COLD SPRING HARBOR SYMPOSIA ON QUANTITATIVE BIOLOGY VOLUME LV COLD SPRING HARBOR SYMPOSIA ON QUANTITATIVE BIOLOGY VOLUME LV The B rain COLD SPRING HARBOR LABORATORY PRESS 1990 COLD SPRING HARBOR SYMPOSIA ON QUANTITATIVE BIOLOGY VOLUME LV 9 1990 by The Cold Spring Harbor Laboratory Press International Standard Book Number 0-87969-059-3 (cloth) International Standard Book Number 0-87969-060-7 (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 XXVI (1961) Cellular Regulatory Mechanisms II (1934) Aspects of Growth XXVII (1962) Basic Mechanisms in Animal Virus Biology III (1935) Photochemical Reactions XXVII1 (1963) Synthesis and Structure of Macromolecules IV (1936) Excitation Phenomena XXIX (1964) Human Genetics V (1937) Internal Secretions XXX (1965) Sensory Receptors V1 (1938) Protein Chemistry XXXI (1966) The Genetic Code VII (1939) Biological Oxidations XXXI1 (1967) Antibodies VIII (1940) Permeability and ~he Nature of Cell Membranes XXXIII (1968) Replication of DNA in Microorganisms IX (1941) Genes and Chromosomes: Structure and Organi- XXXIV (1969) The Mechanism of Protein Synthesis zation XXXV (1970) Transcription of Genetic Material X (1942) The Relation of Hormones to Development XXXVI (1971) Structure and Function of Proteins at the XI (1946) Heredity and Variation in Microorganisms Three-dimensional Level XII (1947) Nucleic Acids and Nucleoproteins XXXVII (1972) The Mechanism of Muscle Contraction XIII (1948) Biological Applications of Tracer Elements XXXVII1 (1973) Chromosome Structure and Function XIV (1949) Amino Acids and Proteins XXXIX (1974) Tumor Viruses XV (1950) Origin and Evolution of Man XL (1975) The Synapse XVI (1951) Genes and Mutations XLI (1976) Origins of Lymphocyte Diversity XVII (1952) The Neuron XLII (1977) Chromatin XVIII (1953) Viruses XL[II (1978) DNA: Replication and Recombination XIX (1954) The Mammalian Fetus: Physiological Aspects of XLIV (1979) Viral Oncogenes Development XLV (1980) Movable Genetic Elements XX (1955) Population Genetics: The Nature and Causes of XLVI (1981) Organization of the Cytoplasm Genetic Variability in Population XLVII (1982) Structures of DNA XXI (1956) Genetic Mechanisms: Structure and Function XLVIll (1983) Molecular Neurobiology XXII (1957) Population Studies: Animal Ecology and De- XLIX (1984) Recombination at the DNA Level mography L (1985) Molecular Biology of Development XXII1 (1958) Exchange of Genetic Material: Mechanism and LI (1986) Molecular Biology of Homo sapiens Consequences LII (1987) Evolution of Catalytic Function XXIV (1959) Genetics and Twentieth Century Darwinism LIII (1988) Molecular Biology of Signal Transduction XXV (1960) Biological Clocks LIV (1989) Immunological Recognition 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 $1.00 per article is paid directly to CCC, 21 Congress St., Salem MA 01970. [0-87969-059-3/90 $1.00 + .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 New York (516)349-1930. FAX: (516)349-1946. Symposium Participants ABELIOVICH, ASA, Dept. of Biology, Massachu- BERG, HOWARD, Dept. of Cellular and Develop- setts Institute of Technology, Cambridge mental Biology, Harvard University, Cam- ADAMS, PAUL, Dept. of Neurobiology and Be- bridge havior, Howard Hughes Medical Institute, BHUSHAN, VIKAS, University of California School State University of New York, Stony Brook of Medicine, San Francisco ADEREM, ALAN, Dept. of Cellular Physiology and Brzzi, EMILIO, Dept. of Brain and Cognitive Sci- Immunology, Rockefeller University, New ences, Massachusetts of Technology, Cam- York, New York bridge AGOSTON, DENES, Laboratory of Cell Biology and BLAKEMORE, COLIN, Dept. of Physiology, Uni- National Institute of Mental Health, Nation- versity of Oxford, England al Institutes of Health, Bethesda, Maryland BLISS, TIMOTHY, Dept. of Neurophysiology and ALBERINI, CRISTINA, Instituto di Chimica, Faculty Neuropharmacology, National Institute for of Medicine, University of Brescia, Italy Medical Research, London, England ALDRICH, RICHARD,Dept. of Neurobiology, Stan- BONHOEFFER, FRIEDRICH,Abteilung Physikalische ford University School of Medicine, Cali- Biologie, Max-Planck Institut, Tubingen, fornia Federal Republic of Germany ALLMAN, JOHN, Dept. of Biology, California In- BONHOEFFER, TOBIAS, Neurobiology Laboratory, stitute of Technology, Pasadena Rockefeller University, New York, New