COLD ON

SPRING HARBOR SYMPOSIA QUANTITATIVE BIOLOGY

VOLUME XVIII

COLD S P R I N G H A R B O R S Y M P O S I A

O N Q U A N T I T A T I V E B I O L O G Y

Founded in 1933

by

REGINALD G. HARRIS Director of the Biological Laboratory

1924 to 1936

The Symposia were organized and managed by Dr. Harris until his death. Their continued use- fulness is a tribute to the soundness of 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 XVIII

VIRUSES

THE BIOLOGICAL LABORATORY COLD SPRING HARBOR, L.I., NEW YORK

1953

COPYRIGHT 1953 BY

THE BIOLOGICAL LABORATORY

LONG ISLAND BIOLOGICAL ASSOCIATION, INC.

All rights reserved. This book may not be reproduced in whole or part except by reviewers for the public press without written permission from the publisher.

PRINTED BY

MONUMENTAL PRINTING COMPANY, BALTIMORE, MARYLAND, U.S.A.

F O R E W O R D

The first paper on the subject of viruses to be included in the Cold Spring Harbor Sym- posia was given at the sixth Symposium, in 1938, by W. M. Stanley and H. S. Loring. Since that time, papers on this topic have been presented at several other sessions, where the main problems under consideration related to genetic mechanisms. This year's Sympo- sium was the first to be devoted entirely to discussion of the fundamental aspects of virus research.

The preliminary plans for this meeting were in progress for several years. The Cold Spring Harbor laboratories have been actively interested in research with viruses ever since 1941, when the T series of phages was used in studies of mutation at the Department of Ge- netics of the Carnegie Institution of Washing- ton. This interest was considerably stimulated when, in the summer of 1945, Max Delbriick organized a course for research workers, on quantitative methods used in research with bacterial viruses. This course has been given at the Biological Laboratory every summer since then, first by Delbriick, then by Mark H. Adams, and in 1952 by A. H. Doermann while Adams was in Europe. Further stimu- lation was provided by the presence of phage workers at the Laboratory during the sum- mers, by several phage conferences held here on Delbr/ick's initiative, and by the addition of A. D. Hershey to the research staff of the Department of Genetics.

Thus the ground was well prepared for a session on viruses. The meeting was originally scheduled for the summer of 1952, but was postponed for a year on account of the phage symposium at Abbaye Royaumont.

This Symposium on Viruses was organized by Max Delbriick, in consultation with Frank L. HorsfalI, Jr., and H. M. Weaver. Its pri- mary purpose was to bring together research workers in this rapidly developing field for discussion of basic problems, and to provide an opportunity for those specializing in bac- terial, animal, and plant viruses to correlate their findings.

[v]

The established procedure of our Symposia has been to schedule two or three invited pa- pers per day and to allow ample time for informal and spontaneous discussion. The scheme was somewhat modified this year. In addition to the twenty authors of scheduled papers, a number of other participants were invited to give prepared discussions, con- cerned primarily with their own research. These discussions are published here as sepa- rate articles.

The last three Symposium volumes were printed by the photo-offset process. This meth. od was used in the hope that publication might be speeded up and also that the cost of pro- duction--and thus the price of the books-- might be decreased. The first and most im- portant of these anticipations was not realized, however; and since, in addition, the quality of the printing was considered less satisfac- tory than in previous volumes, we have re- turned, in this volume, to the typesetting meth- od. We have continued our efforts to com- plete the publication of the proceedings as soon as possible after the Symposium meeting. As part Of this effort, the printer's proofs of all articles in this volume were handled in our editorial office, and no proofs were sent to the authors. In addition to Dr. Katherine Brehme Warren, who served as editor, Dr. Mark H. Adams checked over the proofs.

The Symposium was held from June 5th to June l l th , 1953, and was attended by two hundred and seventy-two scientists. The new lecture hall of the Cold Spring Harbor labora- tories was used for the meetings.

The National Foundation for Infantile Pa- ralysis paid the expenses of invited partici- pants, as well as tile expenses incurred in organizing the meeting; and I wish to acknowl- edge this support with grateful appreciation. I wish also to express appreciation for the continued support of the Carnegie Corpora- tion of New York, whose grant for this year has been used to promote the early publica. tion of this volume.

