Molecular and Cellular Biology

MOLECULAR AND CELLULAR BIOLOGY

VOLUME 7 0 MAY 1987 NUMBER 5

Aaron J. Shatkin, Editor in Chief(1990) Jack Szostak, Editor (1991) N.J. Center for Advanced Massachusetts General Hospital Biotechnology and Medicine Boston Piscataway, N.J. Louis Siminovitch, Editor (1990) Mount Sinai Hospital Robert Tjian, Editor (1991) Harvey F. Lodish, Editor (1991) Toronto, Canada University of California Whitehead Institute for Biomedical Berkeley Research Joan A. Steitz, Editor (1990) Cambridge, Mass. Yale University Harold E. Varmus, Editor (1989) New Haven, Conn. University of California David J. L. Luck, Editor (1987) San Francisco Rockefeller University New York, N.Y.

EDITORIAL BOARD Frederick W. Alt (1987) Jack F. Greenblatt (1988) Steven McKnight (1989) Matthew P. Scott (1989) Arnold J. Berk (1988) Leonard P. Guarente (1988) Janet E. Mertz (1987) Fred Sherman (1988) Alan Bernstein (1987) Christine Guthrie (1989) Robert L. Metzenberg (1988) Arthur Skoultchi (1988) Barbara K. Birshtein (1987) James E. Haber (1987) Robert K. Mortimer (1988) Barbara Sollner-Webb (1989) J. Michael Bishop (1987) Hidesaburo Hanafusa (1989) Paul Neiman (1989) Frank Solomon (1988) Michael R. Botchan (1987) Ari Helenius (1987) Joseph R. Nevins (1987) Karen Sprague (1989) David Botstein (1987) Ira Herskowitz (1987) Carol Newlon (1988) Pamela Stanley (1988) Bruce P. Brandhorst (1987) James B. Hicks (1989) Brad Ozanne (1989) Nat Sternberg (1989) James R. Broach (1988) Alan Hinnebusch (1988) Harvey L. Ozer (1988) Bruce Stillman (1988) Joan Brugge (1988) Michael J. Holland (1987) Mary Lou Pardue (1988) Kevin Struhl (1989) Mario R. Capecchi (1987) Greg Hollis (1987) Carl S. Parker (1987) Bill Sugden (1988) John A. Carbon (1987) Anita K. Hopper (1988) Ira H. Pastan (1988) Lawrence H. Thompson (1988) Marian Carlson (1989) Peter M. Howley (1988) David Patterson (1988) Shirley M. Tilghman (1987) Lawrence A. Chasin (1988) Tony Hunter (1989) Gianni Piperno (1988) Geoffrey Wahl (1989) Nam-Hai Chua (1988) Larry Kedes (1988) John R. Pringle (1988) Jonathan R. Warner (1987) Don W. Cleveland (1987) Robert S. Kerbel (1988) Jean-Paul Revel (1988) Alan M. Weiner (1987) Christopher Coleclough 1987) Daniel Klessig (1989) Daniel B. Rifkin (1988) Harold Weintraub (1988) Terrance G. Cooper (1987) Barbara Knowles (1989) G. S. Roeder (1988) Reed B. Wickner (1988) Elizabeth A. Craig (1988) Marilyn Kozak (1988) Robert G. Roeder (1988) Fred Winston (1988) James E. Dahlberg (1987) Monty Krieger (1989) Naomi E. Rosenberg (1988) Owen Witte (1988) James E. Darneil, Jr. (1988) Susan Lindquist (1987) Norman P. Salzman (1988) Elton T. Young (1987) G. N. Godson (1987) Stuart M. Linn (1989) Paul Schedl (1987) Michael Young (1988) Steve Goff (1989) Paul T. Magee (1988) Randy W. Schekman (1988) Edward Ziff (1988) Michael Green (1988) James Manley (1989) Milton J. Schlesinger (1989) Helen R. Whiteley, Chairman, Publications Board Kirk Jensen, Director ofPublications Linda M. Illig, Managing Editor, Journals Linda M. Illig, Production Editor Molecular and Cellular Biology (ISSN 0270-7306) is devoted to the advancement and dissemination offundamental 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 Department. The journal is published monthly, one volume per year. The nonmember subscription price is $280 per year; single copies are $25. The member subscription price is $43 (foreign $57 [surface rate]) per year; single copies are $8. 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 Publications Department, 1913 I 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 offailure 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 © 1987, American Society for Microbiology. as : / & UNI'VI:t f:( I 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

