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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 access 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, Center for Molecular Medicine, Insti- BIRNBAUMER, LUTZ, National Institute of Environmental tute of Microbiology and Genetics, University of Vi- Health Sciences, National Institutes of Health, Re- enna, Vienna, Austria search Triangle Park, North Carolina BARTEL, DAVID, Dept. of Biology, Whitehead Institute BOCCUNI, PIERNICOLA, Dept. of Molecular Pharmacology for Biomedical Research, Massachusetts Institute of and Therapeutics, Memorial Sloan-Kettering Cancer Technology, Cambridge, Massachusetts Institute, New York, New York

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vi SYMPOSIUM PARTICIPANTS

BOCKLANDT, SVEN, Dept. of Human Genetics, University Spring Harbor Laboratory, Cold Spring Harbor, New of California, Los Angeles York BOND, DIANE, Research and Development, Electrophore- CHANDLER, VICKI, Dept. of Plant Sciences, University of sis Platforms, Applied Biosystems, Foster City, Cali- Arizona, Tucson fornia CHEN, GENGXIN, Cold Spring Harbor Laboratory, Cold BONIFER, CONSTANZE, Dept. of Molecular Medicine, St. Spring Harbor, New York James University Hospital, University of Leeds, CHEN, JACK, Cold Spring Harbor Laboratory, Cold Leeds, United Kingdom Spring Harbor, New York BOURC’HIS, DEBORAH, Dept. of Genetics and Develop- CHEN, XIN, Dept. of Developmental Biology, Stanford ment, Columbia University, New York, New York University School of Medicine, Stanford, California BOYER, LAURIE, Dept. of Biology, Whitehead Institute CHENG, XIAODONG, Dept. of Biochemistry, Emory Uni- for Biomedical Research, Massachusetts Institute of versity School of Medicine, Atlanta, Georgia Technology, Cambridge, Massachusetts CHERUKURI, SRUJANA, Dept. of Cell Biology, Cleveland BRENNAN, JENNIFER, Dept. of Genetics, University of Clinic Foundation, Cleveland, Ohio North Carolina, Chapel Hill CHESS, ANDREW, Dept. of Biology, Whitehead Institute BROMAN, KARL, Dept. of Biostatistics, Johns Hopkins for Biomedical Research, Massachusetts Institute of University, Baltimore, Maryland Technology, Cambridge, Massachusetts BROWN, CAROLYN, Dept. of Medical Genetics, Univer- CHONG, SUYINN, School of Molecular and Microbial Bio- sity of British Columbia, Vancouver, British sciences, University of Sydney, Sydney, New South Columbia, Canada Wales, Australia BROWN, WILLIAM, Dept. of Genetics, Queens Medical CHOO, JUNGHA, Dept. of Biological Sciences, Korea Ad- Centre, University of Nottingham, Nottingham, United vanced Institute of Science and Technology, Daejeon, Kingdom South Korea BUCHBERGER, JOHANNES, Dept. of Cell Biology, Harvard CHUNG, JAE HOON, Dept. of Biological Sciences, Korea Medical School, Boston, Massachusetts Advanced Institute of Science and Technology, Dae- BUNGER, MAUREEN, Lab. of Molecular Carcinogenesis, jeon, South Korea National Institute of Environmental Health Sciences, CIRO, MARCO, Dept. of Experimental Oncology, Euro- National Institutes of Health, Research Triangle Park, pean Institute of Oncology, Milan, Italy North Carolina COCKERILL, PETER, Dept. of Molecular Medicine, St. BUSSEMAKER, HARMEN, Dept. of Biological Sciences, James University Hospital, University of Leeds, Columbia University, New York, New York Leeds, United Kingdom BYSTRICKY, KERSTIN, Dept. of Molecular Biology, Uni- COENRAADS, MONICA, Rett Syndrome Research Founda- versity of Geneva, Geneva, Switzerland tion, Cincinnati, Ohio CABELLO, OLGA, Dept. of Molecular and Cellular Biol- COHEN, AMIKAM, Dept. of Molecular Biology, Hebrew ogy, Baylor College of Medicine, Houston, Texas University Medical School, Jerusalem, Israel CAI, HAINI, Dept. of Cellular Biology, University of COLOT, VINCENT, Unité de Recherche en Génomique Georgia, Athens Végétale, Centre National de la Recherche Scien- CAO, RU, Dept. of Biochemistry and Biophysics, tifique, Evry, France Lineberger Comprehensive Cancer Center, University COMET, ITYS, Institut de Génétique Humaine, Centre Na- of North Carolina, Chapel Hill tional de la Recherche Scientifique, Montpellier, CARBONE, ROBERTA, Dept. of Experimental Oncology, France European Institute of Oncology, Milan, Italy COONROD, SCOTT, Dept. of Genetic Medicine, Weill CARTER, NIGEL, The Wellcome Trust Sanger Institute, Medical College, Cornell University, New York, New Hinxton, Cambridge, United Kingdom York CASLINI, CORRADO, Dept. of Medical Oncology, Fox COROMINAS, MONTSERRAT, Dept. of Genetics, Universitat Chase Cancer Center, Philadelphia, Pennsylvania de Barcelona, Barcelona, Spain CASSEL, SALLIE, Invitrogen Life Technologies, Carlsbad, CURRIE, RICHARD, Center for Scientific Review, National California Institutes of Health, Bethesda, Maryland CEDAR, HOWARD, Dept. of Cellular Biochemistry, Hadas- DAHARY, DVIR, Dept. of Cell Research and Immunology, sah Medical Center, Hebrew University, Jerusalem, Is- Tel Aviv University, Tel Aviv, Israel rael DAI, JUNBIAO, Dept. of Genetics, Development, and Cell CHADWICK, LISA, Institute for Genome Sciences and Pol- Biology, Iowa State University, Ames, Iowa icy, Duke University, Durham, North Carolina DANNENBERG, JAN-HERMEN, Dept. of Medical Oncology, CHAKRAVARTHY, SRINIVAS, Dept. of Biochemistry and Dana-Farber Cancer Institute, Boston, Massachusetts Molecular Biology, Colorado State University, Fort DAS, RAJDEEP, Cold Spring Harbor Laboratory, Cold Collins, Colorado Spring Harbor, New York CHALKER, DOUGLAS, Dept. of Biology, Washington Uni- DAVID, GREGORY, Dept. of Medical Oncology, Dana-Far- versity, St. Louis, Missouri ber Cancer Institute, Boston, Massachusetts CHAN, HUEI, Dept. of Pathology, University of Kentucky, DAVIS, ERICA, Dept. of Genetics, Faculty of Veterinary Lexington Medicine, University of Liège, Liège, Belgium CHAN, SHIRLEY, Dolan DNA Learning Center, Cold DEAN, CAROLINE, Dept. of Cell and Developmental Biol- 00_Symp69_frontmatter 4/21/05 11:36 AM Page vii

