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The Evolution and Flnctional Significance of Stigma

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The Evolution and Flnctional Significance of Stigma THE EVOLUTION AND FLNCTIONAL SIGNIFICANCE OF STIGMA-HEIGHT DIMORPHISM IN FLOWERING PLANTS Angela Marie Baker A thesis subrnitted in conforrnity with the requirements for the degree of Doctorate of Philosophy Graduate Department of Botany University of Toronto O Copyright by Angela Marie Baker 2000 National Library Bibliothèque nationale 1*1 of Canada du Canada Acquisitions and Acquisitions et Bibliographie Services services bibliographiques 395 Wellington Street 395. nie Wellington Ottawa ON K1A ON4 OitawaON K1AW Canada CaMda 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 sell reproduire, prêter, distribuer ou copies of this thesis in rnicroform, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfichelfilm, 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 othenvise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. The evolrttion and functional signifrcance of stigma-height dimorphism in flowering plants Doctor of Philosophy 2000 Angela Marie Baker Department of Botany University of Toronto ABSTRACT Genetic polymorphisms have been widely used as mode1 systems for evolutionaxy studies of adaptation and natural selection. This thesis investigates a distinct plant sexual polyrnorphism - stigma-height dimorphism (SHD) - and is the first comprehensive analysis of the nature of SHD, its taxonomie distribution, and its evolution and functional significance in animal-pollinated plant populations. SHDis characterized by discrete variation in stigma height with anthers positioned simiiarly in flowers of long (L)- and short (S)-styled morphs. The lack of reciprocity between stigmas and anthers in species with SHD distinguishes it from the well-known sexual polymorphism heterostyly. 1 compared floral morphology, incompatibility type and population morph-ratios of taxa with SHD vs. distyly. The systems typically differ in incompatibiiity status and morph ratios, yet SHD, like heterostyly, probably functions to increase proficient polien transfer between floral morphs. 1 examined features of SHD in populations of the Mediterranean geophyte Narcissus. SHD is widespread in the genus making it valuable for studying questions related to the evolution and function of the dimorphism. Self-incompatible populations of N. assoanus had morph ratios ranging from isoplethy (1: 1) to L-biased, while populations of self-compatible N. dubius were L- biased or monomorphic-L. There were no morph-specific differences in the species in outcrossing rates or femde fertility suggesting that differences in pollen transfer and levels of assortative and disassortive mating contribute to the variable morph ratios. A cornputer mating model provided evidence supporting this hypothesis. To investigate the role of sHD in the evolution of distyly, pollen transfer and male fertility were examined in experimental populations of the annual herb Eichhomia paniculata. Pollen transfer between morphs was typicaily greater than among individuals of the same morph. Mating patterns partidly supported the validity of the Lloyd & Webb (1992) model of the evoIution of distyly with SHD as an intermediate stage. The conditions of the model were satisfied when outcross transfers were considered but were met less often when selfing was included. While SHD seems likely to have been an intermediate step in the evolution of heterostyly in some groups, its occurrence in non-heterostylous families indicates that this is not always true. Numerous people have aided in the successful completion of this thesis. I owe many thanks to my thesis supervisor, Spencer Barrett. Spencer has been instrumental in my development as a critical researcher. His enthusiasm, guidance, and willingness to let me take this project in my own direction were very much appreciated. My collaborator, John Thompson, also contributed greatly to the development of this thesis. John's dedication to completing field experiments, even during freezing rain and snow, was much appreciated. John taught me a great deal about the implementation of practical field studies, but also about ail of the wonderful culinary distractions in southem France. My supervisory cornmittee members, Nancy Dengler, Jim Eckenwalder, Tamrny Sage, and bnefly Kermit Ritland provided me with advice and encouragement throughout my time at the University of Toronto. The additional members of my exarnining cornmittee, Anurag Agrawal, Mark Johnston. and James Thomson gave valuable cornrnents and