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Characterization of a Spontaneous Phaseolus Vulgaris Mutant

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Characterization of a Spontaneous Phaseolus Vulgaris Mutant CHARACTERIZATION OF A SPONTANEOUS PHASEOLUS VULGARIS MUTANT WITH THE ABILITY TO SELECTIVELY RESTRICT NODULATION A dissertation presented to the faculty of the College of Arts and Sciences of Ohio University In partial fulfillment of the requirements for the degree Doctor of Philosophy Sarah Laity Bashore August 2006 This dissertation entitled CHARACTERIZATION OF A SPONTANEOUS PHASEOLUS VULGARIS MUTANT WITH THE ABILITY TO SELECTIVELY RESTRICT NODULATION by SARAH LAITY BASHORE has been approved for the Department of Biological Sciences and the College of Arts and Sciences by Allan M. Showalter Professor of Environmental and Plant Biology Benjamin M. Ogles Dean, College of Arts and Sciences Abstract BASHORE, SARAH, Ph.D., August 2006. Molecular and Cellular Biology CHARACTERIZATION OF A SPONTANEOUS PHASEOLUS VULGARIS MUTANT WITH THE ABILITY TO SELECTIVELY RESTRICT NODULATION (195 pp.) Director of Dissertation: Allan M. Showalter A spontaneous Phaseolus vulgaris mutant was isolated that selectively restricts nodulation. This recessive mutation exhibits a phenotype that has never been seen before with beans and is a perfect tool to study the symbiotic relationship and the associated signaling molecules between legumes and Rhizobia. The goal of this dissertation was to characterize the mutant bean’s morphology and nodulation capacity. Rhizobial screening was done to examine how many different strains of Rhizobia were able to nodulate the mutant bean. The bean was examined for phenotypic characteristics and then examined for how the mutation was affecting nodulation. This was done by using green fluorescently labeled bacteria to visualize steps in the nodulation process and by chemical isolation and characterization of the signals involved in forming the symbiosis. This research also examined the overall competitiveness of strains with the ability to nodulate the mutant bean. A final experiment used Tn5 mutagenesis of USDA 2669 to determine if any novel signaling molecules were present in the excluded strain. It was determined that the mutant P. vulgaris had no deleterious phenotypic characteristics and that three strains of Rhizobia, USDA 9017, USDA 9032 and USDA 9041, had the ability to nodulate the mutant. It was also demonstrated that the mutation blocked nodulation before the formation of infection threads and therefore was affecting the plant's perception of the bacterial signal. Approved: Allan M. Showalter Professor of Environmental and Plant Biology Acknowledgments I would like to express my sincere gratitude to Dr. Art. T. Trese, my advisor, my mentor and my friend, who has always stood by me in times of doubt. I would also like to acknowledge Dr. Allan Showalter who served as my dissertation advisor and has helped me through a strange and difficult time. I would also like to acknowledge Dr. Ivan K. Smith and Dr. Marcia J. Kieliszewski for their encouragement and support. I would like to thank Harold Blazier as well as Aaron Matthers, for their technical assistance, and Kurt Hartman for his review of my statistical analysis. I am also grateful for the help and support of the faculty and staff in the Molecular and Cellular Biology Interdisciplinary Program as well as the Plant Biology Department at Ohio University. Financial support from the Molecular and Cellular Biology Program as well as Sigma Xi, Houk Grant and Graduate Student Senate was greatly appreciated. Special thanks goes to my husband, Paul D. Meiss, III and my family, Sam and Marion Bashore, Melissa and Doug Blacksmith, Elizabeth Bashore, Zach Meiss and Lily Meiss who have suffered through this almost as much as I have. They have never stopped supporting and loving me and for that I am eternally grateful and full of love. 6 Table of Contents Page Abstract................................................................................................................................3 Acknowledgements..............................................................................................................5 List of Tables .......................................................................................................................9 List of Figures....................................................................................................................10 List of Abbreviations .........................................................................................................12 Chapter 1. Introduction .....................................................................................................13 1.1 The Symbiotic relationship between Phaseolus vulgaris and Rhizobium sp.........14 1.2 Phaseolus vulgaris L. phaseoli ..............................................................................15 1.3 Genetics of P. vulgaris...........................................................................................19 1.4 Rhizobia sp.............................................................................................................20 1.5 Rhizosphere as a habitat for Rhizobia ...................................................................24 1.6 Genetics of Rhizobia..............................................................................................26 1.7 Flavonoids..............................................................................................................28 1.8 Lipo-chitin oligosaccharides..................................................................................30 1.9 Plant response to LCOs .........................................................................................35 1.10 Plant recognition ..................................................................................................37 1.11 Nodule formation.................................................................................................40 1.12 Nitrogen cycle......................................................................................................44 1.13 Nitrogen fixation by Rhizobia .............................................................................49 1.14 Mutants that restrict nodulation ...........................................................................50 1.15 The mutant P.vulgaris..........................................................................................51 1.16 Objective..............................................................................................................52 Chapter 2. Rhizobial Screening to Examine Nodulation Capacity of Nod- Mutant Phaseolus vulgaris............................................................................................................56 Summary.......................................................................................................................57 Introduction...................................................................................................................58 Materials and Methods..................................................................................................62 Growth of P. vulgaris ..............................................................................................62 Growth of Rhizobium species ..................................................................................63 Inoculation of P. vulgaris ........................................................................................63 Jensen’s reagent .......................................................................................................63 Plant growth.............................................................................................................65 Harvest and identification of positive inoculation...................................................65 Results...........................................................................................................................66 Discussion.....................................................................................................................71 7 Chapter 3. Phenotypic, Developmental and Morphological Differences between a Nodulation of the Nod- P. vulgaris and the Wild-type.....................................74 Summary.......................................................................................................................75 Introduction...................................................................................................................76 Materials and Methods..................................................................................................78 TY media .................................................................................................................78 Analysis of nitrogen content: Elementar C:N Analyzer ..........................................79 Results...........................................................................................................................79 Discussion.....................................................................................................................85 Chapter 4. Mechanism of Nodulation in Nod- P. vulgaris. ..............................................89 Summary.......................................................................................................................90 Introduction...................................................................................................................91 Materials and Methods..................................................................................................94 GFP transformation..................................................................................................94
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