[Advances in Cellular and Molecular Biology of Plants] Cellular and Molecular Biology of Plant Seed Development Volume 4 ||

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<ul><li><p>CELLULAR AND MOLECULAR BIOLOGYOF PLANT SEED DEVELOPMENT</p></li><li><p>Advances in Cellular and Molecular Biology of Plants</p><p>VOLUME 4</p><p>Editor-in-Chief</p><p>Indra K. Vasil, Laboratory ofPlant Cell and Mole cular Biology,University ofFlorida, Gainesville, Florida , USA</p><p>Editorial Advisory Board</p><p>Robert T. Fraley, St. Louis, Missouri. USARobert B. Goldberg, Los Angeles. California. USACharle s S. Levings, III, Raleigh, North Carolina. USARonald L. Phillips, St. Paul. Minnesota, USAJeff Schell , Cologne. Germany</p><p>The titles published in this series are listed at the end of this volume.</p></li><li><p>Cellular andMolecular Biology of</p><p>Plant Seed Development</p><p>Edited by</p><p>BRIAN A. LARKINS</p><p>Department ofPlant Sciences. University ofArizona, Tucson, Arizona, USA</p><p>and</p><p>INDRA K. VASIL</p><p>Laboratory ofPlant Cell and Molecular Biology, University of Florida, Gainesville,Florida , USA</p><p>Springer-Science+Business Media, B.V.</p></li><li><p>A C.I.P. catalogue record for this book is available from the Library of Congress</p><p>ISBN 978-90-481-4878-3 ISBN 978-94-015-8909-3 (eBook)DOl 10.1007/978-94-015-8909-3</p><p>Printed on acid-free paper</p><p>All rights reserved</p><p>1997 Springer Science+Business Media DordrechtOriginally published by Kluwer Academic Publishers in 1997.Softcover reprint of the hardcover Ist edition 1997</p><p>No part of the material protected by this copyright notice may be reproduced orutilized in any form or by any means, electronic or mechanical,including photocopying, recording or by any information storage and retrieval system,without written permission from the copyright owner.</p></li><li><p>Table of Contents</p><p>PrefaceB. Larkins and I.K. Vasil</p><p>Section A - Control of Seed Development</p><p>VII-Vlll</p><p>I . Embryogenesis in Dicotyledonous PlantsR. Yadegari and R. Goldberg 3</p><p>2. Development of the Suspensor: Differentiation, Communication,and Programmed Cell Death During Plant EmbryogenesisB.W. Schwartz, D.M. Vernon, D. Meinke 53</p><p>3. Endosperm Structure and DevelopmentD.A. DeMason 73</p><p>4. Hormonal Regulation of Seed DevelopmentR. Morris 117</p><p>Section B - The Synthesis and Accumulation of Stored Metabolites</p><p>5. The Biochemistry and Cell Biology of Embryo Storage ProteinsN.C. Nielsen, R. Bassiiner, T. Beaman 151</p><p>6. The Prolamin Storage Proteins of Wheat and its RelativesG.Galili 221</p><p>7. The Prolamin Proteins of Maize, Sorghum and CoixC.E . Coleman, J. M. Dannenhoffer, B.A. Larkins 257</p><p>8. The Storage Proteins of Rice and OatD.G. Muench, T. W. Okita 289</p><p>9. The Protease Inhibitors of SeedsK.A . Wilson 331</p></li><li><p>VI Table a/ Contents</p><p>10. Starch Synthesis in the Mai ze Seedi.c. Hannah 375</p><p>II . Synthesis and Storage of Fatty AcidsJ. Browse 407</p><p>12. Accumulation and Storage of Phosphate and MineralsV.Raboy 441</p><p>13. Genetic Regulation of Carbohydrate and Protein Accumulationin SeedsM. Motto , R. Thompson , F. Salamini 479</p><p>Section C - Control of Seed Maturation and Germination</p><p>14. Lea Proteins and the Desiccation Tolerance of Seed sL. Dure</p><p>15. Seed Maturation and Control of DormancyJ. Harada</p><p>Section D - Manipulation of Seeds Through Biotechnology</p><p>525</p><p>545</p><p>16. Biotechnological Approaches to Altering Seed CompositionE. Krebbers, R. Brogli e, B. Hitz, T. Jones , N. Hubbard 595</p><p>Index 635</p></li><li><p>Preface</p><p>The beginnings of human civili zation can be traced back to the time, near-ly 12,000 years ago , when the early humans gradually changed from a lifeof hunting and gathering food , to producing food. This beginning of primi-tive agriculture ensured a dependable supply of food, and fostered the livingtogether of people in groups and the development of society. During thistime, plant seeds were recognized as a valuable source of food and nutrition ,and began to be used for growing plants for food. Ever since, plant seedshave played an important role in the development of the human civilization.Even today, seeds of a few crop species, such as the cereals and legumes, arethe primary source of most human food , and the predominant commodity ininternational agriculture.