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ogy, John Innes Centre, Norwich, United Kingdom FELSENFELD, GARY, Lab. of Molecular Biology, National DEFOSSEZ, PIERRE, Institut Curie, Unité Mixte de Institute of Diabetes and Digestive Kidney Diseases, Recherche, Centre National de la Recherche Scien- National Institutes of Health, Bethesda, Maryland tifique, Paris, France FERRER, JORGE, Dept. of Endocrinology, IDIBAPS, Hos- DIAZ-PEREZ, SILVIA, Dept. of Human Genetics, Univer- pital Clinic de Barcelona, Barcelona, Spain sity of California, Los Angeles FIELDS, SCOTT, Dept. of Immunology, National Jewish DJUPEDAL, INGELA, Dept. of Natural Sciences, Karolinska Medical and Research Center, Denver, Colorado Institutet, University College Södertörn, Huddinge, FIRE, ANDREW, Dept. of Pathology and Genetics, Stan- Sweden ford University School of Medicine, Stanford, Califor- DORMAN, JANICE, Dept. of Epidemiology, University of nia Pittsburgh, Pittsburgh, Pennsylvania FISCHER, SYLVIA, Dept. of Molecular Biology, Mas- DORMANN, HOLGER, Lab. of Chromatin Biology, Rocke- sachusetts General Hospital, Harvard Medical School, feller University, New York, New York Boston, Massachusetts DOSTATNI, NATHALIE, Institut Curie, Unité Mixte de FISCHLE, WOLFGANG, Lab. of Chromatin Biology, Rock- Recherche, Centre National de la Recherche Scien- efeller University, New York, New York tifique, Paris, France FOUREL, GENEVIEVE, Unité Mixte de Recherche, Centre DRGON, TOMAS, Molecular Neurobiology Branch, National de la Recherche Scientifique, Ecole Normale Charles River Laboratories, National Institute on Drug Supérieure, Lyon, France Abuse, Baltimore, Maryland FRAZER, KIMBLE, Div. of Pediatric Hematology and On- DUAN, ZHIJUN, Dept. of Medical Genetics, University of cology, Huntsman Cancer Institute, University of Washington, Seattle Utah, Salt Lake City DUBOWITZ, VICTOR, Neuromuscular Unit, Dept. of Pedi- FRY, CHRISTOPHER, Program in Molecular Medicine, atrics, Imperial College London, London, United School of Medicine, University of Massachusetts, Kingdom Worcester DUNCAN, ELIZABETH, Lab. of Chromatin Biology, Rock- GANN, ALEXANDER, Cold Spring Harbor Laboratory efeller University, New York, New York Press, Woodbury, New York DUNN, REBECCA, Dept. of Molecular Biology, Mas- GASSER, SUSAN, Dept. of Molecular Biology, University sachusetts General Hospital, Harvard Medical School, of Geneva, Geneva, Switzerland Boston, Massachusetts GASZNER, MIKLOS, Lab. of Molecular Biology, National EBERT, JOAN, Cold Spring Harbor Laboratory Press, Institute of Diabetes and Digestive Kidney Diseases, Woodbury, New York National Institutes of Health, Bethesda, Maryland ECCLESTON, ALEX, Nature, Nature Publishing, San Fran- GENDREL, ANNE-VALERIE, Unité de Recherche en cisco, California Génomique Végétale, Centre National de la Recherche ECKHARDT, FLORIAN, Dept. of Life Sciences, Epige- Scientifique, Evry, France nomics AG, Berlin, Germany GEORGES, MICHEL, Dept. of Genetics, Faculty of Veteri- EKWALL, KARL, Dept. of Natural Sciences, Karolinska nary Medicine, University of Liège, Liège, Belgium Institutet, University College Södertörn, Huddinge, GILES, KEITH, Dept. of Biology, Johns Hopkins Univer- Sweden sity, Baltimore, Maryland ELGIN, SARAH, Dept. of Biology, Washington University, GOLDMAN, JOSEPH, Dept. of Molecular Biology, Harvard St. Louis, Missouri Medical School, Boston, Massachusetts EMMONS, SCOTT, Dept. of Molecular Genetics, Albert GOMEZ, JORGE, Dept. of Microbiology and Immunology, Einstein College of Medicine, Bronx, New York Emory University, Atlanta, Georgia ESUMI, NORIKO, Dept. of Ophthalmology, School of GOMOS-KLEIN, JANETTE, Dept. of Biological Sciences, Medicine, Johns Hopkins University, Baltimore, City University of New York, Hunter College, New Maryland York, New York EVANS, JASON, Cold Spring Harbor Laboratory, Cold GONZALO, SUSANA, Molecular Oncology Program, Span- Spring Harbor, New York ish National Cancer Centre, Madrid, Spain EZHKOVA, ELENA, Cold Spring Harbor Laboratory, Cold GOODMAN, LAURIE, Journal of Clinical Investigation, Spring Harbor, New York New York, New York FAN, YUHONG, Dept. of Cell Biology, Albert Einstein GOPINATHRAO, GOPAL, Cold Spring Harbor Laboratory, College of Medicine, Bronx, New York Cold Spring Harbor, New York FARMER, DEBORAH, Gene Regulation and Chromatin GOTO, DEREK, Cold Spring Harbor Laboratory, Cold Unit, Hammersmith Hospital, Clinical Sciences Cen- Spring Harbor, New York tre, Medical Research Council, Imperial College Lon- GOTTSCHLING, DANIEL, Fred Hutchinson Cancer Re- don, London, United Kingdom search Center, Seattle, Washington FEIL, ROBERT, Institute of Molecular Genetics, Centre GREWAL, SHIV, Lab. of Molecular Cell Biology, National National de la Recherche Scientifique, Montpellier, Cancer Institute, National Intitutes of Health, France Bethesda, Maryland FEINBERG, ANDREW, Epigenetics Unit, School of GROSSNIKLAUS, UELI, Institute of Plant Biology, Univer- Medicine, Johns Hopkins University, Baltimore, sity of Zurich, Zurich, Switzerland Maryland GROTH, ANJA, Dept. of Cell Cycle and Cancer, Danish 00_Symp69_frontmatter 4/21/05 11:36 AM Page viii

