THE BIOCHEMISTRY OF GENE EXPRESSION IN HIGHER ORGANISMS THE BIOCHEMISTRY OF GENE EXPRESSION IN HIGHER ORGANISMS THE PROCEEDINGS OF A SYMPOSIUM SPONSORED BY THE INTERNATIONAL UNION OF BIOCHEMISTRY, THE AUSTRALIAN ACADEMY OF SCIENCE AND THE AusTRALIAN BIOCHEMICAL SociETY Edited by J.K.POLLAK Department ofH istology and Embryology, University of Sydney Sydney, Australia and J. WILSON LEE Wheat Research Unit, CSIRO, North Ryde, N.S. W. Australia SPRINGER-SCIENCE+BUSINESS MEDIA, B.V. Library of Congress Catalog Card Number 72-97960 ISBN 978-94-010-2552-2 ISBN 978-94-010-2550-8 (eBook) DOI 10.1007/978-94-010-2550-8 AII rights reserved Copyright © by Springer Science+Business Media Dordrecht 1973 Originally published by Australia & New Zealand Book Co. Pty. Ltd. in 1973 Softcover reprint ofthe hardcover lst edition 1973 No part of this book may be reproduced in any form, by print, photoprint, microfilm, or any other means, without written permission from the publisher Ed ito rial Preface The papers assembled in this volume are based on the symposium on "The Biochemistry of Gene Expression in Higher Organisms" which was held at the University of Sydney from May 14-19, 1972. Many symposia have been held on the control of gene expression in prokaryotes but to date considerably less attention has been paid to eukaryotic organisms. It has been appreciated only recently that some of the information gained from the study of prokaryotes is directly applicable to eukaryotes; however, it is now realized that the principles of the control mechanisms of gene expression in these two classes of organism, differ considerably. This symposium was organized in an effort to bring together workers from widely different fields concerned with gene expression, with the aim of circum scribing the current concepts and speculating on future developments in studies on the mechanisms which control and modulate gene expression, in the widest sense, in eukaryotes. This volume contains all the 36 papers presented at the symposium. In a few instances the sequence of contributions has been changed to provide the reader with a more logical presentation. In addition, three papers which were not actually presented at the symposium, have been included in this volume. These three papers were not read because last-minute hitches prevented speakers from attending. The Editors wish to express their thanks to all the contributors who co-operated by supplying their manuscripts promptly. Unfortunately it is extremely difficult to recreate for the reader of this volume, the same spirit of enthusiasm, bonhomie and stimulation which permeated the meeting and the discussions which lasted well into the nights. The symposium was cosponsored by the International Union of Biochemistry, the Australian Academy of Science and the Australian Biochemical Society to whom the organizers wish to express their gratitude. Particular thanks are also due to Associate Professor J. F. Williams and his committee who so ably handled all of the physical arrangements for the meeting. J. K. PoLLAK J. WILSON LEE v Contents EDITORIAL PREFACE CHROMOSOME STRUCTURE AND THE MANIPULATION AND ANALYSIS OF GENES Chromosome structure and units of function in higher organisms 3 W.J.PEACOCK Transgenosis of bacterial genes from.Escherichia coli to cultures of haploid 21 Lycopersicon esculentum and haploid Arabidopsis thaliana plant- cells CoLIN H. DoY, PETER M. GRESSHOFF and BARRY RoLFE Sequences in genetic nucleic acids 38 F. SANGER Mutations at the end of the iso-1-cytochrome c gene of yeast 56 FRED SHERMAN and JoHN W. STEWART TRANSCRIPTIONAL AND TRANSLATIONAL CONTROL MECHANISMS RNA polymerases and transcriptive specificity in eukaryotic organisms 89 W. J. RUTTER, P. W. MoRRis, M. GoLDBERG, M. PAULE and R. W. MoRRIS The control of gene expression by membrane organization in 105 Saccharoinyces Cerevisiae KEVIN A. WARD, S. MARZUKI and J. M. HASLAM Transcription and translation in mammalian cells 117 S. PENMAN, R. PRICE, S. PERLMAN and R. SINGER Isolated chromatin in the study of gene expression 142 SARAH C. R. ELGIN and J. BoNNER Structural modifications