M. Cresti . G. Cai . A. Moscatelli (Eds.) Fertilization in Higher Plants Springer Berlin Heidelberg New York Barcelona Hong Kong London Milan Paris Singapore Tokyo M. Cresti . G. Cai . A. Moscatelli (Eds.)
Fertilization in Higher Plants Molecular and Cytological Aspects
With 159 Figures, Including 6 Color Plates
Springer Prof. Dr. MAURO CRESTI Dr. GIAMPIERO CAl Dr. ALES SANDRA MOSCATELLI Universita degli Studi di Siena Dipartimento di Biologia Ambientale Via P.A. Mattioli 4 53100 Siena Italia
ISBN-13: 978-3-642-64202-9 e-ISBN-13: 978-3-642-59969-9 DOl: 10.1007/978-3-642-59969-9 Library of Congress Cataloging-in-Publication Data Fertilization in higher plants: molecular and cytological aspects / [edited by1 Mauro Cresti, Giampiero Cai, Alle• sandra Moscatelli. p. em. Includes bibliographical references and index. ISBN-13: 978-3-642-64202-9 1. Fertil• ization of plants. 2. Angiosperms - Molecular aspects. 3. Angiosperms - Cytology. I. Cresti, M. (Mauro) II. Cai, Giampiero, 1963- . III. Moscatelli, Alessandra, 1960- . 571.8'6422 - dC21
This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copy• right Law. © Springer-Verlag Berlin Heidelberg 1999 Softcover reprint ofthe hardcover 1st edition 1999 The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are excempt from relevant protective laws and regulations and therefore free for general use. Production: PRO EDIT GmbH, D-69126 Heidelberg Cover Design: design & production GmbH, D-69121 Heidelberg Cover Illustration: See Chap. 8 Molecular Approach to Female Meiosis in Petunia Hybrida, Porceddu et al. Typesetting: Zechnersche Buchdruckerei, D-67346 Speyer SPIN: 10654542 31/3137-543 210 Preface
Fertilization requires an intimate partner relationship. It takes place at the cellular level: the egg cell interacts with the sperm cell. The peculiar situation in Higher Plants is the Double Fertilization. Each syngamic process in Angiosperms consists of two events: the fusion of the egg cell with one sperm cell resulting in the diploid zygote, and the fusion of embryosac nuclei with the other sperm cell, leading to the production of the triploid endosperm. A prerequisite for the double fertilization of Angiosperms is the synchronuous transport of two sperm cells by the pollen tube. The fertilization in higher plants is not only preceded by the pollination process, with its complicated interactions and with the assistance of pollinating agents (wind, water, animals), but also by a long lasting interaction period between the diploid pistil and the haploid pollen tube, as the carrier of the male material. This interaction phase can be described as the pro gamic phase, which starts with the landing of the pollen grain on the stigmatic surface, and ends with the syngamy, the fusion of the sexual cells. All these coordinated events are well known for a limited number of plant species and have been treated for a long time as a section of plant science called embryology. During the last 40 years it became evident that the physiology of the events, which pre• cede the final fusion of the sexual cells in flowering plants, is of great importance for understanding the fertilization processes and their genetic barriers (incompatibility and incongruity). The fertilization of flowering plants results in the transformation of the zygote into seeds and fruits, the final products of fertilization. Fruit and seed formation are basic processes for the world food supply. Crops, like cereals (e.g. rice, corn, wheat), legums (e.g. soya, peas, beans), fruits (e.g. apples, oranges, almonds, chestnuts), grapes are essential staples to human nutrition, not to mention spices (e.g. pepper, nutmeg). Also, a greater part of the mankind provisions are fats and oils (e.g. olive, corn, oil palm, rape, peanut), semi-luxurious items (e.g. coffee) and fibre delivering plants (e.g. cotton, capoc). The pigments and many natural remedies are the final results of the successful fertilization processes in higher plants. Fundamental understanding and the increased ability to manipulate the gene-controlled stages of fertilization are essential for the survival of mankind. The papers compiled in this book are the written and extended versions of lectures from the European Union Experts which have been given during a Euro Advanced Course. This Euro Course entitled "Sexual Plant Reproduction and Biotechnological Applica• tions: Recent Advances by Molecular Biology, Biochemistry and Morphology" has been supported by the European Commission, and was organized by the undersigned. VI Preface
