Table Of ContentCRC
Handbook of
Flowering
Volume VI
Editor
Abraham H. Halevy
Professor
Department of Ornamental Horticulture
The Hebrew University of Jerusalem
Rehovot
Israel
Boca Raton London New York
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Library of Congress Cataloging-in-Publication Data
(Revised for vol. 6)
CRC handbook of flowering
(Includes bibliographies and indexes.
1. Plants, Flowering of—Handbooks, manuals,
etc. 2. Plants, Cultivated—handbooks, manuals,
etc. I. Halevy, Abraham H., 1927-
II. Handbook of flowering.
SB126.8.C73 1985 635.9 83-21061
ISBN 0-8493-3910-3 (set)
ISBN 0-8493-3911-1 (v. 1)
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FOREWORD
Life would be awfully dull without flowers. The love of flowers is common to all people
all over the world, no matter how different their cultural patterns may be. Artistically,
flowers play an important role in poetry and in decorative arts for their attractive structure,
color and odor in endless variation: for their eternal beauty.
Agriculturally, and perhaps still more so horticulturally, flowers play an essential part in
the production of all crops. This is, of course, most evident in floriculture, where the flower
is the final product. In all fruit crops, whether deciduous or evergreen, and in crops like
grains and the fruit-vegetables, the fruit is the final product, but flowers are the indispensable
introductory step to fruit formation. For the many crops which are grown for their seeds,
including all breeding projects, flowers and fruits cannot be missed for seed formation.
On the other hand, there are crops where flowers and fruits are undesirable — for instance,
those which are grown for their roots, stems, or leaves. However, in seed growing and
breeding of these crops, flowers again are indispensable.
The above implies that control of flowering, whether positively promoting or negatively
preventing flowering, is a cultural measure which has to be applied in the growing of most,
if not all, crops. No wonder that in several cases more or less sophisticated methods for
controlling flowering are rather well known. Quite generally, such empirically developed
methods are genetical and ecological, but an endless variation occurs in the requirements
for flowering among different species and even among the cultivars of one species.
The human mind is not satisfied by answering the question as to which factors influence
flowering — as approached empirically — it also wants to know the physiological background
of the actions of these factors. This is a much more difficult problem, and disciplines like
biophysics, biochemistry, and molecular biology enter the picture. It is self-evident that this
approach has made much less progress than the empirical one. However, it has become
clear that also in this respect an endless variation occurs among different species and that
a general mechanism of action — if it exists — has not been found yet. Be that as it may,
the study of flowering, whether empirical or — on a higher level — physiological, contributes
to the development of the human mind by the attempt to understand some of the mysteries
of Life.
In the foregoing I used "endless variation" no less than three times, and students of
flowering, who concentrate on their own plant(s), run the risk of becoming one-sided. It is,
therefore, highly important to provide them with a catalogue of the flowering behavior of
as many other plants as possible. The present Handbook fills the existing gap by bringing
together our knowledge of the individual cases. The "Table of Contents" illustrates that it
was no simple effort to compose this edition. A. H. Halevy, "Abe" to his friends, has had
the courage and the energy to undertake this task. All those who are interested in flowering
in some or other way owe him their gratitude.
S. J. Wellensiek
PREFACE
Rise up, my love, my fair one, and come away.
For, lo, the winter is past,
The rain is over and gone;
The flowers appear on the earth;
The time of singing is come,
And the voice of the turtledove is heard in our
land;
The fig tree puts forth her green figs,
And the vines in blossom give forth their
fragrance.
(Song of Songs 2:10—13)
From antiquity, poets have expressed humanity's association of flowering with spring,
renewal, singing, beauty, fragrance, and love. This book deals with the more prosaic aspects
of flowering: flower formation and development and the environmental and physiological
factors which regulate them.
Several excellent reviews and books on flowering have been published in the last 25
years. These include Lang's6 and SchwabeV chapters in general encyclopedias of plant
physiology, Evans' opening and concluding chapters in his book,4 the books by Salisbury,8
Vince-Prue," and Bernier et al.1-13 and the several review articles in the Annual Reviews of
Plant Physiology.2-*^-1 •1(U2 With the exception of Evans,4 these authors have presented a
general review of the flowering process and attempted to integrate the data into a unifying
theory. Such unifying theories have generally suffered from the disadvantages noted by
Evans in the preface to his book;4 they deal primarily with the earliest events of the flowering
process, and they are based on data obtained from a small number of "model" plants.
Evans' book, which contains flowering "case histories" of the majority of these species,
includes chapters on only 20 plants, of which only one is a woody plant, two are mono-
cotyledons, and none is a gymnosperm. A great wealth of data on the regulation of flower
formation and development can be found in the practical literature of agriculture, horticulture,
and forestry. Much of this has been often ignored by flowering physiologists. The flowering
process is indeed much more diverse than that revealed in the 20 "model" plants presented
in Evans' book. It was my aim in planning this book to make a more comprehensive view
of the flowering process possible by presenting relevant data from as many plants as possible.