of histones in cultured mammalian cells 164 GEoRGE R. SHEPHERD, BILLIE J. NoLAND, JuLIA M. HARDIN and PAUL BYVOET The role of histones in avian erythropoiesis 177 V. L. SELIGY, G. M. H. ADAMS and J. M. NEELIN Chromosomal components in relation to differentiation of avian red blood 191 cells R. APPELS, R. HARLOW, P. ToLsTOSHEV and J. R. E. WELLS Mechanism of glucocorticoid hormone action and of regulation of gene 206 expression in cultured mammalian cells JoHN D. BAXTER, GUY G. RouSSEAU, STEPHEN J. HIGGINS and GoRDoN M. ToMKINS vii viii Regulation of transcription by glucocorticosteroids 225 C. E. SEKERIS and W. SCHMID A control mechanism in gene expression of higher cells operating at the 242 termination step in protein synthesis I. T. OLIVER The mechanism and control of the biosynthesis of a-lactalbumin by the 260 mammary gland P. N. CAMPBELL Changes in protein synthesis and degradation involved in enzyme 274 accumulation in differentiating liver F. J. BALLARD, M. F. HoPGOOD, LEA RESHEF and R. W. HANSON GENE EXPRESSION AND DEVELOPMENT Obligatory requirement for DNA synthesis during myogenesis 287 erythrogenesis and chondrogenesis H. HoLTZER, R. MAYNE, H. WEINTRAUB and G. CAMPBELL The dependence of gene expression on membrane assembly 305 C. G. DuCK-CHONG andJ. K. PoLLAK Patterns of gene activity in larval tissues of the blowfly Calliphora 320 J. A. THOMSON Hormonal and environmental modulation of gene expression in plant 333 development B. KESSLER DNA and RNA synthesis during growth by cell expansion in Vicia faba 357 cotyledons ADELE MILLERD GENE EXPRESSION IN DIFFERENTIATED CELLS Induction of 8-aminolevulinic acid synthetase in perfused rat liver by 369 drugs, steroids, lead and adenosine-3', 5' -monophosphate A.M. EDWARDS and W. H. ELLIOTT The anaemia-induced reversible switch from haemoglobin A to 379 haemoglobin C in goats and sheep: the two haemoglobins are present in the same cell during the changeover M. D. GARRICK, R. F. MANNING, M. REICHLIN and M. MATTIOLI Gene expression in liver endoplasmic reticulum 393 L. ERNSTER The use of neurological mutants as experimental models 410 P. MANDEL, J. L. NussBAUM, N. NESKOVIC, L. SARLIEVE, E. FARKAs and 0. RoBAIN GENE EXPRESSION IN MITOCHONDRIA AND CHLOROPLASTS The phenomenology of cytoplasinic genetics in yeast: a proposal for an 425 autonomy of Initochondrial membranes and the deterininism of nucleo-cytoplasinic genetic interactions A. W. LINNANE, C. L. BUNN, NEIL HoWELL, P. L. MoLLOY and H. B. LUKINS ix Location of DNAs coding for various kinds of chloroplast proteins 443 S. G. WILDMAN, N. KAWASHIMA, D.P. BouRQUE, FLossiE WoNG, SHALINI SINGH, P. H. CHAN, S. Y. KwoK, K. SAKANO, S. D. KuNG and J. P. THORNBER Nuclear genes controlling chloroplast development in barley 457 KNUD W. HENNINGSEN, J. E. BoYNTON, D. VON WETTSTEIN and N. K. BOARDMAN Gene expression in chloroplasts and regulation of chloroplast differentiation 479 RoBERT M. SMILLIE, N. STEELE ScoTT and D. G. BISHOP Products of chloroplast DNA-directed transcription and translation 504 P. R. WHITFIELD, D. SPENCER and W. BoTTOMLEY GENE EXPRESSION AND THE IMMUNE RESPONSE The relevance of immunology to the biochemistry of gene expression 525 G. J. V. NossAL The reaction of antigen with lymphocytes 532 G. L. ADA, M. G. CooPER and R. LANGMAN Synthesis transport and secretion of immunoglobulin in lymphoid cells 542 FRITZ MELCHERS Molecular and cellular mechanisms of clonal selection 555 GERALD M. EDELMAN Antibody diversification: the somatic mutation model revisited 574 MELVIN COHN An alternate mechanism for immune recognition 593 K. J. LAFFERTY The clonal development of antibody forming cells 606 A. J. CUNNINGHAM Immunoglobulin gene expression in murine lymphoid cells 612 NoEL L. WARNER and ALAN W. HARRis Structure and function of lymphocyte surface immunoglobulin 629 JoHN J. MARCHALONIS, RoBERT E. CoNE, JoHN L. ATWELL and RoNALD T. RoLLEY INDEX 649 LIST OF ABBREVIATIONS Chromosome Structure and the Manipulation and Analysis of Genes 3 Chromosome Structure and Units of Function in Higher Organisms W. J. PEACOCK Division of Plant Industry, Commonwealth