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Germplasm Collection, Characterization, and Enhancement of Eastern Phlox Species

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Germplasm Collection, Characterization, and Enhancement of Eastern Phlox Species GERMPLASM COLLECTION, CHARACTERIZATION, AND ENHANCEMENT OF EASTERN PHLOX SPECIES DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy Graduate School of The Ohio State University By Peter Jeffrey Zale, M.S. Graduate Program in Horticulture and Crop Science The Ohio State University 2014 Dissertation Committee: Dr. Pablo Jourdan, Advisor Dr. Mark Bennett Dr. David Francis Dr. John Freudenstein Copyrighted by Peter J. Zale 2014 ABSTRACT Plants from the genus Phlox have become a staple of gardens worldwide since their introduction to cultivation about 200 years ago, and are admired as versatile garden and landscape plants, container plants, and cut flowers. Their long-lasting, intensive floral displays in a variety of colors, forms, and seasons rank them among the most widely recognized hardy perennials and annuals. The floral beauty has resulted in extensive breeding and selection in three primary species: the annual Phlox drummondii, and the hardy perennials P. paniculata, and P. subulata. Among these three, hundreds of cultivars have been selected over the last 200 years, but the genus includes other species that may have ornamental potential. Phlox L. (Polemoniaceae) consists of ca. 65 species primarily endemic to North America; 20-23 species occur in the eastern U.S.A. and 40-45 in the west. The western taxa occur primarily in arid, mountainous habitats whereas the eastern taxa occur in a different range of diverse ecosystems and habitats. The eastern species are a polymorphic group arranged into 6 subsections within the genus; this group includes the three main cultivated species and also up to 20 related species that exhibit ornamental characteristics, yet are rarely cultivated. Horticultural and botanical interest in Phlox has resulted in a large volume of literature pertaining to the taxonomy, ecology, and cultivation, but there is ii recognition that the genetic diversity of the genus has barely been tapped for ornamental use or as a source of desirable traits, such as disease resistance, and that more widespread availability of diverse germplasm can contribute not only to new cultivated forms but also to a greater understanding of species diversity and interspecific relationships. Such interest has made Phlox a priority genus for germplasm development and conservation at the Ornamental Plant Germplasm Center of The Ohio State University. The work presented here describes the development, characterization and manipulation of such germplasm. The development of a Phlox germplasm collection was initiated in 2010 with an effort to collect seeds and plants of the eastern species from natural populations throughout their distributional range. A total of 187 accessions were collected from wild populations of 22 eastern Phlox species during a series of expeditions to diverse ecosystems throughout the eastern United States. Another 166 accessions were of cultivated origin, acquired from nurseries and individuals; these represented a mix of selections from wild populations, selected seedlings, or hybrids and were used for comparison to wild-collected accessions. The 353 accessions represent the most comprehensive germplasm collection of Phlox to date that include taxa related to the three horticulturally important species. An additional 8 species of western USA Phlox were also acquired as a source of comparative material. This germplasm was characterized in three primary ways. The characterization of the germplasm collection focused on estimation of genome size using flow cytometry, coupled with validation of ploidy estimates by iii chromosome counts. Genome size was surveyed in 287 accessions derived from 29 species (45 taxa); of these, 165 accessions were of wild origin and the rest were cultivated material. The majority of taxa were diploid, but genome size was variable. Genome size consistent with tetraploid and hexaploid levels were found in species which had not been previously reported. Polyploidy in natural Phlox populations seems to occur primarily in populations at the distribution edges suggesting that it may be associated with genetic differentiation, diversification, and adaptation to marginal habitats. The 122 cultivars and hybrids analyzed for genome size were also primarily diploid, but anueploids were discovered in P. paniculata suggesting the use of polyploid breeding in this species. These data have important implications in the conservation, evolution, adaptation, and commercial breeding of Phlox. Characterization of the germplasm collection also included a limited application of microsatellite (SSR) marker analysis to 8 populations, 5 diploid and 3 tetraploid, and 61 