</p><p>Owing to their great importance as food for human s and in internationaltrade , seeds have been a favorite object of study by developmental biologistsand physiologists , nutritionists and chemists. A wealth of useful informationis available on the biology of seeds. However, studies on the molecular biol-ogy of plant seed development are rather recent, and have begun to providecritical new information about the control of seed development, dormancy,germination and storage reserves.</p><p>The age of plant molecular biology began twenty years ago with the isola-tion and characterization ofmRNAs encoding seed storage proteins, followedby the cloning of related genes. The success of these early studie s encour-aged many plant biologists to focus their research on seeds. This was alsobecause seeds are vitally important agricultural products providing much ofthe starch, protein and oil for human and livestock diets, and are good modelsystems to study the biochemical and genetic mechanisms regulating protein ,starch and oil biosynthesis. In addition, seed development embodies severalunique biological processes, such as embryogenesis, dormancy, germination,etc .</p><p>During the past twenty years we have learned a great deal about the bio-chemical and genetic regulation of the processes involved in seed develop-ment. Nevertheless, much remains to be discovered about the physiologicalregulation of embryo development, and the mechanisms leading to seed des-iccation and dormancy, and germination. It is hoped that the rapid evolution</p><p>B.A. Larkins and IX. Vasil (eds.), Cellular and Molecular Bio!ogy ofPlant Seed Development. vi i-vi ii. 1997 Kluwer Academic Publishers,</p></li><li><p>viii Preface</p><p>of genome projects, and the isolation and characterization of mutants affect-ing seed development, will soon unravel these complex processes.</p><p>This volume presents a compilation of chapters describing fundamentalaspects of seed development and maturation by some of the world's leadingexperts. Emerging concepts of embryogenesis, endosperm development andseed maturation and desiccation are discussed in light of recently isolatednovel mutants. The in-depth and up-to-date reviews provide insights into thegenetics, biochemistry and cell biology of metabolic reserve (starch, protein,oil, mineral) synthesis and accumulation. The concluding chapter describesthe practical applications of this knowledge to the manipulation of storagereserve content of seeds by molecular genetic manipulation.</p><p>The human population is projected to double in the next thirty years. Foodproduction, primarily in the form of seeds, must be doubled too during thisperiod to provide food security for the increased population. It is vitallyimportant, therefore, to find ways to further enhance the productivity, qualityand utility of seeds. It is our hope that the information and ideas presented inthese pages will provide the insight and inspiration needed to achieve theseobjectives.</p><p>We thank each of the authors for providing state-of-the-art accounts offascinating and important advances in the cellular and molecular biology ofseed development.