The structure and biological function of histones PDF
Preview The structure and biological function of histones
The Structure and Biological Function of Hist ones Lubomir S. Hnilica The University of Texas M.D. Anderson Hospital and Tumor Institute Texas Medical Center Houston, Texas published by: Boca Raton London New York CRC Press is an imprint of the Taylor & Francis Group, an informa business First published 1972 by CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 Reissued 2018 by CRC Press © 1972 Taylor & Francis Group, LLC. CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works This book contains information obtained from authentic and highly regarded sources. Reasonable efforts have been made to publish reliable data and information, but the author and publisher cannot assume responsibility for the validity of all materials or the consequences of their use. 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Giovanni Fenaroli, Ph.D. BIOENGINEERING University of Milano, Italy David G. Fleming, Ph.D. ThomasE.Furia Case Western Reserve University CIBA-GEIGY Corp. Lester Goodman, Ph.D. National Institutes of Health MARINE SCIENCES F. G. Walton Smith, Ph.D. CHROMATOGRAPHY University of Miami Gunter Zweig, Ph.D. Syracuse University Research Corp. MATERIALS SCIENCE C. T. Lynch, Ph.D. CLINICAL SCIENCES Wright-Patterson Air Force Base Willard R. Faulkner, Ph.D. Vanderbilt University Medical Center MATHEMATICS AND STATISTICS John W. King, M.D., Ph.D. William H. Beyer, Ph.D. Cleveland Clinic Foundation University of Akron Brian Girling, M.Sc., F.I.M.A. ELECTRO-OPTICS The City University, London Robert J. Pressley, Ph .D. Samuel M. Selby, Ph.D., Sc.D. Holobeam Corp. Hiram College ENGINEERING SCIENCES MICROBIOLOGY Ray E. Bolz, D. Eng. Allen I. Laskin, Ph.D. Case Western Reserve University Esso Research and Engineering Co. George L. Tuve, Sc.D. Hubert Lechevalier, Ph.D. THE Chemical Rubber co. Rutgers University ENVIRONMENTAL SCIENCES ORGANIC CHEMISTRY Richard G. Bond, M.S., M.P.H. Saul Patai, Ph.D. University of Minnesota Hebrew University of Jerusalem Conrad P. Straub, Ph.D. Zri Rappoport, Ph.D. University of Minnesota Hebrew University of Jerusalem RADIOLOGICAL SCIENCES TOXICOLOGY Yen Wang. M.D., D.Sc. (Mecl.) Irving Sunshine, Ph.D. University of Pittsburgh Cuyahoga County Coroner's Office, Ohio SPECTROSCOPY Jeanette GrasseDi, M.S. Standard Oil Company (Ohio) W. M. Ritchey, M.S. Case Western Reserve University James W. Robinson, Ph.D. Louisiana State University CRITICAL REVIEWJ OURNALS ANALYTICAL CHEMISTRY MACROMOLECULAR SCIENCE Louis Meites, Ph.D. Eric Baer, Ph.D. Clarkson College of Technology Case Western Reserve University Phillip Geil, Ph.D. BIOCHEMISTRY Case Western Reserve University Gerald Fasman, Ph.D. Jack Koenig, Ph.D. Brandeis University Case Western Reserve University MICROBIOLOGY BIOENGINEERING Allen I. Laskin, Ph.D. David G. Fleming, Ph.D. Esso Research and Engineering Co. Case Western Reserve University Hubert Lechevalier, Ph.D. Rutgers University CLINICAL SCIENCES Willard R. Faulkner, Ph.D. RADIOLOGICAL SCIENCES Vanderbilt University Medical Center Yen Wang, M.D., D.Sc. (Med.) John W. King, M.D., Ph.D. University of Pittsburgh Cleveland Clinic Foundation SOLID STATE SCIENCES ENVIRONMENTAL SCIENCES Richard W. Hoffman, Ph.D. Richard G. Bond, M.S., M.P.H. Case Western Reserve University University of Minnesota Donald E. Schuele, Ph.D. Conrad P. Straub, Ph.D. Bell Telephone Laboratories University of Minnesota TOXICOLOGY FOOD AND NUTRITION Leon Golberg, D.Phil., D.Sc. Thomas E. Furia Albany Medical College of CIBA-GEIGY Corp. Union University THE AUTHOR Lubomir S. Hnilica, Ph.D., is a biochemist and professor of biochemistry at the University of Texas at Houston, M. D. Anderson Hospital and Tumor Institute and at the Graduate School of Biomedical Sciences. Dr. Hnilica received his Ph.D. in organic chemistry from Masaryk University, Brno, Czechoslovakia in 19 52. He subsequently received postdoctoral training at the Czechoslovakian Academy of Science and the Czechoslovakian National Cancer Research Institute in Bratislava. Dr. Hnilica continued his research as a WHO research fellow at the Chester Beatty