individuals from the P. pilosa complex. The data indicated that tested populations of P. pilosa had high genetic diversity and moderate population structuring, typical of outcrossing species with gametophytic self - incompatibility. These preliminary data indicate that P. pilosa is a rich source of genetic diversity for Phlox germplasm collections and enhancement. The manipulation of germplasm involved a series of interspecific hybridization experiments; in two studies there was at least one recurring parent and the other study was performed in a partial diallel. A total of 157 unique hybrids were produced. Interspecific hybridization was most successful among closely related iv species, and the rate of success decreased with increasing phylogenetic distance; however, even crosses between closely related species failed to produce seeds, and indicate that selection of parental taxa is critical. Attempts at recreating previously described hybrid combinations were of limited success, but at least one was successfully resynthesized. Previous reported attempts at interploid crosses failed, but were successful and resulted in morphologically intermediate, aneuploid taxa with intermediate genome sizes to parental taxa. In conclusion, the development of a comprehensive germplasm collection enabled a broad characterization and enhancement of eastern U.S.A Phlox. These initial studies have been necessary to provide baseline information for more focused studies pertaining to future Phlox characterization and hybridization studies. v Dedication This work is dedicated to Simon P. Zale vi ACKNOWLEDGMENTS The sum of my graduate experience has amounted to much more than the research and stories presented in the following pages, and was only possible though interactions with many advisors, colleagues, and friends that enriched the entire experience. I extend thanks to my committee members Dr. Mark Bennett, Dr. David Francis, and Dr. John Freudenstein for faithfully attending committee meetings and providing feedback and ideas. I thank my advisor, Pablo Jourdan for the privileges and autonomy I enjoyed during the research process and the extraordinary priviledge of pursuing my own interests within the context of the OPGC mandate. Many of the germplasm collection trips rank among the finest and most influential experiences of my life. Gratitude is expressed to the staff of the OPGC. In Particular, Susan Stieve, Eric Renze, Russell Eckley, and Steven Haba. Thanks for helping to create an environment conducive to my research. Two student workers, Elizabeth Reeder and Marcus Nichols, were intimately involved in Phlox research and magnitude of what was accomplished would not have been possible without them, and their devotion to thhe numerous facets of the project. My deepest gratitude is expressed to my wife, Kate, for supporting my schooling for the last 7 years. I have been in graduate school longer than we have been married, vii and the completion of this document is also the delivery of a promise made long ago. I thank my family for their unwavering support of my pursuit. Many individuals helped me obtain germplasm samples, permits, and/or provided information, discussions, greenhouse space, and inspiration that were vital the synthesis of this project. Thanks are extended to my friend Dr. Daniel W.H. Robarts, who was a common companion in the field and lab, and linked with many of my best memories of graduate school. I thank Dr. Dan Struve, Charles and Martha Oliver, Dr. Warren Stoutamire, Jason Parrish, Julian Campbell, Ron Determann, Matt Richards, Tom Mitchell, Aaron Floden, Mike Graziano, Marianne Casey, Art Evans, Jim Rodgers, Tony Avent, Carolyn Ferguson, Jim Ault, Shannon Fehlberg, Rick Gardner, Alan Blowers, Bill Martin, Michael Jenkins, Ron Miller, Brandon Sinn, Barrie Francis, Rimmer de Vries, Jim Vent, David Snodgrass. Several institutions contributed to this work in numerous ways and I thank the following; The staff at the The Columbus Metroparks, Atlanta Botanical Garden, The Cincinnati Zoo and Botanical Garden, The Greater Des Moines Botanical Center, Oak Openings preserve, LadyBird Johnson Wildflower Center, Mount Cuba Center, The Ohio State University Herbarium. Many others also helped in the completion of this work and I thank you. viii VITA June 1997 . Olmsted Falls High School 1997 - 2001. B.S. Horticulture, The Ohio State University 2001 - 2007. Nursery Production and Retail Manager. Marvin’s Organic Gardens, Lebanon, Ohio 2007 - 2009. M.S., The Ohio State University 2009 -Present. Ph.D Candidate, The Ohio State University PUBLICATIONS Zale, Peter J.,and P. Jourdan. 2012. Germplasm Development and Preliminary Interspecific Hybridization in Phlox. Acta Horticulturae 953:71-78 Zale, Peter J., Struve, D.K., Jourdan, P., and D.M. Francis. 2011. Rapid assessment of genetic variation for morphological
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