Table Of ContentP. K. Stumpf and E. E. Conn
EDITORS-IN-CHIEF
Department of Biochemistry
and Biophysics
University of California
Davis, California
Volume I The Plant Cell
N. E. Tolbert, Editor
Volume 2 Metabolism and Respiration
David D. Davies, Editor
Volume 3 Carbohydrates: Structure and Function
Jack Pre is s, Editor
Volume 4 Lipids: Structure and Function
P. K. Stumpf, Editor
Volume 5 Amino Acids and Derivatives
B. J. Miflin, Editor
Volume 6 Proteins and Nucleic Acids
Abraham Marcus, Editor
Volume 7 Secondary Plant Products
E. E. Conn, Editor
Volume 8 Photosynthesis
M. D. Hatch and N. K. Boardman, Editors
THE
BIOCHEMISTRY
OF PLANTS
A COMPREHENSIVE TREATISE
Volume 5
Amino Acids and Derivatives
B. J. Miflin, editor
Department of Biochemistry
Rothamsted Experimental Station
Harp enden y England
1980
ACADEMIC PRESS
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COPYRIGHT © 1980, BY ACADEMIC PRESS, INC.
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United Kingdom Edition published by
ACADEMIC PRESS, INC. (LONDON) LTD.
24/28 Oval Road, London NW1 7DX
Library of Congress Cataloging in Publication Data
Main entry under title:
The Biochemistry of plants.
Vol. 5 edited by B. J. Miflin.
Includes bibliographies and index.
CONTENTS: v.l. The plant cell.—v. 2. Metabolism
and respiration.—[etc.] —v. 5. Amino acids and
derivatives.
1. Botanical chemistry. I. Stumpf, Paul Karl,
Date II. Conn, Eric E. III. Miflin, B. J.
QK861.B48 581.19'2 80-13168
ISBN 0-12-675405-5
PRINTED IN THE UNITED STATES OF AMERICA
80 81 82 83 9 8 7 6 5 4 3 2 1
List of Contributors
Numbers in parentheses indicate the pages on which the authors' contributions begin.
J. W. Anderson (203), Department of Botany, La Trobe University, Bun-
doora, Victoria 3083, Australia
Leonard Beevers (115), Department of Botany and Microbiology, University
of Oklahoma, Norman, Oklahoma 73019
J. K. Bryan (403), Department of Biology, Syracuse University, Syracuse,
New York 13210
Anne H. Datko (453), Laboratory of General and Comparative Biochemistry,
National Institute of Mental Health, Bethesda, Maryland 20205
K. J. F. Farnden (65), Department of Biochemistry, University of Otago,
Box 65, Dunedin, New Zealand
P. A. Fentem (271), Department of Botany, University of Manchester, Man
chester M13 9PL, England
D. G. Gilchrist (507), Department of Plant Pathology, University of Califor
nia, Davis, California 95616
John Giovanelli (453), Laboratory of General and Comparative Biochemis
try, National Institute of Mental Health, Bethesda, Maryland 20205
Curtis V. Givan (329), Department of Plant Biology, University of Newcastle-
upon-Tyne, Newcastle-upon-Tyne NEI 7RU, England
Richard H. Hageman (115), Department of Agronomy, University of Illinois,
Urbana, Illinois 61801
A. J. Keys (359), Department of Botany, Rothamsted Experimental Station,
Harpenden, Hertfordshire ALS 2JQ, England
T. Kosuge (507), Department of Plant Pathology, University of California,
Davis, California 95616
xi
xii List of Contributors
F. Larher (609), Laboratoire de Biologie Végétale, Universite de Rennes,
Rennes, France
Peder Olesen Larsen (225), Chemistry Department, Royal Veterinary and
Agricultural University, DK 1871 Copenhagen, Denmark
P. J. Lea (169, 569), Department of Biochemistry, Rothamsted Experimental
Station, Harpenden, Hertfordshire AL5 2JQ, England
A. F. Mann (271), Department of Botany, University of Manchester, Man
chester M13 9PL, England
Mendel Mazelis (541), Department of Food Science and Technology, Uni
versity of California, Davis, California 95616
B. J. Miflin (169, 533, 569), Department of Biochemistry, Rothamsted Ex
perimental Station, Harpenden, Hertfordshire AL5 2JQ, England
S. Harvey Mudd (453), Laboratory of General and Comparative Biochemis
try, National Institute of Mental Health, Bethesda, Maryland 20205
J. G. Robertson (65), Department of Scientific and Industrial Research,
Applied Biochemistry Division, Private Bag, Palmerston North, New
Zealand
G. R. Stewart (271, 609), Department of Botany, University of Manchester,
Manchester M13 9PL, England
John F. Thompson (375), U. S. Plant, Soil and Nutrition Laboratory, Ithaca,
New York 14853
M. G. Yates (1), Unit of Nitrogen Fixation, The Chemical Laboratory,
University of Sussex, Falmer, Brighton BN1 9QQ, England
General Preface
In 1950, James Bonner wrote the following prophetic comments in the
Preface of the first edition of his "Plant Biochemistry'' published by
Academic Press:
There is much work to be done in plant biochemistry. Our understanding of many basic
metabolic pathways in the higher plant is lamentably fragmentary. While the emphasis in
this book is on the higher plant, it will frequently be necessary to call attention to
conclusions drawn from work with microorganisms or with higher animals. Numerous
problems of plant biochemistry could undoubtedly be illuminated by the closer application
of the information and the techniques which have been developed by those working with
other organisms . . .
Certain important aspects of biochemistry have been entirely omitted from the present
volume simply because of the lack of pertinent information from the domain of higher
plants.
The volume had 30 chapters and a total o f490 pages. Many of the biochemi
cal examples cited in the text were derived from studies on bacterial, fungal,
and animal systems. Despite these shortcomings, the book had a profound
effect on a number of young biochemists since it challenged them to enter
the field of plant biochemistry and to correct "the lack of pertinent informa
tion from the domain of higher plants."
Since 1950, an explosive expansion of knowledge in biochemistry has
occurred. Unfortunately, the study of plants has had a mixed reception in
the biochemical community. With the exception of photosynthesis,
biochemists have avoided tackling for one reason or another the incredibly
interesting problems associated with plant tissues. Leading biochemical
journals have frequently rejected sound manuscripts for the trivial reason
that the reaction had been well described in E. coli and liver tissue and thus
was of little interest to again describe its presence in germinating pea seeds!
Federal granting agencies, the National Science Foundation excepted, have
xiii
xiv General Preface
also been reluctant to fund applications when it was indicated that the
principal experimental tissue would be of plant origin despite the fact that
the most prevalent illness in the world is starvation.
The second edition of "Plant Biochemistry'' had a new format in 1965
when J. Bonner and J. Varner edited a multiauthored volume of 979 pages;
in 1976, the third edition containing 908 pages made its appearance. A few
textbooks of limited size in plant biochemistry have been published. In
addition, two continuing series resulting from the annual meetings and sym
posia of phytochemical organizations in Europe and in North America
provided the biological community with highly specialized articles on many
topics of plant biochemistry. Plant biochemistry was obviously growing.
Although these publications serve a useful purpose, no multivolume
series in plant biochemistry has been available to the biochemist trained and
working in different fields who seeks an authoritative overview of major
topics of plant biochemistry. It therefore seemed to us that the time was ripe
to develop such a series. With encouragement and cooperation of Academic
Press, we invited six colleagues to join us in organizing an eight volume
series to be known as kThe Biochemistry of Plants: A Comprehensive
Treatise/' Within a few months, we were able to invite over 160 authors to
write authoritative chapters for these eight volumes.
Our hope is that this Treatise not only will serve as a source of current
information to researchers working in plant biochemistry, but equally impor
tant will provide a mechanism for the molecular biologist who works with£.
coli or the neurobiochemist to become better informed about the interesting
and often unique problems which the plant cell provides. It is hoped, too, the
senior graduate student will be inspired by one or more comments in chap
ters of this Treatise and will orient his future career to some aspect of this
science.
Despite the fact that many subjects have been covered in this Treatise, we
make no claim to have been complete in our coverage nor to have treated all
subjects in equal depth. Notable is the absence of volumes on phytohor-
mones and on mineral nutrition. These areas, which are more closely as
sociated with the discipline of plant physiology, are treated in multivolume
series in the physiology literature and/or have been the subject of specialized
treatises. Other topics (e.g., alkaloids, nitrogen fixation, flavonoids, plant
pigments) have been assigned single chapters even though entire volumes,
sometimes appearing on an annual basis, are available.