M. DEMEREC

L I S T O F P R E V I O U S V O L U M E S

Volume Volume Volume Volume Volume V~lume Volume Volume Volume Volume Volume Volume Volume Volume Volume Volume Volume

I (1933) Surface Phenomena, 239 pp. II (1934) Aspects of Growth, 284 pp. III (1935) Photochemical Reactions, 359 pp. IV (1936) Excitation Phenomena, 376 pp. V (1937) Internal Secretions, 433 pp. VI (1938) Protein Chemistry, 395 pp. VII (1939) Biological Oxidations, 463 pp. VIII (1940) Permeability and the Nature of Cell Membranes, 285 pp. IX (1941) Genes and Chromosomes: Structure and Organization, 315 pp. X (1942) The Relation of Hormones to Development, 167 pp. XI (1946) Heredity and Variation in Microorganisms, 314 pp. XII (1947) Nucleic Acids and Nucleoproteins, 279 pp. XIII (1948) Biological Applications of Tracer Elements, 222 pp. XIV (1949) Amino Acids and Proteins, 217 pp. XV (1950) Origin and Evolution of Man, 425 pp. XVI (1951) Genes and Mutations, 521 pp. XVII (1952) The Neuron, 323 pp.

[vi]

SOME SYMPOSIUM PARTICIPANTS

Top row: M. Chase, A. D. Hershey; F. Jacob, A. Lwoff, L. Siminovitch; F. K. Sanders, E. L. Tatum. S e c o n d row: C. Levinthal, S. E. Luria; W. Hayes, N. D. Zinder; S. Zamenhof, J. D. Watson. T h i r d row: W. M. Stanley, D. Beard, J. W. Beard; M. Demerec, A. Lwoff; J. C. Appleby, R. Weindling. F o u r t h row: B. McClintock, E. M. Witkin, S. Mudd; S. E. Luria, H. D. Hershey; M. H. Adams, W. Weidel, K. G. Lark.

(The pictures were taken by Dr. K. Maramorosch and Dr. K. Brehme Warren.)

SOME SYMPOSIUM PARTICIPANTS

Top row: R. Markham, G. R. Wyatt; L. Szilard, M. Chase, A. H. Doermann; R. D. Hotchkiss, M. Delbriick. Second row: K. B. Warren, C. O. Warren, W. M. Stanley, J. W. Beard, K. Maramorosch; E. S. Anderson, T. F. Ander.

son; N. Visconti, Mrs. M. Delbr/ick, M. Demerec, M. Delbriick. Thirdrow: A. Sabin and discussant; R. C. Williams, D. Beard, T. F. Anderson; G. S. Stent, A. I-I. Doermann. Fourth row: R. Markham, K. Maramorosch; J. W. Beard, F. K. Sanders; W. Hayes, G. Bertani.

LIST OF THOSE ATTENDING OR P A R T I C I P A T I N G IN THE SYMPOSIUM

ABELSON, PHILIP H., Carnegie Institution of Washington, Washington, D. C. ACKERMANN, W. WILBUR, University of Michigan, Ann Arbor, Michigan ADAMS, MARK H., New York University College of Medicine, New York ALEXANDER, HATTIE E., Babies Hospital, New York ALLEN, EMMA G., Veterans Administration Hospital, Brooklyn, New York ALLEN, PETER, Columbia University, New York AMBELLAN, ELISABETH, Columbia University, New York ANDERSON, E. S., Central Enteric Reference Laboratory, London, England ANDERSON, THOMAS F., University of Pennsylvania Hospital, Philadelphia, Pennsyl-

vania APPLEBY, J. CHRISTINE, Rowett Research Institute, Aberdeenshire, Scotland APPLEYARD, RAYMOND K., California Institute of Technology, Pasadena, California BARKSDALE, LANE, New York University, New York BARLOW, JAMES, The Johns Hopkins University, Baltimore, Maryland BARON, L. S., Walter Reed Army Medical Center, Washington, D. C. BARRY, GuY T., Rockefeller Institute for Medical Research, New York BAYLOR, MARTHA BARNES, University of Michigan, Ann Arbor, Michigan BEARD, JOSEPH W., Duke University, Durham, North Carolina BEARD, MRS. JOSEPH W., Duke University, Durham, North Carolina BEISER, SAM M., Rockefeller Institute for Medical Research, New York BENZER, Seymour, Purdue University, Lafayette, Indiana BEBGOLD, G. H., Laboratory of Insect Pathology, Sault Ste. Marie, Ontario, Canada BERNSTEIN, MAURICE, Carnegie Institution of Washington, Cold Spring Harbor, New