</p><p>Brian A. LarkinsIndra K. Vasil</p></li><li><p>Part A</p><p>CONTROL OF SEED DEVELOPMENT</p></li><li><p>1. Embryogenesis in Dicotyledonous Plants</p><p>RAMIN YADEGARI* and ROBERT B. GOLDBERGDepartment ofMolecular. Cell, and Developmental Biology, University ofCalifornia,Los Angeles , CA 90095-1606, USA</p><p>ABSTRACT, Embryogenesis in higher plants establishes the basic shoot-root body pattern,the primary tissue layers, and the meristematic zones of the plant. Continuous differentiationof the meristems is the basis of postembryonic development, the adult phase of the lifecycle . Critical to this process is not only the pattern forming or morphogenetic events takingplace mainly during early embryogenesis, but also a series of cellular and physiologicalprocesses which prepare the maturing embryo for dormancy and germination. Recent geneticand molecular studies in Arabidopsis and other model plants have begun to identify criticalprocesses involved in higher plant embryogenesis. Likewise, Arabidopsis mutations defectivein embryo structure or seedling viability are providing the tools for an analysis of molecularmechanisms responsible for dicot embryogenesis. One critical question is whether cellularinteractions play a role in the formation of embryo pattern, or whether the nearly regularpatterns of cell division observed in many species, including Arabidopsis, are a reflectionof a lineage-dependent mode of cell specification . Analysis of mutations altering cellularpatterns in Arabidopsis embryo indicate that cell-cell interactions most likely take place toestablish cell and tissue layers . Further, there is evidence for inter-regional interactions tocoordinate the overall development of the dicot embryo. However, differentiation processesbased on the activity of cell-autonomous determinants may also operate particularly duringthe earliest zygotic divisions which establish the principal embryonic elements. A secondmajor question concerns the specific gene regulatory mechanisms involved in initiating andmaintain ing differentiation programs within the developing embryo. These and other questionsregarding the underlying processes that control dicot embryogenesis are only beginning to beanswered using a combination of molecular and genetic tools.</p><p>I. Introduction</p><p>New genetic and molecular tools have been used in recent years to dis-sect the mechanisms that control plant embryogenesis. Many genes requiredfor various embryogenic processes in both monocotyledons and dicotyle-dons have been identified using genetic approaches (Meinke, 1985; Clarkand Sheridan, 1991; Mayer et aI., 1991; Johnson et aI., 1994; Hong et aI.,1995). Genetic manipulation of Arabidopsis thaliana by irradiation muta-genesis (Muller, 1963; Usmanov and Muller, 1970), chemical mutagenesis</p><p>* Present Address: Department of Plant and Microbial Biology, University of California,Berkeley, California 94720-3102</p><p>B.A. Larkins and IX. Vasil (eds .). Cellular and Molecular Biology ofPlant Sad Development, 3-5 2. 1997 Kluwer Academic Publishers,</p></li><li><p>4 Ramin Yadegari and Robert B . Goldberg</p><p>(Meinke and Sussex, 1979a,b; Meinke, 1985; Jurgens et a!., 1991; Mayeret aI., 1991, 1993a) and insertional mutagenesis (Errampalli et aI., 1991;Feldmann, 1991; Forsthoefe1 et a!., 1992; Castle et a!., 1993), has identi-fied a large number of zygotic mutants that are defective at different stagesof embryogenesis. These mutants have provided insights into the processesthat perform essential functions during embryogenesis, regulatory as wellas general housekeeping functions. Furthermore, some of these mutationscan be traced back to specific defects during early stages of embryogenesisrevealing the importance of specific cell division patterns and tissue organiza-tions in normal embryo development processes. Both genetic and molecularapproaches have identified genes which are transcribed in specific regionsof the dicot embryo suggesting an underlying prepattem of gene regulatoryprograms involved in embryo tissue and organ development. The correspond-ing regulatory sequences responsible for the region-specific transcription ofthese genes are beginning to be deciphered allowing an entry into the generegulatory pathways involved in embryo pattern specification and develop-ment (see below). In this review we outline the major conceptual insightsthat have been gained from studies ofArabidopsis embryo mutants and geneexpression experiments in other plants that provide new information about theprocesses regulating dicotyledon embryogenesis especially during the earlydevelopmental stages. Recent experimental evidence suggests that a plantembryo has a modular structure and consists of regions which are distinct atthe molecular levels.