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Flower Size-Number Trade-Offs and the Evolution of Floral Display in Animal

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Flower Size-Number Trade-Offs and the Evolution of Floral Display in Animal FLOWER SIZE-NUMBER TRADE-OFFS AND THE EVOLUTION OF FLORAL DISPLAY IN ANIMAL-POLLINATED PLANTS by Anne Catherine Worley A thesis submitted in conformity with the requirements for the degree of Doctor of Philosophy Graduate Department of Botany University of Toronto @ Copyright by Anne Catherine Worley 2000 National Library BiMiottièque nationale du Canada Acquisitions and Acquisitions et Bibliographie Services services bibliographiques 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 seii reproduire, prêter, distribuer ou copies of this thesis in microfonn, vendre des copies de cette thèse sous paper or electronic formats. la fome de microfiche/fiim, 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 substantiai 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 variety of flowers in animai-pollinated plants suggests a history of strong and diverse selection. One neglected aspect of this divenity isjloral display, the size, nurnber, and arrangement of flowers. The evolution of floral display should reflect pollinator-mediated selection and life-history tradesffs imposed by finite resources. Although flower size is expected to Vary negatively with flower number, variable resources could cause positive relations between these traits. Such positive relations may reflect the hierarchical nature of resource allocation, with resources fint divided between somatic gro~thand reproduction' and then fûrther subdivision of reproductive resources. I investigated the evolutionary significance of trade-offs between flower size and number using theoretical' expimental, and comparative approaches. A quantitative-genetic mode1 involving hierarchical allocation indicated that high genetic variation in early allocation masked trade-offs, and altered the direction of evolution in the traits involved. Relations between flower size and number at the individual and population-level were examined in the annual herb, Eichhorniapanicula~o.The size and nurnber of flowers within inflorescences decreased rapidly in response to defoliation and seed production. emphasizing the dynamic nature of floraI allocation. Quantitative-genetic analyses of two glasshouse-grown populations demonstrated heritable variation in flower size and number. Genetic correlations between flower size and number ranged from negative to positive, depending on population and developmental stage. Accounting for variation in resource status removed positive conelations. but did not reveal additional negative correlations. Two generations of artificial selection on flower size and number partially upheld the expectation that sire should increase ai the espense of number. Finally, comparative data from 43 species in the perennial genus. n0rcissrrs. supported flower size-number trade-offs. However, in N. dubiur these traits were positively related, and unrelated when variation in resource status was accounted for. These results illustrated the influence of variable resource levels on traits involved in trade-offs. Flower size and number were negatively correlated when variation in floral traits was high, e.g, among species. However, variable floral resources often caused positive relations between these traits. Thus trade-offs between flower size and number will not always constrain the evolution of floral display, as ofien assumed in theoretical models. The successfûl completion of rny thesis reflects the contribution of many people. 1 owe the greatest debt of gratitude to my supervisor, Spencer Barrett. Spencer's enthusiasm, unflagging energy, and confidence inspired my research in the first place, and helped me cany it through to completion. He is an example to aspire toward. My cornmittee members, Nancy Dengler, David Houle, and bnefly Kemit Ritland provided very usehl advice and encouragement. 1 am especially gratefül to David Houle for his patient reviews of quantitative genetics. The additional members of my examining cornmittees, Susan Mazer, Locke Rowe, Rown Sage, and Tammy Sage, gave valuable feedback that improved the final version of the thesis considerably. Financiai support for the work presented here was provided by a Natural Sciences and Engineering Research Council of Canada (NSERC) gram to Spencer Banen and graduate scholarships from the Ontario government, and the University of Toronto. My greenhouse experiments were large, laborious, and involved the efforts of many beside myself. 