York BERRIE, ALEX, Carnegie Institution of Washington, Cold Spring Harbor, New York BERTANI, G., University of Illinois, Urbana, Illinois BEVAN, E. A., Dartmouth College, Hanover, New Hampshire BINN, LEONARD, University of Michigan, Ann Arbor, Michigan BLACK, FRANK L., Rockefeller Institute for Medical Research, New York BLOMSTRANDT, INGBRITT, Carnegie Institution of Washington, Cold Spring Harbor,

New York BODIA~, DAVID, The Johns Hopkins University, Baltimore, Maryland BOGORAD, LAWRENCE, Rockefeller Institute for Medical Research, New York BOLIN, VERN S., Jensen-Salsbery Laboratories, Inc., Kansas City 10, Missouri BOLTON, ELLIS, Carnegie Institution of Washington, Washington, D. C. BONNER, DAVID, Yale University, New Haven, Connecticut BOREK, BLANCHE, Veterans Administration Hospital, Brooklyn, New York BOREK, ERNEST, Veterans Administration Hospital, Brooklyn, New York BOVARNICK, MARIANNA R., Veterans Administration Hospital, Brooklyn, New York BOWEN, GEORGE H., Oak Ridge National Laboratory, Oak Ridge, Tennessee BOLD, THEODORE, National Foundation for Infantile Paralysis, New York BRAUN, WERNER, Camp Detrick, Frederick, Maryland BRIODY, B. A., Hahnemann Medical College, Philadelphia, Pennsylvania BRITTEN, ROY, Carnegie Institution of Washington, Washington, D. C. BROCUM, J. C., Amherst College, Amherst, Massachusetts BROWN, ARTHUR, State University of New York, Brooklyn, New York BROWN, GEORGE B., Sloan-Kettering Institute for Cancer Research, New York

[ix]

x LIST OF PARTICIPANTS

BRYSON, VERNON, The Biological Laboratory, Cold Spring Harbor, New York BURNET, F. M., The Walter and Eliza Hall Institute, Melbourne, Australia BURGI, ELIZABETH, Cornell University, Ithaca, New York BURTON, KENNETH, University of Chicago, Chicago, Illinois BUSH, MILTON, T., Vanderbilt University, Nashville, Tennessee CHADWICK, CONSTANCE, Carnegie Institute of Washington, Cold Spring Harbor, New

York CHASE, MARTHA, Oak Ridge National Laboratory, Oak Ridge, Tennessee CLARKE, DELPmNE, Rockefeller Institute for Medical Research, New York CODZA, HELEN, Beth Israel Hospital, New York COHEN, SEYMOUR S., Children's Hospital of Philadelphia, Philadelphia, Pennsylvania COLOWICK, MARYDA SWANSTROM, Baltimore, Maryland COMMONER, BARRY, Washington University, St. Louis, Missouri COWEN, NAOMI, Yale University, New Haven, Connecticut COWIE, DEAN B., Carnegie Institute of Washington, Washington, D. C. COWLES, PHILIP B., Yale University, New Haven, Connecticut CROrUTT, L. E., Indiana University, Bloomington, Indiana DAS, NERMAL, Carnegie Institution of Washington, Cold Spring Harbor, New York DAVIDSON, HARRIET, The Biological Laboratory, Cold Spring Harbor, New York DAVIS, BERNARD D., Cornell University, Medical College, New York DEICHES, HELEN, The Biological Laboratory, Cold Spring Harbor, New York DELAMATER, EDWARD D., University of Pennsylvania, Philadelphia, Pennsylvania DELBRi]CK, MAX, California Institute of Technology, Pasadena, California DEMEREC, M., Carnegie Institution of Washington, and The Biological Laboratory,