This includes the majority of the cultivated plants on which such information is available:
field crops, fruits, vegetables, ornamentals, industrial plants, and forest trees, not only of
the temperate regions, but also of subtropical and tropical climates. To accomplish this goal
I have invited scientists from all over the world to contribute chapters on specific plants or
groups of plants. Many of the authors have not only reviewed the available literature, but
have also included previously unpublished data. Many of the chapters present the first general
review of the flowering process in their specific subject area.
The book deals with all aspects of flowering, including juvenility and maturation, flower
morphology, flower induction, and morphogenesis to anthesis. Flower morphogenesis has
been taken to include also development of individual flower parts, sex expression, and flower
malformations. When possible, the authors have attempted to present information on all
stages of the flowering process. In many cases, however, this has not been feasible, since
little or nothing is known about some of the stages. In most cases the "flowering story" is
terminated at anthesis. In some plants, however, flower structure and anthesis are directly
related to pollination, and in these cases pollination is also included. In some commercial
food crops the description is also extended somewhat beyond anthesis to include important
factors in crop production. In many cultivated plants, mostly ornamentals, practical methods
for manipulation of flowering are included.
The length of the individual chapters and the emphasis on specific aspects depends in
most cases on the availability of experimental information and not on the importance of the
plant as a crop or the significance of the physiological stage described. Some important
economic plants are absent from this book since little or no information is available on their
flowering. In some cases most of the chapter is devoted to a single aspect of flowering,
such as juvenility, flower induction, flower development, sex expression, cleistogamy,
development of certain flower parts, or flower opening. Some chapters concentrate on
physiological aspects, others on ecological, morphological or genetic ones. Other aspects
are covered only briefly or even absent, not because they are not important but because they
have not been studied in detail.
I am well aware that this book is far from being a comprehensive encyclopedia of flowering,
I would greatly appreciate comments from readers on errors found in articles, missing
information, and plants not included in the book, whose flowering process have been
documented.
The handbook consists of six volumes:
• Volume I — contains general chapters on groups of plants, and individual chapters
on plants beginning with the letter A
• Volume II — contains plants of letters B to E
• Volume III — contains plants of letters F to O
• Volume IV — contains plants of letters P to Z
• Volume V — is a supplementary volume and includes chapters on plants not included
in Volumes I to IV, listed in alphabetical order.
• Volume VI — a second supplementary volume
The merit of the book rests upon the work of the individual authors. I am grateful to them
for their efforts, cooperation, and forbearance. I would like to thank my colleagues in the
Editorial Board who helped me to select the authors and to review the chapters, and the
many other colleagues who helped in reviewing (and sometimes rewriting) specific chapters.
I hope that this handbook will serve as a reference and source book for scientists interested
in the flowering process of particular plants, and will draw their attention to the lack of
information on important aspects of the flowering process in many important plant species.
I also hope that the wealth of information accumulated here will be useful in future attempts
to synthesize general theories of the physiology of flowering.
REFERENCES
1. Bernier, G., Kinet, J. M., and Sachs, R. M., The Physiology of Flowering, CRC Press, Boca Raton,
Fla., 1981, Vol. 1, 149; Vol. 2, 231.
2. Chouard, P., Vernalization and its relation to dormancy, Annu. Rev. Plant Physiol., 11, 191—238, 1960.
3. Doorenbos, J. and Wellensiek, S. J., Photoperiodic control of floral induction, Annu. Rev. Plant Physiol.,
10, 147—184, 1959.
4. Evans, L. T., Ed., The Induction of Flowering. Some Case Histories, Macmillan, Melbourne, 1969, 328.
5. Evans, L. T., Flower induction and the flowering process, Annu. Rev. Plant Physiol., 22, 365—394,
1971.
6. Lang, A., The physiology of flower initiation, in Encyclopedia of Plant Physiology, Vol. 15 (Part I),
Ruhland, W., Ed., Springer-Verlag, Berlin, 1965, 1380—1563.
7. Salisbury, F. B., The Flowering Process, Pergamon Press, Oxford, 1963, 234.
8. Salisbury, F. B., Photoperiodism and the flowering process, Anna. Rev. Plant Physiol., 12, 293—326,
1961.
9. Schwabe, W. W., Physiology of vegetative reproduction and flowering, in Plant Physiology, A Treatise,
Steward, F. C., Ed., Vol. VI-A, Academic Press, New York, 1971, 233^11.
10. Searle, N. E., Physiology of Flowering, Anna. Rev. Plant Physiol., 16, 97—118, 1965.
11. Vince-Prue, D., Photoperiodism in Plants, McGraw Hill, London, 1975, 444.
12. Zeevaart, J. A. D., Physiology of flower formation, Annu. Rev. Plant Physiol., 27, 321—348, 1976.
13. Kinet, J. M., Sachs, R. M., and Bernier, G., The Physiology of Flowering, Vol. 3, CRC Press, Boca
Raton, Fla., 1985, 274.
