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A STREAMLINED Synthetlc OCTOPLOID SYSTEM THAT A STREAMLINED SYNTHETlC OCTOPLOID SYSTEM THAT EMPHASIZES FRAGARIA VESCA AS A BRIDGE SPEClES A Thesis Presented to The Faculty of Graduate Studies of The University of Guelph by ROBERT HAROLD BORS In partial fuifilment of requirements for the degree of Doctor of Philosophy Robert Bors O June, 2000 National Library Bibliothèque nationale l*m of Canada du Canada Acquisitions and Acquisitions et Bibliographie Services services bibliographiques 395 Wellington Street 395, nie Wellington OttawaON KIAON4 Ottawa ON K1 A ON4 Canada Canada The author has granted a non- L'auteur a accordé une licence non exclusive licence allowing the exclusive permettant à la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or sel1 reproduire, prêter, distribuer ou copies of this thesis in microform, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/film, de reproduction sur papier ou sur format électronique. The author retains ownership of the L'auteur conserve la propriété du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantial extracts fiom it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. ABSTRACT A STREAMLINEO SYNTHETIC OCTOPLOID SYSTEM T HAT EMPHASIZES FRAGAR~AVESCA AS A BRIDGE SPECIES Robert Harold Bors Advisor: 3. A. Sullivan University of Guelph. 2000 The synthetic octoploid (SO) system is a method of combining diploid, tetraploid and hexaploid Fragaha species into hybrid octoploids for introgression into octoploid strawberry cultivars of F.x ananassa. I n th is stud y, (diplo id x diploid) and (diploid x hexaploid) crosses were used to create tetraploids followed by (tetraploid x tetraploid) crosses to create SOS from a wide germplasm base. Techniques refined for the SO system included Row cytometry protocols for identification of ploidy levels, in vitro germination and in vitro delivery of colchicine using the dropper method. Of the 36 species combinations investigated, 20 resulted in viable F, plants, 13 bloomed and produced seeds, and seven did not bloom. When nine species were intercrossed at the diploid level, the best crossability occurred with F.pentaphylla and F.vesca; each fomed fertile Ç, hybrids with five other species. No sig nificant differences in interspecific crossability were found among F-vesca subspecies bractacea, americana, vesca and var. sempeflorens when pollinated with Enilgerensis, F.nubico/a, Epentaphylla, and Eviridis. These hybrids were treated with colchicine to create tetraploids. Tetraploids were also created from Fmoschata (6x) hybridized with F.viridis or F mbicola. The crossing strategy at the tetraploid level ernphasized F. vesca as a bridge species for F.nilgerrensis. Enubicola, F.pentaphy/la, and F.viridis. A collection of 1O4 tetraploids inct uding (F. vesca x diploid) hybrids, F. vexa (4x), F.onentalis, (F.nubicola x Fmoschata), (Fmoschata x F.vindis), [F.orientalis x, (Fmoschata x diploid species)] and (F.viridis x F.nubicola) were intercrossed and doubled with colchicine to create SOS. When the (F.vesca x diploid species) colchi-tetraploids were intercrossed, only 37% fruit set with an average of three seedslflower. When these colchiploids were pollinated with tetraploids derived from F.orientaîis or (Emoschata x diploid species), 52% fruit set and 10 seeds/flower were obtained. A total of 192 SOSwere created from 25 combinations of species. Of the 98 Sots planted in 1997, 28 produced fruit in 1998. The seven species successfully incorporated into SOSwere F.vesca, F.nilgerrensis, F.nubicola, F.pentaphylla, F-viridis, F. orientalis, and F.moschata. The results suggests Fragana species are closely related and have great potential for breeding at the diploid and tetraploid levels and for introgression into 8x material. ACKNOWLEDGEMENTS I wish to express my sincere appreciation to my wife, Loretta, and my children, Kirk, Aurora, and Nicholas who made many sacrifices to allow me to continue rny studies. Their support and enthusiasm was invaluable. I gratefully acknowledge my advisor, Dr. J. Alan Sullivan for his guidance, support. and encouragement over the entire period of my Ph.D. program. I am tremendously indebted to committee member Dr. K. Kasha whose Cytogenetics course formed the theoretical backbone on which this thesis was inspired. I thank my advisory cornmittee members Drs. D. Wolyn and T. Michaels for their valuable suggestions and encouragement. I acknowledge Sue Couling and Ron Dutton for assisting the care and culture of field and greenhouse plants. The technical expertise of Jan Brazelot, Dr. Peter Pauls. and Shawn Rogers is appreciated in utilizing flow cytometry in this thesis. TABLE OF CONTENTS ACKNOWLEDGMENTS ..................................i TABLE OF CONTENTS ................................. ii LIST OF TABLES ......................................vi LIST OF FIGURES .....................................x LIST OF APPENDICES ................................xii CHAPTER 1: LITERATURE REVIEW ......................1 INTRODUCTION ........................................... 