Cold Spring Harbor, New York DEMEREC, ZLATA, Carnegie Institution of Washington, Cold Spring Harbor, New York DIXON, JUNE, Carnegie Institution of Washington, Cold Spring Harbor, New York DOERMANN, A. H., Oak Ridge National Laboratory, Oak Ridge, Tennessee DOTY, PAUL, Harvard University, Cambridge, Massachusetts DULBECCO, R., California Institute of Technology, Pasadena, California DUNNE, HOWARD W., Camp Detrick, Frederick, Maryland EDELMAN, ABRAHAM, Huntington, New York EHRLICH, HENRY L., Rensselaer Polytechnic Institute, Troy, New York EISENSTARK, A., Kansas State College, Manhattan, Kansas ELLISON, SOLON A., Columbia University, New York EPSTEIN, HERMAN T., University of Pittsburgh, Pittsburgh, Pennsylvania EPSTEIN, RICHARD, University of Rochester, Rochester, N. Y. ESPANA, CARLOS, E. R. Squibb and Sons, New Brunswick, New Jersey EVANS, AUDREY, Rockefeller Institute for Medical Research, New York EVERSOLE, ILSE, Stanford University, Stanford, California EVERSOLE, RUSSELL, Stanford University, Stanford, California FENNER, FRANK, The Australian National University, Canberra, Australia FISHER, H. W., University of Michigan, Ann Arbor, Michigan FLANAGAN, ARTHUR D.; Lederle Laboratories, Pearl River, New York FLUKE, DONALD, Brookhaven National Laboratory, Upton, New York FODOR, ANDREW, New York University, New York FORRO, FREDERICK, Brookhaven National Laboratory, Upton, New York Fox, MAURICE S., University of Chicago, Chicago, Illinois FRANKEL, J. W., New York City FRANKLIN, RICHARD M., Yale University, New Haven, Connecticut

LIST OF PARTICIPANTS xi

FRASER, DEAN, University of California, Berkeley, California FRASER, DOROTHY KEPPEL, California Institute of Technology, Pasadena, California FRENCH, R. C., University of Toronto, Toronto, Canada FRIEDMAN, MURRAY, Yale University, New Haven, Connecticut GAJDUSEK, W. CARLETON, Walter Reed Army Medical Center, Washington, D. C. GAREN, ALLEN, Carnegie Institution of Washington, Cold Spring Harbor, New York GAYLORD, WILLIAM H., Yale University, New Haven, Connecticut GERONIMUS, LIP PMAN, The Biological Laboratory, Cold Spring Harbor, New York GIBBS, MARTIN, Brookhaven National Laboratory, Upton, New York GILVARG, CHARLES, Tuberculosis Research Laboratory, U. S. Public Health Service,

New York GITTERMAN, CHARLES, Merck & Co., Inc., Rahway, New Jersey GITTERMAN, SELMA, Rutgers University, New Brunswick, New Jersey GOEBEL, WALTHER F., Rockefeller Institute for Medical Research, New York GONZALEY, ELSA L., Columbia University, New York GOODGAL, SOL H., The Johns Hopkins University, Baltimore, Maryland GOTLIEB, TAMAR, Public Health Research Institute, New York COTS, JOSEPH S., University of Pennsylvania, Philadelphia, Pennsylvania GRAFF, SAMUEL, Columbia University, New York GRAHAM, A. F., University of Toronto, Toronto, Canada GRANOFF, ALLAN, Public Health Research Institute, New York GRAY, ALAN, Children's Hospital, Philadelphia, Pennsylvania GREEN, HOWARD, University of Chicago, Chicago, Illinois GROMAN, NEAL B., University of Washington, Seattle, Washington GROSS, S. R., Columbia University, New York GROUPE, VINCENT, Rutgers University, New Brunswick, New Jersey HAFF, RICHARD F., Western Reserve University, Cleveland, Ohio HARTMAN, PHILIP E., University of Pennsylvania, Philadelphia, Pennsylvania HAYES, WILLIAM, Postgraduate Medical School of London, London, England HEMMERLEY, JEAN, Carnegie Institution of Washington, Cold Spring Harbor, New

