Light and Plant Development

497 Pages·1976·18.378 MB·English

by  H. Smith (Auth.)

Published Proceedings of Previous Easter Schools in Agricultural Science
SOIL ZOOLOGY*
Edited by D.K.McE. Kevan (Butterworths, London, 1955)
THE GROWTH OF LEAVES*
Edited by F.L. Milthorpe (Butterworths, London, 1956)
CONTROL OF THE PLANT ENVIRONMENT*
Edited by J.P. Hudson (Butterworths, London, 1957)
NUTRITION OF THE LEGUMES*
Edited by E.G. Hallsworth (Butterworths, London, 1958)
THE MEASUREMENT OF GRASSLAND PRODUCTIVITY*
Edited by J.D. Ivins (Butterworths, London, 1959)
DIGESTIVE PHYSIOLOGY AND NUTRITION OF THE RUMINANT*
Edited by D. Lewis (Butterworths, London, 1960)
NUTRITION OF PIGS AND POULTRY*
Edited by J.T. Morgan and D. Lewis (Butterworths, London, 1961)
ANTIBIOTICS IN AGRICULTURE*
Edited by M. Woodbine (Butterworths, London, 1962)
THE GROWTH OF THE POTATO*
Edited by J.D. Ivins and F.L. Milthorpe (Butterworths, London, 1963)
EXPERIMENTAL PEDOLOGY*
Edited by E.G. Hallsworth and D.V. Crawford (Butterworths, London, 1964)
THE GROWTH OF CEREALS AND GRASSES*
Edited by F.L. Milthorpe and J.D. Ivins (Butterworths, London, 196 5)
REPRODUCTION IN THE FEMALE MAMMAL*
Edited by G.E. Lamming and E.C. Amoroso (Butterworths, London, 1967)
GROWTH AND DEVELOPMENT OF MAMMALS
Edited by G.A. Lodge and G.E. Lamming (Butterworths, London, 1968)
ROOT GROWTH*
Edited by W.J. Whittington (Butterworths, London, 1968)
PROTEINS AS HUMAN FOOD
Edited by R.A. Lawrie (Butterworths, London, 1970)
LACTATION
Edited by J.R.Falconer (Butterworths, London, 1971)
PIG PRODUCTION
Edited by D.J.A. Cole (Butterworths, London, 1972)
SEED ECOLOGY
Edited by W. Hey decker (Butterworths, London, 1973)
HEAT LOSS FROM ANIMALS AND MAN: ASSESSMENT AND CONTROL
Edited by J.L. Monteith and L.E. Mount (Butterworths, London, 1974)
MEAT
Edited by D.J.A. Cole and R.A. Lawrie (Butterworths, London, 1975)
PRINCIPLES OF CATTLE PRODUCTION
Edited by Henry Swan and W.H. Broster (Butterworths, London, 1976)
* These titles are now out of print
Light and Plant
Development
H. SMITH
Department of Physiology and Environmental Studies
University of Nottingham School of Agriculture
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First published 1976
ISBN 0 408 70719 4
© The several contributors named in the list of contents, 1976
LIBRARY OF CONGRESS CATALOGING IN PUBLICATION DATA
Easter School in Agricultural Science, 22d, University
of Nottingham, Eng., 1975.
Light and plant development.
Bibliography: p.
Includes index.
1. Plants, Effect of light on -Congresses I . Smith,
Harry. II. Title.
QK757.E26 1975 581. Γ9Ί53 76-10178
ISBN 0-408-70719-4
Printed in England by Cox & Wyman Ltd, London, Fakenham and Reading
PREFACE
This volume presents the Proceedings of the 22nd University of
Nottingham Easter School in Agricultural Science which was held at
Sutton Bonington from April 7th to 10th, 1975. Each Easter School
is intended to be a forum in which a specific problem of particular
importance to agriculture can be discussed by specialists from science,
commerce, industry and practical agriculture. In most cases, the
immediacy of the chosen topics for practical agriculture is clearly
evident; in others, the subject areas are as yet of potential importance
only to agricultural productivity, 'light and Plant Development' comes
into the latter category. In planning and organising this Easter School,
I have endeavoured to produce a volume which will provide a fully
comprehensive and up-to-date summary of the control of growth and
development of higher plants by light. Certain aspects, however, have
been omitted, notably photosynthesis, which obviously could take up
several volumes in its own right. Similarly, phototropism receives scant
attention, although Chapter 2 contains the first report of really good
evidence on the chemical nature of the photoreceptor. The concentra­
tion of virtually the whole of the volume on phytochrome may seem,
at first glance, unbalanced, but it does reflect realistically our relatively
high level of understanding of phytochrome compared with our almost
complete ignorance of other photoreceptors. It is hoped that this
volume will assist crop scientists to come to grips with a topic which
is still almost totally studied at the fundamental level, but which never­
theless has great potential importance for crop production.
