Dipak; and in Mount Vernon: Emily (for being like a sister to me), Karen, Marianne, Sacha, Jeff, Lucas, and Paul. To my Filipino community and friends in Pullman, for helping me get through homesickness and survive being away from home: Ate Janet and family, Medy and Dan, Greg and Yeyet, Anne, Teddy, Walter, Jeff, Chrome, James, and Joseph. To Barb, the person I ran to when I needed strength to get through a difficult day, and for always putting difficult situations in perspective. My family, for giving me the freedom to explore life on my own, and allowing me to take a very ambitious path, with assurance of their love every step of the way. To my parents, Dado and Vivian whom I love dearly; my siblings, Vida, May, and Ryan; my niece Milcah and nephew Chio, thank you for all your love, prayers, and support. All glory and praise to the Almighty God, for without Him all is in vain. iv MANAGEMENT OF DAMPING-OFF CAUSED BY PYTHIUM SPP. IN ORGANIC VEGETABLE PRODUCTION IN THE PACIFIC NORTHWEST Abstract by Ana Vida C. Alcala, Ph.D. Washington State University July 2013 Chair: Lindsey J. du Toit Vegetable production is an important sector of the organic food industry in the Pacific Northwest USA. Significant acres of organic vegetable crops, particularly processing green pea and sweet corn, are grown in the semi-arid Columbia Basin of central Washington and north central Oregon, where Pythium damping-off can cause losses in early spring when cool and wet soil conditions are conducive to the disease. In this study, 37 certified organic fields were surveyed in the Columbia Basin for Pythium species, from which 305 isolates were baited and identified to 19 species. Pathogenicity tests of isolates of each species on pea in cool and wet soil conditions revealed isolates of 9 species were pathogenic, with differences in virulence among species and among isolates within species. Pythium ultimum (24.6% of the 305 isolates), P. irregulare group 1 (15.1%), and P. abappressorium (4.9%) were the most prevalent pathogenic species. Real-time PCR assays detected P. ultimum in 100% of the 37 fields compared to 78% for P. abappressorium, and 57% for P. irregulare group 1. Given the lack of highly effective seed treatments for damping-off control in organic production, organic seed and drench treatments were evaluated in five pea field trials in the v Columbia Basin, and two pea trials plus one sweet corn trial in maritime western Washington in 2011-12. Nordox seed treatment and seed priming demonstrated the greatest potential for damping-off control. Pea seed priming with 16 h of seed soaking + 10 h of air drying optimized priming for rapid emergence. In two field trials in 2012, combining Nordox seed treatment with seed priming using biochar as an alternative to air-drying, was promising for damping-off control. Seed exudates produced during germination are measured using electrical conductivity (EC). The EC levels of 17 pea seed lots (six cultivars) significantly affected emergence and susceptibility to Pythium damping-off. A negative linear relationship was demonstrated between EC level and emergence, with a stronger regression at higher inoculum levels of P. ultimum. Measuring the EC of pea seed lots, and quantifying Pythium inoculum in soil using real-time PCR assays may enable organic pea growers to assess accurately the risk of damping-off. vi TABLE OF CONTENTS Page ACKNOWLEDGEMENTS………………………………………………………………. iii ABSTRACT……………………………………………………………………………… v LIST OF APPENDICES………………………………………………………………… x LIST OF TABLES……………………………………………………………………….. xi LIST OF FIGURES……………………………………………………………………… xiii CHAPTER 1. Literature review 1.1. Organic vegetable production in the Pacific Northwest..…….……….…. 1 1.2. Organic pea production in the PNW…………..……………...…………. 3 1.3. Damping-off in organic production …………….………………………. 5 1.3.1. Pythium spp………………………………………………….……. 7 1.3.2. Isolation, identification, and quantification of Pythium spp.……… 12 1.3.3. Damping-off management in organic crop production.……..……. 15 1.4. Electrolyte leakage ………………………...…………………………..... 22 1.5. Conclusion and research needs…..……...……………………………..... 24 1.6. Literature cited……...…………………………...………………………. 25 2. Survey of Pythium species associated with damping-off of pea in certified organic fields in the Columbia Basin of central Washington 2.1. Introduction……………………………………………………………... 45 2.2. Materials and methods………………………………………………….. 52 2.2.1. Fields surveyed……………………………………………………. 52 vii 2.2.2. Pathogenicity tests……………………………………………….... 56 2.2.3. Quantification of Pythium spp. by real-time PCR assays………… 62 2.3. Results………………………………………………………………….. 65 2.3.1. Survey of Pythium spp…………………………………………… 65 2.3.2. Pathogenicity tests………………………………………………... 68 2.3.3. Quantification of Pythium spp. by real-time PCR assays……….... 73 2.4. Discussion……………………………………………………………… 77 2.5. Literature cited………………………………………………………… 87 3. Evaluation of seed and drench treatments for the control of damping-off in organic pea production in the Pacific Northwest 3.1. Introduction…………………………………………………………….. 128 3.2. Materials and methods…………………………………………………. 135 3.2.1. 2011 Field trials…………………………………………………. 135 3.2.2. Seed priming……………………………………………………… 144 3.2.3. 2012 Field trials…………………………………………………... 145 3.2.4. Data analyses……………………………………………………… 151 3.3. Results…………………………………………………………………... 152 3.3.1. 2011 Field trials…………………………………………………... 152 3.3.1.1. Environmental data……………………………………... 152 3.3.1.2. Pea trials………………………………………………… 153 3.3.1.3 Sweet corn trial………………………………………….. 158 3.3.2. Optimization of the priming protocol for pea seeds……………… 159 3.3.3. 2012 Field trials……………………………………………....…... 163 viii 3.3.3.1. Environmental data……………………………………... 163 3.3.3.2. Seed electrical conductivity (EC) test and germination assays for 2012 trials……………………………………. 165 3.3.3.3. Ephrata and Mount Vernon trials..…………………..…. 167 3.3.3.4. Royal City trials………………….….…....……………. 171 3.4. Discussion……………………………………………....……………… 174 3.5. Literature cited………………………………………………………… 185 4. Influence of electrical conductivity (EC) of pea on Pythium damping-off 4.1. Introduction…………………………………………………………….. 228 4.2. Materials and methods…………………………………………………. 232 4.2.1 Growth chamber trials…………………………………………….. 232 4.2.2. Exudate bioassay…………………………………………………. 236 4.3. Results………………………………………………………………….
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