York HENLE, WERNER, Children's Hospital, Philadelphia, Pennsylvania HERRIOTT, ROGER M., The Johns Hopkins University, Baltimore, Maryland HERSHEY, A. D., Carnegie Institution of Washington, Cold Spring Harbor, New York HIRST, GEORGE K., Public Health Research Institute, New York HORSFALL, FRANK L., Hospital of the Rockefeller Institute, New York HOTCHIN, JOHN E., The Johns Hopkins University, Baltimore, Maryland HOTCHKISS, ROLLIN D., Rockefeller Institute for Medical Research, New York HUGHES, INGE n., University of Pennsylvania, Philadelphia, Pennsylvania HUNTER, MARY E., University of Pennsylvania, Philadelphia, Pennsylvania JACOB, FRANCOIS, Institut Pasteur, Paris, France JACOB, MIRIAM, New York University, New York JAKOBY, WILLIAM B., Yale University, New Haven, Connecticut JESAITIS, M. A., Rockefeller Institute for Medical Research, New York KAISER, DALE, California Institute of Technology, Pasadena, California KAPLAN, Louis, Sloan-Kettering Institute for Cancer Research, New York KARR, HELEN V., Public Health Research Institute, New York KAUFMANN, B. P., Carnegie Institution of Washington, Cold Spring Harbor, New York KEMP:F, J. E., University of Illinois, Chicago, Illinois KEYNAN, A., Columbia University, New York

xii LIST OF PARTICIPANT5

KLEIN, MORTON, Temple University, Philadelphia, Pennsylvania KLEINSCHMIDT, W. J., Eli Lilly and Co., Indianapolis, Indiana KOPPELMAN, RAY, University of Chicago, Chicago, Illinois KOPROWSKI, H.. Lederle Laboratories, Pearl River, New York KOZLOFF, LLOYD M., University of Chicago, Chicago, Illinois KRAUSS, MARJORIE R., New York University, New York KREGZDE, J., Jewish Sanatorium and Hospital for Chronic Diseases, Brooklyn, New

York KUHN, DAISY A., University of Pennsylvania, Philadelphia, Pennsylvania LABRUM, EDGAR L., Carnegie Institution of Washington, Cold Spring Harbor, New

York LANE, DOROTHY, Rockefeller Institute for Medical Research, New York LANNI, FRANK, University of Illinois, Urbana, Illinois LANNI, YVONNE THERY, University of Illinois, Urbana, Illinois LANNING, MABY, Children's Hospital, Philadelphia, Pennsylvania LARK, KARL GORDON, New York University, New York LENNOX, EDWIN S., University of Michigan, Ann Arbor, Michigan LESTER, GABRIEL, Chas. Pfizer & Co., Brooklyn, New York LEVINTHAL, CYRUS, University of Michigan, Ann Arbor, Michigan LIED, MARGARET, California Institute of Technology, Pasadena, California LIEF, FLORENCE, Children's Hospital, Philadelphia, Pennsylvania LIND, PATRICIA E., The Walter and Eliza Hall Institute, Melbourne, Australia LIu, OSCAR C., Children's Hospital, Philadelphia, Pennsylvania LIVELY, ETHELYN, University of Illinois, Urbana, Illinois LONDON, MORRIS, Columbia University, New York LURIA, S. E., University of Illinois, Urbana, Illinois LWOFF, ANDRE, Institut Pasteur, Paris, France MACDOWELL, E. C., Carnegie Institution of Washington, Cold Spring Harbor, New

York MACKAL, ROY P., University of Chicago, Chicago, Illinois MACLENI'qAN, JOI-IN D., Columbia University, New York MANDEL, BENJAMIN, Public Health Research Institute, New York MANDELL, JOSEPH D., California Institute of Technology, Pasadena, California MANGUN, GEORGE H., Warner Institute for Therapeutic Research, New York MANSON, LIONEL A., Western Reserve University, Cleveland, Ohio MARAMOROSCH, KARL, Rockefeller Institute for Medical Research, New York MARKHAM, ROY, Cambridge Uniyersity, Cambridge, England MARMUR, J., Rockefeller Institute for Medical Research, New York MCCLAIN, MARY E., University of California, Berkeley, California McCLELLAND, LAURELLA, Merck Institute for Therapeutic Research, Rahway, New

Jersey McCLINTOCK, BARBARA, Carnegie Institution of Washington, Cold Spring Harbor,

New York MCDONALD, MARGARET, Carnegie Institution of Washington, Cold Spring Harbor,

