Subcellular Biochemistry 5 Volume ADVISORY EDITORIAL BOARD J. ANDRE Laboratoire de Biologie Cellulaire, 4 Faculte des Sciences, 91 Orsay, France D. L. ARNON Department of Cell Physiology, Hilgard Hall, University of California, Berkeley, California 94720, USA J. BRACHET Laboratoire de Morphologie Animale, Faculte des Sciences, Universite Libre de Bruxelles, Belgium J. CHAUVEAU Institut de Recherches Scientifiques sur Ie Cancer, 16 Avenue Vaillant Couturier, 94 Ville Juif, Boite Postale 8, France C. de DUVE Universite de Louvain, Louvain, Belgium and The Rockefeller University, New York, NY 10021, USA M. KLINGENBERG Institut flir Physiologische Chemie und Physikalische Biochemie, Universitiit Miinchen, Goethestrasse 33, Miinchen 15, Germany A. LIMA-de-FARIA Institute of Molecular Cytogenetics, Tornavagen 13, University of Lund, Lund, Sweden O. LINDBERG The Wenner-Gren Institute, Norrtullsgatan 16, Stockholm, VA, Sweden V. N. LUZIKOV A. N. Belozersky Laboratory for Molecular Biology and Bioorganic Chemistry, Lomonosov State University, Building A, Moscow 117234, USSR • H. R. MAHLER Chemical Laboratories, Indiana University, Bloomington, Indiana 47401, USA M. M. K. NASS Department of Therapeutic Research, University of Pennsylvania School of Medicine, Biology Service Building, 3800 Hamilton Walk, Philadelphia, Pennsylvania 19104, USA A. B. NOVIKOFF Department of Pathology, Albert Einstein College of Medicine, Yeshiva University, Eastchester Road and Morris Park Avenue, Bronx, NY 10461, USA R. N. ROBERTSON Research School of Biological Sciences, P. O. Box 475, Canberra City, A.C.T. 2601, Australia P. SIEKEVITZ The Rockefeller University, New York, NY 10021, USA F. S. SJOSTRAND Department of Zoology, University of California, Los Angeles, Califor nia 90024, USA A. S. SPIRIN A. N. Bakh Institute of Biochemistry, Academy of Sciences of the USSR, Leninsky Prospekt 33, Moscow V-71, USSR D. von WETTSTEIN Department of Physiology, Carlsberg Laboratory, Gl. Carlsbergvej 10, DK-2500, Copenhagen, Denmark V. P. WHITTAKER Abteilung flir Neurochemie, Max-Planck Institut fLir Biophysikalische Chemie, D-3400 Gottingen-Nikolausberg, Postfach 968, Germany A Continuation Order Plan is available for this series. A continuation order will bring delivery of each new volume immediately upon pl.lblication. Volumes are billed only upon actual shipment. For further information please contact the publisher. Subcellular Biochemistry 5 Volume Edited by Donald B. Roodyn University College London London, England PLENUM PRESS • NEW YORK AND LONDON ISBN-13: 978-1-4615-7944-1 e-ISBN-13: 978-1-4615-7942-7 DOl: 10.1 007/ 978-1-4615-7942-7 This annual series is a continuation of the journal Sub-Cellular Biochemistry, Volumes 1 to 4 of which were published quarterly from 1972 to 1975 © 1978 Plenum Press. New York Softcover reprint of the hardcover 1s t edition 1978 All rights reserved No part of this book may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher Contributors Michel Bergeron Departement de Physiologie, Universite de Montreal, Montreal, Quebec, Canada B. B. Biswas Department of Biochemistry, Bose Institute, Calcutta 700009, India D. E. Buetow Department of Physiology and Biophysics, University of Illinois, Urbana, Illinois 61801, U.S.A. Philip Hohmann Department of Experimental Biology, Roswell Park Memorial Institute, Buffalo, New York 14263, U.S.A. S. K. Malhotra Biological Sciences Electron Microscopy Laboratory, University of Alberta, Edmonton, Canada T6G 2E9 Hilton H. Mollenhauer Veterinary Toxicology and Entomology Research Laboratory, USDA, ARS, College Station, Texas 77840, U.S.A. D. James Morrt~ Department of Medicinal Chemistry and Pharmacognosy and Department of Biological Sciences, Purdue University, West Lafayette, Indiana 47907, U.S.A. Gordon Parry Biodynamics Laboratory, University of California, Berke ley, California 94720, U.S.A. Pranab Roy Department of Biochemistry, Bose Institute, Calcutta 700009, India Pierre Sassier Departement de Physiologie, Universite de Montreal, Montreal, Quebec, Canada W. M. Wood Department of Physiology and Biophysics, University of Illinois, Urbana, Illinois 61801, U.S.A. v Aims and Scope SUBCELLULAR BIOCHEMISTRY aims to bring together work on a wide range of topics in subcellular biology in the hope of stimulating progress towards an integrated view of the cell. In addition to dealing with conven tional biochemical studies on isolated organelles, articles published so far and planned for the future consider such matters as the genetics, evolution, and biogenesis