EMBRYOLOGY OF FLOWERING PLANTS TERMINOLOGY AND CONCEPTS Embryology of Flowering Plants Terminology and Concepts Volume 2 Seed EDITED BY T.B. BATYGINA Science Publishers Enfield (NH) Plymouth, UK CIP data will be provided on request SCIENCE PUBLISHERS An Imprint of Edenbridge Ltd., British Channel Islands, UK Post Office Box 699 Enfield, New Hampshire 03748 United States of America Internet site: http://www.scipub.net [email protected] (marketing department) [email protected] (editorial department) [email protected] (for all other enquiries) ISBN 13: 978-1-57808-263-6 (hbk) © 2006, Copyright reserved All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without prior permission. This book is sold subject to the condition that it shall not, by way of trade or otherwise, be lent, re-sold, hired out, or otherwise circullated without the publisher's prior consent in any form of binding or cover other than that in which it is published and without a similar condition including this condition being imposed on the subsequent purchaser. Published by Science Publishers, Enfield, NH, USA An Imprint of Edenbridge Ltd. V.L. Komarov Botanical Institute Russian Academy of Sciences DEPARTMENT OF EMBRYOLOGY AND REPRODUCTIVE BIOLOGY RUSSIAN FOUNDATION FOR BASIC RESEARCH Contributors T.B. Batygina (editor), E.V. Andronova, G.M. Anisimova, O.V. Antipova, E.A. Bragina, V.B. Brukhin, M. Cresti, M.F. Danilova, O. Erdelska, W.E. Friedman, E.N. Gerassimova- Navashina, G. Jurgens, O.P. Kamelina, O.A. Khvedynich, H. Kieft, S.N. Korobova, L.V. Kovaleva, L.G. Krasnikov, A.A.M. Van Lammeren, Y.-Q. Li, F. Ma, S.S. Medvedev, D.W. Meinke, T.N. Naumova, S.G. Nawaschin, Z.I. Nikiticheva, N.V. Obrucheva, T.A. Plyushch, V.V. Polevoy, L.M. Pozdova, M.V. Razumova, B. Rodkiewicz, S.D. Russell, T.S. Salamatova, J.H.N. Schel, I.I. Shamrov (secretary and editorial board member), S.M. Szcziparev, M.P. Solntseva, E. Szczuka, E.S. Teryokhin, G.E. Titova, V.E. Vasilyeva (editorial board member), W.L.H. Van Veenendaal, M.A. Vishnyakova, T.D. Vyshenskaya, J.L. Van Went, X.L. Ye, E.C. Yeung, S.Y. Zee, N.A. Zhinkina, G.Ya. Zhukova (editorial board member) To S.G. Nawaschitt the discoverer of double fertilization in angiosperms (100th anniversary) Seeds are the germ of life, a beginning and an end, the fruit of yesterday's harvest and the promise of tomorrow's. O.L. Freeman Preface S.G. Nawaschin's discovery stimulated numerous investigations into plant fertilization, not only in Angiospermae but also in Gymnospermae. One of his most talented pupils—E.N. Gerassimova-Navashina—developed new notions about this process, specifically the idea of fertilization types. Her investigations also became classical. Their embryological school traditions are continued nowadays in different laboratories of the world and, primarily, in the Department of Embryology and Reproductive Biology, Komarov Botanical Institute, Russian Academy of Sciences (head of the leading Scientific High School of the Russian Federation, T.B. Batygina). The present staff of the Komarov Botanical Institute, Russian Academy of Sciences, considered it a pleasant duty to include in this second volume of Embryology of Flowering Plants the history of double fertilization discovery and some works of S.G. Nawaschin, published in Russian and German journals, which are bibliographical rarities. These investigations have not lost their significance over time because they contain detailed descriptions of the double fertilization process in classical objects. The recent tendency towards consolidation of embryology with genetics, physiology and biochemistry is obvious. Progress in molecular biology has strengthened these relationships. Earlier the main genetical method was hybridological analysis, which enables recording these or those alternative features, but nowadays the attention of geneticists is focusing more and more on questions of genetical feature expression during ontogenesis, regularities of caryogamy, plasmogamy and cytokinesis, as well as histogenesis and organogenesis. For many geneticists in the world the centre of attention has become the study of mutations which affect the first developmental stages of male and female generative structures and embryogenesis (for example, the disturbance of asymmetrical initiation of the first wall in the zygote). This permits revealing the action mechanism of genes in the processes of development and organogenesis. Successful models for studying mutations have been found (Arabidopsis thaliana, Zea mays). Unfortunately, specialists in the area of studying inheritance of features do not adequately consider the fundamental achievements of embryology. However, it is classical embryology, in fact, with such representatives as K. Schnarf, R. Soueges, viii D.A. Johansen, P. Maheshwari, B.M. Johri, J.S. Mestre, J.L. Guignard, A. Lebegue, S.G. Nawaschin, M.S. Yakovlev, E.N. Gerassimova-Navashina a.o., which has made significant contributions to understanding the main regularities of the origin and early stages of individual development. This book pays great attention to the morphological and functional aspects of the fertilization process and embryogenesis, the main subjects of investigation by embryologists of St. Petersburg Academy of Science for many decades. Besides, it is important that the very peculiar type of sexual reproduction closely connected in flowering plants with seed reproduction, has permitted such plants 'to conquer' the Earth and create the basis for the development and flourishing of our civilization. Just the seeds are the main nutritive source for mankind, which is why it is impossible to overevaluate the significance of scientific progress in their study. Together with the traditional ideas of seed development, the original notions on the problems of origin, correlations and evolutionary