New York McHuGII, ROSEMARY, Francis Delafield Hospital, New York MEAD, MRS. L. M., Huntington, New York MEDILL, MARY, University of Pennsylvania, Philadelphia, Pennsylvania MELNICK, JOSEPH L., Yale University, New Haven, Connecticut MILLER, C. ARDEN, University of Kansas, Kansas City, Kansas

LIST OF PARTICIPANTS xiii

MITSUIIASI-II, S., Tuberculosis Research Laboratory, U. S. Public Health Service, New York

MOORE, A., Sloan-Kettering Institute for Cancer Research, New York MOSER, HERMANN, Carnegie Institution of Washington, Cold Spring Harbor, New

York MUDD, STUART, University of Pennsylvania, Philadelphia, Pennsylvania MUNK, KLAUS, Max Planck Institut, Tfibingen, Germany NAYLOR, H. B., Cornell University, Ithaca, New York NOVlCK, AAaON, University of Chicago, Chicago, Illinois OSTER, GERALD, Polytechnic Institute of Brooklyn, Brooklyn, New York PAEGE, LYNN M., Brookhaven National Laboratory, Upton, New York PAGET, OLIVER, The Biological Laboratory, Cold Spring Harbor, New York PALLADE, GEORGE, Rockefeller Institute for Medical Research, New York PAPPENI.IEIMER, A. M., JR., New York University, New York PARKER, ROBERT F., Western Reserve University, Cleveland, Ohio PAUCKER, KURT, Children's Hospital, Philadelphia, Pennsylvania POLLARD, ERNEST, Yale University, New Haven, Connecticut PORTER, K. R., Rockefeller Institute for Medical Research, New York PRINCE, A. M., Western Reserve University, Cleveland, Ohio PucK, THEODORE T., University of Colorado Medical Center, Denver, Colorado RANDALL, CHARLES C., Vanderbilt University, Nashville, Tennessee REBACK, JOHN F., University of Notre Dame, Notre Dame, Indiana REICI-IMAN, ELI, Harvard University Cambridge, Massachusetts REINER, JOHN M., Delafield Hospital, New York RICKENBERG, H. V., Yale University, New Haven, Connecticut ROBERTS, RICHARD, Carnegie Institution of Washington, Washington, D. C. ROELKE, MRS. CHARLES, Glastonbury, Connecticut ROPER, JOSEPH A., California Institute of Technology, Pasadena, California BE ROTHSCHILD, B., Columbia University, New York ROTMAN, BORIS, University of Illinois, Urbana, Illinois ROWEN, ROBERT, New York University, New York RUBIN, HARRY, University of California, Berkeley, California SABIN, ALBERT B., University of Cincinnati, Cincinnati, Ohio SAGER, RUTH, Rockefeller Institute for Medical Research, New York SANDERS, F. K., University Museum, Oxford, England SAVAGE, GEORGE M., The Upjohn Company, Kalamazoo, Michigan SAZ, ARTHUR K., National Institutes of Health, Bethesda, Maryland SCI-IAErrER, PIERRE A., Columbia University, New York SCHtAFFER, WERNER, Max Planck Institut, Tfibingen, Germany SCHERER, WILLIAM F., University of Minnesota, Minneapolis, Minnesota SCHLESINGER, n. WALTER, Public Health Research Institute, New York SCHWARTZ, BENJAMIN, Warner Institute for Therapeutic Research, New York SCOTT, MRS. DWIGHT MCNAIR, Children's Hospital, Philadelphia, Pennsylvania SEGAL, WILLIAM, Public Health Research Institute, New York SELDEEN, MURRAY J., New York University, New York SIMINOVITCH, LOUIS, Institut Pasteur, Paris, France SINSHEIMER, ROBERT L., California Institute of Technology, Pasadena, California SKAAR, PALMER D., University of Wisconsin, Madison, Wisconsin SOMMER, A., Delafield Hospital, New York STAHL, FRANKLIN W., University of Rochester, New York