viii SYMPOSIUM PARTICIPANTS

Cancer Society, Copenhagen, Denmark Woodbury, New York GRUNSTEIN, MICHAEL, Dept. of Biological Chemistry, IOSHIKHES, ILYA, Dept. of Biomedical Informatics, Ohio University of California, Los Angeles State University, Columbus, Ohio GUEGLER, KARL, Applied Biosystems, Foster City, Cali- IRVINE, DANIELLE, Dept. of Chromosome Research, Mur- fornia doch Childrens Research Institute, Melbourne, Aus- GUENTHER, MATTHEW, Whitehead Institute for Biomedi- tralia cal Research, Massachusetts Institute of Technology, ISHINO, FUMITOSHI, Medical Research Center, Tokyo Cambridge, Massachusetts Medical and Dental University, Tokyo, Japan HAIG, DAVID, Dept. of Organismic and Evolutionary Bi- ISHOV, ALEXANDER, Dept. of Anatomy and Cell Biology, ology, Botanical Museum, Harvard University, Cam- University of Florida, Gainesville bridge, Massachusetts IWAMOTO, KAZUYA, Lab. for Molecular Dynamics of HAJKOVA, PETRA, Gurdon Institute, Wellcome Trust, Mental Disorders, Brain Science Institute, RIKEN, Cancer Research U.K., University of Cambridge, Wako, Japan Cambridge, United Kingdom JAENISCH, RUDOLF, Dept. of Biology, Whitehead Institute HALL, IRA, Cold Spring Harbor Laboratory, Cold Spring for Biomedical Research, Massachusetts Institute of Harbor, New York Technology, Cambridge, Massachusetts HAN, JEFFREY, Dept. of Molecular Biology and Genetics, JAYARAMAN, LATA, Dept. of Oncology, Bristol-Myers School of Medicine, Johns Hopkins University, Balti- Squibb, Princeton, New Jersey more, Maryland JELINEK, TOM, Upstate Biotechnology, Inc., Lake Placid, HANNON, GREGORY, Cold Spring Harbor Laboratory, New York Cold Spring Harbor, New York JENUWEIN, THOMAS, Research Institute of Molecular HARRIS, ABIGAIL, Abcam, Ltd., Cambridge, United King- Pathology, Vienna Biocenter, University of Vienna, dom Vienna, Austria HE, LIN, Cold Spring Harbor Laboratory, Cold Spring JIA, SONGTAO, Lab. of Molecular Cell Biology, National Harbor, New York Cancer Institute, National Institutes of Health, HE, YUEHUI, Dept. of Biochemistry, University of Wis- Bethesda, Maryland consin, Madison JIMENEZ, JOSE, Dept. of Cellular and Molecular Physiol- HEARD, EDITH, Mammalian Developmental Epigenetics ogy, Joslin Diabetes Center, Boston, Massachusetts Group, Institut Curie, Unité Mixte de Recherche, Cen- JOHNSTONE, KAREN, Dept. of Molecular Genetics and Mi- tre National de la Recherche Scientifique, Paris, crobiology, University of Florida, Gainesville France JORDAN, ALBERT, Dept. of Gene Regulation, Centre de HECHT, MERAV, Dept. of Cell Biochemistry and Human Regulació Genòmica, Barcelona, Spain Genetics, Hebrew University, Jerusalem, Israel JOSHI-TOPE, GEETA, Cold Spring Harbor Laboratory, HELD, WILLIAM, Dept. of Molecular and Cellular Biol- Cold Spring Harbor, New York ogy, Roswell Park Cancer Institute, Buffalo, New KAJI, KEISUKE, Institute for Stem Cell Research, Univer- York sity of Edinburgh, Edinburgh, Scotland, United King- HENIKOFF, STEVEN, Howard Hughes Medical Institute, dom Fred Hutchinson Cancer Research Center, Seattle, KAKKAD, REGHA, Center for Molecular Medicine, Insti- Washington tute of Microbiology and Genetics, Vienna Biocenter, HIRATANI, ICHIRO, Dept. of Biochemistry and Molecular University of Vienna, Vienna, Austria Biology, Upstate Medical University, State University KAKUTANI, TETSUJI, Dept. of Integrated Genetics, Na- of New York, Syracuse tional Institute of Genetics, Shizuoka, Japan HOEK, MAARTEN, Cold Spring Harbor Laboratory, Cold KALANTRY, SUNDEEP, Dept. of Genetics, University of Spring Harbor, New York North Carolina, Chapel Hill HOHN, BARBARA, Friedrich Miescher Institute for KAMBERE, MARIJO, Dept. of Molecular Biology and Bio- Biomedical Research, Basel, Switzerland chemistry, Wesleyan University, Middletown, Con- HORIKOSHI, MASAMI, Lab. of Developmental Biology, In- necticut stitute of Molecular and Cellular Biosciences, Univer- KAMPMANN, MARTIN, Lab. of Cell Biology, Rockefeller sity of Tokyo, Tokyo, Japan University, New York, New York HOULARD, MARTIN, Département de Biologie Joliot- KANNANGANATTU, PRASANTH, Cold Spring Harbor Labo- Curie, Service de Biochimie et de Génétique Molécu- ratory, Cold Spring Harbor, New York laire, CEA de Saclay, Gif-sur-Yvette, France KANNO, MASAMOTO, Dept. of Immunology, Hiroshima HSU, DUEN-WEI, Dept. of Biochemistry, Oxford Univer- University, Hiroshima, Japan sity, Oxford, United Kingdom KAPOOR, AVNISH, Dept. of Plant Sciences, University of HSU, MEI, Dept. of Microbiology and Immunology, Dart- Arizona, Tucson mouth Medical School, Lebanon, New Hampshire KARPF, ADAM, Dept. of Pharmacology and Therapeutics, HUA, SUJUN, Dept. of Genetics, Yale University School Roswell Park Cancer Institute, Buffalo, New York of Medicine, New Haven, Connecticut KASSCHAU, KRISTIN, Center for Gene Research and INAMOTO, SUSUMU, Dept. of Biotechnology, Institute of Biotechnology, Oregon State University, Corvallis, Research and Innovation, Chiba, Japan Oregon INGLIS, JOHN, Cold Spring Harbor Laboratory Press, KATO, MASAOMI, Dept. of Integrated Genetics, National 00_Symp69_frontmatter 4/21/05 11:36 AM Page ix

