Plant Biochemistry PDF
Preview Plant Biochemistry
Contributors T. Akazawa Arthur L. Karr Peter Albersheim Joel L. Key Leonard Beevers Bessel Kok A. A. Benson Tsune Kosuge James Bonner Abraham Marcus R. W. Breidenbach Ph. Matile J. K. Bryan B. A. Notion R. H. Burris Roderic B. Park J. E. Gander Jack Preiss M. D. Hatch Peter H. Quail Peter K. Helper D. W. Rains E. J. Hewitt Ziva Reuveny David Tuan-Hua Ho P. K. Stumpf D. P. Hucklesby Erhard Stutz Alice Tang Jokela J. E. Varner Lloyd G. Wilson Plant Biochemistry Third Edition Edited by James Bonner California Institute of Technology Joseph E. Varner Washington University ACADEMIC PRESS New York · San Francisco · London A Subsidiary of Harcourt Brace Jovanovich, Publishers COPYRIGHT © 1976, BY ACADEMIC PRESS, INC. 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, INC. Ill Fifth Avenue, New York, New York 10003 United Kingdom Edition published by ACADEMIC PRESS, INC. (LONDON) LTD. 24/28 Oval Road, London NW1 Library of Congress Cataloging in Publication Data Bonner, James Frederick, Date ed. Plant biochemistry. Includes bibliographies. 1. Botanical chemistry. I. Varner, J. E. II. Title. QK861.B6 1976 581.l'92 76-21693 ISBN 0-12-114860-2 PRINTED IN THE UNITED STATES OF AMERICA List of Contributors Numbers in parentheses indicate the pages on which the authors' contributions begin. T. AKAZAWA (381) Research Institute for Biochemical Regulation, School of Agriculture, Nagoya University, Chikusa, Nagoya, Japan PETER ALBERSHEIM (225) Department of Chemistry, University of Colorado, Boulder, Colorado LEONARD BEEVERS (771) Department of Botany and Micro biology, University of Oklahoma, Norman, Oklahoma A. A. BENSON (65) Scripps Institution of Oceanography, University of California at San Diego, La Jolla, California JAMES BONNER (3, 37) Division of Biology, California Institute of Technology, Pasadena, California R. W. BREIDENBACH (91) Plant Growth Laboratory, Department of Agronomy and Range Science, University of California, Davis, California J. K. BRYAN (525) Department of Biology, Syracuse University, Syracuse, New York R. H. BURRIS (887) Department of Biochemistry, University of Wisconsin, Madison, Wisconsin J. E. GANDER (337) Department of Biochemistry, College of Bio logical Sciences, University of Minnesota, St. Paul, Minnesota M. D. HATCH (797) Division of Plant Industry, Commonwealth Scientific and Industrial Research Organization, Canberra City, Australia PETER K. HELPER (147) Department of Biological Sciences, Stan ford University, Stanford, California E. J. HEWITT (633) Plant Physiology and Research Station, Uni versity of Bristol, Long Ashton, England DAVID TUAN-HUA HO (713) Department of Biology, Washington University, St. Louis, Missouri xiii xiv LIST OF CONTRIBUTORS D. P. HUCKLESBY (633) Plant Physiology and Research Station, University of Bristol, Long Ashton, England ALICE TANG JOKELA (65) Department of Microbiology, San Diego State University, San Diego, California ARTHUR L. KARR (405) Department of Plant Pathology, Univer sity of Missouri-Columbia, Columbia, Missouri JOEL L. KEY (463) Botany Department, University of Georgia, Athens, Georgia BESSEL ΚΟΚ (845) Martin Marietta Laboratories, Baltimore, Maryland TSUNE KOSUGE (277) Department of Plant Pathology, University of California, Davis, California ABRAHAM MARCUS (507) The Institute for Cancer Research, Philadelphia, Pennsylvania Ph. MATILE (189) Department of General Botany, Swiss Federal Institute of Technology, Zurich, Switzerland Β. A. NOTTON (633) Plant Physiology and Research Station, Uni versity of Bristol, Long Ashton, England RODERIC B. PARK (115) Department of Botany, University of California, Berkeley, California JACK PREISS (277) Department of Biochemistry and Biophysics, University of California, Davis, California PETER H. QUAIL (683) Research School of Biological Sciences, Australian National University, Canberra, Australia D. W. RAINS (561) Department of Agronomy and Range Science, University of California, Davis, California ZIVA REUVENY* (599) MSU/ERDA Plant Research Laboratory, Michigan State University, East Lansing, Michigan P. K. STUMPF (427) Department of Biochemistry and Biophysics, University of California, Davis, California ERHARD STUTZf (15) Department of Biological Sciences, North western University, Evanston, Illinois J. E. VARNER (714) Department of Biology, Washington University, St. Louis, Missouri LLOYD G. WILSON (599) MSU/ERDA Plant Research Laboratory, Michigan State University, East Lansing, Michigan * Present address : Biology Division, Oak Ridge National Laboratory, Oak Ridge, Tennessee. f Present address : Laboratoire de Physiologie Végétale et Biochemie, Université de Neuchâtel, Neuchâtel, Switzerland. Preface This treatise is intended for the advanced