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00_Symp69_frontmatter 4/21/05 11:36 AM Page i COLD SPRING HARBOR SYMPOSIA ON QUANTITATIVE BIOLOGY VOLUME LXIX 00_Symp69_frontmatter 4/21/05 11:36 AM Page ii www.cshl-symposium.org Institutions that have purchased the hardcover edition of this book are entitled to online access to the companion Web site at www.cshl-symposium.org. Please contact your institution's library to gain ac- cess to the Web site. The site contains the full text articles from the 2004 Symposium and the Sym- posia held in 1998–2003 as well as archive photographs and selected papers from the 61-year history of the annual Symposium. Institutions requiring assistance with activating their online accounts should contact Kathy Cirone, CSHL Press Subscription Manager, at [email protected]. 00_Symp69_frontmatter 4/21/05 11:36 AM Page iii COLD SPRING HARBOR SYMPOSIA ON QUANTITATIVE BIOLOGY VOLUME LXIX Epigenetics www.cshl-symposium.org Meeting Organized by Bruce Stillman and David Stewart COLD SPRING HARBOR LABORATORY PRESS 2004 00_Symp69_frontmatter 4/21/05 11:36 AM Page iv COLD SPRING HARBOR SYMPOSIA ON QUANTITATIVE BIOLOGY VOLUME LXIX ©2004 by Cold Spring Harbor Laboratory Press International Standard Book Number 0-87969-729-6 (cloth) International Standard Book Number 0-87969-731-8 (paper) International Standard Serial Number 0091-7451 Library of Congress Catalog Card Number 34-8174 Printed in the United States of America All rights reserved COLD SPRING HARBOR SYMPOSIA ON QUANTITATIVE BIOLOGY Founded in 1933 by REGINALD G. HARRIS Director of the Biological Laboratory 1924 to 1936 Previous Symposia Volumes I (1933) Surface Phenomena XXXIV (1969) The Mechanism of Protein Synthesis II (1934) Aspects of Growth XXXV (1970) Transcription of Genetic Material III (1935) Photochemical Reactions XXXVI (1971) Structure and Function of Proteins at the Three- IV (1936) Excitation Phenomena dimensional Level V (1937) Internal Secretions XXXVII (1972) The Mechanism of Muscle Contraction VI (1938) Protein Chemistry XXXVIII (1973) Chromosome Structure and Function VII (1939) Biological Oxidations XXXIX (1974) Tumor Viruses VIII (1940) Permeability and the Nature of Cell Membranes XL (1975) The Synapse IX (1941) Genes and Chromosomes: Structure and Organization XLI (1976) Origins of Lymphocyte Diversity X (1942) The Relation of Hormones to Development XLII (1977) Chromatin XI (1946) Heredity and Variation in Microorganisms XLIII (1978) DNA: Replication and Recombination XII (1947) Nucleic Acids and Nucleoproteins XLIV (1979) Viral Oncogenes XIII (1948) Biological Applications of Tracer Elements XLV (1980) Movable Genetic Elements XIV (1949) Amino Acids and Proteins XLVI (1981) Organization of the Cytoplasm XV (1950) Origin and Evolution of Man XLVII (1982) Structures of DNA XVI (1951) Genes and Mutations XLVIII (1983) Molecular Neurobiology XVII (1952) The Neuron XLIX (1984) Recombination at the DNA Level XVIII (1953) Viruses L (1985) Molecular Biology of Development XIX (1954) The Mammalian Fetus: Physiological Aspects of De- LI (1986) Molecular Biology of Homo sapiens velopment LII (1987) Evolution of Catalytic Function XX (1955) Population Genetics: The Nature and Causes of Genetic LIII (1988) Molecular Biology of Signal Transduction Variability in Population LIV (1989) Immunological Recognition XXI (1956) Genetic Mechanisms: Structure and Function LV (1990) The Brain XXII (1957) Population Studies: Animal Ecology and LVI (1991) The Cell Cycle Demography LVII (1992) The Cell Surface XXIII (1958) Exchange of Genetic Material: Mechanism and Con- LVIII (1993) DNA and Chromosomes sequences LIX (1994) The Molecular Genetics of Cancer XXIV (1959) Genetics and Twentieth Century Darwinism LX (1995) Protein Kinesis: The Dynamics of Protein XXV (1960) Biological Clocks Trafficking and Stability XXVI (1961) Cellular Regulatory Mechanisms LXI (1996) Function & Dysfunction in the Nervous System XXVII (1962) Basic Mechanisms in Animal Virus Biology LXII (1997) Pattern Formation during Development XXVIII (1963) Synthesis and Structure of Macromolecules LXIII (1998) Mechanisms of Transcription XXIX (1964) Human Genetics LXIV (1999) Signaling and Gene Expression in the Immune System XXX (1965) Sensory Receptors LXV (2000) Biological Responses to DNA Damage XXXI (1966) The Genetic Code LXVI (2001) The Ribosome XXXII (1967) Antibodies LXVII (2002) The Cardiovascular System XXXIII (1968) Replication of DNA in Microorganisms LXVIII (2003) The Genome of Homo sapiens Front Cover (Paperback): Identical twins, New York City, 2003. ©Randy Harris. Authorization to photocopy items for internal or personal use, or the internal or personal use of specific clients, is granted by Cold Spring Harbor Laboratory Press, provided that the appropriate fee is paid directly