MOLECULAR AND CELLULAR BIOLOGY VOLUME 3 * NUMBER 12 * DECEMBER 1983 Aaron J. Shatkin, Editor in Chief (1985) Roche Institute of Molecular Biology Nutley, N.J. Harvey F. Lodish, Editor (1986) Louis Siminovitch, Editor (1985) Massachusetts Institute of Technology Hospital for Sick Children Cambridge, Mass. Toronto, Canada David J. L. Luck, Editor (1987) Paul S. Sypherd, Editor (1985) Rockefeller University University of California New York, N.Y. Irvine, Calif. EDITORIAL BOARD Renato Baserga (1985) Ira Herskowitz (1984) Daniel B. Rifkin (1985) Alan Bernstein (1984) Larry Kedes (1985) Robert G. Roeder (1985) J. Michael Bishop (1984) Marilyn Kozak (1985) James E. Rothman (1984) Joan Brugge (1985) Elias Lazarides (1985) Phillip A. Sharp (1985) Breck Byers (1985) John B. Little (1985) Fred Sherman (1985) John A. Carbon (1984) William F. Loomis, Jr. (1985) Pamela Stanley (1985) Lawrence A. Chasin (1985) Paul T. Magee (1985) Joan A. Steitz (1985) Nam-Hai Chua (1985) Robert L. Metzenberg (1985) James L. Van Etten (1985) Terrance G. Cooper (1984) Robert K. Mortimer (1985) Jonathan R. Warner (1984) James E. Darneil, Jr. (1985) Harvey L. Ozer (1985) Robert A. Weinberg (1984) Gary Felsenfeld (1985) Mary Lou Pardue (1985) I. Bernard Weinstein (1985) Norton B. Gilula (1985) Mark Pearson (1985) Harold Weintraub (1985) James E. Haber (1984) Jeremy Pickett-Heaps (1985) Reed B. Wickner (1985) Benjamin Hall (1985) Robert E. Pollack (1985) Leslie Wilson (1985) Ari Helenius (1984) Keith R. Porter (1985) Edward Ziff (1985) Susan A. Henry (1985) John R. Pringle (1985) Helen R. Whiteley, Chairman, Publications Board Walter G. Peter Ill, Director, Publications Linda M. Illig, Managing Editor, Journals Eleanor S. Tupper, Production Editor Molecular and Cellular Biology (ISSN 0270-7306) is devoted to the advancement and dissemination of fundamental knowledge concerning the molecular biology of eucaryotic cells, of both microbial and higher organisms. Instructions to authors are published in the January issue each year; reprints are available from the editors and the Publications Office. The journal is published monthly, one volume per year. The nonmember subscription price is $84 per year; single copies are $9. The member subscription price is $22 (foreign $26 [surface rate]) per year; single copies are $7. Correspondence relating to subscriptions, reprints, defective copies, availability of back issues, lost or late proofs, disposition of submitted manuscripts, and general editorial matters should be directed to the ASM Publication Office, 1913 1 St., NW, Washington, DC 20006 (area 202 833-9680). Claims for missing issues from residents of the United States, Canada, and Mexico must be submitted within 3 months after publication of the issues; residents of all other countries must submit claims within 6 months of publication of the issues. Claims for issues missing because of failure to report an address change or for issues "missing from files' will not be allowed. Second-class postage paid at Washington, DC 20006, and at additional mailing offices. POSTMASTER: Send address changes to Molecular and Cellular Biology, ASM, 1913 I St., NW, Washington, DC 20006. Made in the United States of America. Copyright © 1983, American Society for Microbiology. i*: A lt(1' All Rights Reserved. The code at the top of the first page of an article in this journal indicates the copyright owner's consent that copies of the article may be made for personal use or for personal use of specific clients. This consent is given on the condition, however, that the copier pay the stated per-copy fee through the Copyright Clearance Center Inc., 21 Congress St., Salem, MA 01970, for copying beyond that permitted by Sections 107 and 108 of the U.S. Copyright Law. 