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Reproductive Biology of the Perennial Desert Shrub, Thymelaea Hirsuta: Studies in Gender, Self-Fertility and Architecture

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Reproductive Biology of the Perennial Desert Shrub, Thymelaea Hirsuta: Studies in Gender, Self-Fertility and Architecture Reproductive Biology of the Perennial Desert Shrub, Thymelaea hirsuta: Studies in Gender, Self-fertility and Architecture by Rita Morbia A Thesis Submitted to the Faculty of Graduate Studies and Research through the Department of Biological Sciences in Partial Fulfillment of the Requirements for the Degree of Master of Science at the University of Windsor Windsor, Ontario, Canada "1 996 Rita Morbia National tibrary Bbliothèque nationale I*I ofCada du Canada Acquisitions and Acquisitions et Bibliographie Services seMces bibliographiques 395 WeUington Saeet 395, nie Wellingtori OttawaON K1AW ûttawaON KIA 0134 canada cana& The author has granted a non- L'auteur a accordé une licence non exclusive licence ailowing the exclusive permettant à la National Libraiy of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or seIl reproduire, prêter, distn'buer ou copies of this thesis in microfom, vendre des copies de cette thèse sous paper or electronic formats. la fome de microfiche/ih, de reproduction sur papier ou sur format élecîronique. 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 thése ni des extraits substantiels rnay be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. The modular construction of plants allows them to display significant phenotypic plasticity in both morphology and sex expression. Thymelaea hirsuta (Thymelaeaceae) is a xerophytic evergreen shrub cornmon to the Mediterranean region and known to dernonstrate significant variability in sex expression. A series of observational and experimental studies were carried out to investigate the nature of this variability in greenhouse-raised plants. Seeds were obtained from Egyptian populations, and following seedling establishment, an irrigation treatment was applied. Plants were watered every 3 days, 6 days or 9 days. A survey of two year old flowering plants (n = 305) showed significant bimodality in sex expression. Quantitative gender estimates indicated that individual plants tended to be either more male or more female. Water treatment had no significant effect upon estirnates of gender, number of male flowers, number of fernale flowers, total number of flowers or an index of sexual lability. A controlled pollination study revealed significant differences in seed mass among geitonogarnous, half-sib and outcrossed treatments, resuking in an estimate of inbreeding depression (6) of 0.23. Nonsignificant differences in fruit set were also observed. In general, selfing resulted in less fit progeny. To examine the effect of water treatment on various morphological ("architectural") parameters, repeated measurements were taken every two months on a subset of 202 plants for a total of ten months. Water treatment, time and the interaction effect between water treatment and time al1 significantly affected plant architecture, particularly height. Male, fernale and hermaphrodite flowers were compared for differences in position. There were significant differences among male, female and hermaphrodite flowers with respect to plant height, vertical distribution and branch order, suggesting that male flowers and female flowers may have different iii morphological optima. These studies indicate that selection for the evolution of dioecy in Thymelaea hirsuta may involve a combination of selection for outcrossing and resource or pollinator efficiency. ACKNOWLEDGMENTS I would like to thank Dr. Jon Lovett Doust for his support and encouragement over the course of the past two years. I would also like to thank the members of rny cornmittee, Dr. P. Sale and Dr. S. Paul. I would like to acknowledge the other members of rny lab who have helped me tremendously in terms of moral support as well as advice. These people include Ruben Boles, Cynthia Lokker, Tarun Mal, Dave SUS~O,Adam Ekler, Kelly Potter, Jun Yang and Maciej Biernacki. Andrea Risk and Sue Roe, I also thank for their encouragement and friendship. I would also like to thank the following faculty members: Dr. Lesley Loven Doust, Dr. Jan Ciborowski and Dr. O. Thomas. Many undergraduate assistants have helped me enormously in the collection and processing of data, including Lynda Welch, Harinder Singh, Kirsten Gyetvai, Nadine Jarrett and Paula Davis. I would like to thank Ali El- Keblawy for bringing the Thymelaea hirsuta seeds frorn Egypt, without which there would be no thesis. Mr. Daniel Boles made beautiful drawing of Thymelaea hirsuta for my thesis, and I am grateful for his effort. I would also like to thank rny parents and sisters for their tolerance and patience, if not actual outright encouragement. I would especially like to thank my friend, Y. Zalik who was never more than a phone cal1 away. TABLE OF CONTENTS ABSTRACT ....................................... iii ACKNQWLEDGEMENTS ............................... v LIST OF TABLES .................................... viii LIST OF FIGURES ................................... xi CHAPTER 1 . GENERAL INTRODUCTION Plant Sex Expression ............................. 