student or professional worker in the plant sciences. It is directed to the biochemist who desires information in areas of biochemistry that are unique to plants, for ex ample, cell wall matters, photosynthesis, or nitrogen fixation, or who is interested in the degree to which plants share biochemical pathways found in other organisms. This work will also be valuable to plant biolo gists in general. Biochemistry can and does contribute to the under standing and solution of the problems involved in many of the more specialized aspects of plant biology—taxonomy, morphology, ecology, horticulture, agronomy, phytopathology, to name a few. We believe this book can help students and research workers in these diverse fields by providing them with a ready source of biochemical information directly applicable to plants. Finally, we feel that it can be used successfully as a text in plant biochemistry courses. The student in such a course would need some background in organic chemistry, but previous study of bio chemistry would not necessarily be required. We have tried to present each topic comprehensively in the sense that we have started with general principles and ended with the current state of the subject. We hope that the reader, after having studied a topic in this book, will find himself qualified to go into his laboratory and start investigations possessing the latest knowledge available in that field. To assist the research worker we have included references per tinent to the original literature; to assist the student we have also in cluded suggestions for more general reading on each topic. The student without previous knowledge of biochemistry will find such reading desirable—perhaps necessary. In the eleven years that have elapsed since the publication of the first edition of "Plant Biochemistry" there have been, of course, advances in all areas of this subject. Some are due to the increasing emphasis of plant physiology on plant biochemistry as a means of better understand- xv xvi PREFACE ing the individual physiological processes; others to the understanding of subjects not comprehended eleven years ago. Examples of the latter can be found throughout the volume. We trust this edition will prove useful to its readers. We thank our colleagues for their contributions and intellectual support of this volume. We are also indebted to the staff of Academic Press for their continuous and skillful help in the preparation of this work. James Bonner Joseph Varner 1 Cell and Subcell JAMES BONNER I. Introduction 3 II. The Subcellular Components of the Plant Cell . .. 3 III. The Logic of Cell Life 6 IV. Methods of Cell Fractionation 8 V. Separation of the Golgi Apparatus 13 VI. Cell as a Community 13 General References 14 I. Introduction One of the most powerful generalizations of biochemistry is that cells of all kinds and of all creatures possess the same, rather small, number of kinds of subcellular components. These subcellular entities are similar between the different kinds of cells, not only in their morphology and submicroscopic structure but also in chemical composition and most im portantly in chemical function, each kind contributing its own mite to the overall functioning of the cell. The untangling of the biochemical pathways of metabolism and the development of our understanding of the strategy of life has been due very largely to the technology developed since approximately 1950 which has made it possible to separate several subcellular components from one another and to identify the enzyme sys tems associated with each. We turn our attention, therefore, first to the subcellular components of the plant cell. II. The Subcellular Components of the Plant Cell The principal subcellular components of the plant cell, and those with which biochemistry is principally concerned, are the nucleus, the chloroplast, the mitochondria, the lysosomes and other vacuoles, the ribosomes, messenger RNA, and the individual soluble enzymes. Table I summarizes the number of each of these kinds of entities found in a typical or average cell. The vast majority of plant cells contains, of 3 4 JAMES BONNER course, one nucleus, although many instances of multinucleate cells are known—even in higher plants, for example, latex vessels or the sieve tubes, cells that are multinucleate by virtue of dissolution of transverse cell walls. Chloroplasts in the photosynthetic portion of the plant number in general a few tens—50 is the rough average per cell. To this number should be added the proplastids from which the mature chloroplasts arise, but since today we still have no good estimate of the number of proplas tids to be found in a typical cell, we will not consider them further. The proplastids are found in the nonphotosynthetic