xiv LIST OF PARTICIPANTS

STANLEY, W. M., Univevsity of California, Berkeley, California STEIGMAN, ALEX J., University of Louisville, Louisville, Kentucky STENT, GUNTHER S., University of California, Berkeley, California ST. LAWRENCE, PATRICIA, Yale University, New Haven, Connecticut STREISINGER, GEORGE, University of Illinois, Urbana, Illinois STUARD, BARBARA, The Biological Laboratory, Cold Spring Harbor, New York SUSKIND, S. R., Yale University, New Haven, Connecticut SYVERTON, J. T., University of Minnesota, Minneapolis, Minnesota SZILARD, LEO, University of Chicago, Chicago, Illinois SZYBALSKI, WACLAW, The Biological Laboratory, Cold Spring Harbor, New York TATUM, E. L., Stanford University, Stanford, California TEEGARDEN, L. ELIZABETH, University of Michigan, Ann Arbor, Michigan TESSMAN, ETHEL, Yale University, New Haven, Connecticut TREFEERS, HENRY P., Yale University, New Haven, Connecticut TRESAR, WALTER C., Lederle Laboratories, Pearl River, New York TUTTLE, ALICE L., Yale University, New Haven, Connecticut UMBREIT, WAYNE W., Merck Institute for Therapeutic Research, Rahway, New Jersey VAN VUNAKIS, HELEN, The Johns Hopkins University, Baltimore, Maryland VlSCONTI, NICCOLO, Carnegie Institution of Washington, Cold Spring Harbor, New

York VOGT, MARGUERITE, California Institute of Technology, Pasadena, California YON BORSTEL, ROBERT, Carnegie Institution of Washington, Cold Spring Harbor, New

York WALLACE, BRUCE, The Biological Laboratory, Cold Spring Harbor, New York WANG, T. P., The Johns Hopkins University, Baltimore, Maryland WARREN, CHARLES 0., The Commonwealth Fund, New York WARREN, KATHERINE BREHME, Hofstra College, Hempstead, New York WATSON, J. D., The Cavendish Laboratory, Cambridge, England WEBB, MICHAEL, Rutgers University, New Brunswick, New Jersey WEIDEL, WOLFHARD, Max Planck-Institut fur Biologie, Tiibingen, Germany WEINDLING, RICHARD, Lederle Laboratories, Pearl River, New York WEISBLAT, D. I., The Upjohn Company, Kalamazoo, Michigan WELSH, J. N., New York University, New York WERNER, G. H., State University, New York WEYER, J., Yale University, New Haven, Connecticut WILLIAMS, ROBLEY C., University of California, Berkeley, California WISSEMAN, CHARLES L., Walter Reed Army Medical Center, Washington, D. C. WITKIN, E., Carnegie Institution of Washington, Cold Spring Harbor, New York WOLLMAN, ELIE L., Institut Pasteur, Paris, France WYATT, G. R., Laboratory of Insect Pathology, Sault Ste. Marie, Canada YANOFSKY, CHARLES, Yale University, New Haven, Connecticut YAmV, H., Tuberculosis Research Laboratory, U. S. Public Health Service, New York YAVERBAUM, SIDNEY, University of Pennsylvania, Philadelphia, Pennsylvania YAW, KATHERINE, University of Delaware, Newark, Delaware YOUNGNER, J. S., The Municipal Hospital, Pittsburgh, Pennsl~lvania ZAHLER, STANLEY i. , University of Illinois, Urbana, Illinois ZAMENHOF, STEPHEN, Columbia University, New York ZINOER, NORTON D., Rockefeller Institute for Medical Research, New York

C O N T E N T S

DELBRUCK, M. Introductory remarks about the program .................. 1

VIRUS IN THE VEGETATIVE STATE AND ITS MATURATION

DOERMANN, A.H. The vegetative state in the life cycle of bacterio- phage: evidence for its occurrence, and its genetic characteriza- tion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

LEVINTHAL, C. Recombination in phage: its relationship to heterozy- gosis and growth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

FRASER, DOROTHY, and DULBECCO, R. A genetic analysis of the fac- tors controlling the h character in bacteriophage T3 . . . . . . . . . . . . . . . . 15

APPLEBY, J. CHRISTINE. Genetic recombination experiments with influenza virus ... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

BURNET, F. M., and LIND, PATRICIA, E. Influenza virus recombina- tion: experiments using the de-embryonated egg technique . . . . . 21

HIRST, G.K. Intracellular reactions between two types of influenza virus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

LEVINTHAL, C., and FISHER, H.W. Maturation of phage and the evi- dence of phage precursors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29

