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COLD SPRING HARBOR SYMPOSIA ON QUANTITATIVE BIOLOGY VOLUME XXVI COLD SPRING HARBOR SYMPOSIA ON QUANTITATIVE BIOLOGY Founded in 1933 by REGINALD G. HARRIS Director o/ the Biological Laboratory 1924 to 1936 The Symposia were organized and managed by Dr. Harris until his death. Their continued use- /ulness is a tribute to the soundness o/ his vision. The Symposium volumes are published by the Long Island Biological Association as a part of the work of the Biological Laboratory Cold Spring Harbor, L.I., New York COLD SPRING HARBOR SYMPOSIA ON QUANTITATIVE BIOLOGY VOLUME XXVI Cellular Regulatory Mechanisms THE BIOLOGICAL LABORATORY COLD SPRING HARBOR, L.I., NEW YORK 1961 COPYRIGHT 1962 BY THE BIOLOGICAL LABORATORY LONG ISLAND BIOLOGICAL ASSOCIATION, INC. Library of Congress Catalog Number: 34-8174 All rights reserved. This book may not be reproduced in whole or in part except by reviewers for the public press without written permission from the publisher PRINTED BY WAVERLY PRESS, INC., BALTIMORE,MARYLAND, U.S.A. FOREWORD During the past twenty years, research in genetics and biochemistry, while superficially directed toward an understanding of different aspects of cell behavior, has been moving to a union which appears to have been completed in the presen- tations at this symposium on Cell Regulatory Mechanisms. Sessions were devoted to the role of the genetic material as an information code in protein synthesis and the mechanism of information transfer from DNA to the site of synthesis. Major attention was devoted to cellular mechanisms governing the rates of enzyme ac- tivity and enzyme synthesis. While most of the research presented dealt with microorganisms, the significance of these efforts for questions of normal and ab- normal growth and development in multicellular forms is quite apparent. I wish to express our appreciation to Bernard D. Davis, Eugene W. Knox, Jacques Monod, Van R. Potter, Gordon M. Tomkins, and H. Edwin Umbarger, whose generous and thoughtful advice provided the basis for organization of this program, and to H. A. Itano, Erwin Chargaff, A. D. Hershey, Elliot Volkin, Fritz Lipmann, Joseph F. Gots, Edward L. Tatum, H. O. Halvorson, Efraim Racker, Van R. Potter, Claude Villee, Eugene W. Knox, and Leo Szilard, who served as chairmen of the various sessions. The meetings this year were held from the 4th through the 12th of June, and were attended by approximately 200 participants, including 25 from abroad. In addition to the support of the Long Island Biological Association, I am pleased to acknowledge the support of this program by The National Institutes of Health, U.S. Public Health Service, National Science Foundation, Rockefeller Foundation, United States Atomic Energy Commission, and The United States Air Force under Grant AF-AFOSR-61-73; monitored by The Air Force Office of Scientific Research of the Air Research and Development Command. This year our editor was assisted in the preparation of the volume by Ruth Schalet, Eleanor Chovnick and Hildegarde Mukai. We are grateful to them and to the staff members of the Biological Laboratory who assisted in the difficult task of indexing the volume. Arthur Chovnick, Laboratory Director Editor: Leonora Friseh LIST OF PREVIOUS VOLUMES Volume I (1933) Surface Phenomena, 239 pp. Volume II (19341 Aspects of Growth, 284 pp. Volume III (1935) Photochemical Reactions, 359 pp. Volume IV (1936) Excitation Phenomena, 376 pp. Volume V (1937) Internal Secretions, 433 pp. Volume VI (1938) Protein Chemistry, 395 pp. Volume VII (1939) Biological Oxidations, 463 pp. Volume VIII (1940) Permeability and the Nature of Cell Membranes, 285 pp. Volume IX (1941) Genes and Chromosomes: Structure and Organization, 315 pp. Volume X (1942) The Relation of Hormones to Development, 167 pp. Volume XI (1946) Heredity and Variation in Microorganisms, 314 pp. Volume XII (1947) Nucleic Acids and Nucleoproteins, 279 pp. Volume XIII (1948) Biological Applications of Tracer Elements, 222 pp. Volume XIV (1949) Amino Acids and Proteins, 217 pp. Volume XV (1950) Origin and Evolution of Man, 425 pp. Volume XVI (1951) Genes and Mutations, 521 pp. Volume XVII (1952) The Neuron, 323 pp. Volume XVIII (1953) Viruses, 301 pp. Volume XIX (1954) The Mammalian Fetus: Physiological Aspects of Development, 225 pp. Volume XX (1955) Population Genetics: The Nature and Causes of Genetic Varia- bility in Populations, 346 pp. Volume XXI (1956) Genetic Mechanisms: Structure and Function, 392 pp. Volume XXII (1957) Population Studies: Animal Ecology and Demography, 437 pp. Volume XXIII (1958) Exchange of Genetic Material: Mechanisms and Consequences, 450 pp. Volume XXIV (1959) Genetics and Twentieth Century Darwinism, 321 pp. Volume XXV (1960) Biological Clocks, 524 pp. SOME