</p><p>II. General Features of Embryogenesis Are Similar in Higher Plants</p><p>In flowering plants (angiosperms), double fertilization of the egg cell and thepolar nuclei (within the central cell) by sperm nuclei produces a diploid zygoteand a triploid endosperm, respectively (Esau, 1977; Raven et al., 1992) . Asa differentiated organ , the endosperm is present during seed developmentand provides nutrients for either the developing embryo, the germinatingseedling, or both (Lopes and Larkins, 1993). The zygote, on the other hand ,develops into an embryo and will give rise to the body plan of the matureplant (sporophyte) after seed germination. Angiosperm embryos contain twoprimary organ systems- the axis and the cotyledon (Raven et a!., 1992) (Fig-ure I). These organs have distinct developmental fates and are composed ofthree basic, or primordial , tissue layers - protoderm, procambium, and groundmeristem - which will become the epidermal, vascular, and parenchyma tis-sues of the young seedling, respectively (Esau, 1977; Raven et a!., 1992).The axis, or hypocotyl-radicle region of the embryo, contains the shoot androot meristems, and will give rise to the mature plant after seed germina-tion (Figure 1). The root meristem will give rise to only one organ, the root,while the shoot meristem will produce, directly or indirectly, all the vegeta-tive and reproductive organs of the mature plant. By contrast, the cotyledon</p></li><li><p>POST-FERTILIZATION</p><p>Embryogenesis in dicotyledonous plants 5</p><p>GLOBULAR/HEART TRANSITION</p><p>HEARTEMBRYO</p><p>TRANSITIONEMBRYO</p><p>IHs</p><p>uPd</p><p>tG-CELL GL08ULAREP EMBRYO</p><p>2/4-CELL 8-CELLEP EP</p><p>PROEMBRYO</p><p>~ ~: .,~j:P ~ZYGOTE HELL</p><p>.....--------------,1'I ------- - - - - - - - - --,</p><p>ORGAN EXPANSION AND MATURATION</p><p>Gm</p><p>Pc</p><p>SC</p><p>PdA</p><p>RM</p><p>TORPEDO EMBRYO WALKING-STICK EMBRYO MATURE EMBRYO</p><p>-CE</p><p>SC</p><p>En</p><p>Fig. I . A generalized overview of dicot embryogenesis. Schematic representations of embry-onic stages are based on light microscopy studies of Arabidopsis (Mansfield and Briarty, 1991,1992; Mayer et al., 1991) and Capsella (Schulz and Jensen, 1968a, b) embryo development.For a comprehens ive description of the stages of Arabidopsi s embryo development refer toJUrgens and Mayer (1994). Abbreviations: T, terminal (apical) cell; B, embryo basal cell; EP,embryo proper ; S, suspensor; Be, suspensor basal cell; Pd, protoderm; u, upper tier; I, lowertier; Hs, hypophy sis; Pc, procambium; Gm, ground meristem; C, cotyledon; A, axis; MPE,micropylar end; CE, chalazal end; SC, seed coat; En, endosperm; SM, shoot meristem; RM,root meristem.</p><p>functions primarily in accumulation of food reserve s that are utilized by theseedling for growth and development after germination, becomes photos yn-thet ically acti ve during the seedling stage, and senesces shortly after theseedling emerge s from the so il (Figure I). That is, during embryogenesis, theco tyledon mobil izes food reserves and then switches roles during seedlingdevelopment to break down these reserves prior to the emergence of leaves,allowing the plant to become photosynthetically active. In many higher plants,incl uding Arabidopsis, the embryo might be photosynthetically active prior</p></li><li><p>6 Ramin Yadegariand Robert B. Goldberg</p><p>TABLE 1</p><p>Major events of flowering plant embryogenesis.</p><p>Post-fertilizationlproembryo</p><p>Apical and basal cell differentiation</p><p>Formation of sus...</p></li></ul>

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