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Institute of Genetics, Mishima, Japan Medical School, Boston, Massachusetts KATO, TADAFUMI, Lab. for Molecular Dynamics of Men- KUCEROVA, MARTINA, Dept. of Biological Sciences, tal Disorders, Brain Science Institute, RIKEN, Wako, Hunter College, City University of New York, New Japan York, New York KAUFMAN, PAUL, Dept. of Molecular and Cell Biology, KURATOMI, GO, Lab. for Molecular Dynamics of Mental Lawrence Berkeley National Laboratory, University of Disorders, Brain Science Institute, RIKEN, Wako, California, Berkeley Japan KE, QINGDONG, Dept. of Environmental Medicine, New KURODA, MITZI, Harvard-Partners Center for Genetics York University, Tuxedo, New York and Genomics, Harvard University, Boston, Mas- KELLY, WILLIAM, Dept. of Biology, Emory University, sachusetts Atlanta, Georgia KWON, HYOCKMAN, Dept. of Bioscience and Biotechnol- KELSEY, GAVIN, Developmental Genetics Programme, ogy, Hankuk University of Foreign Studies, Yongin, The Babraham Institute, Cambridge, United Kingdom Seoul, South Korea KENNEDY, SCOTT, Dept. of Molecular Biology, Mas- LABOURIER, EMMANUEL, Dept. of Research and Develop- sachusetts General Hospital, Harvard Medical School, ment, Ambion, Inc., Austin, Texas Boston, Massachusetts LAI, ANGELA, Johnson and Johnson Research, Straw- KIDNER, CATHERINE, Cold Spring Harbor Laboratory, berry Hills, Australia Cold Spring Harbor, New York LAKEY, NATHAN, General and Administrative Offices, KIKYO, NOBUAKI, Stem Cell Institute, University of Min- Orion Genomics, St. Louis, Missouri nesota, Minneapolis LAKOWSKI, BERNARD, Dept. of Neuroscience, Nematode KIM, JOHN, Dept. of Molecular Biology, Massachusetts Genetics Group, Institut Pasteur, Paris, France General Hospital, Harvard Medical School, Boston, LANDE-DINER, LAURA, Dept. of Cellular Biochemistry, Massachusetts Hebrew University, Jerusalem, Israel KIM, JOOMYEONG, Div. of Genome Biology, Lawrence LEE, JEANNIE, Dept. of Molecular Biology, Mas- Livermore National Laboratory, Livermore, California sachusetts General Hospital, Harvard Medical School, KIM, MI YOUNG, Dept. of Molecular Biology and Genet- Boston, Massachusetts ics, Cornell University, Ithaca, New York LI, MO, Dept. of Cellular Biology, University of Georgia, KINGSTON, ROBERT, Dept. of Molecular Biology, Mas- Athens sachusetts General Hospital, Harvard Medical School, LI, YING, Dept. of Cellular and Developmental Biology, Boston, Massachusetts Fox Chase Cancer Center, Philadelphia, Pennsylvania KLAR, AMAR, Lab. of Gene Regulation and Chromosome LIM, JAE-HWAN, Lab. of Metabolism, National Cancer Biology, National Cancer Institute, Frederick, Mary- Institute, National Institutes of Health, Bethesda, land Maryland KLAUTKY, TRUDEE, Field Application Specialist, Invitro- LIPPMAN, ZACHARY, Cold Spring Harbor Laboratory, gen, Shrewsbury, Massachusetts Cold Spring Harbor, New York KLOSE, ROBERT, Wellcome Trust Centre for Cell Biol- LIPSICK, JOSEPH, Dept. of Pathology, Stanford University, ogy, University of Edinburgh, Edinburgh, Scotland, Stanford, California United Kingdom LIU, LIANG, Dept. of Biology, University of Alabama, KLYMENKO, TETYANA, Gene Expression Programme, Eu- Birmingham ropean Molecular Biology Laboratory, Heidelberg, LOPES, ALEXANDRA, Dept. of Population Genetics, Germany IPATIMUP, Instituto de Patologia e Immunologia KOH, MINGSHI, Dept. of Biological Sciences, National Molecular da Universidade de Porto, Porto, Portugal University of Singapore, Singapore, China LOUIS, EDWARD, Dept. of Genetics, University of Leices- KONFORTI, BOYANA, Nature Structural & Molecular Bi- ter, Leicester, United Kingdom ology, New York, New York LOWE, SCOTT, Cold Spring Harbor Laboratory, Cold KOUSKOUTI, ANTIGONE, Dept. of Molecular Genetics, In- Spring Harbor, New York stitute of Molecular Biology and Biotechnology, LU, XIANGDONG, Dept. of Biochemistry, University of Foundation for Research and Technology, Herakleion, North Carolina, Chapel Hill Crete, Greece LUBITZ, SANDRA, Dept. of Biotechnology, Dresden Uni- KOWENZ-LEUTZ, ELIZABETH, Dept. of Tumor Develop- versity of Technology, Dresden, Germany ment and Differentiation, Max Delbrück Center for LUSTIG, ARTHUR, Dept. of Biochemistry, Health Sciences Molecular Medicine, Berlin, Germany Center, Tulane University, New Orleans, Louisiana KRAUS, W. LEE, Dept. of Molecular Biology and Genet- LYLE, ROBERT, Dept. of Genetic Medicine and Develop- ics, Cornell University, Ithaca, New York ment, School of Medicine, University of Geneva, KRIAUCIONIS, SKIRMANTAS, Wellcome Trust Centre for Geneva, Switzerland Cell Biology, University of Edinburgh, Edinburgh, MADHANI, HITEN, Dept. of Biochemistry and Biophysics, Scotland, United Kingdom University of California, San Francisco KROGAN, NEVAN, Dept. of Medical Genetics, C.H. Best MAGGERT, KEITH, Dept. of Biology, University of Utah, Institute, University of Toronto, Toronto, Ontario, Salt Lake City Canada MAJUMDER, SADHAN, Dept. of Molecular Genetics, M.D. KSANDER, BRUCE, Dept. of Ophthalmology, Harvard Anderson Cancer Center, University of Texas, Houston 00_Symp69_frontmatter 4/21/05 11:36 AM Page x

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MAJUMDER, SARMILA, Dept. of Molecular and Cellular Netherlands Biochemistry, Ohio State University, Columbus, Ohio MONTGOMERY, NATHAN, Dept. of Genetics, University of MANDIYAN, VALSAN, Dept. of Pharmacology, Yale Uni- North Carolina, Chapel Hill versity, New Haven, Connecticut MOONEY, MYESHA, Dept. of Microbiology, Emory Uni- MANGONE, MARCO, Cold Spring Harbor Laboratory, Cold versity, Atlanta, Georgia Spring Harbor, New York MORRIS, KEVIN, Dept. of Medicine, University of Califor- MANIATAKI, ELISAVET, Dept. of Pathology, University of nia at San Diego, La Jolla Pennsylvania, Philadelphia MOTTAGUI-TABAR, SALIM, Center for Genomics and MANN, CARL, Service de Biochimie et de Génétique Bioinformatics, Karolinska Institutet, Stockholm, Swe- Moléculaire, CEA de Saclay, Gif-sur-Yvette, France den MARIN BIVENS, CARRIE, Dept. of Research, Jackson Lab- MUCHARDT, CHRISTIAN, Expression Génétique Molecu- oratory, Bar Harbor, Maine laire, Centre National de la Recherche Scientifique, In- MARTIENSSEN, ROBERT, Cold Spring Harbor Laboratory, stitut Pasteur, Paris, France Cold Spring Harbor, New York MUNGALL, ANDREW, The Wellcome Trust Sanger Insti- MASUI, OSAMU, Dept. of Nuclear Dynamics and Genome tute, Hinxton, Cambridge, United Kingdom Plasticity, Institut Curie, Unité Mixte de Recherche, MURCHISON, ELIZABETH, Cold Spring Harbor Laboratory, Centre National de la Recherche Scientifique, Paris, Cold Spring Harbor, New York France MUTSKOV, VESCO, Lab. of Molecular Biology, National MATOUK, CHARLES, Institute of Medical Sciences, Uni- Institute of Diabetes and Digestive and Kidney Dis- versity of Toronto, Toronto, Ontario, Canada eases, National Institutes of Health, Bethesda, Mary- MCCANN, JENNIFER, Dept. of Molecular Medicine, Ot- land tawa Health Research Institute, Ottawa, Ontario, MYAKISHEV, MAX, Lab. of Metabolism, National Cancer Canada Institute, National Institutes of Health, Bethesda, MELLO, CRAIG, Program in Molecular Medicine, Cancer Maryland Center, School of Medicine, University of Mas- NAGASE, HIROKI, Dept. of Cancer Genetics, Roswell Park sachusetts, Worcester Cancer Institute, Buffalo, New York MELNICK, ARI, Albert Einstein College of Medicine, NAKATANI, YOSHIHIRO, Dept. of Cancer Biology, Dana- Bronx, New York Farber Cancer Institute, Harvard Medical School, MEYER, BARBARA, Dept. of Molecular and Cellular Biol- Boston, Massachusetts ogy, Howard Hughes Medical Institute, University of NARAYAN, SANTOSH, Dept. of Pharmacology and Neuro- California, Berkeley science, University of North Texas Health Science MEYER, CLIFFORD, Dept. of Biostatistics, Dana-Farber Center, Fort Worth, Texas Cancer Institute, Cambridge, Massachusetts NISHIO, HITOMI, Dept. of Pediatrics, Mount Sinai School MEYER, ERIC, Laboratoire de Génétique Moléculaire, of Medicine, New York, New York Unité Mixte de Recherche, Centre National de la NOLAN, CATHERINE, Dept. of Zoology, University Col- Recherche Scientifique, Ecole Normale Supérieure, lege of Dublin, Dublin, Ireland Paris, France NOWACKI, MARIUSZ, Laboratoire de Génétique Molécu- MIAO, FENG, Dept. of Diabetes, Beckman Research Insti- laire, Unité Mixte de Recherche, Centre National de la tute, City of Hope, Duarte, California Recherche Scientifique, Ecole Normale Supérieure, MIN, JINRONG, Cold Spring Harbor Laboratory, Cold Paris, France Spring Harbor, New York NUNEZ, SABRINA, Cold Spring Harbor Laboratory, Cold MISHRA, NILAMADHAB, Dept. of Internal Medicine, Wake Spring Harbor, New York Forest University School of Medicine, Winston-Salem, OHTSUBO, HISAKO, Institute of Molecular and Cellular North Carolina Biosciences, University of Tokyo, Tokyo, Japan MISHRA, RAKESH, Centre for Cellular and Molecular Biol- OKADA, YUKI, Dept. of Biochemistry and Biophysics, ogy, Hyderabad, India University of North Carolina, Chapel Hill MITO, YOSHIKO, Dept. of Basic Science, Fred Hutchinson OMHOLT, STIG, Center for Integrative Genetics, Agricul- Cancer Research Center, Seattle, Washington tural University of Norway, Aas, Norway MITTELSTEN SCHEID, ORTRUN, Gregor Mendel Institute of ONISHI, MEGUMI, Dept. of Cell Biology, Harvard Medical Molecular Plant Biology, Vienna, Austria School, Boston, Massachusetts MIZUNO, YOSUKE, Center for Genomics and Bioinformat- OONO, KIYOHARU, Research Center for Environmental ics, Karolinska Institutet, Stockholm, Sweden Genomics, Kobe University, Kobe, Japan MO, XIANMING, Dept. of Tumor Development and Differ- ORDWAY, JARED, Dept. of Biomarkers, Orion Genomics, entiation, Max Delbrück Center for Molecular St. Louis, Missouri Medicine, Berlin, Germany ORLANDO, VALERIO, Dept. of Epigenetics and Genome MOHAN, KOMMU, Centre for Human Genetics, Bangalore, Reprogramming, Dulbecco Telethon Institute, Insti- India tute of Genetics and Biophysics, CNR, Naples, Italy MOHD-SARIP, ADONE, Dept. of Molecular and Cell Biol- ORTIZ, BENJAMIN, Dept. of Biological Sciences, Hunter ogy, Sylvius Laboratory, Centre for Biomedical Genet- College, City University of New York, New York, ics, Leiden University Medical Centre, Leiden, The New York 00_Symp69_frontmatter 4/21/05 11:36 AM Page xi