to the Copyright Clearance Center (CCC). Write or call CCC at 222 Rosewood Drive, Danvers, MA 01923 (508-750-8400) for information about fees and regulations. Prior to photocopying items for educational classroom use, contact CCC at the above address. Additional information on CCC can be obtained at CCC Online at http://www.copyright.com/ All Cold Spring Harbor Laboratory Press publications may be ordered directly from Cold Spring Harbor Laboratory Press, 500 Sunnyside Boulevard, Woodbury, NY 11797-2924. Phone: 1-800-843-4388 in Continental U.S. and Canada. All other locations: (516) 422-4100. FAX: (516) 422-4097. E-mail: [email protected]. For a complete catalog of all Cold Spring Harbor Laboratory Press publications, visit our World Wide Web Site http://www.cshlpress.com/ Web Site Access: Institutions that have purchased the hardcover edition of this book are entitled to online access to the companion Web site at www.cshl-symposium.org. For assistance with activation, please contact Kathy Cirone, CSHL Press Subscription Manager, at [email protected]. 00_Symp69_frontmatter 4/21/05 11:36 AM Page v Symposium Participants AAPOLA, ULLA, Institute of Medical Technology, Univer- BARTHOLOMEW, BLAINE, Dept. of Biochemistry, South- sity of Tampere, Tampere, Finland ern Illinois University School of Medicine, Carbon- ADAMS, CHRISTOPHER, Dept. of Genomics, Invitrogen dale, Illinois Life Technologies, Carlsbad, California BARTOLOMEI, MARISA, Dept. of Cell and Developmental AIZAWA, YASUNORI, Johns Hopkins University, Balti- Biology, School of Medicine, University of Pennsyl- more, Maryland vania, Philadelphia ALLIS, C. DAVID, Lab. of Chromatin Biology, Rocke- BATISTA, PEDRO J., Program in Molecular Medicine, In- feller University, New York, New York stituto Gulbenkian de Ciência-Portugal and School of ALLISON, ANNE, Dept. of Microbiology, University of Medicine, University of Massachusets, Worcester Virginia, Charlottesville BAUER, MATTHEW, Div. of Biological Sciences, Univer- ALLSHIRE, ROBIN, Wellcome Trust Centre for Cell Biol- sity of Missouri, Columbia ogy, University of Edinburgh, Edinburgh, Scotland, BAULCOMBE, DAVID, John Innes Centre, Norwich, United United Kingdom Kingdom ALMEIDA, RICARDO, Institute of Cellular and Molecular BEAUDET, ARTHUR, Dept. of Molecular and Human Ge- Biology, University of Edinburgh, Edinburgh, Scot- netics, Baylor College of Medicine, Houston, Texas land, United Kingdom BECKER, PETER, Adolf Butenandt Institute, University of ALMOUZNI, GENEVIEVE, Institut Curie, Unité Mixte de Munich, Munich, Germany Recherche, Centre National de la Recherche Scien- BEIER, VERENA, Dept. of Functional Genome Analysis, tifique, Paris, France German Cancer Research Center, Heidelberg, Ger- ALTSCHULER, STEVEN, Center for Genomic Research, many Harvard University, Cambridge, Massachusetts BELOZEROV, VLADIMIR, Dept. of Biochemistry and AMARIGLIO, NINETTE, Dept. of Pediatric Hematology and Molecular Biology, University of Georgia, Athens Oncology, Sheba Medical Center, Tel Hashomer, Is- BENDER, JUDITH, Dept. of Biochemistry and Molecular rael Biology, School of Public Health, Johns Hopkins Uni- AN, WENFENG, Dept. of Molecular Biology and Genetics, versity, Baltimore, Maryland Johns Hopkins University, Baltimore, Maryland BENDER, LAUREL, Dept. of Biology, Indiana University, ANTAO, JOSE, Dept. of Molecular Biology, Harvard Med- Bloomington, Indiana ical School, Boston, Massachusetts BERGER, SHELLEY, Dept. of Molecular Genetics, Wistar ARAI, YOSHIO, Dept. of Radiation Oncology, Veteran’s Institute, Philadelpha, Pennsylvania Administration of Pittsburgh, University of Pittsburgh, BERNARDS, RENÉ, Div. of Molecular Carcinogenesis, The Pittsburgh, Pennsylvania Netherlands Cancer Institute, Amsterdam, The Nether- ARROYAVE, RANDY, Dept. of Biological Sciences, City lands University of New York, Hunter College, New York, BESTOR, TIMOTHY, Dept. of Genetics and Development, New York Columbia University, New York, New York AUGER, ANDREANNE, Centre de recherche de l’Hôtel- BIBIKOVA, MARINA, Dept. of Molecular Biology, Illu- Dieu de Québec, Laval University, Québec, Canada mina, Inc., San Diego, California AVITAHL-CURTIS, NICOLE, Models of Disease, Novartis BICKMORE, WENDY, Dept. of Chromosome Biology, Hu- Institutes for Biomedical Research, Cambridge, Mas- man Genetics Unit, Medical Research Council, Edin- sachusetts burgh, Scotland, United Kingdom AVRAM, DORINA, Center for Cell Biology and Cancer Re- BIRCHLER, JAMES, Div. of Biological Sciences, Univer- search, Albany Medical College, Albany, New York sity of Missouri, Columbia BALCIUNAITE, EGLE, Dept. of Pathology, School of BIRD, ADRIAN, Wellcome Trust Centre for Cell Biology, Medicine, New York University, New York, New University of Edinburgh, Edinburgh, Scotland, United York Kingdom BARLOW, DENISE,