of cell structures, bioenergetics, membrane structure and functions, and interactions between cell compartments, particularly between mitochondria and cytoplasm and between nucleus and cytoplasm. Articles for submission should be sent to Dr. D. B. Roodyn, Depart ment of Biochemistry, University College London, Gower Street, London WCIE 6BT, U.K., and are best sent in the period February to April inclusive of each year. There are no rigid constraints as to the size of the articles and in general they should be between 9,000 and 36,000 words, with an optimum size of about 20,000 words. Although articles may deal with highly specialized topics, authors should try as far as possible to avoid specialist jargon and to make the article as comprehensible as possible to the widest range of biochemists and cell biologists. Full details of the preparation of manuscripts are given in a comprehensive Guide for Con tributors which is available from the Editor or Publishers on request. vii Preface The transition from the quarterly Sub-Cellular Biochemistry to the annual SUBCELLULAR BIOCHEMISTRY is a good opportunity to restate the aims and scope of this publication. They were originally given (in Volume 1 No. 1) as follows: This review and essay journal ... brings together work on a wide range of topics in sub-cellular biochemistry in the hope of stimulating progress towards an integrated view of the cell. It deals with the biochemistry and general biology of nuclei, mitochondria, lysosomes, peroxisomes, chloroplasts, cell membranes, ribosomes, cell sap, flagellae and other specialized cell components. In addition to articles dealing with conventional biochemical studies on sub-cellular struc tures, the journal publishes articles on the genetics, evolution and biogenesis of cell organelles, bioenergetics, membrane behaviour and the interaction between cell structures, particularly between nucleus and cytoplasm. The first four volumes (in the quarterly format) fulfilled many, but not all, of these stated aims, and it is hoped that further articles in the new annual series will soon fill any deficiencies in the range of topics covered. Over the years we have intentionally not interpreted the title of the publication in a too literal sense. Although we have included specific articles on individual subcellular fractions (and certainly hope to do so again) the publication is definitely not only concerned with studies on the biochemistry of isolated cell fractions. The primary target is the "integrated view of the cell." Ultimately this must mean bringing together the results from a very wide range of different techniques and disciplines. Just as we have not been too literal in our interpretation of the title, so we have not been too strict in the format and arrangement of the articles published. They have extended all the way from very short review letters to massive reviews sometimes taking up a whole issue. Similarly some of the articles have been essentially factual and descriptive, whereas others have endeavored to be controversial or even polemical. We hope that these ix x Pr€fac€ tendencies will continue in the annual series. In this issue, although the layout of the various articles has been standardized to have a uniform appearance throughout the book, the articles vary greatly in size and we trust that authors in future will feel quite free to organize their articles as the subject under consideration dictates, free from any constraints imposed by the publication.* The articles published in this volume well illustrate the general aims and direction of SUBCELLULAR BIOCHEMISTRY. We start with a massive review on "The Mitochondrial Translation System" by D. E. Buetow and W. M. Wood. Such has been the progress in this burgeoning field that a review covering the recent research in the topic in a comprehen sive way must have the necessary elbow room, and we are happy to provide it. The use of large and extensive tables is a convenient way of collating and systematizing studies on subcellular fractions and the authors have made skillful use of this technique. Another interesting aspect of this article is that although the topic is superficially a "straightforward" one, dealing with one specific property of mitochondria, in fact all sorts of questions of evolution and the interaction between cell compartments are raised; this has occurred in many of the articles previously published-the nature of modern cell biochemistry is such that it is becoming increasingly difficult to treat subjects adequately