Research Institute in England. His present research concerns proteins of the cell nucleus; the biochemistry, structure, and metabolism of nucleohistones, the interaction of nuclear proteins with nucleic acids; and the biochemical mechanisms of gene regulations in eukaryotes. PREFACE Although the discovery of histones is approaching its first centennial, their exact biological role remains unknown. The explosion of interest in histones witnessed during the last decade has resulted in numerous publications in this area. Since the growing number of scientific papers concerning these proteins now amounts to several thousand, only the more recent references are discussed in detail, their selection reflecting the topics treated in the individual chapters. Because another group of chromatin proteins, the nonhistone or acidic nuclear proteins, is rapidly gaining interest among scientists, a brief discussion of them has been included in the last chapter. In general, this book endeavors to present an analysis of the current knowledge of the structure, properties, and possible functions of histones. The broad scope of this topic prevents the discussion of certain individual aspects in any great detail. For this purpose, some of the more specialized review articles should be consulted. Hopefully, the present volume will contribute to a more general understanding of histone biochemistry and will provide stimulation, as well as a source of references, to the student of the cell nucleus and its functions. L S. Hnilica Houston, Texas TABLE OF CONTENTS Chapter 1 Historic Background . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 Chapter 2 Chemical Properties of Histones . 3 Isolation Procedures . 3 Histone Heterogeneity . . . . 5 General Occurrence and Characterization of Histones . 5 Heterogeneity and Nomenclature .6 Quantitative Assays . . . . . . . . . . . . . . 10 Histone Fractionation . . . . . . . . . . . . . 11 Chemical Composition and Structure of Histone Fractions 16 The Very Lysine-Rich His tones F 1 {I) . . . . . . . 16 The Moderately Lysine-Rich Histone F2b (llb2) ~0 The Slightly Lysine-Rich (Arginine-Lysine-Rich) Histone F2a2 (llbl) 21 The Glycine and Arginine-Rich Histone F2al (N) 23 The Alanine and Arginine-Rich Histone F3 (III) 24 Tissue and Species Specificity of His tones 27 Similarities and Differences 27 Histones in Nucleated Erythrocytes 30 Protamines . . . . . . . . . 35 Basic Proteins in Male Gametes . 38 Basic Proteins in Female Gametes 42 Chapter 3 Molecular Properties of Histones 47 Molecular Weights of Histones 47 Histone Aggregation 47 Physicochemical Determinations 48 The Conformation of Histones 49 Chapter 4 Histone Metabolism 57 Histone Biosynthesis . . . . . . 57 Localization of Histone Synthesis 57 Rates of Histone Biosynthesis 60 Histone Turnover . . . . . . . . ·. 64 Regulation of Histone Synthesis 66 Temporal Relationships Between DNA and Histone Synthesis 67 Basic Protein Synthesis During Gametogenesis and Embryogenesis 69 Spermatogenesis 69 Embryogenesis . . . . . . . . . . . . . . . . . . . . 71 Chapter 5 Metabolic Modifications of Histones 79 Acetylation 79 Phosphorylation 84 Methylation 88 Thiolation . . . 90 Ribosylation of Nuclear Proteins 92 Other Modifications of Histones 92 Chapter 6 Histone Interactions with DNA 93 In vitro Complexes of DNA with Other Molecules 93 Chromatin ............... . 97 The Isolation and Composition of Chromatin 98 Chromatin Structure . . . . . . . . 99 DNA-Histone Interactions in Chromatin . 109 Chapter 7 Histone Interactions with Other Molecules . 125 Interactions with Proteins, RNA, and Other Polyanions . . . . 125 Histone Interactions with Steroid Hormones and Carcinogens . 128 Chapter 8 Histones in Genetic Restriction . 133 Quantitative Restriction . . 133 In vitro RNA Synthesis . 133 In vitro DNA Synthesis . 135 Chromatin as a Template for in vitro RNA Synthesis . 136 Qualitative Restriction . . . . . . . . . . . . . . 140 Chapter 9 Non-Histone Proteins of Chromatin . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151 Chapter 10 Conclusions and Perspectives .161 Acknowledgments .167 References .168
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