Table Of ContentEMBRYOLOGY 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