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OZER, JOSEF, Dept. of Pharmacology, School of Moléculaire, Unité Mixte de Recherche, Centre Na- Medicine, Boston University, Boston, Massachusetts tional de la Recherche Scientifique, Ecole Normale PADDISON, PATRICK, Cold Spring Harbor Laboratory, Supérieure, Paris, France Cold Spring Harbor, New York PROBST, ALINE, Dept. of Plant Biology, University of PAGAN, JULIA, Dept. of Cancer and Cell Biology, Geneva, Geneva, Switzerland Queensland Institute of Medical Research, Brisbane, PRUITT, ROBERT, Dept. of Botany and Plant Pathology, Australia Purdue University, West Lafayette, Indiana PALLAORO, MICHELE, Dept. of Biochemistry, Institute for PTASHNE, MARK, Dept. of Molecular Biology, Memorial Research in Molecular Biology, Merck, Pomezia, Italy Sloan-Kettering Cancer Institute, New York, New PAN, QI, Dept. of Molecular Biology, Massachusetts York General Hospital, Harvard Medical School, Boston, RAKYAN, VARDHMAN, Dept. of Epigenomics, The Well- Massachusetts come Trust Sanger Institute, Hinxton, Cambridge, PARO, RENATO, Zentrum für Molekulare Biology, Uni- United Kingdom versity of Heidelberg, Heidelberg, Germany RAMACHANDRAN, ILENG, Cold Spring Harbor Laboratory, PARRY, DEVIN, Dept. of Molecular Biology, Mas- Cold Spring Harbor, New York sachusetts General Hospital, Harvard Medical School, RAMASWAMY, AMUTHA, Dept. of Biomedical Informat- Boston, Massachusetts ics, Ohio State University, Columbus, Ohio PARTHUN, MARK, Dept. of Molecular and Cellular Bio- RANDO, OLIVER, Bauer Center for Genomics Research, chemistry, Ohio State University, Columbus, Ohio Harvard University, Cambridge, Massachusetts PARTRIDGE, JANET, Dept. of Biochemistry, St. Jude Chil- RAUSCHER, FRANK, Dept. of Molecular Genetics, Wistar dren’s Research Hospital, Memphis, Tennessee Institute, Philadelphia, Pennsylvania PASZKOWSKI, JERZY, Dept. of Plant Biology, University RECHAVI, GIDEON, Sheba Cancer Research Center, Sheba of Geneva, Geneva, Switzerland Medical Center, Tel-Hashomer, Israel PELISSON, ALAIN, Institut de Génétique Humaine, Centre REDDI, PRABHAKARA, Dept. of Cell Biology, University National de la Recherche Scientifique, Montpellier, of Virginia, Charlottesville France REESE, KIMBERLY, Dept. of Cell and Developmental Bi- PENNELL, ROGER, Ceres, Inc., Malibu, California ology, Howard Hughes Medical Institute, University PETERSON, CRAIG, Program in Molecular Medicine, Med- of Pennsylvania, Philadelphia ical Center, School of Medicine, University of Mas- REIK, WOLF, Developmental Genetics Programme, The sachusetts, Worcester Babraham Institute, Cambridge, United Kingdom PETRIE, VICTORIA, Dept. of Biochemistry, St. Jude Chil- REINBERG, DANNY, Dept. of Biochemistry, Howard dren’s Research Hospital, Memphis, Tennessee Hughes Medical Institute, UMDNJ-Robert Wood PFEFFER, PETER, Dept. of Reproduction, Agresearch Johnson Medical School, Piscataway, New Jersey Crown Research Institute, Hamilton, New Zealand RICE, JUDD, Dept. of Biochemistry and Molecular Biol- PHAM, ANH-DUNG, Dept. of Biochemistry, Howard ogy, University of Southern California, Los Angeles, Hughes Medical Institute, UMDNJ-Robert Wood California Johnson Medical School, Piscataway, New Jersey RICHARDS, ERIC, Dept. of Biology, Washington Univer- PIEDRAHITA, JORGE, Dept. of Molecular Biomedical Sci- sity, St. Louis, Missouri ences, North Carolina State University, Raleigh, North RIDDIHOUGH, GUY, Science, American Association for Carolina the Advancement of Science, Washington, D.C. PIEN, STEPHANE, Dept. of Plant Systems Biology, VIB, RIDDLE, NICOLE, Div. of Biological Sciences, University University of Gent, Gent, Belgium of Missouri, Columbia PILLUS, LORRAINE, Dept. of Biology, University of Cali- RINE, JASPER, Dept. of Molecular and Cellular Biology, fornia at San Diego, La Jolla University of California, Berkeley PIRROTTA, VINCENZO, Dept. of Zoology, University of RIVERA, ISABEL, Dept. of Biochemistry, Faculty of Phar- Geneva, Geneva, Switzerland macy, University of Lisbon, Lisbon, Portugal PLASTERK, RONALD, Hubrecht Laboratory, Netherlands ROBERTS, CHARLES, Dept. of Pediatric Oncology, Dana- Institute for Developmental Biology, Utrecht, The Farber Cancer Institute, Boston, Massachusetts Netherlands ROLLINS, ROBERT, Dept. of Genetics and Development, POGACIC´, VANDA, Friedrich Miescher Institute for Columbia University, New York, New York Biomedical Research, Basel, Switzerland RONEMUS, MICHAEL, Cold Spring Harbor Laboratory, v POLLOCK, MILA, Cold Spring Harbor Laboratory Library, Cold Spring Harbor, New York Cold Spring Harbor, New York ROSEBROCK, ADAM, Dept. of Molecular Genetics and Mi- POOT, RAYMOND, Marie Curie Research Institute, Oxted, crobiology, State University of New York, Stony United Kingdom Brook POZZI, SILVIA, Dept. of Cancer Genetics, Roswell Park ROSSMANN, MARLIES, Cold Spring Harbor Laboratory, Cancer Institute, Buffalo, New York Cold Spring Harbor, New York PRASANTH, SUPRIYA, Cold Spring Harbor Laboratory, ROSSON, GARY, Lineberger Comprehensive Cancer Cen- Cold Spring Harbor, New York ter, University of North Carolina, Chapel Hill PRIOLEAU, MARIE-NOELLE, Laboratoire de Génétique ROZENBERG, JULIAN, Lab. of Metabolism, National Can- 00_Symp69_frontmatter 4/21/05 11:36 AM Page xii