as well as in the photo synthetic portions of the plant—for example, in roots. Mitochondria characteristically occur in the plant cell in the order of the 100's—500 to 1000 being a typical number. Lysosomes and other vacuoles are present in about equal numbers. Ribosomes, the next smaller category of particles, occur in vastly greater numbers than chloroplasts or mitochondria. A growing functional plant cell might perhaps contain a few hundred thousand ribosomes, although this number varies greatly with age, state of activity, and so on. The bulk of the cytoplasmic protein, the portion to which we refer in general as the nonparticulate cytoplasm, is of course composed of enzyme molecules, in fact, a great number of kinds of enzyme molecules. The total number of enzyme molecules in a typical cell would be of the order of 1,000,000,000. These consist of several thousand, perhaps 1000 to 10,000, different species of enzyme molecules, each qualified to catalyze one specific kind of chemical reac tion. A typical plant cell might then contain 1,000,000,000 enzyme mole cules of 10,000 different kinds, 100,000 being representative of each of the 10,000 different species present. The proportion that any individual kind of enzyme molecule constitutes of the total soluble cytoplasmic pro terin often departs widely, however, from the average 0.001%, which would be expected on the basis of the above calculation. Thus, we know that particular kinds of enzyme molecules in particular kinds of cells may constitute from a few tenths to as much as several percent of total soluble protein. As a general rule, however, we must expect, because there are so many kinds of enzyme molecules, that each one will constitute but a small proportion of the total, and it is not surprising, therefore, that in the purification of enzymes, enrichments of 10,000-fold or more are not uncommonly needed to achieve pure material. We have referred above to messenger RNA as a typical component of the plant cell. Messenger RNA may indeed be isolated and character ized by methods to be considered in a later chapter. For the time being, it may be noted that messenger RNA may most easily and characteristi cally be detected in the plant cell by virtue of its interaction with the ribosome. Ribosomes interact with and attach to messenger RNA, and 1. CELL AND SUBCELL 5 since a single messenger RNA strand may simultaneously bind many ribosomes, the great bulk of the ribosomes of the plant cell are often detected as large aggregates of so-called polysomes. Transfer RNA is characterized by small molecular size like the soluble enzymes and is a component of the nonparticulate cytoplasmic material. For this reason, it is often known in the literature of the cell as soluble RNA, although the preferred name is tRNA. The entire assemblage of subcellular structure as outlined above is, of course, contained within the membrane system, and we might properly include the membrane system as one of the most characteristic of cellular components. This membrane system comprises not only the protoplasmic membrane itself but also the membranes surrounding the nucleus, chloro plasts, and vacuole, as well as the membrane elements of the mitochon dria, the endoplastic reticulum, and other structures outlined below. The plant cell is characterized also by the cell wall external to the protoplas mic membrane, and in a sense the wall might, too, be considered as a subcellular component characteristic of the plant cell. The plant cell contains still further subcellular systems. These en tities are, of course, important subcellular components, but they are either less universal or less understood than those enumerated in Table I. They include, for example, lysosomes that contain hydrolytic enzymes and con duct the autolysis of injured or aging cells. They include also the Golgi apparatus that concerns itself with concentration, chemical modification, and secretion of enzymes and substrates that are to be secreted into the outside of the cell (as for example, cell wall-forming materials). The glyoxysomes are particularly well known in germinating fatty seeds, where they are responsible for the transformation of fatty acids into sugar precursors. Peroxisomes are responsible for photorespiration. Into this category also we must place the microtubules and related structures responsible for photoplasmic streaming, and also we may place the TABLE I Numbers and Sizes of Subcellular Particles of Various Classes Present in a Typical Plant Cell Subcellular particle Diameter Number per cell Nucleus 5-20 μΐη 1 Chloroplasts 5-20 Aim 50-200 5 Mitochondria 1-5 μ m 500-2000 8 Ribosomes 250 Â 5-50 Χ ΙΟ Enzyme molecules 20-100 Â 5-50 Χ ΙΟ
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