Abraham H. Halevy
PREFACE
Volume VI
This final second supplementary volume of the Handbbook of Flowering is the largest of
this treatise, containing 84 chapters by 80 authors. As with Volume V, some of the included
chapters were planned for earlier volumes and some were commissioned specifically for this
volume. In a few cases, I invited colleagues to contribute new chapters on plants already
previously described (Anigozanthus, Trifolium, Zantedeschia), when I realized that important
information had not been included in earlier chapters.
This volume includes a major contribution by Anton Lang on Nicotiana and four major
chapters on coniferous forest trees, as well as others on important crops and model plants
of flowering physiology.
The six volumes of this encyclopedia include 380 chapters on specific plants or groups
of plants, written by more than 250 authors. Since these plants were arranged three times
in alphabetical order in Volumes I to IV, V, and VI, and many plants were described in
general chapters according to categories (e.g., Forage Plants), we have included a compre-
hensive taxonomic index of all the plants mentioned in Volumes I through VI.
Although this Handbook is the most comprehensive book on flowering published to date,
I am well aware that it does not contain all the relevant available information on the subject.
Almost 20 years have passed since the publication of Evans' book and this final volume
of the present Handbook. I sincerely hope that a second, better, and more comprehensive
edition of the Handbook of Flowering will be published in less time.
I have worked on this book for more than 8 years. Many colleagues have helped me carry
out this undertaking, for which I am very grateful. I would especially like to thank Mr. and
Mrs. Carl Pearlstein and their family of San Francisco, who so generously supported the
research on flowering in my laboratory during the last 10 years. Without their continuous
help and interest, it would have been impossible for me to continue my research activities
while working on the book.
Abraham H. Halevy
THE EDITOR
Abraham H. Halevy, Ph.D., is Professor of Horticulture and Plant Physiology at the
Hebrew University of Jerusalem, Rehovot, Israel.
Dr. Halevy obtained his M.Sc. and Ph.D. degrees from the Hebrew University in 1953
and 1958, respectively.
Dr. Halevy was a Research Fellow at the U.S. Department of Agriculture Research Center
at Beltsville, Md. (1958 to 1959) and has been a Visiting Professor at Michigan State
University (1964 to 1965) and the University of California, Davis (1970 to 1971, 1982 to
1984, and 1986).
He has twice received the Alex Laurie award of the American Scoiety of Horticultural
Science, and was recently (1983) nominated as a fellow of the society.
Dr. Halevy is the Editor-in-Chief of the Israel Journal of Botany and the Flowering
Newsletter. He has published over 200 scientific papers in international journals, and has
been a guest invited lecturer in numerous symposia around the world.
His current research interests are the regulation of flowering and of flower senescence.
CONTRIBUTORS
Aage S. Andersen, Ph.D. Osvaldo H. Caso, Dr. in Natural
Department of Horticulture Sciences
Agricultural University Centre de Ecofisiologia Vegetal
Copenhagen CONICET
Denmark Buenos Aires
Argentina
Vera M. Andrade, Dr. in Sciences
Department of Applied Biology
Michael St. John Clowes, D.Phil.
UNESP
Tea Research Foundation
Jaboticabal, Sao Paulo
Mulanje
Brazil
Malawi
Allan M. Armitage, Ph.D.
Lila Cohen, Ph.D.
Department of Horticulture
Department of Botany
University of Georgia
Tel Aviv University
Athens, Georgia
Tel Aviv
Israel
Yolande Arnaud
Laboratoire du Physiologie
du Developpement des Plantes Anthony J. Davy, Ph.D.
Universite Pierre et Marie Curie School of Biological Sciences
(Paris VI) University of East Anglia
Paris Norwich
France England
Tova Arzee, Ph.D.
August A. De Hertogh, Ph.D.
Department of Botany
Department of Horticultural Science
Tel Aviv University
North Carolina State University
Tel Aviv
Raleigh, North Carolina
Israel
Amos Blumenfeld, Ph.D. Tom J. de Jong, Ph.D.
Department of Subtropical Horticulture Department of Population Biology
Volcani Center University of Leiden
Agricultural Research Organization Leiden
Bet Dagan Netherlands
Israel
Nativ Dudai
Bertrand Boeken, Ph.D.
Experiment Station, Newe Ya'ar
Department of Biology and
Agricultural Research Organization
Jacob Blaustein Institute
Haifa
for Desert Research
Israel
Sede Boqer Campus
Israel
Rivka Dulberger, Ph.D.
J. Br0ndum-Pedersen Department of Botany
Research Centre for Horticulture G.S. Wise Center for Life Sciences
Institute of Glasshouse Crops Tel Aviv University
Arslev Ramat Aviv
Denmark Israel
