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Fusarium Graminearum and Its Trichothecene Chemotype Diversity

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Fusarium Graminearum and Its Trichothecene Chemotype Diversity THE CONTINUING BATTLE BETWEEN WHEAT AND FUSARIUM GRAMINEARUM: UNDERSTANDING THE MOLECULAR PHYLOGENETIC RELATIONSHIPS, CHEMOTYPE DIVERSITY AND TRICHOTHECENE BIOSYNTHESIS GENE EXPRESSION PATTERNS By CHAMI CHATHURANGI AMARASINGHE A Thesis submitted to the Faculty of Graduate Studies of The University of Manitoba In Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY Department of Plant Science University of Manitoba Winnipeg, Manitoba Canada Copyright ©2016 by CHAMI CHATHURANGI AMARASINGHE ABSTRACT Fusarium head blight (FHB) continues to threaten the economic sustainability of wheat and barley production in Canada and worldwide. The overall goal of this thesis is to expand our current knowledge of the FHB pathogen, Fusarium graminearum and its trichothecene chemotype diversity. Continuous monitoring of trichothecene chemotypes may well inform on the potential risk and the type of Fusarium populations present in a given region. Fusarium populations in Winnipeg and Carman, Manitoba were examined using chemotype as a marker in the field. Rapid expansion of the 3-acetyldeoxynivalenol (3-ADON) chemotype was observed in Winnipeg and Carman. 3-ADON chemotype is consistently found at high frequencies over the previously common 15-acetyldeoxynivalenol (15-ADON) chemotype, suggesting that the shift in pathogen populations is continuing. This study provides the first evidence on the presence of nivalenol (NIV) producing F. cerealis strains in winter wheat in Manitoba, Canada. Therefore, discovery of NIV producing F. cerealis in wheat poses a serious concern for the wheat industry in Canada. Phylogenetic, chemotypic, phenotypic, and pathogenic abilities of 150 strains of F. graminearum species complex (FGSC) from eight countries were investigated. Type and amount of trichothecenes produced by a strain are key factors in determining the level of aggressiveness of that strain regardless of its species origin. The sequence variations of TRI8 gene in different species in the FGSC were examined as Fusarium species may produce different types of trichothecenes depending on differences in the core trichothecene (TRI) cluster genes. The TRI8 haplotypes did group according to chemotype rather than by species, indicating that 3-ADON, 15-ADON and NIV chemotypes have a single evolutionary origin. Comparison of TRI gene expression demonstrated that accumulation of TRI transcripts was higher in 3-ADON producing strains compared to 15-ADON and NIV strains. The presence of masked mycotoxins ii deoxynivalenol-3-glucoside (D3G) in food and feed is an increasing concern. Canadian spring wheat cultivars inoculated with different chemotypes produce D3G upon Fusarium infection and moderately resistant/intermediate cultivars showed higher D3G/DON ratio compared to susceptible cultivars. iii ACKNOWLEDGEMENTS I gratefully acknowledge my supervisor Dr. Dilantha Fernando for his valuable guidance, mentorship, and advice. His strong support and continuous encouragement motivated me throughout my study leading to successful completion of my Ph. D. project. Also I would like to thank Dr. Fernando, for accepting me for the Fusarium project in his lab. His advice, own experience and guidance helped me to get through so many difficult situations. I was fortunate to have an advisor like you in my life. I extend my sincere appreciation to Drs. Anita Brûlé-Babel, Barbara Sharanowski and Michele Piercey-Normore for serving as my Ph.D. committee members and for their invaluable guidance, support, and the critical review of the thesis. Also I would like to thank my external examiner Dr. Ravindra Chibbar for his valuable comments and revisions on my thesis. I wish to thank Dr. Sheryl Tittlemier (Canadian Grain Commission, Winnipeg) and Dr. Senay Simsek (North Dakota State University, Fargo, USA) for providing facilities to carry out mycotoxin analysis in their labs. I would like to acknowledge Drs. Thomas Miedaner (Universitaet hohenheim, Stuttgrat, Germany), Malgorzata Jedryczka (Polish Academy of Science, Pozan, Poland), Philip Jennings (The food and environment research agency, York, UK), Pawan Kumar Singh (CIMMYT, Mexico), Silvia Barcellos Rosa (Melhoramento de Sementes Ltd. , Brazil), Kemal Kazan (CISRO plant industry, Brisbane, Australia), Altus Viljoen (Stellenbosch University, Stellenbosch , South Africa), Rubella Goswami (North Dakota State University, Fargo, USA), Nancy J. Alexander (United States Department of Agriculture, Peoria, Illinois, USA), Sussane Vogelgsang (Federal Department of Economic Affairs FDEA, Zurich, Switzerland), Jianhua Wang (Huazhong agricultural University, Hubei, China), Tapani Yli-Mattila (University of iv Turku, Turku, Finland), Friday