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DNA Damage Recognition PDF

869 Pages·2006·14.472 MB·English
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DK3110_title 8/10/05 10:26 AM Page 1 DNA Damage Recognition edited by Wolfram Siede University of North Texas Health Science Center Fort Worth, Texas, U.S.A. Yoke Wah Kow Emory University School of Medicine Atlanta, Georgia, U.S.A. Paul W(cid:2) Doetsch Emory University School of Medicine Atlanta, Georgia, U.S.A. New York London DK3110_Discl.fm Page 1 Monday, August 15, 2005 1:41 PM Published in 2006 by Taylor & Francis Group 270 Madison Avenue New York, NY 10016 © 2006 by Taylor & Francis Group, LLC No claim to original U.S. Government works Printed in the United States of America on acid-free paper 10 9 8 7 6 5 4 3 2 1 International Standard Book Number-10: 0-8247-5961-3 (Hardcover) International Standard Book Number-13: 978-0-8247-5961-2 (Hardcover) This book contains information obtained from authentic and highly regarded sources. Reprinted material is quoted with permission, and sources are indicated. A wide variety of references are listed. Reasonable efforts have been made to publish reliable data and information, but the author and the publisher cannot assume responsibility for the validity of all materials or for the consequences of their use. No part of this book may be reprinted, reproduced, transmitted, or utilized in any form by any electronic, mechanical, or other means, now known or hereafter invented, including photocopying, microfilming, and recording, or in any information storage or retrieval system, without written permission from the publishers. For permission to photocopy or use material electronically from this work, please access www.copyright.com (http://www.copyright.com/) or contact the Copyright Clearance Center, Inc. (CCC) 222 Rosewood Drive, Danvers, MA 01923, 978-750-8400. CCC is a not-for-profit organization that provides licenses and registration for a variety of users. For organizations that have been granted a photocopy license by the CCC, a separate system of payment has been arranged. Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation without intent to infringe. Library of Congress Cataloging-in-Publication Data Catalog record is available from the Library of Congress Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com Taylor & Francis Group is the Academic Division of T&F Informa plc. Preface The topics of this book are the various molecular mechanisms that are involved in the process of DNA damage recognition as the initial step of DNA damage repair and of other related responses, such as damage tolerance and cell cycle checkpoint regulation. The authors were asked to provide review-type articles designed with the researcher in the field in mind. But sufficient introductory comments were requested so that a non-expert or an interested advanced student with some back- groundknowledgecanfollow.Whilewedidnothaveanarrowdefinitionofdamage recognitioninmind,acompletedescriptionofcellularDNArepairmechanismswas certainlynotourgoal.However,lookingatthescopeofthisimpressivecollectionof in-depth reviews, it almost happened... Inthebeginningofthebook,certaintheoreticalaspectsofdamagerecognition that are common themes throughout the book have been addressed. How can one imagine that recognition proteins find their rare targets? How can we envisage pro- teinmovement—byrandomdiffusionor‘‘patrolling’’alongDNA?Anotherequally important topic addresses proteincooperation that enhances recognition specificity. ThereisclearlyanemphasisonstructuralaspectsofDNAdamagerecognition throughoutthebook.Whereversuchinformationisavailable,itisexplainedindetail how protein/DNA damage contacts are being accomplished and which types of structuralfeaturesorconsequencesofDNAdamagearebeingprobedbytherecog- nition apparatus in order to permit a distinction from undamaged DNA. In this fashion,damagerecognitionisaddressedwithinthemajorpathwaysofDNArepair, i.e., simple damage reversal, nucleotide excision repair, base excision repair, mismatch repair, recombinational repair, and DNA endjoining. We interpret the pathways of damage tolerance as a consequence of a combined recognition/accom- modation process and thus we have also included chapters on translesion synthesis. DNA recognition steps occur at several levels within a single repair pathway and this complexity has been considered. How are repair intermediates being recognized? How is a repair intermediate handed over to the next player? If there is a competition between different mechanisms: how is a pathway choice accom- plished?Whilewedonotemphasizedownstreamreactions,thestrategyoforiginat- ing a transmissible downstream signal will be addressed wherever appropriate, especially in the context of regulatory responses. WewereequallyinterestedinputtingspecificaspectsofDNAdamagerecogni- tion in the cellular context. Certain chapters provide the necessary backdrop by