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COLD SPRING HARBOR SYMPOSIA ON QUANTITATIVE BIOLOGY VOLUME LVIII COLD SPRING HARBOR SYMPOSIA ON QUANTITATIVE BIOLOGY VOLUME LVIII DNA and Chromosomes COLD SPRING HARBOR LABORATORY PRESS 1993 COLD SPRING HARBOR SYMPOSIA ON QUANTITATIVE BIOLOGY VOLUME LVIII 1993 by The Cold Spring Harbor Laboratory Press International Standard Book Number 0-87969-065-8 (cloth) International Standard Book Number 0-87969-066-6 (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 xxvIII (1963) Synthesis and Structure of Macromolecules II (1934) Aspects of Growth XXIX (1964) Human Genetics III (1935) Photochemical Reactions XXX (1965) Sensory Receptors IV (1936) Excitation Phenomena XXXI (1966) The Genetic Code V (1937) Internal Secretions XXXII (1967) Antibodies VI (1938) Protein Chemistry XXXIII (1968) Replication of DNA in Microorganisms VII (1939) Biological Oxidations XXXIV (1969) The Mechanism of Protein Synthesis VIII (1940) Permeability and the Nature of Cell Membranes XXXV (1970) Transcription of Genetic Material IX (1941) Genes and Chromosomes: Structure and Organiza- XXXVI (1971) Structure and Function of Proteins at the tion Three-dimensional Level X (1942) The Relation of Hormones to Development XXXVII (1972) The Mechanism of Muscle Contraction XI (1946) Heredity and Variation in Microorganisms XXXVIII (1973) Chromosome Structure and Function XII (1947) Nucleic Acids and Nucleoproteins XXXIX (1974) Tumor Viruses XIII (1948) Biological Applications of Tracer Elements XL (1975) The Synapse XIV (1949) Amino Acids and Proteins XLI (1976) Origins of Lymphocyte Diversity XV (1950) Origin and Evolution of Man XLII (1977) Chromatin XVI (1951) Genes and Mutations XLIII (1978) DNA: Replication and Recombination XVII (1952) The Neuron XLIV (1979) Viral Oncogenes XVIII (1953) Viruses XLV (1980) Movable Genetic Elements XIX (1954) The Mammalian Fetus: Physiological Aspects of XLVI (198l) Organization of the Cytoplasm Development XLVII (1982) Structures of DNA XX (1955) Population Genetics: The Nature and Causes of XLVIII (1983) Molecular Neurobiology Genetic Variability in Population XLIX (1984) Recombination at the DNA Level XXI (1956) Genetic Mechanisms: Structure and Function L (1985) Molecular Biology of Development XXII (1957) Population Studies: Animal Ecology and De- LI (1986) Molecular Biology of Homo sapiens mography LII (1987) Evolution of Catalytic Function XXIII (1958) Exchange of Genetic Material: Mechanism and LIII (1988) Molecular Biology of Signal Transduction Consequences LIV (1989) Immunological Recognition XXIV (1959) Genetics and Twentieth Century Darwinism LV (1990) The Brain XXV (1960) Biological Clocks LVI (1991) The Cell Cycle XXVI (1961) Cellular Regulatory Mechanisms LVII (1992_) The Cell Surface XXVII (1962) Basic Mechanisms in Animal Virus Biology Front Cover (Paperback): The scaffold/loop model can explain chromosome bands. This extended scaffold/loop model displays the unfolded, hypothetical chromatid fiber on the left; this fiber serves to explain the more compact metaphase chromosome on the right. (For details, see Saitoh and Laemmli, p. 757, this volume.) Back Cover (Paperback): Electron density of the mixed sequence nucleosome core particle. (For details, see Richmond et al., p. 268, this volume.) 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 for libraries and other users registered with the Copyright Clearance Center (CCC) Transactional Reporting Service, provided that the base fee of $5.00 per article is paid directly to CCC, 21 Congress St., Salem MA 01970. [0-87969-065-8/93 $5.00 + .00]. This consent does not extend to other kinds of copying, such as copying for general distribution, for advertising or promotional purposes, for creating new collective works, or for resale. All Cold Spring Harbor Laboratory Press publications may be ordered directly from Cold Spring Harbor Laboratory Press, 10 Skyline Drive, Plainview, New York 11803. Phone: 1-800-843-4388 in Continental U.S. and Canada. All other locations (516)349-1930. FAX: (516)349-1946. Symposium Participants AGUINAGA, PILAR, Comprehensive Sickle-Cell BERMAN, JUDITH, Dept. of Botany, University of Center, Meharry Medical College, Nashville, Minnesota, St. Paul Tennessee BERNARDI, GIORGIO, Dept. of Molecular Genet- ALBERTS, BRUCE, Dept. of Biochemistry and Bio- ics, Institut Jacques Monod, Paris, France physics, University of California, San Fran- BERRY, MEERA, Lab. of Gene Structure and Ex- cisco pression, National Institute for Medical Re- ALBERTSON, DONNA, Molecular Genetics Unit, search, The Ridgeway, London, United MRC Laboratory