1 FRAGARIA SPECIES ...................................1 INTERSPECIFIC CROSSABILITY IN FRAGARIA ................. 7 Crossability arnong diploid species ...................... 14 lncompatibility mechanisms in diploid Fragah ............. 17 Crossability arnong polyploids ........................... 19 Outcrossing mechanisms in polyploids ................... 22 Crossability among dipioids and polyploids ................ 24 Surnmary of interspecific crossability ..................... 26 GENOMlC INVESTIGATIONS ................................ 26 Karyotyping ..........................................27 Investigation of chloroplast DNA ......................... 28 Investigations of nuclear DNA ...........................29 Isozyme studies ..................................... 30 Chromosome pairing in diploid hybrids .................... 32 Chromosome pairing in tetraploid hybrids .................. 32 Chromosome pairing in octoploid cultivars ................. 33 Genomic designations proposed for Fragaria species ........ 34 Genomic investigations and introgression strategy ........... 35 Pivotal Genome Theory ............................... 37 ADVANTAGES OF USlNG FRAGARIA VESCA AS A BRIDGE SPEClES ................................................39 Introduction ......................................... 39 F.vesca is widely adapted and rnay be a progenitor species ... 39 F . vesca is self-compatible ............................. 41 Some forms of F. vesca are adapted to cultivation ........... 42 Fragana vesca var . sempeflorens has useful traits that facilitate breeding ..................................... 42 Fruitquality ......................................... 43 Lack of inbreeding depression .......................... 43 F . vesca is amenable to chromosome doubling ............. 44 F. vexa cytoplasrn .................................. 44 F-vesca as a bridge species ............................45 INTROGRESSION OF FRAGARIA SPECIES USlNG SYNTHETIC OCTOPLOID SYSTEMS .............................. 46 Crossing strategies to create SOS ........................47 lmproving SO production ........................ 55 lmproved strategy for SO production ..................... 57 CHAPTER 2: GENERAL MATERIALS AND METHODS ....... 59 Introduction .............................................. 59 ln vitro germination ........................................ 59 Colchicine treatment for chromosome doubling ................... 60 Acclimatization of seedlings ..................................61 Handling of parent plants for crossing ..........................63 Floral induction ............................................ 63 Pollination ............................................... 64 Organization of crosses .....................................66 Seed collection and storage .................................. 66 Data collection and analysis ................................. 66 Visual screening of plants following colchicine treatment ........... 67 Flow cytometry protocol ..................................... 68 Flow cytometry standards and the estimation of ploidy level .........74 CHAPTER 3: FRAGARIA SPECIES DIFFER IN DNA CONTENT 78 ABSTRACT ..............................................78 INTRODUCTlON ........................................ 79 MATERIALS AND METHODS ................................ 80 RESULTS AND DISCUSSION ............................... 83 CONCLUSION ............................................ 89 CHAPTER 4: HYBRlDlZATlON OF FRAGARIA VESCA SUBSPECIES AS FEMALE PARENTS WlTH F.NlLGERRENSIS, F*NUBICOLA. F.PENTAPHYLLA AND F*VIRIDIS rn m a 91 ABSTRACT ..............................................91 INTRODUCTION ..........................................92 Advantages of using Evesca as a bridge species ........... 93 Advantages of using F.vesca as a female parent ............ 95 Evidence that F .vesca subspecies have diverged ........... 97 Goals of this study .................................... 98 MATERIAL AND METHODS .................................98 Experimental design and data analysis ................... 102 RESULTS ..............................................103 Fruit set (%) and seeds/fiower ......................... 104 Seeds with embryos and embryo germination ............. 109 Differences in intraspecific crosses ......................109
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