Table Of ContentAdvances in
BOTANICAL RESEARCH
incorporating Advances in Plant Pathology
Editor-in-Chief
J. A. CALLOW School of Biological Sciences, University
of Birmingham, Birmingham, UK
Editorial Board
J. H. ANDREWS University of Wisconsin-Madison,
Madison, USA
H. G. DICKINSON University of Oxford, Oxford, UK
M. KREIS Universitk de Paris-Sud, Orsay, France
R. M. LEECH University of York, York, UK
R. A. LEIGH Rothamsted Experimental Station,
Harpenden, UK
E. LORD University of California, Riverside, USA
D. J. READ University of Shefield, Shefield, UK
I. C. TOMMERUP CSIRO, Perth, Australia
Advances in
BOTANICAL RESEARCH
incorporating Advances in Plant Pathology
Classic Papers
edited by
J. A. CALLOW
School of Biological Sciences,
University of Birmingham,
Birmingham, UK
VOLUME 27
1997
ACADEMIC PRESS
San Diego London Boston New York Sydney Tokyo Toronto
This book is printed on acid-free paper
Copyright @ 1997 by ACADEMIC PRESS
All rights reserved.
No part of this publication may be reproduced or transmitted in any form
or by any means electronic or mechanical, including photocopy,
recording, or any information storage and retrieval system, without
permission in writing from the publisher.
Academic Press
525 B Street, Suite 1900, San Diego, California 92101-4495, USA
http://www.a pnet .com
Academic Press Limited
24-28 Oval Road, London NW1 7DX, UK
http:llwww.hbuk.co.uWapl
A catalogue record for this book is available from the British Library
ISBN 0-12-005927-4
Typeset by Keyset Composition, Colchester, Essex
Printed in Great Britain by Hartnolls Limited, Bodmin, Cornwall
97 98 99 00 01 02 EB 9 8 7 6 5 4 3 2 1
CONTRIBUTORS TO VOLUME 27
D. BOULTER, Department of Biological Sciences, University of Durham,
South Road, Durham DHI 3LE, UK
W. W. CARMICHAEL, Department of Biological Sciences, Wright State
University, Dayton, Ohio 45435, USA
R. R. D. CROY, Department of Biological Sciences, University of Durham,
South Road, Durham DHI 3LE, UK
C. J. HOWE, Department of Biochemistry and Cambridge Centre for
Molecular Recognition, University of Cambridge, Cambridge CB2 1 QW,
UK
R. KUNZE , Ludwig- Maximilians- Universitat Munchen, Institut fur Genetik
und Mikrobiologie, Maria- Ward-Str. IA, 80638 Miinchen, Germany
T. LARKUM, School of Biological Sciences, University of Sydney, NSW
2006, Australia
W.-E. LONNIG, Max-Planck-Institut fur Zuchtungsforschung, Carl-von-
Linnt-Weg 10, 50829 Koln, Germany
J. A. RAVEN, Department of Biological Sciences, University of Dundee,
Dundee DD1 4HN, UK
H. SAEDLER, Max-Planck-Institut fur Zuchtungsforschung, Carl-von-
Linnt-Weg 10, 50829 Koln, Germany
PREFACE
The first volume of Advances in Botanical Research appeared in 1963. Since
then over 150 articles have been published in 27 volumes. The series has been
consistently well received with a high IS1 rating in the plant science sector
and has always been concerned to commission articles of substance, without
the tight page limits often imposed by other review series, in which the
foundations of a topic can be laid, hypotheses elaborated, and original
evidence brought to bear. The idea for the present volume arose from a
citations analysis of the articles published since the start of the series. The
editorial board considered which of the subject areas represented by the ‘top
ten’ most cited articles had advanced significantly. From this short-list
invitations were issued to five authors to revisit the original topic of their
reviews, up-dating and extending it as appropriate. All accepted. The result
is this ‘Classic Papers’ volume.