Finally, we wish to thank all our colleagues for their enthusiastic coopera
tion in bringing these eight volumes so rapidly into fruition. We are grateful
to Academic Press for their gentle persuasive pressures and we are indebted
to Ms. Barbara Clover and Ms. Billie Gabriel for their talented assistance in
this project.
P. K. Stumpf
E. E. Conn
Preface to Volume 5
This book is intended to cover those aspects of plant biochemistry relating
to the metabolism of amino acids with particular emphasis on amino acids
present in proteins. The treatment of the subject is in three parts. First, the
metabolism of sulfur and nitrogen from the inorganic sources available to
plants through to their incorporation into organic compounds is discussed; in
this part considerable emphasis has been placed on the process of nitrogen
fixation because of the large amount done in this field over recent years and
its potential importance in an energy restricted world. Second, the synthesis
of amino acids is considered according to the families in which they are
normally grouped. Because many papers in the literature show evidence of a
general lack of knowledge of the chemistry of amino acids (autoclaving
glutamine is one of the editor's pet hates) this second section is preceded by
a general discussion of amino acid chemistry. Also, throughout this section
the authors have attempted to answer the questions "where in the cell do the
reactions take place?' ' and "how are the pathways regulated?" as well as
dealing with the formal biochemistry of their topic. Finally, the third section
of the book tries to place the preceding chapters into the context of more
general aspects of plant metabolism by discussing the general catabolism of
amino acids, the movement of nitrogenous compounds around the healthy,
whole organism, particularly the interdependence of the various plant parts,
and finally the behavior (and probably the very important role) of amino
acids and related compounds in plants subject to various forms of stress.
Throughout the book I have tried to reduce duplication both between
chapters and between volumes in the treatise while preserving the right of
authors to view the subject in their own individual ways and trying to ensure
that each chapter and the volume as a whole stand as a coherent entity.
xv
xvi Preface to Volume 5
Insofar that the readers feel that the book fails in this regard or conversely
fails to cover the subject it is the editor and not the individual authors who
must bear the blame.
Finally in this brief preface I would like to thank all the authors for the
cooperation throughout this project and for the excellence of their contribu
tions. I have found reading and editing their contributions educative and
enjoyable. I would also like to thank Peter Lea and Michael Kirkman for
help and advice with various aspects of editing this volume, my family for
their tolerance and understanding, and lastly pay tribute to all the very hard
secretarial work, particularly on my own chapters, performed by Gillian
Watson and Susan Wilson.
B. J. Miflin
Biochemistry of Nitrogen 1
Fixation
M. G. YATES
I. Introduction 1
II. Biochemistry of Nitrogenase 3
A. Assay and Purification 3
B. Nomenclature 4
C. Components of Nitrogenase Systems 5
D. Metal Clusters in Nitrogenase Proteins 11
E. Third Component Proteins in Nitrogenase 20
F. Mechanism of Nitrogenase Activity 20
G. Discussion 34
III. Physiological Aspects of Nitrogen Fixation 35
A. Electron Transport to Nitrogenase 35
B. Protection of Nitrogenase against Damage by 0 44
2
C. Role of Leghemoglobin 50
D. Role of Hydrogenase 50
E. Regulation of Nitrogenase Activity in Vivo 53
F. Regulation of Nitrogenase Synthesis 55
IV. Conclusions 56
References 57
I. INTRODUCTION
After many years of scientific controversy, biological nitrogen fixation was
established as fact by the definitive experiments of Hellriegel and Wilfarth in
the 1880s (see Wilson, 1940). To the best of present-day knowledge this life
process is confined to, though widespread amongst, prokaryotic organisms:
it is found in both facultative and obligate anaerobes, obligate aerobes, au-
totrophs, chemoautotrophs and heterotrophs, photosynthetic bacteria, and
blue-green algae (Cyanobacteria). Early suggestions that fungi fix nitrogen
The Biochemistry of Plants, Vol. 5 1
Copyright © 1980 by Academic Press, Inc.
All rights of reproduction in any form reserved.
ISBN 0-12-675405-5