SYMPOSIUM PARTICIPANTS Top row: R. Quayle, H. Holzer, H. Kornberg, H. E. Umbarger; M. Schaechter, F. Neidhardt; T. F. Anderson, S. Brenner, J. Watson, A. D. Hershey, F. Jacob, F. Stahl. Second row: A. Koch, V. Potter; H. Kalckar, 0. MaalCe, Miss Maal~e, F. Lipmann; B. Nisman, S. Spiegelman. Third row: G. Streisinger, S. Benzer; F. Jacob, L. Szilard; D. Perrin, J. Changeux, F. Jacob. Bottom row: A. Novick, P. Margolin; C. Yanofsky, N. Franklin, J. D. Watson, N. H. Horowitz, F. J. de Serres; T. F. Anderson, J. Monod, A. Garen. SOME SYMPOSIUM PARTICIPANTS Top row: H. Halvorson, B. D. Davis, J. Hurwitz, S. Luria; E. Englesberg, B. P. Kaufmann; M. Yarmolinsky, G. Buttin. Second row: A. Chovnick, E. Glassman; H. Vogel, R. Vogel, L. Gorini; J. D. Watson, E. Lennox, L. Szilard, H. Eagle. Third row: D. Shemin, J. Waller, G. Cohen; M. Meselson, S. Benzer; E. Freese, B. Strauss, M. Demeree. Bottom row: B. Magasanik, W. Maas, M. Umbarger, A. Magasanik; I. Zabin, G. Tomkins; E. Racker, B. Chance. (Photographs by Robert P. Beaumont and Ilse Schwinck) List of Those Attending the Symposium ADLER, J., Department of Biochemistry, University of CHAMPE, SEWELL, Department of Biological Sciences, Wisconsin, Madison, Wisconsin Purdue University, W. Lafayette, Indiana AKEY,ALICE, Adelphi College, Garden City, New York CHANCE, BRITTON, Johnson Foundation, University of ANDERSON, RICHMOND I~., Rockefeller Foundation, Pennsylvania, Philadelphia, Pennsylvania New York City CtL4NGEUX, JEAN-PIERRE, Service de Bioehimie Cellu- ANDERSON,T. F., Institute for Cancer Research, Phila- laire, Institut Pasteur, Paris, France delphia, Pennsylvania CHARGAFF, ERWIN, Columbia University, New York BAKEMEIER,RICHARD, Dept of Medicine, University of City Rochester, Rochester, N. Y. CHARGAFF, ERWIN, MRS., c/o Dr. Erwin Chargaff BENZER, SEYMOUR,Department of Biological Sciences, CHOVNICK, ARTHUR, Long Island Biological Associa- Purdue University, Lafayette, Indiana tion, Cold Spring Harbor, New York BERNHEIMER, HARRIET, State University of New COHEN, GEORGES, Centre National de la Recherche York, Downstate Medical Center, Brooklyn, New Scientifique, Gif-sur-Yvette, France York COHEN, GEORGES,MRS., C/O Dr. Georges Cohen BLUME, DORIS, Long Island Biological Association, COLEMAN, DOUGLAS L., R. B. Jackson Laboratories, Cold Spring Harbor, New York Bar Harbor, Maine BOYER, SAMUEL H., Department of Medicine, Johns COLLAZO,ANA M. DIAL, Department of Biology, Uni- Hopkins Hospital, Baltimore, Maryland versity of Puerto Rico, Rio Piedras, Puerto Rico BRENNER, SYDNEY,M. R. C. Unit for Molecular Biol- COWIE, DEAN B., Department of Terrestrial Magnet- ogy, Cavendish Laboratory, Cambridge, England ism, Carnegie Institution of Washington, Wash- BRESCH, CARSTEN,Abt. fiir Mikrobiologie, Botanisehes ington, D. C. Institut, Universitat KSln, KSln-Lindenthal, DAs, C. C., Department of Genetics, Carnegie Institu- Germany tion of Washington, Cold Spring Harbor, New BRESCH, CARSTEN,MRS., e/o Dr. Carsten Breseh York BRITTEN, ROY J., Department of Terrestrial Magnet- DAVIS, BERNARD D., Department of Bacteriology & ism, Carnegie Institution of Washington, Wash- Immunology, Harvard Medical School, Boston, ington, D. C. Massachusetts BROWN, DONALDD., Department of Embryology, Car- DEMEREC, M., Brookhaven National Laboratory, Up- negie Institution of Washington, Baltimore, Mary- ton, New York land DINTZIS, HOWARD M., Biology Department, Massa- BURGI, ELIZARETtt,Department of Genetics, Carnegie chusetts Institute of Technology, Cambridge, Institution of Washington, Cold Spring Harbor, Massachusetts New York DOI, RoY HIROSHI, Department of Microbiology, Uni- BURR, W. W., JR., Biology Branch, Division of Biology versity of Illinois, Urbana, Illinois & Medicine, U. S. Atomic Energy Commission, DREYER, WILLIAM J., National Heart Institute, Be- Washington, D. C. thesda, Maryland BUTTIN, GERARD, Service de Biochimie Cellulaire, In- EAGLE, HARRY, National Institutes of Health, Be- stitut Pasteur, Paris, France thesda, Maryland CAMPBELL, A., Department of Biology, University of EAGLE, HARRY,MRS., C/O Dr. Harry Eagle Rochester, Rochester, New York ECHOES, HARRISON H., Department of Biochemistry, CAMPBELL,A., MRS., C/O Dr. A. Campbell University of Wisconsin, Madison, Wisconsin CANFIELD, ROBERT E., National Heart Institute, Na- tional Institutes of Health, Bethesda, Maryland ECKER, RICHARD E., Department of Microbiology, CAR~ART, ANNE, Department of Genetics, Carnegie College of Medicine, University of Florida, Gaines- Institution of Washington, Cold Spring Harbor, ville, Florida New York EHRENSTEIN, GUNTHER VON, Rockefeller Institute, CARLSON, ELOF A., Department of Zoology, University New York City of California, Los Angeles, California ENGLESBERG, ELLIS, Department of Biological Sci- CASPARI, ERNST, Department of Biology,