The original article ‘Biochemistry of storage protein synthesis and deposi-
tion in the developing legume seed’ was written by D. Boulter in 1981. The
intervening 16 years has seen two main developments which compelled the
choice of this article for an update by Boulter and his co-worker R. Croy;
viz. recent advances in 3-D structural analysis, and the advent of methods
for the manipulation of genes for biotechnological purposes.
N. Kerby and J. A. Raven wrote their original article ‘Transport and
fixation of inorganic carbon by marine algae’ 11 years ago. In this up-date,
Raven considers not only the significant advances in our understanding of
the biochemical mechanisms involved, but also puts this new information into
evolutionary and biogeochemical contexts.
Since W. W. Carmichael wrote his article on ‘Cyanotoxins’ in 1986, there
has been a major expansion in research programmes on this topic as the
negative impacts of these toxins on water quality and aquaculture industries
has been realized. Carmichael reports on recent advances in the under-
standing of the chemistry, biosynthesis, regulation, mechanism of action,
molecular genetics and toxicology of these toxins.
A. W. D. Larkum and J. Barrett commented in 1981 that understanding
of light-harvesting processes in algae tended to be dominated by models
derived from the much more uniform photochemical systems of higher plants.
Algae exhibit a much greater diversity in their photochemical apparatus and
light-harvesting strategies and the up-dated article by Larkum and C.J . Howe
reviews the impact of molecular genetics (to which many algae are especially
amenable) on our understanding of the structure and regulation of algal
photosystems.
xxiv PREFACE
The major focus of the 1986 article on plant transposable elements (P.
Nevers, N. Shepherd and H. Saedler) was the role played by these mobile
genetic elements in generating variegation, especially in floral organs. The
authors commented at the time that the field was becoming increasingly
important because, in addition to illuminating studies on molecular develop-
ment, there was a growing realization that transposable elements are
important in evolution. They also speculated that transposable elements
would become increasingly important as tools, in the cloning of genes known
only for their phenotype, and in the genetic manipulation of crop plants for
biotechnological purposes. Since 1986 the field has burgeoned and the
up-dated article by R. Kunze, H. Saedler and W.-E. Lonnig comprehensively
reviews important advances in the occurrence of transposable elements and
the mechanism of transposition, but major sections on their role in evolution
and their use as molecular tools, amply justify the faith of the earlier
article.
J. A. Callow
ABBREVIATIONS
ACh acetylcholine
AnTx-a anatoxin-a
CAM crassulacean acid metabolism
CF putative nucleic-acid-binding function
CHO Chinese hamster ovary
CLO cyanobacteria-like organisms
CNS central nervous system
CTP cyanobacteria toxin poisoning
CY cylindrospermopsin
DCR 0.35 kb GC-rich first exon fragment
DSP diarrhoetic shellfish poison
ESTs expressed sequence tags
FAB fast atom bombardment
HABs harmful algal blooms
IOC International Oceanographic Commission
LTRs long terminal repeats
MAC maximum acceptable concentration
MITES miniature inverted-repeat transposable elements
NLS nuclear localization sequences
NMR nuclear magnetic resonance
ORF open reading frame
PBPs ph ycobiliproteins
PBS primer binding site
PCOC photorespiratory carbon dioxide cycle
PCR polymerase chain reaction
PDB Belemnite from a Cretaceous Pee-Dee formation
PEPCK phosphoenol pyruvate carboxykinase
PFD photon flux density
PLC primary liver cancer
PPIA protein phosphatase inhibition assays
PPT polypurine tract
PSP paralytic shellfish poison
RSSs recombination signal sequences
RT reverse transcriptase
RUBISCO ribulose bisphosphate carboxylase oxygenase
TEs transposable elements
TIR terminal inverted repeat sequences
xxvi ABBREVIATIONS
TNPD putative EnlSpm transposase
TPase transposase
TSDs target site duplications
UCR GC-rich 0.2 kb promoter region
uv ultraviolet
VLPS virus-like particles
The Structure and Biosynthesis of Legume Seed Storage
Proteins: A Biological Solution to the Storage of Nitrogen in
Seeds
D . BOULTER and R . R . D . CROY
Department of Biological Sciences. University of Durham. South
Road. Durham DHl 3LE. UK
I . Introduction .............................................................................. 2
I1 . What are Legume Seed Storage Proteins? ..................................... 4