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. Author Index Antequera, Francisco, 2287 Harris, John F., 2271, 2280 Parada, Luis F., 2298 Held, William A., 2232 Hodgson, Clague P., 2221 Barclay, Stephen L., 2117 Reich, Nancy C., 2143 Benham, F. J., 2259 Richter, J. D., 2131 Innis, Jeffrey W., 2203 Robertson, M., 2211 Bernstein, Alan, 2180, 2191 Rogers, J., 2166 Bostock, C., 2211 Rudnicki, Michael A., 2271 Jones, N. C., 2131 Christie, S., 2211 Jones-Villeneuve, Elizabeth Santos, Tomas, 2287 M. V., 2271 Sanwal, B. D., 2166 Joyner, Alexandra L., 2180, 2191 Scott, Walter A., 2203 Dahlberg, J. E., 2211 Shatkin, Aaron J., 2172 Simonsen, Christian C., 2250 Knopf, John L., 2232 Smith, Gale E., 2156 Edwards, Mary K. S., 2280 Smith, James H., 2151 Elder, Paula K., 2221 Smith, L. D., 2131 Eliceiri, George L., 2151 Levine, Arnold J., 2143 Summers, Max D., 2156 Ernst, Heinrich, 2172 Levinson, Arthur D., 2250 Lund, E., 2211 Tamame, Mercedes, 2287 Temin, Howard M., 2241 Filipowicz, Witold, 2172 Foster, Douglas N., 2221 McBurney, Michael W., 2271, Fraser, M. J., 2156 2280 Villanueva, Julio R., 2287 Melier, Erna, 2117 Mitchen, J. L., 2211 Gallagher, John F., 2232 Watanabe, Shinichi, 2241 Getz, Michael J., 2221 Weeks, D. L., 2131 Gilfix, B., 2166 Ono, Tetsuya, 2221 Weinberg, Robert A., 2298 Goodfellow, P. N., 2259 Oren, Moshe, 2143 Wiles, M. V., 2259 ACKNOWLEDGMENT The following have served as invited special reviewers for the journal during 1983, and their help is greatly appreciated. George Acs Herbert Ennis George Khoury Bruce Adams Ronald Evans David Knipe Eileen Adamson Phillip Feigelson Barbara Knowles Irene Andrulis James Feramisco Monty Krieger Herbert N. Arst, Jr. John Fessler F. Lacroute Susan Astrin David Finkelstein C. J. Lai Corrado Baglioni Jane Flint Alan Lambowitz Andrew Ball Wayne Flintoff Gayle Lamppa David Baltimore William Frazier Ronald Laskey Amiya Banerjee Karl Freeman Robert Lawther Heinz Baumann Robert Friedman Paul Lefebvre Robert Benbow Alice Fulton Michael Leibowitz Thomas Benjamin Joseph Gall Peter Lengyel Marian Beremand Alan Garen Arnold Levine Susan Berget J. I. Garrels Susan Lindquist Merton Bernfield Patricia Gearhart Vic Ling Kenneth Berns Peter Geiduschek Stuart Linn Elizabeth Blackburn Raymond Gesteland Elwood Linney Peter Blumberg Eli Gilboa Peter Linsley Keith Bostian Harold Ginsburg David Livingstone Michael Botchan Yakov Gluzman Paul Lizardi David Botstein Steve Goff Peter Lomedico Ted Bradley Allan Goldberg Vivian Mackay M. Breitenbach E. G. Gollub Robert Malone Martin Breitman Martin Gorovsky James Manley J. R. Broach Michael Gottesman Thomas Manney Thomas Broker Frank Graham Kenneth Marcu Jeremy Bruenn Steve Grant Steve Martin Joseph Bryan Mark Groudine George Marzluf Manuel Buchwald Leonard Guarente Michael Matthews Howard Bussey Rad Gupta Michael McBurney Ron Butow Dean Hamer Roderick McInnes Judith Campbell Hidesaburo Hanafusa Steve McKnight Mario Capecchi Roger Hand Janet Mertz