givingup-to-datereviewsofcertainpathways.However,alsochromosomestructure, iii iv Preface DNA structure, and sequence context, which may affect DNA damage recognition positivelyornegatively,arebeingaddressed.Theinterferencewithcellularprocesses such as replication and transcription is discussed for several examples since such interference as a consequence of DNA damage may influence, aide, or even initiate a recognition process. Allthatisleftistothankourauthorsfortheirhardworkandsuperbcontribu- tions. We would like to acknowledge the help of Stacy Harman Holloway. We are also indebted to Anita Lekhwani, Moraima Suarez, Joseph Stubenrauch, and their colleagues at Taylor & Francis Books for pursuing the idea and providing such excellent editorial support. We also thank the National Institute of Environmental Health Sciences for supporting a research collaboration amongst the three editors as well as two other authors of individual chapters that made this book possible (‘‘Cellular Responses to Genotoxic Stress’’ Program Project ES11163). Wolfram Siede Yoke Wah Kow Paul W. Doetsch Contents Preface . . . . iii Contributors . . . . xvii PART I. MECHANISMS OF DAMAGERECOGNITION: THEORETICAL CONSIDERATIONS . .. .. .. . .. .. .. .. .. .. .. . .. .. .. .. .. .. . .. .. 1 1. Dynamics of DNA Damage Recognition . . . . . . . . . . . . . . . . . . . . . 3 Eleanore Seibert, Roman Osman, and J. B. Alexander Ross 1. Introduction . . . . 3 2. Role of DNA Flexibility in Sequence-Dependent Activity of UDG . . . . 4 3. Opening and Bending Dynamics of G(cid:1)U Mismatches in DNA . . . . 8 4. Conclusions . . . . 13 References . . . . 15 2. In Search of Damaged Bases . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 R. Stephen Lloyd, A. K. McCullough, and M. L. Dodson 1. Introduction . . . . 21 2. Mechanism for an Increased Rate of Target Site Location . . . . 21 3. In Vitro Evidence for Processive Nicking Activity of DNA Glycosylases . . . . 23 4. Discovery and Significance of In Vivo Processive Nicking Activity by T4-pdg . . . . 26 5. DNA Bending as a Potential Prerequisite for Nucleotide Flipping . . . . 27 6. Mechanisms of Nucleotide Flipping . . . . 29 7. Specificity of Glycosylase Binding Sites and Catalytic Activities . . . . 30 References . . . . 31 v vi Contents 3. Increased Specificity and Efficiency of Base Excision Repair Through Complex Formation . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 Karen H. Almeida and Robert W. Sobol 1. Introduction . . . . 33 2. DNA Lesion Recognition and Removal . . . . 36 3. Strand Incision . . . . 42 4. Gap Filling and Religation . . . . 46 5. XRCC1 Coordination . . . . 49 6. Long-Patch Repair . . . . 52 7. Emerging Subpathways . . . . 54 8. Conclusions . . . . 54 References . . . . 55 PART II. UVDAMAGEAND OTHERBULKY DNA-ADDUCTS . .. .. .. .. .. . . 65 4. Structure and Properties of DNA Photoproducts . . . . . . . . . . . . . . 67 John-Stephen Taylor 1. Introduction . . . . 67 2. Cyclobutane Pyrimidine Dimers . . . . 69 3. Other Dimer-Related Products . . . . 79 4. (6–4) Products . . . . 81 5. Dewar Photoproduct . . . . 86 6. Spore Photoproduct . . . . 88 7. TA(cid:2) Product . . . . 89 8. Conclusions . . . . 90 References . . . . 90 5. Damage Recognition by DNA Photolyases . . . . . . . . . . . . . . . . . . 95 Gwendolyn B. Sancar 1. Overview of Photolyases . . . . 95 2. The Nature of the Substrates . . . . 96 3. Characterization of Substrate Binding and Discrimination by Photolyases . . . . 97 4. Interactions at the Photolyase–Photoproduct Interface: The Molecular Basis for Substrate Binding and Discrimination . . . . 98 5. Substrate Binding In Vivo . . . . 105 6. Summary and Future Directions . . . . 107 References . . . . 107 6. Damage Recognition by the Bacterial Nucleotide Excision Repair Machinery . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111 Deborah L. Croteau, Matthew J. DellaVecchia, Milan Skorvaga, and Bennett Van Houten 1. Introduction . . . . 111 2. Diversity of DNA Lesions Recognized . . . . 113 3. The Proteins and Their Structural Domains . . . . 114 Contents vii 4. Reaction PathwayforDamageDetection and Processing . . . . 123 5. DNA Damage Recognition within the Biological Context of the Cell . . . . 130 6. Similarities in Damage Recognition and Verification Between Bacterial and Eukaryotic Nucleotide Excision Repair Systems . . . . 133 References . . . . 133 7. Recognition of DNA Damage During Eukaryotic Nucleotide Excision Repair . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139 Hanspeter Naegeli 1. Introduction . . . . 139 2. Nucleotide Excision Repair Substrates . . . . 139 3. Eukaryotic NER Reaction . . . . 140 4. Subunits of the Eukaryotic NER Machinery . . . . 142 5. Stepwise Assembly of the Mammalian NER Recognition Complex . . . . 143 6. A Preassembled Repairosome in Yeast? . . . . 144 7. RoleofDamagedDNABindinginDamageRecognition . . . . 146 8. Recognition of Bulky Lesions During Transcription-Coupled DNA Repair . . . . 