111. Why More than One Storage Protein? .......................................... 7
IV . Nomenclature .......................................................................... 10
V. ................................................................................ 11
y and Subunit Structure ............................................. 11
B . Secondary. Tertiary and Quaternary Structure ........................ 16
VI Legumins ................................................................................ 25
A . Primary and Subunit Structure ............................................. 25
B . Secondary. Tertiary and Quaternary Structure ........................ 29
VII . The 2s Albumins and Globulins ................................................. 32
A . Primary and Subunit Structure ............................................. 32
B . Secondary. Tertiary and Quaternary Structure ........................ 33
VIII . Synthesis. Processing. Intracellular Targeting and Deposition of
Storage Proteins ....................................................................... 34
IX . Storage Proteins as Part of the Process of Seed Formation ............. 36
x . Storage Protein Genes .............................................................. 40
A . Vicilin Genes ................. ............................................ 40
B . Legumin Genes .............. ............................................ 40
C . Structure and Regulation of Expression ................................ 41
Advances in Botanical Research Vol . 27 Copyright 0 1997 Academic Press Limited
ISBN (1-12-005927-4 All rights of reproduction in any form reserved
2 D. BOULTER & R. R. D. CROY
D. Why Multiple Copies of Storage Protein Genes? .................... 43
E. Structural Similarities between Vicilin and Legumin - Evidence
for a Common Ancestral Gene ............................................ 44
XI. Structural Features and Constraints Related to Function ................. 45
A. Conserved Structures in Vicilin and Legumin Proteins ............. 45
B. Hydration and Packing ....................................................... 51
C. Deamidation and Nitrogen Mobilization ................................ 54
XII. Biotechnology: Genetic Engineering of Storage Proteins for
Improved Characteristics ..................................................... 56
A. Introduction ..................................................................... 56
B. The Enabling Technologies ................................................. 57
C. What Modifications? .......................................................... 59
D. Strategies for Nutritional Improvement of Storage Proteins ....... 60
XIII. Conclusions ............................................................................. 69
Acknowledgements ................................................................... 70
References .............................................................................. 70
I. INTRODUCTION
In a paper written for Advances in Botanical Research in 1981 entitled
“Biochemistry of storage protein synthesis and deposition in the developing
legume seed” (Boulter, 1981), it was concluded that legume seed storage
proteins would continue to be a source of important research problems for
some time to come. This has indeed proved to be so and, as a consequence,
this review is an account of many of these research findings.
Faced with this task, the authors have not attempted to cite all the relevant
literature, especially as there are already excellent recent accountsheviews
available on legume seed storage protein structure (Wright, 1987, 1988;
Lawrence et al., 1990,1994; Shewry, 1995; Shewry et al., 1995), biosynthesis
and intracellular transport (Chrispeels, 1991; Muntz et al., 1993; Casey et
al., 1993; Nielsen et ad., 1995). Instead, we have emphasized the biology of
this structural solution to the storage of nitrogen in seeds as compared to
other strategies such as unusual (non-protein) amino acids. This biological
function dictates and constrains most of the structural, biosynthetic, intracel-
lular transportation and deposition characteristics detailed in this paper.
Inevitably, in not attempting an encyclopaedic coverage, the review has a
personal bias.
The importance of legume storage proteins as a major food protein
resource world wide has always attracted a significant research effort by plant
scientists. In addition, at various times, legume storage proteins or their
encoding genes have been among the few plant experimental materials at
the cutting edge of plant science research. At other times, their lack of