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cer Institute, National Institutes of Health, Bethesda, SI, KAUSIK, Dept. of Neurobiology and Behavior, Maryland Columbia University, New York, New York RUGG-GUNN, PETER, Dept. of Surgery, University of SIDDIQUI, KHALID, Cold Spring Harbor Laboratory, Cold Cambridge, Cambridge, United Kingdom Spring Harbor, New York RUSCHE, LAURA, Dept. of Biochemistry, Duke Univer- SIDHU, RAVINDER, Bone and Joint Research Unit, St. sity, Durham, North Carolina Bartholomew’s Hospital, London, United Kingdom RUSK, NICOLE, Nature Methods, New York, New York SIGOVA, ALLA, Dept. of Biochemistry and Molecular RUSSANOVA, VALYA, National Institute of Child Health Pharmacology, School of Medicine, University of and Human Development, National Institutes of Massachusetts, Worcester Health, Bethesda, Maryland SKIPPER, MAGDALENA, Nature Reviews Genetics, Lon- RUVKUN, GARY, Dept. of Molecular Biology, Mas- don, United Kingdom sachusetts General Hospital, Harvard Medical School, SKOULTCHI, ARTHUR, Dept. of Cell Biology, Albert Ein- Boston, Massachusetts stein College of Medicine, Bronx, New York SAAVEDRA, RAUL, Scientific Review Branch, National SLAWSON, ELIZABETH, Dept. of Biology, Washington Institute of Neurological Disorders and Stroke, Na- University, St. Louis, Missouri tional Institutes of Health, Bethesda, Maryland SMITH, ANDREW, Cold Spring Harbor Laboratory, Cold SACCHI, NICOLETTA, Dept. of Cancer Genetics, Roswell Spring Harbor, New York Park Cancer Institute, Buffalo, New York SMITS, GUILLAUME, IRIBHM, Université Libre de Brux- SANDERS, STEVEN, Gurdon Institute, Wellcome Trust, elles, Brussels, Belgium Cancer Research U.K., University of Cambridge, SOLTER, DAVOR, Dept. of Developmental Biology, Max Cambridge, United Kingdom Planck Institute for Immunobiology, Freiburg, Ger- SASAKI, HIROYUKI, Div. of Human Genetics, National In- many stitute of Genetics, Mishima, Japan SOUTOGLOU, EVANTHIA, National Cancer Institute, Na- SASSONE-CORSI, PAOLO, Institut de Génétique et de Bi- tional Institutes of Health, Bethesda, Maryland ologie Moléculaire et Cellulaire, Illkirch, France SPADA, FABIO, Sars International Centre for Marine SATO, NORIKO, Tokyo Metropolitan Institute of Medical Molecular Biology, University of Bergen, Bergen, Science, Tokyo, Japan Norway SCHILDKRAUT, CARL, Dept. of Cell Biology, Albert Ein- SPARMANN, ANKE, Dept. of Microbiology, State Univer- stein College of Medicine, Bronx, New York sity of New York, Stony Brook SCHLEGEL, ROBERT, Dept. of Oncology, Novartis Insti- SPECTOR, DAVID, Cold Spring Harbor Laboratory, Cold tutes for BioMedical Research, Cambridge, Mas- Spring Harbor, New York sachusetts SPECTOR, MONA, Cold Spring Harbor Laboratory, Cold SCHOENHERR, CHRISTOPHER, Dept. of Cell and Structural Spring Harbor, New York Biology, University of Illinois, Urbana SRIDHARAN, RUPA, Howard Hughes Medical Institute, SCHOLZ, CHRISTIAN, Dept. of Hematology and Oncology, University of California, Los Angeles Charité Campus Virchow-Klinikum, Berlin, Germany SRINIVASAN, LAKSHMI, Dept. of Animal Biology, School SCHONES, DUSTIN, Cold Spring Harbor Laboratory, Cold of Veterinary Medicine, University of Pennsylvania, Spring Harbor, New York Philadelphia SEISER, CHRISTIAN, Dept. of Medical Biochemistry, Vi- STANCHEVA, IRINA, Dept. of Biochemistry, University of enna Biocenter, University of Vienna, Vienna, Austria Edinburgh, Edinburgh, Scotland, United Kingdom SELKER, ERIC, Institute of Molecular Biology, University STEBBINS, MICHAEL, Nature Genetics, New York, New of Oregon, Eugene York SHARP, JUDITH, Dept. of Biochemistry and Biophysics, STEWART, DAVID, Meetings and Courses Programs, Cold University of California, San Francisco Spring Harbor Laboratory, Cold Spring Harbor, New SHARP, PHILLIP, Center for Cancer Research, Mas- York sachusetts Institute of Technology, Cambridge, Mas- STILLMAN, BRUCE, President and CEO, Cold Spring Har- sachusetts bor Laboratory, Cold Spring Harbor, New York SHEN, CHANG-HUI, Dept. of Biology, College of Staten STRAHL, BRIAN, Dept. of Biochemistry and Biophysics, Island, City University of New York, Staten Island, University of North Carolina, Chapel Hill New York STRUNNIKOV, ALEXANDER, Lab. of Gene Regulation and SHIBAHARA, KEI-ICHI, Dept. of Integrated Genetics, Na- Development, National Institute of Child Health and tional Institute of Genetics, Mishima, Japan Human Development, National Institutes of Health, SHINKAI, YOICHI, Dept. of Cell Biology, Institute for Bethesda, Maryland Virus Research, Kyoto University, Kyoto, Japan SU, RUEY-CHYI, Dept. of Microbiology, Immunology, SHOVLIN, TANYA, Dept. of Biomedical Sciences, Univer- and Molecular Genetics, Howard Hughes Medical In- sity of Ulster, Coleraine, Londonderry, Northern Ire- stitute, University of California, Los Angeles land, United Kingdom SURANI, AZIM, Gurdon Institute, Wellcome Trust, Cancer SHUREIQI, IMAD, Dept. of Clinical Cancer Prevention, Research U.K., University of Cambridge, Cambridge, M.D. Anderson Cancer Center, University of Texas, United Kingdom Houston SUSSMAN, HILLARY, Genome Research, Cold Spring Har- 00_Symp69_frontmatter 4/21/05 11:36 AM Page xiii