147 9. Bipartite Substrate Discrimination in the GGR Pathway . . . . 148 10. XPC–hHR23B as a Sensor of Defective Base Pairing . . . . 149 11. Transcription Factor IIH as a Sensor of Defective Deoxyribonucleotide Chemistry . . . . 151 12. Role of XPA–RPA in Integrating Different Recognition Signals . . . . 153 13. Damage-Specific Recruitment of XPG and XPF–ERCC1 . . . . 155 14. Regulation of the Damage Recognition Process . . . . 155 15. Conclusions . . . . 158 References . . . . 159 8. Interactions of the Transcription Machinery with DNA Damage in Prokaryotes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165 Isabel Mellon 1. General Overview . . . . 165 2. The Behavior of RNA Polymerase Complexes with Different Types of DNA Damage . . . . 168 3. The Behavior of RNA Polymerase Complexes at Lesions and NER . . . . 170 4. The Behavior of RNA Polymerase Complexes at Lesions and BER . . . . 173 5. Summary and Future Directions . . . . 175 References . . . . 175 viii Contents 9. DNA Repair in Actively Transcribed Genes in Eukaryotic Cells . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181 Moon-shong Tang 1. Introduction . . . . 181 2. Heterogeneity of DNA Repair . . . . 182 3. Methods for Detecting TCR and GGR . . . . 185 4. DNA Repair in Transcriptionally Active Genes in Different Organisms . . . . 188 5. Models of TCR in Eukaryotic Cells . . . . 193 6. Effect of Different Kinds of DNA Damage on TCR . . . . 194 References . . . . 195 10. Chromatin Structure and the Repair of UV Light-Induced DNA Damage . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201 Fritz Thoma 1. Introduction . . . . 201 2. Nucleosomes: Heterogeneity in a Conserved Structure . . . . 202 3. Dynamic Properties of Nucleosomes Regulate DNA Accessibility . . . . 203 4. Damage Tolerance of Nucleosomes . . . . 208 5. Repair of Nucleosomes by Photolyase . . . . 209 6. Repair of Nucleosomes by NER . . . . 211 7. Site-Specific Repair in Nucleosome and Damage Recognition . . . . 214 8. Chromatin Remodeling and DNA Repair . . . . 214 9. Conclusions . . . . 216 References . . . . 216 11. The Ultraviolet Damage Endonuclease (UVDE) Protein and Alternative Excision Repair: A Highly Diverse System for Damage Recognition and Processing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223 Paul W. Doetsch, Vladimir Beljanski, and Binwei Song 1. Introduction . . . . 223 2. Discovery and Initial Characterization of S. pombe UVDE . . . . 224 3. Recognition and Processing of UV Photoproducts . . . . 226 4. Recognition and Processing of Platinum G-G Diadducts . . . . 227 5. Recognition and Processing of Abasic Sites . . . . 227 6. Modified Bases not Recognized by UVDE . . . . 229 7. Recognition and Processing of Base–Base Mismatches . . . . 230 8. Recognition and Processing of Insertion–Deletion Loops . . . . 231 9. Subsequent Steps Following UVDE-Initiated Alternative Excision Repair . . . . 232 10. Schizosaccharomyces pombe UVDE Homologs . . . . 233 11. Conclusions . . . . 234 References . . . . 234 Contents ix 12. Structural Aspects of Pt-DNA Adduct Recognition by Proteins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239 Uta-Maria Ohndorf and Stephen J. Lippard 1. Background . . . . 239 2. Introduction . . . . 239 3. Structural Consequences of Platinum-Binding to Double-Stranded DNA . . . . 240 4. Recognition of cis-DDP-1,2 Intrastrand Cross-Link by Cellular Proteins . . . . 245 5. Summary and Outlook . . . . 254 References . . . . 255 13. Structural Aspects of Polycyclic Aromatic Carcinogen-Damaged DNA and Its Recognition by NER Proteins . . . . . . . . . . . . . . . . 263 Nicholas E. Geacintov, Hanspeter Naegeli, Dinshaw J. Patel, and Suse Broyde 1. Introduction . . . . 263 2. Metabolism of PAH to Diol Epoxides and Formation of Stereoisomeric DNA Adducts . . . . 265 3. Methods . . . . 267 4. PAH–DNA Adducts: Conformational Motifs . . . . 269 5. Insights into the Structural Motifs at the Nucleoside Adduct Level Derived from Computational Approaches . . . . 273 6. PAH–DNA Adduct Conformational Motifs and NER . . . . 274 7. Structural Differences Between Bay and Fjord Stereoisomeric PAH-N6–Adenine Adducts and Correlations with NER Susceptibilities . . . . 276 8. Computational Analysis . . . . 279 9. Conclusions . . . . 289 References . . . . 290 PART III.NON-BULKY BASE DAMAGE .. .. .. .. .. .. .. . .. .. .. .. .. .. . .. 297 14. Structural Features of DNA Glycosylases and AP Endonucleases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 299 Joy L. Huffman, Ottar Sundheim, and John A. Tainer 1. The Base Excision Repair Pathway . . . . 299 2. DNA Glycosylase Structural Families . . . . 300 3. Specific Mechanisms for Recognition of Damage . . . . 303 4. AP Endonucleases . . . . 313 5. Emerging Questions . . . . 315 References . . . . 315 15. Repair of Oxidized Bases . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 323 Yoke Wah Kow 1. Biological Consequences of Oxidative Damage . . . . 323

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