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Antagonistic Co-Evolution Between a Plant and One of Its Parasites Is Commonly Portrayed As an Arms Race (Ref)

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Antagonistic Co-Evolution Between a Plant and One of Its Parasites Is Commonly Portrayed As an Arms Race (Ref) DEVELOPMENT AND CHARACTERIZATION OF WHEAT GERMPLASM FOR RESISTANCE TO STEM RUST UG99 IN WHEAT A Dissertation Submitted to the Graduate Faculty of the North Dakota State University Of Agriculture and Applied Science By Qijun Zhang In Partial Fulfillment of the Requirements For the Degree of DOCTOR OF PHILOSOPHY Major Department: Plant Science December 2013 Fargo, North Dakota North Dakota State University Graduate School Title DEVELOPMENT AND CHARACTERIZATION OF WHEAT GERMPLASM FOR RESISTANCE TO STEM RUST UG99 IN WHEAT By Qijun Zhang The Supervisory Committee certifies that this disquisition complies with North Dakota State University’s regulations and meets the accepted standards for the degree of DOCTOR OF PHILOSOPHY SUPERVISORY COMMITTEE: Steven S. Xu Chair Xiwen Cai Justin D. Faris Timothy L. Friesen Shaobin Zhong Approved: 12/20/13 Richard D. Horsley Date Department Chair ABSTRACT World wheat production is currently threated by stem rust (caused by Puccinia graminis f. sp. tritici) Ug99 race (TTKSK). The ongoing global effort to combat Ug99 is focusing on the identification and deployment of Ug99-resistant genes (Sr) into commercial cultivars. The objectives of this study were to identify TTKSK-effective Sr genes in untapped durum and common wheat germplasm and introgression of TTKSK-effective Sr genes from tetraploid wheat (Triticum turgidium) and Aegilops tauschii into hexaploids through production of synthetic hexaploid wheat (SHW). For identification of TTKSK-effective Sr genes, 177 durum and common wheat cultivars and lines were first evaluated using three highly virulent races TTKSK, TRTTF, and TTTTF and 71 cultivars and lines with TTKSK resistance were identified. The TTKSK-resistant cultivars and lines were then evaluated using six local races and the molecular markers that are diagnostic or tightly linked to the known TTKSK-effective Sr genes. The race specification and marker analysis showed that several previously deployed TTKSK-effective Sr genes such as Sr2, Sr24 and Sr42 were present in some of the cultivars and lines. A number of resistant cultivars and lines derived from wheat relatives such as Thinopyrum ponticum, Th. elongatum, Th. intermedium, and Ae. speltoides may carry novel Sr genes. For SHW development, 200 new SHW lines were developed by crossing 181 tetraploid wheat accessions to 14 Ae. tauschii accessions. Sixty-six of the new SHW lines, 14 previously-developed SHW lines, and their parents were evaluated for resistance to TTKSK, TRTTF, TTTTF and six other races and genotyped using molecular markers linked to the known genes in T. dicoccum and Ae. tauschii. The evaluation data showed that 44 SHW lines were resistant to TTKSK. The race specification and marker analysis showed that Sr2 from T. dicoccum and Sr33 from Ae. tauschii were present in some of the SHW lines and a number of SHW lines have novel genes conferring iii TTKSK resistance. The durum and wheat cultivars and lines and SHW lines with known and novel Sr genes conferring resistance to TTKSK will be useful resources for improving wheat resistance to TTKSK and other emerging races of stem rust. iv ACKNOWLEDGEMENTS I express the deepest thanks to my major adviser, Dr. Steven S. Xu, for his encouragement, support, and guidance during my graduate study in the past five years. He spent numerous hours to guide me in my thesis research and preparation. I also express my deepest appreciation to my co-advisor, Dr. Xiwen Cai, for his support and guidance in this study and to my graduate committee, Dr. Justin D. Faris, Dr. Timothy L. Friesen, and Dr. Shaobin Zhong for their assistance and reviewing this dissertation. I am highly grateful to the Department of Plant Science and North Dakota State University for providing the opportunity for me in pursuit of the Ph.D degree and to Dr. Richard D. Horsley, Dr. Michael C. Edwards, and Mrs. Eileen Buringrud for their help and assistance. I am thankful to USDA-ARS for funding the Graduate Research Assistantship. My thanks also go to Charles and Linda Moses for providing a scholarship. I express my deepest appreciations to Drs. Yue Jin, Matthew N. Rouse, Daryl L. Klindworth, Timothy L. Friesen, and Dr. Shaobin Zhong for their assistance in stem rust testing, to Drs. Shiaoman Chao, Lili Qi, Justin D. Faris, and Yunming Long for marker analysis, Drs. Chao-Chien Jan and Zhao Liu for the GISH protocol, and Dr. Elias M. Elias for providing durum wheat germplasm. I extend my sincere thanks to Jason E. Axtman, Danielle J. Holmes, Mary M. Osenga, Richard M. Sonju, Rachel McArthur, and Stan Stancyk for technical assistance. My thanks also go to Drs Zhixia Niu, Chenggen