transformations of seed structures are presented in this book. Data are drawn not only for angiospermous, but also for gymnospermous plants. New data in experimental embryology, dealing with the morphogenesis of reproductive structures, specifically development of the somatic embryo, enable a discussion of the new notion of asexual reproduction—embryoidogeny. New classifications of the developmental types of main seed structures, worked over mainly by researchers of the Botanical Institute, are presented here. Investigations on ecological embryology, the chlorophyllous embryo and its autonomy for example are generalized. New types of embryogenesis and endospermogenesis are also described. The task of the authors was generalization of the theoretical notions on which classical and experimental data were based, concerning the development of flower generative organs, the process of seed formation and germination, and the system of seed reproduction as a whole. This is especially urgent given the need to preserve rare and vanishing plant species, to create a seed bank and to elaborate new biotechnologies for multiplication of avaluable plant forms and varieties associated with the problems of seed reproduction. Unification of embryological terms and notions will promote further development of ideas about the reproductive processes occurring during plant ontogenesis. The first volume of the three-volume edition Embryology of Flowering Plants. Terminology and Concepts was published in English in 2002. It was devoted to the common flower structure and structure of the main generative organs—the anther and ovule. The present book is the second volume of this edition. It consists of five parts and reflects the processes which take part in seed formation. The authors took into account the long-ago matured broadening of plant embryology borders, the science which turns from descriptive into causal discipline. Besides traditional chapters—'Fertilization', 'Embryo' and 'Endosperm'—there are chapters on the nature of such storage tissue as perisperm, the seed coat structure, and also perspectives on genetic analysis of plant embryogenesis and features of post-seminal development, including seed dormancy and germination. In preparing the book we used such well-known monographs as F. Netolitzky, Anatomie der Angiospermen-Samen (1926); K. Schnarf, Embryologie der Angiospermen. Archegoniaten (1929) and Vergleichende Embryologie der Angiospermen (1931); R. Soueges, Les Lois du Developpement (1937) and Embryogenie et Classification (1939); P. Maheshwari, ix An Introduction to the Embryology of Angiosperms (1950); D.A. Johansen, Plant Embryology (1950); P.A. Baranov, The History of Plant Embryology (1955); A.L. Takhtajan, Foundations of Evolutionary Morphology of Angiosperms (1964), Outline of the Classification of Flowering Plants (1980) and System Magnoliophyta (1987); V. A. Poddubnaya-Amoldi, General Embryology of Angiosperms (1964b) and Cytoembryology of the Angiosperms: Principles and Perspectives (1976); G. Davis, Systematic Embryology of Angiosperms (1966); T.B. Batygina, Wheat Embryology (1974); T.B. Batygina, The Grain of Cereals (Atlas); E.S. Teryokhin, Parasitic Flowering Plants. The Evolution of Ontogenesis and the Mode of Life (1977); E. Comer, The Seeds of Dicotyledons (1976); K. Esau, Anatomy of Seed Plants (1977); B.M. Johri (ed.), Embryology of Angiosperms (1984); T.B. Batygina and M.S. Yakovlev (eds.), Comparative Embryology of Flowering Plants (1981, 1983, 1985, 1987, 1990); V. Raghavan, Experimental Embryogenesis in Vascular Plants (1976) and Embryogenesis in Angiosperms: A Developmental and Experimental Study (1986); M.F. Willson, Plant Reproductive Ecology (1983); B.M. Johri, K. Ambegaokar, P.S. Srivastava Comparative Embryology of Angiosperms (1992) and others. For more precise definition of terms meaning the following dictionaries were used: M.C. Cooke, A Manual of Botanic Terms [1873?]; B.D. Jackson, A Glossary of Botanic Terms (1916); N.N. Zabinkova and M. E. Kirpichnikov, Latin-Russian Dictionary for Botanists (1957); M.E. Kirpichnikov and N.N. Zabinkova, Latin-Russian Dictionary for Botanists (1977), Biological Encyclopedic Dictionary (1986, 1989); The Oxford English Dictionary (1989) and others. In this encyclopedic edition the principles of constructing the dictionary with monographic description of embryonic structures and processes are combined. However, unlike traditional dictionary structure, the terminological articles in every part of the book are disposed not in alphabetical order, but according to theme so as to create an integrated picture of each main seed structure. A subject index is given at the end of the book. Terminological articles comprise the main part of the text and include: the definition and semantics of the terms, their history, and major data about the origin, development, functions and classification of the structure described. In a number of cases, questions concerning evolutionary transformations and the character of distribution of embryological features among flowering plants are discussed. The second group of articles combines conceptional, and hence the most complex articles with a discussion of questions of modern angiosperm embryology. To these are related a number of texts taken from native plant embryology classics, which have preserved their acuity to our time. These are the articles by S.G. Nawaschin and E.N. Gerassimova-Navashina. Some cases of difference between author terms and terms accepted in this edition are footnoted. Most of the terminological and conceptional articles are complemented with illustrations such as drawings, microphotographs (LM, TEM, SEM), schemes and diagrams. Complete bibliographical data are given in the References at the end of the volume. T. B. Batygina Z. I. Nikiticheva