Scientific and Industrial Research Organization, Canberra, A.C.T., Australia. It is not my intention in this paper to present an exhaustive review of what is known of the structure of the eukaryotic chromosome; rather I thought in this conference I would emphasize the role of cytological and cytogenetic approaches to the problem, and demonstrate that they are a necessary adjunct to biochemical and molecular analyses. In the first part of the paper, in which I will consider the possibility that the eukaryotic chromosome is similar to the bacterial chromo some in having one continuous DNA molecule, I will largely discuss a cytological experiment I carried out in association with Dr. Sheldon Wolff (University of California, San Francisco) and Dr. Dan Lindsley (University of California, San Diego). In the second part of the paper I will largely confine myself to a discus sion of the chromosomes of Drosophila melanogaster, a species in which recent genetic work, when considered with molecular data, has posed a problem which is perhaps the fundamental issue in chromosome research-what is the unit of functional organization in the chromosome? IS THE CHROMOSOME A CONTINUOUS DNA MOLECULE? The possibility that chromosomes of higher organisms may contain one con tinuous DNA molecule was first indicated by the autoradiographic analysis of chromosome duplication by Taylor et al. ( 1957). In this simple experiment root tip cells of Vicia faba, the broad bean, were provided with tritiated thymidine ( [3H]dT) during the DNA replication period of one cell cycle. Some of the cells were then examined by autoradiography at the next metaphase; others were per mitted to proceed through a further DNA replication period in which the isotope was omitted, and examined at the succeeding metaphase-the second metaphase after incorporation of [3H]dT. In the first metaphase both sister chromatids of each chromosome were labelled whereas in the second division, in each chromo- 4 CHROMOSOME STRUCTURE; GENE MANIPULATION AND ANALYSIS some, only one of the two sister chromatids was labelled. An example of the second metaphase labelling pattern is shown in Fig. 1 which shows a cell from a Chinese Hamster tissue culture. The patterns found in the two successive meta phases imply that chromosomal DNA followed a semi-conservative mode of duplication with the unreplicated chromosome (chromatid) containing two sub units extending along its length. FIGURE 1 Autoradiogram of a Chinese Hamster cell. This cell is at the second metaphase after incorporation of ['H)dT and shows segregation of label over sister chromatids. Most chromosomes are labelled along their full length. The arrow indicates a chromosome with one terminal sister chromatid exchange; the double arrow indicates a chromosome with three exchanges. An exception to the labelling pattern described above is the finding of some chromosomes in the second metaphase with labelling over both sister chromatids. Peacock ( 1963) described such isolabelling in Vicia faba and pointed out that it may indicate that more than two strands existed along a chromosome, although in general the strands were parcelled together into two subunits. This possibility was in concert with other classes of cytological evidence which suggested a multineme structure to chromosomes (Wolff, 1969). Isolab elling has been found in a number of organisms ( Deaven and Stubblefield, 1969; Darlington and Haque, 1969) and although it is certainly not a technical artifact as Thomas ( 1971) has asserted, it does not provide critical evidence for the existence of more than two subunits in the unreplicated chromosome. Peacock ( 1963) recognized that auto radiographic resolution could not exclude the possibility that isolabelling resulted from a number of breakage and reunion events between sister chromatids, and both Dupraw ( 1968) and Comings ( 1971) have suggested other ways in which isolabelling is compatible with only two chromatid subunits, given exchanges between sister chromatids. Sister chromatid exchanges are certainly visible in
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