Obanor (CISRO Plant Industry, Brisbane, Australia) and Julie Toussaint-Ferreyrolle (Arvalis Institut du Végétal, INRA, France) for generously providing Fusarium strains for my research. Also I would like to thank technicians, Ms. Paula Parks, Ms. Mary Meleshko, Mr. Roger Larios, Mr. Lorne Adams, Mr. Doug Durninn, Ms. Eppie Austria and research associates Dr. Maria Stoimenova and Dr. Arvind Hirani (Dept. of Plant Science, University of Manitoba) for their assistance in both lab and field experiments. I wish to thank all the members of support and administration staff of the Department of Plant Science, University of Manitoba for their kindness and support during my studies. Special thanks to my past and present lab mates for their company, especially my dear friends, Ru Li and Xuehua. A friendly chat with you was priceless for me to share my happy and difficult times. I would like to express my gratitude to the Canadian Wheat Board and the University of Manitoba Graduate Fellowship for funding my doctoral scholarships. Also I would like to acknowledge the funding provided by Agri-Food Research and Development Initiative (ARDI) Manitoba and Western Grains Research Foundation (WGRF). Last but not least, I wish to thank my Amma, Thaththa, Sajeewa and my sister for their continuous support, patience and encouragement. v DEDICATION Every challenging work needs encouragment, support & love from those who were very close to our hearts My Ever Loving Parents, My Sister, My Husband Sajeewa & Our Cute Little Baby Girl Naylie You were there for me every step of this challenging journey. You are the pillars of my success !!! vi TABLE OF CONTENTS ABSTRACT ................................................................................................................................... ii ACKNOWLEDGEMENTS ........................................................................................................ iv DEDICATION.............................................................................................................................. vi LIST OF TABLES ...................................................................................................................... xii LIST OF FIGURES ................................................................................................................... xiii LIST OF APPENDICES ........................................................................................................... xvi FOREWORD............................................................................................................................. xvii CHAPTER 1 .................................................................................................................................. 1 GENERAL INTRODUCTION .................................................................................................... 1 CHAPTER 2 .................................................................................................................................. 6 LITERATURE REVIEW ............................................................................................................ 6 2.1 The Disease- Fusarium head blight ....................................................................................... 6 2.2 The Host - Wheat .................................................................................................................. 7 2.2.1 Wheat production in Canada .......................................................................................... 7 2.3 The Pathogen- Fusarium graminearum- A brief overview .................................................. 8 2.4 Taxonomy of the genus Fusarium ........................................................................................ 9 2.5 Life cycle and symptoms of FHB ....................................................................................... 12 2.5.1 Pathways of Fusarium infection in wheat .................................................................... 15 2.6 Fusarium mycotoxins- Trichothecenes ............................................................................... 17 2.6.1 Trichothecene biosynthesis........................................................................................... 19 2.6.2 Biological activity of trichothecenes in animals........................................................... 23 2.6.3 Biological activity of trichothecenes in plants ............................................................. 25 2.6.4 Role of deoxynivalenol in Fusarium pathogenicity ..................................................... 26 2.6.5 Detoxification of DON in plants .................................................................................. 27 2.7 Chemotypes of F. graminearum species complex .............................................................. 29 2.7.1
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