Chu, Guotai Yu, Zhaohui Liu, Zengcui Zhang, Gongjun Shi, Guojia Ma, Aifeng Liu, and Gang Feng for friendship and insightful discussion. I express my deepest gratitude to my parents and my sisters for their support and encouragement. I express my deepest appreciation to my wife for her support and help during the preparation of my thesis. v TABLE OF CONTENTS ABSTRACT ................................................................................................................................... iii ACKNOWLEDGEMENTS ............................................................................................................ v LIST OF TABLES ....................................................................................................................... viii LIST OF FIGURES ....................................................................................................................... ix LIST OF APPENDIX TABLES ..................................................................................................... x CHAPTER 1. GENERAL INTRODUCTION ............................................................................... 1 Literature Cited .................................................................................................................. 2 CHAPTER 2. LITERATURE REVIEW ........................................................................................ 4 Wheat and Wheat Gene Pools.............................................................................................4 Exploitation of Wheat Gene Pools for Germplasm Improvement ......................................6 Wheat Stem Rust...............................................................................................................13 Ug99 Lineage Races and Their Control ...........................................................................24 Literature Cited ................................................................................................................ 32 CHAPTER 3. EVALUATION AND MOLECULAR CHARACTERIZATION OF WHEAT LINES AND CULTIVARS FOR STEM RUST UG99 RESISTANCE DERIVED FROM RELATIVES OF WHEAT ................................................................................. 46 Abstract ............................................................................................................................. 46 Introduction ....................................................................................................................... 47 Methods and Materials ...................................................................................................... 50 Results ............................................................................................................................... 53 Discussion ......................................................................................................................... 61 Literature Cited ................................................................................................................. 83 CHAPTER 4. DEVELOPMENT AND CHARACTERIZATION OF SYNTHETIC HEXAPLOIDS FROM UNDER-EXPLOITED TETRAPLOIDS AS A NEW RESOURCE FOR STEM RUST UG99 RESISTANCE ...................................................90 vi Abstract ............................................................................................................................ 90 Introduction ...................................................................................................................... 91 Methods and Materials ..................................................................................................... 95 Results .............................................................................................................................. 99 Discussion ...................................................................................................................... 105 Literature Cited .............................................................................................................. 120 APPENDIX ................................................................................................................................. 127 vii LIST OF TABLES Table Page 2.1. An international nomenclature system designating P. graminis f. sp. tritici (Pgt) races ...... 15 2.2. Chromosome location and original source of Sr genes ..........................................................19 3.1. Wheat cultivars/breeding lines and their parents characterized for their resistance to stem rust ......................................................................................................................................... 72 3.2. Avirulence/virulence on the North America differentials for nine stem rust races used in this study ................................................................................................................................ 75 3.3.
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