Mary Case Leland Hartwell Mathew Meselson Francis Castellino Vincent Hascall, Jr. Mark Meuth Sewell Champe William Haywood Malcolm Moore Clarence Chan Joyce Heckman Steve Mount Lan Bo Chen T. L. Helser Richard Mulligan Rex Chisholm Ellen Henderson Paul Neiman Albert Chung David Henner Yale Nemerson Stanley Cohen James Hicks Walter Neupert Christopher Coleclough David Hirsch Joseph Nevins Jonathan Cooper M. J. Holland Carol Newlon Nicholas Cowan Anita Hopper Maynard Olson Brian Cox J. E. Hopper Yasuji Oshima William Crane Stephen Howell Brad Ozanne George Cross Peter Howley Richard Palmiter Lloyd Culp Ann Hubbard Thoru Pederson Samuel Dales Joel Huberman Hugh Pelham Kathleen Danna Tony Hunter Sheldon Penman Ronald Davis Marcello Jacobs-Lorena D. Perlman Rowland Davis Alan Jacobson P. S. Perlman Carol Dieckmann Warren Jelinek Sidney Pestka Robert Dottin Edward Johnson Thomas Petes Gideon Dreyfuss Grace Ju Magnus Pfahl Thomas Easton Robert Kamen James Pipas Argiris Efstratiadis Kathleen Karrer David Prescott Ellis Englesberg Randy Kaufman Murray Rabinowitz Uttam Rajbhandary Mark Shotwell Robert Tjian Peter Ray Mohammed Siddiqui Paul Torrence Steve Reed Carolyn Silflow La-Chee Tsui Richard Rifkind Samuel Silverstein George vandeWoude Jasper Rine Richard Singer Harold Varmus G. S. Roeder Anna Skalka Alexander Varshavsky Bernard Roizman Arthur Skoultchi Inder Verma Michael Rosbash Martin Snider Jan Vilcek Jack Rose Davor Solter Robert Wagner Naomi Rosenberg Patricia Spear Eugenia Wang M. Rosentraus George Stark David Ward Jeffrey Ross Nat Stemnberg Donald Weeks Joan Ruderman Bernard Strauss Martin Weigert Frank Ruddle Sidney Strickland Eric Weinberg H. Ruis Kiranur Subramanian Alan Weiner David Sabatini Ako Sugino Irving Weissman Robert Schimke Jack Szostak Sherman Weissman Milton Schlesinger Igor Tamm Curtis Wilkerson David Schlessinger John Taylor Fred Winston Bartholomew Sefton Peter Tegtmeyer Dyann Wirth Pravinkumar Sehgal Larry Thompson Ronald Worton Ganes Sen Jeremy Thorner Keith Yamamoto Thomas Shenk W. Timberlake Men-Chao Yao Michael Sherman D. J. Tipper Michael Young INDEX TO DATE OF ISSUE Month Date of Issue Pages January 6 January 1983 1-147 February 4 February 1983 149-296 March 3 March 1983 297-494 April 30 March 1983 495-756 May 3 May 1983 757-965 June 2 June 1983 967-1162 July 28 June 1983 1163-1332 August 5 August 1983 1333-1543 September 2 September 1983 1545-1685 October 10 October 1983 1687-1887 November 2 November 1983 1889-2115 December 2 December 1983 2117-2301 STATEMENT OF OWNERSHIP MANAGEMENT AND CIRCULATION IA TITLE OF PUBLICATION IS. PUUCAlTON NO. 2. OATE OF FILING Molecular and Cellular Biology I i i i i i 30 September 1983 3. FREQUENCY OF ISUE 3A. NO.OF ISSUNSP LISHEO 25. ANNUAL SUSCRWPTIOFI LI.Y PtIClE Monthly 12 $22 abr, $84 nonabr 4. COMPLETE MAILING ADORES Of INOt OFfCEOF PtLICATION (et, CSy. aMety,Syawe.eZScZok f'oatPAneJ 1913 I Street, N.W., Washington, D.C. 20006 5 COMPLETE MAILING ADORES Of THE HEADOUANTEtS OF GENERAL WEINE#D OFFICES OF THE PUBLISHER (NeotePte, 1913 I Street, N.W., Washington, D.C. 20006 6. FULL NAMES ANO COMPLETE MAILING ADOREO OF PUBLISHER. EOITOR. ANDMANAGING EDITOR (NIi mUSrI TNorDb.E.Mk PUNLISHER (N... ACet*_ .p AeEA Amlerican Society for Microbiology. 1913 I Street, N.W., Washington, D.C. 20006 EDITOR N_..WtCap_ I's Adew Director of Publications W lter G.