in narrow terms. This effect is shown in the second article by P. Hohmann on the role of the HI histones. At one level the articles asks a very specific question: what is the function of a particular type of histone? However, in order to answer this the author is lead to consider broad questions of chromosomal control and function. We are pleased to include the author's suggestions as to the role of HI histones in order to stimulate further discussion and research in this interesting topic. The next article, by P. Sassier and M. Bergeron, deals with cell proliferation in the small intestine epithelium. At first sight this might appear to a rather "physiological" subject for our pUblication. It turns out that the intestinal epithelial system is an excellent model for studying cells at various stages of the cell cycle. A fundamental strategic problem in all biochemical research is the choice of tissue. One wonders how many well-intentioned experiments on rat liver or E. coli have not been particularly fruitful, not because the techniques were inadequate, but because the choice of starting material posed insuperable problems for that particular problem. Thus the more we advertise the particular technical advantages of different biological systems (particularly those that are not "The inclusion offul! reference citations, including the title of the reference, is a bonus that we ask of authors; it is quite optional and we are grateful to those authors who go to the trouble to produce such lists. However, we fully understand the position of those who find it too difficult. Preface xi often used) the more we can help our colleagues to marry the right tissue with the right problem. The next article, by B. B. Biswas and Pranab Roy on "Plant Growth Substances as Modulators of Transcription," illustrates another aspect of the dangers of imbalance in biochemical research. Although there is a massive literature on the fundamental biochemical and cellular effects of hormones in animals, the effects of plant hormones has not received anywhere near this sort of attention. This reflects a general tendency in which biochemists appear to tum more readily to animal or microbial systems than to plants. Whether the reasons are primarily educational or simply that few laboratories have the facilities for the growth of plants under reasonably controlled conditions, it is nevertheless striking how poorly current biochemical research makes use of the extraordinary profu sion of the plant kingdom, making up as it does a good proportion of the total biomass of the planet. It is hoped that the article by Biswas and Roy will help to correct this situation. The next two articles complement each other. "Molecular Structure of Biological Membranes: Functional Characterization" by S. K. Malhotra deals with the very important question of how one membrane differs from another. Some years ago (under the now famous stimulus of the exposition of the "fluid mosaic model" by Singer and Nicolson) all the talk was on "membranes." It seemed that every other paper and textbook dealt in some way or another with "biomembrane research." Unfortunately what may be true for one membrane may not be true for another. The wide spread citation in the literature of the term "unit membrane" (originating from J. D. Robertson) mistakenly lead many to think that the cell is some sort of continuum of more or less identical membranes, differing here and there by the apposition of a few granules or ribosomes, or perhaps by some local "thickening." However, there is now no doubt at all that while different membranes possess certain common features, there are also very considerable differences between them. The problem is therefore to estab lish with certainty precisely what is general to all membranes and con versely what are the precise and specific properties of a given membrane. Malhotra's article is an important stimulus to discussion of this very fundamental question. G. Parry's article on "Membrane Assembly and Turnover" is primar ily concerned with problems of biosynthesis and assembly and the question of membrane identity is secondary. Nevertheless it does arise since it is, of course, impossible to understand precisely how membranes are assembled if we do not know how one membrane differs from another. There is an interacting process in research on membrane structure and biogenesis: the more we know about the fine organization of individual types of membrane, the more meaningful can be our experiments on biogenesis. Conversely, an