I am gratified to be able to say that the Easter School, and the
Annual European Photomorphogenesis Symposium which followed
(April 10th- 12th), were highly successful. This was due principally to
patient and conscientious organisation by the Conference Secretary,
Miss Edna Lord, who carried an extremely heavy burden for several
weeks without complaint — I am deeply grateful to her. I also wish
to record my appreciation for the assistance given by Mrs. Laurel Dee
and Mrs. Carol Stanton, who typed the abstracts, Mrs. Clarice Ingram,
Mrs. Lena Parker, Mrs. Valerie Blunt and Mr John Blunt, who organised
the visual aids, and Miss Audrey Evans, Mr Geoffrey Holmes, Mr Sandy
Giles and Dr Chris Johnson, who looked after various aspects of parti­
cipant welfare. In planning the two meetings I was advised by Professor
J.L. Monteith, Dr W.R. Briggs and Dr Daphne Vince-Prue, to all of
whom I am grateful.
Special thanks should also go to the Chairmen of the Sessions, all
of whom accepted my invitation to take on this task at very short
Preface
notice; they were Professor J.L. Monteith, Dr W.R. Briggs, Professor
H. Mohr, Professor W. Haupt, Professor A.W. Galston and Dr W.S.
Hillman. I should also like to express my gratitude to Plant Protection
Ltd., Bracknell, for a very welcome financial contribution, and to the
Royal Society, the Agricultural Research Council and the British Council,
who separately assisted three of the overseas contributors to travel to
Britain.
Finally, I should like to thank the 128 participants from 20 countries
who, by their friendliness, created a superb atmosphere for the communi­
cation of ideas.
H. SMITH
To
H.A. BORTHWICK and S.B. HENDRICKS
By the unanimous approval of the participants at the 22nd University
of Nottingham Easter School in Agricultural Science, this volume is
dedicated to the late Dr Harry A. Borthwick and to Dr Sterling B.
Hendricks, whose joint inspiration and perseverance over more than 35
years have laid the basis of our present knowledge and understanding
of photomorphogenesis.
1
H.A. BORTHWICK AND S.B. HENDRICKS - PIONEERS OF
PHOTOMORPHOGENESIS1
WINSLOW R. BRIGGS
Department of Plant Biology, Carnegie Institution of Washington,
Stanford, California 94305, U.S.A.
The participants in this 22nd Easter School take great pleasure in
dedicating this volume to Dr Harry A. Borthwick and Dr Sterling B.
Hendricks in honour of their monumental contributions to the field to
which this School is devoted. Their early interest in flowering led
them into a remarkable series of studies, the results of which are
fundamental to most of what is discussed in this volume. Their
research was carried out at the Pioneering Laboratory of Plant Physio­
logy at the Plant Industry Station of the United States Department of
Agriculture; they and their many colleagues made it one of the most
distinguished laboratories in the world, and unique in the Department
of Agriculture.
While it is not the purpose of these short paragraphs to review all
of their work, it is appropriate to list some of their major accomplish­
ments in photomorphogenesis. In 1946, in one of a series of studies
with Parker, they carefully examined the spectral sensitivity of inhibition
of flowering by a light break in the middle of the night in the two
short-day plants Biloxi soybean and cocklebur (Parker et al, 1946).
The results were the first action spectrum for a phytochrome effect,
and showed an action maximum near 665 nm. Two years later came
an action spectrum for night interruption in Wintex barley, showing
for the first time flowering promotion in a long-day plant under non-
inductive photoperiod (Borthwick, Hendricks and Parker, 1948). The
action spectrum was essentially identical with that for the short-day
plants, and constituted the first evidence that the same photoreceptor
might regulate flowering in both types of photoperiodically sensitive
plants. The next year (Parker et al, 1949) they were joined by
F.W. Went to measure the action spectrum for leaf enlargement and
stem growth inhibition in dark-grown pea seedlings. They thus
obtained the first of many by now familiar action spectra for vegeta­
tive responses to phytochrome phototransformation. A year later
(Parker, Hendricks and Borthwick, 1950) appeared another study on a
long-day plant, Hyoscyamus. The work was really a condensed version
1 CIW-DPB Publication No.553
1
2 Borthwick and Hendricks - pioneers of photomorphogenesis
of the Wintex barley paper, but it is significant in that the authors
suggest for the first time a phycocyanin-like pigment as the photo-
receptor-a full 15 years before the suggestion was verified experimen­
tally.