Lectures were given in the Conference Center of the University of Siena, the Certosa di Pontignano, whereas the practical courses and demonstrations took place in the Labor• atories of the Dipartimento di Biologia Ambientale - UniversWt degli Studi di Siena. This compilation is a snapshot of this distinctive activity in a field of great impor• tance. The Authors have full responsibility for their papers, which have not been re• viewed or edited. We thank all the Speakers and the Instructors of the Course for their contributions, the Participants for their enthousiastic approach, the Technical Staff for their devoted assistance and the Publisher for the fast production of these proceedings. Our hope is that this picture of the present state of the art demonstrates the fast progress made in this field of research, especially when compared with the earlier pro• ceedings of the 1990 workshop entitled "Sexual Plant Reproduction" (Springer, Berlin, Heidelberg, New York, 1991). In addition, it became evident that pollen, pollen tubes and egg cells are excellent ex• amples for plant cell research, especially suitable for investigations on cell tip growth and polarization, signal transduction, channel and ion flux activity, gene expression, cytoskeleton and cell wall structure, biosynthesis and accumulation of specific sub• stances. This basic research and its results are important for future steps toward manipula• tion of the fertilization processes in higher plants. The manipulation will not only con• cern the testing of inhibiting drugs, but also the effects of pesticides used in agriculture, horticulture, viniculture and arboriculture. The future application of basic information gained on the steps of fertilization process will contribute to the promotion of genetic engineering in plants, in order to transform and improve crop plants. Cell culture and suspension culture are already methods, where basic science and biotechnology have met. Pistil activation is the next step to be handled for the improve• ment of important economic fruit production. The role of special genes and mutations will offer insight into the regulation of early zygotic embryogenesis in higher plants, and also into the effects of externally applicated phytohormones and their influence on the control of maturation of zygotic embryos. Basic knowledge of the induction pro• cesses, local control and patterning of cell division, especially the meiotic processes, which are essential for the formation of the sexual cells, are necessary for the progress of biotechnology in sexual plant reproduction.
Siena, Spring 1998 Mauro Cresti Contents
Polyamines and Gene Expression of Biosynthetic Enzymes in Sexual Plant Reproduction ...... N. Bagni, A. Tassoni, M. Franceschetti 1.1 Polyamines in Higher Plants: An Overview 1.2 The Role of Polyamines in Pollen 5 1.3 Concluding remarks 7 References 9
Carotenoid Biosynthesis in Plant Reproductive Organs: Regulation and Possible Functions ...... 13 G.Giuliano 2.1 Biosynthesis 13 2.2 Regulation . 15 2.3 Possible Functions 18 References 19
3 Lipid Accumulation and Related Gene Expression in Gametophytic and Sporophytic Anther Tissues 23 P. Piffanelli, D. J. Murphy 3.1 Introduction ...... 23 3.2 Lipid Biosynthetic Pathways in Anther Tapetal and Pollen Cells 24 3.3 Lipid Accumulation in Anther Tapetal Cells ...... 28 3.4 Intracellular Lipid Accumulation in Developing Pollen Grains 35 3.5 Conclusions and Future Prospects 38 References ...... 39
4 Sex Determination or Sex Dimorphism? On Facts and Terminology ...... 45 A. Lardon, C. Delichere, F. Moneger, I. Negrutiu 4.1 Introduction ...... 45 4.2 Sexual Dimorphism in Maize - A summary 46 4.3 Sexual Dimorphism in White Campion 46 4.4 Conclusions 50 References 51 VIII Contents
5 Meiosis .... 53 T. Schwa rza cher 5.1 Introduction . 53 5.2 Terminology . 54 5.3 Synaptonemal Complex 55 5.4 Recombination -Yeast Model 57 5.5 Recombination Nodules 57 5.6 Homologous Chromosome Pairing 58 5.7 Chiasma Distribution ...... 60 5.8 Future Outlook ...... 61 5.9 Cytological Methods for Looking at Meiotic Chromosomes 62 References ...... 65