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bor Laboratory Press, Woodbury, New York VERDEL, ANDRE, Dept. of Cell Biology, Harvard Medical SWIGUT, TOMASZ, Lab. of Molecular Vertebrate Embry- School, Boston, Massachusetts ology, Rockefeller University, New York, New York VINCENZ, CLAUDIUS, Dept. of Biological Chemistry, SYMER, DAVID, Lab. of Immunobiology, National Cancer Howard Hughes Medical Institute, School of Institute, Frederick, Maryland Medicine, University of Michigan, Ann Arbor TACHIBANA, MAKOTO, Lab. of Mouse Models, Institute VINSON, CHARLES, Lab. of Metabolism, National Insti- for Virus Research, Kyoto University, Kyoto, Japan tutes of Health, Bethesda, Maryland TAHILIANI, MAMTA, Dept. of Pathology, Center for Blood WAGLE, MAHENDRA, Lab. of Developmental Neurobiol- Research, Harvard Medical School, Boston, Mas- ogy, Temasek Life Sciences Laboratory, Singapore, sachusetts China TAIPALE, MIKKO, Gene Expression Programme, Euro- WALSH, MARTIN, Dept. of Pediatrics and Human Genet- pean Molecular Biology Laboratory, Heidelberg, Ger- ics, Mount Sinai School of Medicine, New York, New many York TAKEDA, SHIN, Dept. of Plant Biology, University of WANG, DUO, Dept. of Molecular Biology, Massachusetts Geneva, Geneva, Switzerland General Hospital, Harvard Medical School, Boston, TAM, PATRICK, Embryology Unit, Children’s Medical Massachusetts Research Institute, Westmead, Sydney, Australia WANG, SONG, Dept. of Biological Sciences, Stanford TAMADA, HIROSHI, Stem Cell Institute, University of University, Stanford, California Minnesota, Minneapolis WANG, YANMING, Lab. of Chromatin Biology, Rocke- THOMPSON, JEFFREY, Dept. of Biology, Denison Univer- feller University, New York, New York sity, Granville, Ohio WARBURTON, PETER, Dept. of Human Genetics, Mount TIAN, YANAN, Dept. of Veterinary Physiology and Phar- Sinai School of Medicine, New York, New York macology, Texas A&M University, College Station, WEBSTER, KYLIE, Dept. of Genetics and Bioinformatics, Texas Walter and Eliza Hall Institute of Medical Research, TIMMERMANS, MARJA, Cold Spring Harbor Laboratory, Parkville, Victoria, Australia Cold Spring Harbor, New York WEST, ADAM, Div. of Cancer Sciences and Molecular TOMPA, RACHEL, Dept. of Biochemistry and Biophysics, Pathology, Western Infirmary, University of Glasgow, University of California, San Francisco Glasgow, Scotland, United Kingdom TORRES, HECTOR, National Institute of Environmental WEST, KATHERINE, Div. of Cancer Sciences and Molecu- Health Sciences, National Institutes of Health, Re- lar Pathology, Western Infirmary, University of Glas- search Triangle Park, North Carolina gow, Glasgow, Scotland, United Kingdom TSIKITIS, MARY, Dept. of Pathology, School of Medicine, WHITEHEAD, NEDRA, Applied Sciences Branch, Centers New York University, New York, New York for Disease Control, Atlanta, Georgia TSUCHIMOTO, SUGURU, Institute for Molecular and Cellu- WICKNER, REED, Lab. of Biochemistry and Genetics, Na- lar Biosciences, University of Tokyo, Tokyo, Japan tional Institute of Diabetes and Digestive and Kidney TUTEJA, JIGYASA, Dept. of Physiological and Molecular Diseases, National Institutes of Health, Bethesda, Plant Biology, University of Illinois, Urbana Maryland TUZON, CREIGHTON, Dept. of Biochemistry and Molecu- WILHELM, BRIAN, The Wellcome Trust Sanger Institute, lar Genetics, Health Sciences Center, Universiy of Hinxton, Cambridge, United Kingdom Colorado, Denver WILLIAMSON, CHRIS, Mammalian Genetics Unit, Medical URA, KIYOE, Div. of Gene Therapy Science, Graduate Research Council, Harwell, Oxfordshire, United King- School of Medicine, Osaka University, Osaka, Japan dom VALER, MARC, Dept. of Research and Development and WILSON, JON, Dept. of Protein Structure, National Insti- Marketing, Agilent Technologies, Waldbronn, Ger- tute for Medical Research, London, United Kingdom many WITKOWSKI, JAN, Banbury Center, Cold Spring Harbor VAN HOUDT, HELENA, Dept. of Plant Systems Biology, Laboratory, Cold Spring Harbor, New York VIB, University of Gent, Gent, Belgium WONG, LEE, Dept. of Chromosome Research, Murdoch VAN LOHUIZEN, MAARTEN, Div. of Moleular Genetics, Children’s Research Institute, Royal Children’s Hospi- The Netherlands Cancer Institute, Amsterdam, The tal, Parkville, Victoria, Australia Netherlands WU, HAO, Dept. of Molecular and Medical Pharmacol- VAN LOO, KAREN, Dept. of Molecular Animal Physiol- ogy, University of California, Los Angeles ogy, University of Nijmegen, Nijmegen, The Nether- WU, JIANG, Dept. of Pathology and Developmental Biol- lands ogy, Howard Hughes Medical Institute, Stanford Uni- VAN RIETSCHOTEN, JOHANNA, Dept. Molecular Cell Biol- versity, Stanford, California ogy and Immunology, VU Medical Center, Amster- WYSOCKA, JOANNA, Lab. of Chromatin Biology, Rocke- dam, The Netherlands feller University, New York, New York VAUGHN, MATTHEW, Cold Spring Harbor Laboratory, XIANG, TIANHAO (TIM), Dept. of Reproductive Technol- Cold Spring Harbor, New York ogy, ViaGen, Inc., Austin, Texas VAURY, CHANTAL, Université D’Auvergne, Clermont- XU, GUOLIANG, Shanghai Institute for Biological Sci- Ferrand, France ences, Chinese Academy of Sciences, Shanghai, China 00_Symp69_frontmatter 4/21/05 11:36 AM Page xiv

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XUAN, ZHENYU, Cold Spring Harbor Laboratory, Cold YUSA, KOSUKE, Dept. of Social and Environmental Spring Harbor, New York Medicine, Osaka University, Osaka, Japan YANG, SEUNG KYOUNG, Dept. of Biological Sciences, Ko- ZAMORE, PHILLIP, Dept. of Biochemistry and Molecular Bi- rea Advanced Institute of Science and Technology, Dae- ology, School of Medicine, University of Massachusetts, jeon, South Korea Worcester YANG, XIANG-JIAO, Dept. of Medicine, McGill University, ZHANG, FENG, Dept. of Genetics, Development, and Cell Montréal, Québec, Canada Biology, Iowa State University, Ames, Iowa YE, XIAOFEN, Dept. of Molecular Oncology, Fox Chase ZHANG, YI, Dept. of Biochemistry and Biophysics, Cancer Center, Philadelphia, Pennsylvania Lineberger Comprehensive Cancer Center, University of YILDIRIM, EDA, Lab. of Signal Transduction, National In- North Carolina, Chapel Hill stitute of Environmental Health Sciences, National Insti- ZHANG, ZHIGUO, Dept. of Biochemistry and Molecular Bi- tutes of Health, Research Triangle Park, North Carolina ology, Mayo Clinic, Rochester, Minnesota YOKOYAMA, KAZUNARI, Gene Engineering Division, ZHAO, KEJI, National Heart, Lung, and Blood Institute, Na- RIKEN, Tsukuba Science City, Ibaraki, Japan tional Institutes of Health, Bethesda, Maryland YOUNG, DALLAN, Dept. of Biochemistry and Molecular Bi- ZHONG, GAN-YUAN, Dept. of Agriculture & Nutrition, Pio- ology, University of Calgary, Calgary, Alberta, Canada neer Hi-Bred International, Johnston, Iowa 00_Symp69_frontmatter 4/21/05 11:36 AM Page xv