1 Gender Quantification ............................ 10 . Mating Systems ................................ 13 Materna1 and Paternal Influences on Seed Production ....... 17 Secondary Sex Characteristics ...................... 25 Evolution of Dioecy .............................. 30 Plant Architecture ............................... 40 Th ymelaea hrisuta i) General Description ....................... 47 ii) Sex Expression ..........................54 iii) Evolution of Dioecy in the Thymelaeaceae ....... 61 2 . GENDER STUDY Objective ..................................... 64 Materials and Methods ............................ 66 Results ...................................... 72 Discussion .................................... 98 3. SELF-FERTILITY STUDY Objective .................................... 109 Materials and Methods ........................... 110 Results ..................................... 113 Discussion ................................... 129 4 . ARCHITECTURE STUOY Objective ................................. .. 134 Materials and Methods ........................... 135 Results ...................................... 138 Discussion ................................... 157 GENERAL DISCUSSION ............................... 165 LlTERATURE ClTED ..................................170 vii LIST OF TABLES Chapter 1 Table 1.1 Summary of gender phenotypes reported for Thymelaea hirsuta ................................... 55 Table 1-2. Description of gender phenotypes in Tbymelaea hirsuta, according to El-Keblawy et al. (1995) ..................... 60 Chapter 2 Table 2.1. Mean gender value and variance of male and femala "clusters" of Thymelaea hirsuta, using al1 flowering plants, and only intermediate gender values ............................. 76 Table 2.2. Summary of gender phenotypes for two year old plants of T'ymelaea hirsuta, raised under greenhouse conditions ... 77 Table 2.3. Results of MANOVA for total number of flowers produced, total number of female flowers produced, total number of male flowers produced, gender value, sex lability index, days to first flowering and length of flowering period ....... 78 Table 2.4. Mean ( *SE) number of days to first flowering and the affects of water treatment and site in Thymelaea hirsuta ...... 79 viii Table 2.5.a) Results of Chi-Square analyses to determine if proportions of male, fernale dichogamous and labile phenotypes differ significantly for field (from El-Keblawy et al. 1995) versus greenhouse populations ................. 80 Table 2.5.b) Results of Chi-Square analyses to determine if proportions of male, fernale dichogarnous and labile phenotypes differ significantly for field (from Ramadan et al. 1994) versus greenhouse populations .............................. 8 1 Table 2.6. Results of maximum likelihood ANOVA from multiway frequency analysis testing associations among water treatrnent, maternai gender and water treatment ............. 82 Chapter 3 Table 3.1. Number of pollinations of each cross type performed on flowering plants of Thymelaea hirsuta .................. 1 17 Table 3.2. Results of ANOVA for effects of pollination treatment on proportion fruit set (plant as replication unit) .............. 1 19 Table 3.3. Results of ANOVA for effects of pollination treatment on seed mass (plant as replication unit) ................... 120 Table 3.4. Results of ANOVA for effects of pollination treatment on fruit set (flower as replication unit) .................... 121 Table 3.5. Results of ANOVA for effects of pollination treatment on seed mass (flower as replication unit) ................... 122 Chapter 4 Table 4.1. Results of repeated measures MANOVA testing the effects of water treatment and cohort (time) on height, primary length, secondary length, tertiary length, number of secondary branchas and number of primary branches ................... 141 Table 4.2. Results of repeated measures MANOVA testing the effects of water treatment, and cohort (time) on height, tota! length O? five primary branches (including all higher orders) and total nurnber of branches on primary branches ................... 142 LIST OF FIGURES Chapter 1. Figure 1.1. Fitness (reproductive success) of either sex, as a function of resource allocation ........................... 31 Figure 1.2. Conditions for stability of an hermaphroditic
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