6 Regulation of Gene Expression During Pollen Development 69 G. J. Wullems, J. A. M. Schrauwen 6.1 Pollen Development as a Model System to Study Cell Differentiation 69 6.2 Cellular Localization of mRNA of the Microspore Specific Gene ntm19 72 6.3 Analysis of Microspore Specific Promoters in Transgenic Tobacco 73 6.4 Pollen Coat Oleosins ...... 74 6.5 Expression of the Late Pollen Specific Gene ntp303 During Development and Pollen Tube Growth 75 References ...... 76
7 Double Fertilization in Flowering Plants: Origin, Mechanisms and New Information from in vitro Fertilization 79 J. -E. Faure 7.1 Introduction ...... 79 7.2 Origin of Double Fertilization ...... 79 7.3 Mechanisms of Fertilization Form in Planta Observation 80 7.4 New Information from in vitro Fertilization 84 7.5 Conclusions 86 References ...... 87
8 Molecular Approach to Female Meiosis in Petunia Hybrido 91 A. Porceddu, Ch. Moretti, S. Sorbolini, S. Guiderdone, L. Lanfaloni, F. Lorenzetti, M. Pezzotti 8.1 Introduction ...... 91 8.2 Callose Deposition During Megasporogenesis in Petunia Flowers ...... 92 8.3 Cell Division, Flower Development and Macrosporogenesis 92 8.4 Seeking Petunia Meiotic Genes by Yeast Trans-complementation Analysis 97 References ...... 98 Contents IX
9 Homomorphic Self-Incompatibility in Flowering Plants 101 D. de Nettancourt 9.1 Early Work, Definition and Importance of SI 101 9.2 Different SI Systems ...... 101 9.3 Recognition and Rejection Mechanisms 102 904 The Origin of SI Systems ...... 106 9.5 Relationships of S-Proteins to Other Plant Functions 107 9.6 Contributions of SI Research to Plant Breeding and Biotechnology 108 9.7 Conclusions 109 References · .... 110
10 Cell Death of Self-Incompatible Pollen Tubes: Necrosis or Apoptosis? ...... 113 A. Geitmann 10.1 Introduction...... 113 10.2 Different Ways of Cells to die ...... 115 10.3 Different Ways to the rejection of Pollen Tubes 117 lOA Conclusions l30 References ...... 132
11 Aspects of the Cell Biology of Pollination and Wide Hybridization ...... 139 J. S. Heslop-Harrison 11.1 Introduction ...... l39 11.2 Pollen Desiccation at the Time of Dehiscence 140 11.3 Pollen Viability ...... 141 11.4 The Stigma and Style, and Interactions With the Pollen Grain and Pollen Tube 141 11.5 Conclusions 143 References · ...... 144
12 Pollen Coat Signals With Respect to Pistil Activation and Ovule Penetration in Gasteria Verrucosa (Mill.) H. Duval 145 M. T. M. Willemse 12.1 Introduction ...... 145 12.2 Ovular Incompatibility 145 12.3 Pollen Coat Substances 146 1204 Pollen Coat Signals 147 12.5 Pollen Coat Proteins . . 148 12.6 Ovule Activation .... 149 12.7 Micropylar Penetration by the Pollen Tube 151 12.8 Final Remark 155 References · ...... 156 x Contents
13 Tapetum and Orbicules (Ubish Bodies): Development, Morphology and Role of Pollen Grains and Tapetal Orbicules in Allergenicity ...... 157 G. EL -G haza Iy 13.1 Introduction...... 157 13.2 Localization of Allergens in Pollen Grains and Orbicules 160 13.3 Tapetum and Orbicules . . . . 160 13.4 Allergenicity of Pollen Grains 169 References ...... 171
14 Development and Substructures of Pollen Grains Wall 17S G. EL-Ghazaly 14.1 Introduction...... 175 14.2 Material and Methods ...... 176 14.3 Development of Pollen Grains Wall 177 14.4 The Substructures (Exine Subunits) of Pollen Grains and Spores 194 References ...... 196
15 Mechanisms of Microspore Polarity and Differential Cell Fate Determination in Developing Pollen ...... 201 D.Twell 15.1 Pollen Cell Lineage and Development ...... 201 15.2 The Importance of Asymmetric Division for Differential Cell Fate 202 15.3 Models of Differential Cell Fate Determination ...... 203 15.4 The Determination and Expression of Microspore Polarity 204 15.5 Genetic Screening for Polarity and Cell Fate Mutants 208 15.6 Perspective 213 References ...... 214
16 Genetic Control of Pollen Development and Function 217 M. Sari-Gorla, M. E. Pe 16.1 Introduction .... 217 16.2 Pollen Development 218 16.3 Pollen Function .. 225 16.4 Concluding Remarks 229 References ...... 230
17 The Use of the Vibrating Probe Technique to Study Steady Extracellular Currents During Pollen Germination and Tube Growth 235 A. M. Shipley, J. A. Feijo 17.1 Introduction...... 235 17.2 The Vibrating Voltage Probe .. . 237 17.3 The Vibrating Ion-Selective Probe 239 Contents XI
17.4 The Vibrating Probe and Plants 243 17.5 The Vibrating Probe Technology 246 17.6 Conclusions ...... 249 References 250
18 The Role of Calcium and Associated Proteins in Tip Growth and Orientation ...... 253 R. Malh6 18.1 [Ca2+lc in Pollen Tube Growth and Reorientation 253 18.2 The Role of Calmodulin ...... 260 18.3 The Role of IP3 and Protein Kinases in Tube Growth and Reorientation 263 18.4 Exocytosis and Apical Growth 265 18.5 Future Prospects 267 References ...... 268