HENLE, WERNER. Developmental cycles in animal viruses . . . . . . . . . . . . . 35 SANDERS, F .K . Evidence of a multiplication cycle in mouse enceph-

alomyelitis virus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 MARAMOROSCH, KARL. Do developmental stages occur in the repro-

ductive cycle of aster-yellows virus? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51 SCrILESINGER, R. WALTER. The relation of functionally deficient

forms of influenza virus to viral development ......................... 55 MELNICK, JOSEPH L., and GAYLORD, WILLIAM H. Intracellular de-

velopment of pox viruses .......................................................... 61

PROVIRUS

BERTANI, G. Lysogenic v e r s u s lyric cycle of phage multiplication .... 65 LIEB, MARGARET. Studies on lysogenization in E s c h e r i c h i a co l i . . . . . . . . 71 HAYES, W. The mechanism of genetic recombination in E s c h e r i c h i a

c o l i . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 APPLEYARD, R . K . Segregation of lambda lysogenicity during bac-

terial recombination in E . coI i K-12 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95 DELAMATER, EDWARD D. The mitotic mechanism in bacteria ........... 99

THE TRANSITION FROM PROVIRUS TO VEGETATIVE VIRUS

JACOB, FRANCOIS, and WOLEMAN, ELIE L. Induction of phage de- velopment in lysogenic bacteria .............................................. 101

WATSON, J, D., and CRICK, F. H.C. The structure of DNA ............... 123 WYATT, G. R. The quantitative composition of deoxypentose nucleic

acids as related to the newly proposed structure .... . . . . . . . . . . . 133

[xv]

xvi CONTENTS

THE TRANSITION FROM THE INFECTIVE TO THE VEGETATIVE STATE

HERSHEY, A . D . Functional differentiation within particles of bac- teriophage T2 ........................................................................... 135

MARKHAM, RoY. Chemistry of some functional components of viruses 141 PUCK, THEODORE T. The first steps of virus invasion ....................... 149 WEIDEL, WOLFHARD, Phage receptor systems of E. coli B . . . . . . . . . . . . 155

STRUCTURE OF VIRUSES

LANNI, FRANK, and LANNI, YVONNE THERY. Antigenic structure of bacteriophage ......................................................................... 159

SCHXFFER, WERNER, and MuNK, KLAUS. Studies on the complement- fixing-antigen of the fowl plague virus .... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169

LARK, KARL G., and ADAMS, MARK H. The stability of phages as a function of the ionic environment .......................................... 171

WILLIAMS, RORLEY C. The shapes and sizes of purified viruses as determined by electron microscopy ......................................... 185

ANDERSON, THOMAS F. The morphology and osmotic properties of bacteriophage systems ...........................................................

JESAITIS, MARGERIS, and GOEBEL, WALTHER F. The interaction be- 197 tween T4 phage and the specific lipocarbohydrate of phase II Sh. sonnei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205

BIOCHEMICAL STUDIES OF VIRUS INFECTIONS

KOZLOFF, LLOYD M. Origin and fate of bacteriophage material ........ 207 COHEN, SEYMOUR S. Studies on controlling mechanisms in the met-

abolism of virus-infected bacteria ............................................ 221

HOST-CONTROLLED VARIATIONS OF VIRUSES

LURIA, S.E. Host-induced modifications of viruses ............................ 237 BOWEN, G. H. Studies of ultraviolet irradiation phenomena--an

approach to the problems of bacteriophage reproduction ............ 245 STENT, GUNTHER. Mortality due to radioactive phosphorus as an

index to bacteriophage development ........................................... 255 ZINDER, NORTON D. Infective heredity in bacteria ............................ 261 BARON, L.S . Genetic transfer by means of Vi Phage lysates ............ 271

VIRUSES IN TISSUE CULTURE

DULBECCO, RENATO, and VOGT, MARGUERITE. Some problems of animal virology as studied by the plaque technique .............................. 273

YOUNGNER, J. S. Some observations on the behavior of poliomyelitis viruses in tissue culture ......................................................... 281

SYVERTON, JEROME T., and SCHERER, WILLIAM F. Applications of strains of mammalian cells to the study of animal viruses .......... 285

FENNER, FRANK. Host parasite relationships in myxomatosis of the Australian wild rabbit ............................................................ 291