ALTMAN, JENNIFER, Saunders Scientific Publica- York tions, London, England BOWER, JAMES, Dept. of Biology, California Insti- ANDERSEN, PER, Institute of Neurophysiology, tute of Technology, Pasadena University of Oslo, Norway BRANDWEIN, HARVEY, Pall Corporation, Glen ANDERSEN, RICHARD, Dept. of Brain and Cogni- Cove, New York tive Sciences, Massachusetts Institute of BREFORT, GEORCES, Research Directorate, Technology, Cambridge Rhone-Poulenc/Sante, Antony, France ANDERSON, DAVID, Dept. of Biology, California BucK, L1NDA, Howard Hughes Medical Institute, Institute of Technology, Pasadena Columbia University College of Physicians & AXLE, RICHARD, Institute of Cancer Research, Surgeons, New York, New York Columbia University College of Physicians & BUELL, GARY, Pharmacia Genetics, La Jolla, Surgeons, New York, New York California BADING, HILMAR, Dept. of Neurobiology and BURROUS, MARY, Dept. of Neurobiology, Rocke- Molecular Genetics, Harvard Medical feller University, New York, New York School, Boston, Massachusetts BuscH, CHRISTOPI~ER, Dept. of Medical Re- BARABAN, JAY, Dept. of Neur0science, Johns search, Max-Planck Institut, Heidelberg, Hopkins University School of Medicine, Bal- Federal Republic of Germany timore, Maryland BUTMAN, JOHN, Dept. of Biology, Washington BARGMANN, COR1, Dept. of Biology, Massachu- University, St. Louis, Missouri setts Institute of Technology, Cambridge BYRNE, JOHN, Dept. of Neurobiology and BARINAGA, MARCIA, Science, Berkeley, California Anatomy, University of Texas Medical BAYLOR, DENIS, Dept. of Neurobiology, Stanford School, Houston University School of Medicine, California CADD, GARY, Dept. of Pharmacology, Howard BEGEMANN, MARTIN, Dept. of Molecular and De- Hughes Medical Institute, University of velopmental B~ology, Rockefeller Universi- Washington, Seattle ty, New York, New York CAPORALE, LYNN, Dept. of Industrial and Aca- BEKKERS, JOHN, Dept. of Molecular Neurobiol- demic Relations, Merck Sharp & Dohme Re- ogy, Sulk Institute, La Jolla, San Diego, search Laboratories, West Point, Pennsyl- California vania BENZER, SEYMOUR, Dept. of Biology, California CARDOZO, DAVID, Dept. of Neurobiology, Har- Institute of Technology, Pasadena vard Medical School, Boston, Massachusetts vi SYMPOSIUM PARTICIPANTS CATE, JOHN, Depts. of Pathology and Laboratory National Institute of Mental Health, Bethes- Medicine, Medical University of South da, Maryland Carolina, Charleston DIAMOND, JEFF, Dept. of Ophthalmology, Uni- CEPKO, CONSTANCE, Dept. of Genetics, Harvard versity of California, San Francisco Medical School, Boston, Massachusetts DOWMAN, JOHN, Dept. of Neurosciences, Medical CHANDA, PRANAB, Depts. of Microbiology and Research Council, London, England Biotechnology, Wyeth-Ayerst Laboratories, DUFFY, CHARLES, Laboratory for Sensorimotor Philadelphia, Pennsylvania Research, National Eye Institute, National CHANGEUX, JEAN-PIERRE, Dept. of Molecular Institutes of Health, Bethesda, Maryland Neurobiology, Institut Pasteur, Paris, France DUHAMEL, JEAN-RENE, National Eye Institute, CHAVEZ-NORIEGA, LAURA, Salk Institute, La National Institutes of Health, Bethesda, JoUa, San Diego, California Maryland CHEN, JOSEPH, Depts. of Physiology and Bio- EDELMAN, GERALD, Neurosciences Research Pro- physics, New York University Medical Cen- gram, Rockefeller University Neurosciences ter, New York Institute, New York, New York CHU, HUNG-MING, Neuroscience Program, Har- ERKMAN, LINDA, Dept. of Biochemistry, Sciences vard University, Boston, Massachusetts II, The University, Geneva, Switzerland CHUN, JEROLD, Whitehead Institute, Cambridge, ESSEN, LARS-OLIVER, Institute of Neurobiology, Massachusetts University of Heidelberg, Federal Republic CLOTHIAUX, EUGENE, Center of Neuroscience and of Germany Physics, Brown University, Providence, FAGG, GRAHAM, Ciba-Geigy Ltd., Basel, Swit- Rhode Island zerland COHEN-CORY, SUSANA, Dept. of Neurobiology, FEKETE, DONNA, Dept. of Genetics, Harvard Rockefeller University, New York, New Medical School, Boston, Massachusetts York FINGER, WOLFGANG, Universitat Tubingen, Fed- COLBY, CAROL, Laboratory for Sensorimotor Re- eral Republic of Germany search, National Eye Institute, National In- FISCHBACH, GERALD, Depts. of Anatomy and stitutes of Health, Bethesda, Maryland Neurobiology, Washington University CONSTANTINE-PATON, MARTHA, Neuroscience School