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MOLECULAR CHARACTERIZATION of PHOTOPERIODIC FLOWERING in STRAWBERRY (Fragaria SP.)

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MOLECULAR CHARACTERIZATION of PHOTOPERIODIC FLOWERING in STRAWBERRY (Fragaria SP.) MOLECULAR CHARACTERIZATION OF PHOTOPERIODIC FLOWERING IN STRAWBERRY (Fragaria SP.) By PHILIP JACOB STEWART A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA May 2007 1 ©2007 Philip J. Stewart 2 To my parents, who always believed in me, and taught me to love knowledge and the natural world. To the memory of my grandfather, Jackson Lee Stewart, Jr., who introduced me to agriculture, and who will forever be my image of the American farmer. To my children, Zea and Adina, who have been a constant inspiration and joy to me, and reminders of the things in life that are truly important. And, most of all, to my wife, Cynthia, who has been unwavering in her faith in my abilities, in her willingness to sacrifice for my academic and professional pursuits, and, most importantly, in her love. 3 ACKNOWLEDGMENTS I would like to take this opportunity to thank at least a few of the many people who have have assisted me along the path to this degree. First, I would like to offer my love and gratitude to my parents, Lee and Faith Stewart, and my in-laws, Barton and Teri Cowan, all of whom have provided continuous support, both emotional and financial, throughout the course of my graduate studies. None of this would have been possible without them. Secondly, I would like to thank those that who have provided invaluable assistance in my academic and professional development, in particular my graduate advisors, Kevin Folta and Craig Chandler. Their help was critical to the work described here. I am also grateful for the assistance and suggestions offered by the other members of my committee: Paul Lyrene, Natalia Peres, and José Chaparro, all of whom helped to shape this study, and Daniel Sargent, of East Malling Research, U.K., for the use of his diploid mapping population. Much thanks also go to Dawn Bies and Maureen Clancy, for technical assistance, and also to my fellow students in the Folta lab: Denise Tombolato, Stefanie Maruhnich, Thelma Madzima, and Wendy Gonzalez. I’ve enjoyed working with all of you and wish you much success as you finish your studies and move into professional life. And, finally, the greatest debt of gratitude is owed my beautiful wife, Cynthia, who has not only been tolerant but encouraging as I have dragged her from place to place across the country in pursuit of this dream. Thank you so much—I hope it was worth it. 4 TABLE OF CONTENTS page ACKNOWLEDGMENTS...........................................................................................................4 LIST OF TABLES......................................................................................................................8 LIST OF FIGURES ....................................................................................................................9 ABSTRACT .............................................................................................................................11 CHAPTER 1 INTRODUCTION .............................................................................................................12 2 LITERATURE REVIEW...................................................................................................14 The genus Fragaria ...........................................................................................................14 Polyploidy in Strawberries .................................................................................................15 Flowering Habits in Strawberry .........................................................................................16 Short-Day Strawberries...............................................................................................16 Remontant Strawberries ..............................................................................................18 Factors Affecting Expression Of Photoperiodic Flowering.................................................23 Temperature Effects On Flowering .............................................................................23 Vernalization Requirements ........................................................................................24 Juvenility and Plant Age Effects..................................................................................25 Growth Pattern Differences Between Plants Of Differing Flowering Habits.......................26 Inheritance Of Flowering Habit In Strawberry ...................................................................27 Molecular Markers For Flowering Habit ............................................................................32 Molecular Control Of Flowering In Arabidopsis ................................................................33 Components Upstream of CO......................................................................................34 Components Downstream of CO.................................................................................35 The CONSTANS-LIKE Gene Family .........................................................................36 Components of the Flowering Pathways are Conserved Among Species .....................38 Using Model Systems To Understand Development In Rosaceae ................................40 3 CHARACTERIZATION OF CONSTANS-LIKE GENES IN Fragaria ..............................44 Introduction .......................................................................................................................44 Results...............................................................................................................................46 Cloning And Identification Of Four CONSTANS-LIKE Genes Of Fragaria...............46 FrCO/FrCOL Gene Structure......................................................................................48 Southern blot analysis of FrCO Copy Number............................................................50 Mapping Of Fragaria CO And COL Genes In A Diploid Mapping Population ...........50 Comparison Of Strawberry Genotypes Under Varying Photoperiods...........................51 Allele-Specific Expression Patterns Differ Betwen Short-Day And Day-Neutral Genotypes ...............................................................................................................52 5 Expression Of Other Genes Under Different Photoperiods ..........................................53 Some FraCOL2 Transcripts Contain An Unspliced Intron...........................................54 Discussion .........................................................................................................................54 The Co Gene Family In Fragaria (Rosaceae)..............................................................54 Evolutionary Relationships Among FrCO Sequences..................................................58 FrCO Expression ........................................................................................................61 Expression of Other COL Genes .................................................................................65 Materials and Methods.......................................................................................................68 Plant Material .............................................................................................................68 Plant Growth Conditions.............................................................................................69 Cloning of FraCO.......................................................................................................70 Identification of COL Genes in Genebank...................................................................70 Gene Structure Characterization..................................................................................70 Southern Blot Analysis ...............................................................................................71 Genetic Linkage Mapping COL genes in Fragaria......................................................71 Extraction of Nucleic Acids ........................................................................................72 Reverse Transcription and RT- PCR ...........................................................................73 4 IDENTIFICATION AND CHARACTERIZATION OF FLOWERING-RELATED GENES IN Fragaria AND OTHER ROSACEAE .............................................................93 Introduction .......................................................................................................................93 Results...............................................................................................................................94 Identification of Rosaceae Flowering Gene Homologs ................................................94 FrSOC1 Expression ....................................................................................................97 FrTFL1 and FrLFY Expression...................................................................................97 Discussion .........................................................................................................................98 Rosaceae MADS-Box Genes ......................................................................................98 The FT/TFL1 Gene Family in Rosaceae.................................................................... 101 Other Flowering Genes ............................................................................................. 105 FrSOC1, FrLFY, and FrTFL1 Expression
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