19 Measuring Ion Channel Activity During Polar Growth of Pollen Tubes 271 F. Armstrong, R. Benkert, F.-W. Bentrup, G. Obermeyer 19.1 Pollen and Plasma Membrane Ion Transporter . . . 271 19.2 Release of Tip Protoplasts With Laser Microsurgery 272 19.3 Results and Discussion 274 19.4 Conclusion 280 References ...... 281
20 The Rheological Properties of the Pollen Tube Cell Wall 283 A. Geitmann 20.1 Introduction ...... 283 20.2 The Plant Cell Wall - Rheology and Expansion in Growing Cells 284 20.3 Pollen Tube Cell Wall ...... 285 20.4 Pulsating Growth in Tip Growing Cells ... 288 20.5 Turgor and Tip Growth ...... 294 20.6 The Biological Function of Pulsating Growth 295 20.7 Summary. 296 References ...... 297
21 Actin Filament- and Microtubule-Based Motor Systems: Their Concerted Action During Pollen Tube Growth 303 A. Moscatelli, G. (ai, M. (resti 21.1 The Male Gametophyte During Fertilization 303 21.2 Cytoplasmic Organization 303 21.3 The AFs-Myosin System 305 21.4 MTs and MT-Motor Proteins 307 21.5 Concluding Remarks 312 References ...... 313 XII Contents
22 The Pollen Tube Oscillator: Towards a Molecular Mechanism ofTip Growth? ...... 317 lA. Feij6 22.1 Introduction ...... 317 22.2 Oscillations in Pollen ...... 317 22.3 Why are Oscillations Important? 320 22.4 The Molecular Basis of the Pollen Tube Oscillator 322 22.5 Case Study: Lilium Oscillations ...... 324 22.6 Towards a Molecular Model of Tip Growth? 333 References ...... 334
23 Fertilization and Zygotic Embryo Development in Vitro ...... 337 E. Kranz, J. Kumlehn, T. Dresselhaus 23.1 Fertilization ...... 337 23.2 Zygotic Embryo Development ...... 340 23.3 Gene Expression after in Vitro Fertilization 344 23.4 Conclusions and Prospects 347 References ...... 348
24 MADS Box Genes Controlling Ovule and Seed Development in Petunia ... 351 L. Colombo, G. C. Angenent 24.1 Introduction...... 351 24.2 Ovule Development ...... 351 24.3 MADS Box Genes and the ABC Model 353 24.4 MADS Box Genes Controlling Ovule Development in Petunia ...... 353 24.5 The Ovule: A Separate Floral Organ ...... 355 24.6 Other MADS Box Genes Expressed in the Ovule . 356 24.7 FBP7 and FBPll are Required for Correct Seed Development 357 24.8 Conclusions and Perspectives 358 References ...... 359
25 Domains of Gene Expression in Developing Endosperm ...... 361 H.-A. Becker, G. Hueros, M. Maitz, S. Varotto, A. Serna, R. D. Thompson 25.1 Origin of Endosperm Tissue ...... 361 25.2 Stages in Cereal Endosperm Development 362 25.3 Mutants Affecting Endosperm Development 364 25.4 Regulation of Gene Expression in the Endosperm 366 25.5 Manipulation of Endosperm Gene Expression In Vitro 370 25.6 Endosperm Development and Function in Arabidopsis 372 References ...... 373 Contents XIII
26 Advances on the Study of Sexual Reproduction in the Corc-Tree (Quercus suber L), Chestnut (Castanea sativa Mill.) and in Rosaceae (Apple and Almond) ...... 377 J. A. Feijo, A. C. Certal, L. Boavida, I. van Nerum, T. Valdiviesso, M. M. Oliveira, W. Broothaerts 26.1 Introduction ...... 377 26.2 The Cork-Tree (Quercus suber L.) 378 26.3 The Chestnut (Castanea sativa Mill.) 383 26.4 Self-Incompatibility in the Rosaceae 388 26.5 Conclusions 393 References ...... 393
27 Pollen as Food for Humans and Animals and as Medicine ...... 397 H. F. Linskens 27.1 Pollen Everywhere ...... 397 27.2 Chemical Composition of Pollen 398 27.3 Pollen as Food for Humans 398 27.4 Pollen as Medicament . . . 400 27.5 Pollen as Feed for Animals 402 References ...... 402
28 Field Release of Transgenic Virus Tolerant Tomatoes 405 S. Valanzuolo, M. M. Monti, M. Colombo, G. Cassani 28.1 Introduction ...... 405 28.2 CMV Resistance: The Biotechnological Approach 406 28.3 Technogen Experiment With Satellite RNA 407 28.4 Discussion 410 References ...... 411
29 Grapevine Biotechnology Coming on the Scene ...... 413 R. Vignani, M. Scali, M. Cresti 29.1 Introduction ...... 413 29.2 Biotechnology for a Better Understanding of Grapevine Biology 414 29.3 Grapevine Cell Cultures as a Tool for Genetic Manipulation and for the Production of Secondary Metabolites 416 29.4 Biotechnology for Genetic Improvement and Maintenance of the Grapevine Germoplasm . . . . 417 29.5 Conclusions and Future Perspectives 422 References...... 423
Subject Index 427
Color Plates . 439