They then turned their attention to light-promoted lettuce seed
germination (Borthwick et al, 1952), a system first discovered by
Flint and McAlister in the mid 1930s at the Smithsonian Institution.
An action spectrum for promotion of germination showed a maximum
at 665 nm, suggesting the same photoreceptor to be involved. However,
since the lettuce seed system was known to be reversible by far-red
light, they also obtained the first action spectrum for far-red reversal.
The same paper reported repeated photoreversibility following several
sequential alternating red and far-red treatments, and the authors
proposed for the first time a single photoreversible pigment. A second
paper on lettuce seed germination (Borthwick et al, 1954) explored
the system in greater detail, demonstrating that the light reactions in
both directions were independent of temperature, and showing for the
first time escape from photoreversibility with time after far-red but
not after red treatment.
Meanwhile, Borthwick, Hendricks and Parker (1952) asked the
suddenly obvious question: since the action spectrum for promotion
of lettuce seed germination was identical with those measured for the
other effects, could these other effects be reversed by far-red light as
well? Using cocklebur seedlings grown under day lengths that were
sufficiently short for flowering, they showed that far-red light com­
pletely cancelled the effect of red light in inhibiting flower formation.
The action spectrum for this far-red reversibility was roughly the same
as that for lettuce seed germination. Photoreversibility was thus not
just a peculiar property of certain varieties of lettuce seeds.
Other workers at Beltsville then demonstrated similar photoreversible
control of flavone formation in tomato skins (Piringer and Heinze,
1954) and promotion of leaf enlargement and epicotyl elongation and
inhibition of hypocotyl elongation in bean seedlings (Downs, 1955).
Then Hendricks, Borthwick and Downs (1956) showed both with
Pinto bean internodal elongation and Lepidium and lettuce seed germ­
ination that the photochemistry followed first-order kinetics for both
red and far-red photoresponses. By this time, the Beltsville group was
convinced that a single photoreversible pigment was involved, and they
set out to find it.
Their success in detecting phytochrome spectrophotometrically in
plant tissue and achieving preliminary isolation in buffer is documented
in their often cited 1959 paper (Butler et al, 1959), a paper that
marked the beginning of in vitro work with phytochrome. It also
presented instrumentation techniques which are now widely used, not
only for phytochrome but also for a wide variety of other types of
spectral studies. There followed from Beltsville the first action spec­
trum for phytochrome phototransformation in vitro (Butler, Siegelman
and Hendricks, 1964), the first partial purification technique for
Winsiow R. Briggs 3
phytochrome (Siegelman and Firer, 1964), establishment of the chromo-
phore as a bilitriene similar to allophycocyanin (Siegelman and Hendricks,
1965; Siegelman, Turner and Hendricks, 1966), and numerous other
papers on phytochrome reactions both in vivo and in vitro. The
demonstration (Fondeville, Borthwick and Hendricks, 1966) that leaflet
closing in Mimosa was under rapid phytochrome control gave strong
impetus to the already existing suspicion that phytochrome was some­
how acting in association with a membrane system. This evidence,
summarised by Hendricks and Borthwick (1967), was integrated into a
useful model for phytochrome action (Borthwick et al., 1969).
Borthwick retained an active interest in phytochrome (Borthwick,
1972a,b) until his death on May 21st, 1974. Hendricks continues a
vigorous pursuit of the problem of seed germination (Hendricks and
Taylorson, 1975), where new ground is again being broken.
Among other things, what emerges from this listing of contributions
is the extraordinary breadth of inquiry that Borthwick and Hendricks
applied to plant photomorphogenesis. Their approaches ranged from
whole plant physiology through organ physiology, biochemistry, photo­
chemistry and biophysics. We commend them for this breadth, and
for the durability, magnitude and insight of their research.
References
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Shropshire, W. Jr. Academic Press, New York
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