of Molecular Biology, Kingdom Cambridge, United Kingdom BIANCHI, ALLESSANDRO, Rockefeller University, ALFANO, CHRISTINE,Dept. of Biochemistry, Stan- New York, New York ford Medical Center, Stanford, California B1ESSMANN, HARALD, Developmental Biology ANAND, RAKESH, Zeneca, V.I.M., Macclesfield, Center, University of California, Irvine Cheshire, United Kingdom BIRD, ADRIAN, Institute of Cell and Molecular ANTEQUERA, FRANCISCO, Dept. of Cell and Molec- Biology, Medical Research Council, Univer- ular Biology, Edinburgh University, Scot- sity of Edinburgh, Scotland, United Kingdom land, United Kingdom BIRD, LOUISE, Dept. of Biochemistry, Oxford AZORIN, F., Investigacion y Desarrollo, Consejo University, United Kingdom Superior de lnvestigaciones Cientifica, Bar- BLACKBURN, ELIZABETH, Dept. of Microbiology celona, Spain and Immunology, University of California, BACCHETTI, SILVIA, Dept. of Pathology, McMas- San Francisco ter University, Hamilton, Ontario, Canada BLENNOW, ELIZABETH,Dept. of Clinical Genetics, BAJALICA, SVETLANA, Dept. of Clinical Genetics, Karolinska Hospital, Stockholm, Sweden Karolinska Institute, Stockholm, Sweden BONNER, WILLIAM, NCI, National Institutes of BAR, ISABELLE, Dept. of Physiology, Facult6 de Health, Bethesda, Maryland M6decin, FUNDP Medical School, Namur, BORST, PIET, Dept. of Molecular Biology, Nether- Belgium lands Cancer Institute, Amsterdam BAVYKIN, SERGEI, Dept. of Biological Chemistry, BOTCHAN, MICHAEL, Dept. Molecular and Cel- University of California, Davis lular Biology, University of California, Ber- BECK, ANITA, Dept. of Molecular Biotechnology, keley University of Washington, Seattle BOURBONNII~RE, MARTIN, Dept. of Neurology and BEDOYAN, JIRAIR, Biophysics Research Division, Neurosurgery, McGill University, Montreal, University of Michigan, Ann Arbor Quebec, Canada BEHN-KRAPPA, ANNETT, Dept. of Virology, Uni- BOYES, JOAN, Dept. of Molecular Biology, versity of Cologne, Germany NIDDK, National Institutes of Health, BELL, STEPHEN, Cold Spring Harbor Laboratory, Bethesda, Maryland Cold Spring Harbor, New York BRENNER, SYDNEY, Scripps Research Institute, La BELLON, STEVEN, Dept. of Molecular Biophysics Jolla, California and Dept. of Molecular Biol- and Biochemistry, Yale University, New ogy, Molecular Genetics Unit, MRC Labora- Haven, Connecticut tory, Cambridge, United Kingdom BELMAAZA, ABDELLAH, Dept. of Research and BRESNICK, EMERY, Dept. of Molecular Biology, Development, Canadian Red Cross Society, National Institutes of Health, Bethesda, Montreal, Quebec, Canada Maryland BENGAL, EYAL, Dept. of Molecular Biology and BREWER, BONITA, Dept. of Genetics, University Virology, Salk Institute for Biological of Washington, Seattle Studies, San Diego, California BROWN, KAREN, Dept. of Biochemistry, Oxford BERG, BARBARA, Dept. of Molecular Genetics University, United Kingdom and Cell Biology, University of Chicago, Il- BROWN,WILLIAM, Dept. of Biochemistry, Oxford linois University, United Kingdom V vi SYMPOSIUM PARTICIPANTS BUCAN, MAJA, Dept. of Psychiatry, University of COHEN, DANIEL, Centre d'Etude du Polymor- Pennsylvania, Philadelphia phisme Humain, Paris, France BURGOYNE, L.A., School of Biological Sciences, COUNTER, CHRIS, Dept. of Pathology and Bio- Flinders University of South Australia, chemistry, McMaster University, Hamilton, Adelaide Ontario, Canada BURLEY, STEPHEN, Dept. of Molecular Biophysics, COUNTRYMAN, JILL, Dept. of Molecular Bio- Rockefeller University, New York, New York physics and Biochemistry, Yale University, BUSTIN, M~CHAEL, Lab. of Molecular Car- New Haven, Connecticut cinogenesis, NCI, National Institutes of CRANE-ROBINSON, COLYN, Dept. of Biophysics, Health, Bethesda, Maryland University of Portsmouth, United Kingdom CARBON, JOHN, Dept. of Biological Sciences, Uni- CREMER, THOMAS, Dept. of Human Genetics, versity of California, Santa Barbara University of Heidelberg, Germany CARDOSO, MARIACRISTINA, Dept. of Cardiology, CROSS, SALLY,Dept. of Cell and Molecular Biolo- Children's Hospital, Boston, Massachusetts gy, University of Edinburgh, Scotland, CAREY, NESSA, Dept. of Anatomy, Charing Cross United Kingdom and Westminister Medical School, London, CROTHERS, DONALD, Dept. of Chemistry, Yale United Kingdom University, New Haven, Connecticut CARLSON, MARIAN, Dept. of Human Genetics, CUISSET, LAURENCE, Institut Cochin de Gen- Columbia University, New York, New York etique Moleculaire, Paris, France CARR, KIMBERLY,Nature, London, United King- DALRYMPLE, MICHAEL, Pharmaceutical Proteins, dom Ltd., Roslin, Scotland, United Kingdom CARTER, KENNETH, Dept. of Cell
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