First row: R. Plasterk; A. Auger, D. Young, H. Van Houdt; D. Allis, E. Bernstein Second row: E. Selker; P. Sharp, H. Cedar; E. Louis, D. Gottschling, S. Henikoff; E. Heard Third row: Y. Fan, G. Fourel; P. Sassone-Corsi, B. Stillman; B. Ortiz, G. Felsenfeld Fourth row: A. Surani, P. Hajkova; V. Pirrotta, J. Paszkowski, M. Timmermans 00_Symp69_frontmatter 4/21/05 11:37 AM Page xvi

First row: L. Goodman; S. Emmons, G. Ruvkun, B. Meyer; E. Richards, D. Baulcombe, I. Stancheva Second row: M. Macari demonstrating the online Oral History Project; S. Gasser, C. Schildkraut Third row: E. Yildirim, S. Cherukuri, B. Meyer; H. Sussman, M. Bartolomei, J. Lee Fourth row: G. Kelsey, E. Davis; P. Overbeek, O. Cabello; I. Hall, J. Birchler 00_Symp69_frontmatter 4/21/05 11:37 AM Page xvii

First row: S. Kennedy, E. Meyer; D. Haig; R. Pruitt; H. Sasaki; C. Peterson, S. Berger Second row: C. Mann, P. Kaufman; G. Rechavi, A. Cohen; M. Grunstein, A. Bird Third row: D. Spector, D. Stewart, P. Sassone-Corsi; P. Cockerill, C. Williamson; U. Grossniklaus, Y. Lazebnik Fourth row: Airslee Wine and Cheese; O. Rando, S. Altschuler 00_Symp69_frontmatter 4/21/05 11:37 AM Page xviii

First row: C. Tuzon, K. Ekwall; A. Pelisson, Z. Lippman Second row: B. Hohn, G.-Y. Zhong; W. Bickmore; J. Partridge, L. Gann Third row: Symposium Cocktails on Blackford Lawn 00_Symp69_frontmatter 4/21/05 11:37 AM Page xix

First row: Lunchtime on Blackford Lawn Second row: M. Gaszner; G. Almouzni, Y. Nakatani; S. Lubitz, K. Webster Third row: J. Watson, R. Wickner; K. Giles 00_Symp69_frontmatter 4/21/05 11:37 AM Page xx

First row: Brunch chez Blackford; K. West, M.-N. Prioleau Second row: P. Sherwood, A. Klar, J. Hicks; V. Pogacic,v ′ V. Beier Third row: J. Birchler, R. Martienssen; Grace closed circuit TV 00_Symp69_frontmatter 4/21/05 11:37 AM Page xxi

First row: P. Kannanganattu, V. Ellison; D. Barlow, R. Lyle; V. Chandler, J. Bender Second row: A Chess; Y. He, C. Dean; W. Kelly, V. Colot; S. Elgin Third row: D. Gottschling, R. Jaenisch; P. Zamore, D. Bartel; A. Skoultchi, A. Harris Fourth row: Summary (D. Gottschling) 00_Symp69_frontmatter 4/21/05 11:37 AM Page xxii

First row: Y. Okada, H. Chan; T. Bestor, X. Cheng; B. Strahl Second row: Y. Arai; A. Eccleston, R. Kingston, M. van Lohuizen; W. Reik; N. Sato Third row: J. Brennan; M. Hoek, D. Goto; R. Rollins, N. Rusk, W. Held, G. Smits Fourth row: A. Feinberg, M. Stebbins; D. Solter, J. Witkowski, A. Surani; G. Rosson 00_Symp69_frontmatter 4/21/05 11:37 AM Page xxiii

Foreword

Until recently, the general view of heredity has been seen through the lens of DNA. Indeed, the 2003 Symposium on “The Genome of Homo sapiens” contributed to that view by empha- sizing the importance of DNA sequence and its origins. But increasingly, investigators are ex- ploring a set of secondary phenomena that give rise to heritable changes in gene function that oc- cur without a change in the underlying DNA sequence—epigenetic mechanisms such as DNA methylation, histone acetylation, imprinting, RNA interference, gene silencing, and paramuta- tion. A growing body of evidence indicates that epigenetic changes are important contributors to the pathogenesis of disease in humans, animals, and plants and may lie at the heart of many important gene–environment interactions. And so it seemed timely to hold a Symposium explicitly devoted to “Epigenetics.” Previous Symposia that have in part examined the role of the macromolecular context in which the primary genetic information is found include the 1941 Symposium on “Genes and Chromo- somes: Structure and Organization,” which emphasized a biophysical approach to these structures; the two closely separated Symposia that examined “Chromosome Structure and Function” (1973) and “Chromatin” (1977), at which latter meeting the nature of the nucleosome was un- veiled; and the 1993 Symposium on “DNA and Chromosomes,” by which time the human genome project with its focus on the primary sequence was well underway. The 69th Sympo- sium, however, was the first to fully explore the heritable aspects to these and related biochem- ical phenomena. The field of epigenetics as we know it today was prominently introduced at the 1951 Sympo- sium on “Genes and Mutations.” There, Ed Lewis presented data on position effect variegation in Drosophila, a phenomenon that has played an important role in the history of the field. Equally importantly, Barbara McClintock presented her ideas about heterochromatin and movable genetic elements, the so-called Ac-Ds system in maize that opened up understanding of transposi- tion and its links to gene silencing and formation of heterochromatin. Some 53 years later, the 69th Symposium witnessed a rather complete molecular description of her ideas, including links to RNAi. The 69th Symposium ran for 5 days and included 460 participants with 68 oral presentations and 210 poster presentations. We wish to thank Dr. David Haig for his superb Dorcas Cummings Memorial public lecture presenting a theoretical approach to epigenetics, on the subject of The Divided Self—Brains, Brawn and the Superego. We also wish to thank the first night speakers, Drs. Davor Solter, Barbara Meyer, David Allis, and Andrew Fire for their superb overview presentations, and we particularly thank Dr. Daniel Gottschling for agreeing to summarize the Meeting. We thank Jenna Williams and Mary Smith in the Meetings and Courses office for efficiently running the meeting and John Inglis and his staff at the Cold Spring Harbor Laboratory Press, particularly Joan Ebert, Kathleen Bubbeo, and Danny deBruin, for publishing both the on-line and printed versions of the meeting manuscripts. Essential funds to run this meeting were obtained from the National Cancer Institute, a branch of the National Institutes of Health. In addi- tion, financial help from the corporate benefactors, sponsors, affiliates, and contributors of our meetings program is essential for these Symposia to remain a success and we are most grateful for their continued support.

Bruce Stillman David Stewart March 2005

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