1 owe a special debt to Bill Cole, who cheerfully helped with everything from lab work to data enuy. 1 am particularly gratefûl to Taline Sarkissian, Stephen Wright. Ho Sang Yoon. Leigh Howes, Doreen Chung, Dominik Halas and Linley Jesson. who al1 joined me in countlsss long, hot houn of tending and measuring plants. Bruce Hall. Andrew Peuie. and Karl Wimmi provided greenhouse camaraderie. ensured that trays and soi1 magically appeared when thep were needed, and that plants, clippings, and pests disappeved when they were not. Finally. the study of Narcissus dubius would not have ken completed without the hard work of Angela Baker and John Thompson. Fnends and lab mates provided much support and entertainment. academic and othenvise. Philosophical discussions, great meals, and the odd expedition out of Toronto with Andrea Case and Pat Lorch were essential throughout my time here. 1 thanli Brendan Larson for his reverence for nature and the best birding trips ever, and Linley Jesson for her irreverence in general. interest in othen, and joie de vive. Angie Baker broadened my horizons with new aspects of city life and Sean Graham provided the thoughtful perspective of a senior colleague. Alison Stuart displayed enviable confidence and enthusiasm for academic life. 1 will remember them. and many others, with fondness. My family's support was most essential. My parents, Ray and Elizabeth Worley, have encouraged perseverance and high standards throughout my academic career, as well as fostenng my interest in natural history. My parents-in-law, Bogdan and Jaga Czaykowski, were very supportive and enjoyed discussions about my blooming flowers. My husband, Piotr Czaykowski, suppIied the greatest support with his constant love and friendship. Piotr's belief in me, wiIIingness to let growing seasons govern our lives, and capacity for enjoyment continue to e~chmy work and life. Results -_-------_---* * --.-------------------.-.----.----------------------------------------*--- * --.--* * -------..---- Genetic Vanances and Covariances--__----&---- ----------------- .--a---------------- -----------------.--.--.---- Direction of Evolutio"----_-_.__--------------.------.------* --------.--------*-* .------.-----------.-.-----.--.----.--.-- Rate of Evolution*_-----_--_------------------.------------------------------.----------- * ---.--*- *--*A* ----*---------. .----- Time to Trade-off-*-*.--- * ----- * ------.------.------a-- * ------------------------------------------.-----.--* ---- ------------ Revealing Tde-off~through Artificial Selection-----------------------.-----.--------.-----------.---- 4. EVOLUTIONOF FLORALDISPLAY IN EICHHO~~VIAP,-t.V~CL'L-IT:-f: GESETIC AXD ENVIRONMENTAL VARIATIONIN FLOWER SIZE AND NUMBER 66 Smmq.-.--..-..--....-.--.-...---.----------- * -----**-*...----.-*--.-*---- *.* *-*-- * .-----.---..-.------.--* .---.--..--------.*------- 67 Introduction...-.--....-..--....---.-..-.-----.--.---.---..-..----.-.--.----.------* * ------------*----.-..**..-...---------.---------*. vii Metho& -_-__---*.-*-.* -*----- * -.------.-----------* ---------------------.--------- * ---.--*-* --.---.-.-*--------- ----- ------- -------- ---- Experimental Design and Data Collection ----------a-------------------- .-.--------- ---.------.-.---. ------- First Generation........................................ * ------- * ----------.----*-* -*-*a-.------ --- --------------- ------ Second GeneratioR*----- *---*--*-*---*--* -------.--------.--------.-------.-.--..-.-----.----*-.-* ..*----* ---------- Data Anabsis-.---------------- *---* -----.---------------- * .---------------------.---.-.-.---.---------------.---.--.-------.-- Phenotypic Relations.---- * --------.----------.--.----------.--------------.-.--- * ------*--* ----** a.------.--- * ---.-- Genet;~Parameters -----A.--------------- * -------.---.-------------.---------.-----.--------------------*- *-.- *-* ---* Results -----* -*-*---*-*--.--------------------.---- * ---------------.--.---.--* --------.-.---*--.-*.-*-*---. * -.-a*--. -me-.--- - --.-.-.-----. Variation in Floral Traits Indices of Plant Six-_---._--.-------.------.--- ** . Population Different1ation ----------------.--*- * .-.-.* ---*-*--*.--.--. **-.------------------.---.-.---.---.-- Temporal Differences arnong Inflorescences and Generations*-----*-.-----...---..-..---. &ritable Vanation *- * ----*---*-.------------.-..--- - ----.-----.-----.---- Relations beb-een Floral Traits and Indices of Plant Size .-.--------_-.~.--------.---..-.-.--.---.- Phenotypic Relations----.*------*----*-------.-----..---.--.--------------.----...-.-----*--.---- - -.--a------ -me---- Genetic md Envi~ocmentalCorrelations
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