suggestions that improved the final version of this thesis. This research was generously funded by a Natural Sciences and Engineering Research Council of Canada (NSERC) grant to Spencer Barrett, and by graduate scholarships from NSERC, the Ontario government (Ontario Graduate Scholarship, Ontario Graduate Scholarship in Science and Technology), and the University of Toronto. During the day-to-day tedium that represents the life of a grad student, it was great to have Bill Cole around. No matter what problem I faced, Bill was willing to help.. .I am particularly grateful for the early-morning Eichhornia emasculation sessions. Bruce Hall, Andrew Petrie, and Karl Wimmi also were a great help in deaiing with dl things Eichhornia and always stopped by to Say hello during weekend watering sessions. I also owe thanks to Jannice Friedman, Taline Sarkissian, and Stephen Wright who frequently helped me with mundane tasks without cornplaint. Sandy Speller and Tarnar Mamourian always managed to help cut through the red tape even when swamped with other commitments. Department-mates provided much support, both academically and socially. In the early years, Phillip Awadalla, Sean Graham, and Tracy Solomon were cornpanions for cocktails, enjoyable meals followed by stress-relieving workouts at the gym, numerous movies, and great conversation. Andrea Case, Anne Worley, John Pannell, and Brendon Larson made the Barrett lab an interesting place to be. In particular, Andrea and Anne were great lab mates.. Andrea because of her enthusiasm and Anne for her determination. In later years. Linley Jesson, Dave Kubien, Marcel Dorken, and Steve Griffin were great academic colleagues and friends. We have had many enlightening conversations about science and numerous topics beyond. In particular, 1 thank Linley Jesson for knowing what 1 was thinking even when 1 didn't know myself. Because of this, she was always around to supply timely advice related to al1 aspects of my life. Dave Kubien knew exactly when to twist my rubber arrn to get me to the pub and Marcel Dorken usuall y managed to drag me to the ElMo to see Tchort even though they're a metal band- 1 also wish to acknowledge my friends and farniiy who constantly reminded me of the fact that there was a world outside of the Earth Sciences building. Vik, Andrew, and Sarah provided much-needed Saturday night distractions and philosophical discussions. Phink, St. Dave, and Jurgen provided places to escape from al1 things botanical. Jeff was always encouraging and 1-11 miss the long discussions and bike rides in the Don Valley. Support from my family was essentiai.. .my morn and dad, Cindy and Larry Baker, have always supported and encouraged my academic endeavours (and they even stopped asking, "When will you be done?" fairly early on); my aunt, Bev Machniak, showed me that it is possible to persevere under any circumstance; and my grandfather. Matthew, encouraged me to appreciate the elegance of flowers. My partner. Dr. David Hay, supplied unconditional love. support, fnendship, and has helped me in many ways. r ABSTRACT ACWOWLEDCEMENTS TABLEOFCONTENTS..........................................................v i LISTOFTABLES.............................................................. xi LISTOFFIGURES............................................................ xiiia.. LISTOFAPPENDICES......................................................... xiv LISTOF ABBREVIATIONSAND SYMBOLS.......................................... xv GENERAL~NTRODUC~ON........................................................ 1 S~YLARPOLYMORPH~SMS........................................................2 FUNCTIONALINTERPRETATIONS OF FLORAL DESIGN ................................... -5 EVOLU~ONARYRELATIONS OF STIGMA-HEIGHTDIMORPHISM AND DISTYLY................ 7 The Charlesworth and Charlesworth (1979)model ............................. 7 The Lloyd and Webb (1992a.b)model ....................................... .9 TheRichards(l998)model............................................... 11 POLLENL~ATION AND POUEN TRANSFERMODELS ................................. 11 RESEARCH~BECTIVES...................-.....................................l 3 LNTRODU~ON............................................................... 20 SEX-ORGANPOS~ON LN SPECIES WITH STIGMA-HEIGHTDMORPHISM VS. DISTYLY......... -20 A cornparison of reciprocal herkogamy in stigma-height dimorphiic vs. distylous species .................................................. 25 FLORALMORPHOLOGY OF SPECIESwrr~ STIGMA-WGHT DIMORPHISM vs . DISTYLY........ -32 Corolla morplzology and pollinators
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