Aaronson, Stuart, 1984 Galliot, Brigitte, 2003 Maroni, Gustavo, 1710 Sexton, Julianne, 1602 Allen, James M., 1710 Ganea, Doina, 1656 Martin, Frank, 1797 Shahan, K., 1938, 1947 Amouyel, P., 1995 Gaudray, Patrick, 1740 Martinson, Harold G., 1917 Sherman, David H., 1865 Anderson, Stephen K., 1978 Gentry, Larry E., 1585 Marzluf, George A., 1691 Sherr, Charles J., 1673 Arthur, Richard R., 1978 Gibbs, Carol P., 1978 Matsukage, Akio, 2012 Shi, Y., 1947 Askew, David, 1673 Gilmartin, M., 1938, 1947 Max, Edward E., 1815 Shih, Thomas Y., 1999 Ayares, David, 1656 Gimble, Jeffrey M., 1815 May, Strat, 1673 Shio, Helen, 1848 Ayres, Kathleen, 1823 Gottesfeld, Joel M., 1612 McClure, Daniel S., 1592 Simantov, Rabi, 1830 Graber, Mark, 1984 McPhaul, Michael, 1841 Slater, Michael R., 1906 Backendorf, Claude, 2024 Groffen, John, 1955 Meinkoth, Judy L., 1740 Small, Gillian M., 1848 Baltimore, David, 1989 Miller, Maureen, 1759 Small, Michael B., 1638 Barreau, Christian, 1725 Haenlin, Marc, 2003 Mitsunobu, Fumihiro, 1776 Sokabe, Hirofumi, 1873 Begue, A., 1995 Hagag, Nabil, 1984 Mohier, Eliane, 2003 Sommer, Bernd, 1807 Begueret, Joel, 1725 Hardy, Kenneth J., 1967 Montano, X., 1782 Spizz, Gwendolyn, 1973 Berg, Paul, 1841 Harris, Curtis C., 2031 Mowry, Kimberly L., 1663 Stam, Kees, 1955 Berger, Shelley L., 1681 Hattori, Seisuke, 1999 Stanners, Clifford P., 1623 Bishop, J. Michael, 1638, 1697 Hay, Nissim, 1638 Nahon, Jean-Louis, 1856 Stavenhagen, Jeffrey, 1716 Bosselman, Robert A., 1797 Hayashi, Yuko, 2012 Nakagawa, Terry Y., 1731 Stehelin, D., 1995 Brash, Douglas E., 2031 Heisterkamp, Nora, 1955 Nakamura, Shun, 1999 Steitz, Joan A., 1663 Braun, Richard, 1933 Hemenway, Cynthia, 1716 Nakkim, Eric, 1740 Stevens, Priscilla Wilkins, Bruszewski, Joan, 1797 Hirose, Fumiko, 2012 Neuman, Wilma, 1823 1751 Buford, Edward S., 1961 Hochman, Paula S., 1865 Nicolson, Margery, 1797 Stone, James, 1697 Buonocore, Linda, 1759 Howard, Bruce H., 1894 Nishimoto, Yoshio, 2012 Subler, Mark, 1740 Howard, Tazuko H., 1894 Nojima, Hiroshi, 1873 Swanson, Maurice S., 1731 Campbell, Colin R., 1656 Hoy, Cynthia A., 2007 Nonet, Michael, 1602 Sypherd, Paul S., 1925 Carrino, John J., 1933 Hsu, Rou-Yin, 1797 Nusse, Roel, 2019 Carroll, Susan M., 1740 Hu, Sylvia, 1797 Cartier, Mireille, 1623 Huber, Brigitte T., 1865 Olson, Eric N., 1973 Tanaka, Akio, 1978 Cartwright, Christine A., 1830 Hunter, Tony, 1830 Otto, Edward, 1710 Tang, Wei Jen, 1681 Chang, Mildred W.-M., 1623 Thielmann, Heinz W., 2024 Chekuri, Lavanya, 1656 Ihle, James N., 1673 Padmanabhan, Raji, 1894 Thompson, J. Ryan, 1764 Chepelinsky, Ana B., 1807 Imanaka, Tsuneo, 1848 Palmiter, Richard D., 1646, Thompson, Larry H., 2007 Chung, Stephen, 1823 Iyer, A., 1629 1710 Tizard, Richard, 1865 Clanton, David J., 1999 Payne, Paul A., 1973 Toyoshima, Kumao, 1776 Coll, J., 1995 Jakobovits, Edward, 1697 Perrot, Michael, 1725 Tratner, Isabelle, 1856 Cornelisse, Cees, 2019 Perryman, M. Benjamin, 1973 Triezenberg, Steven J., 1681 Cosgrove, Dominic E., 1592 Kaina, Bernd, 2024 Peterlin, B. Matija, 1967 Twardzik, Daniel R., 1585 Cox, G. Stanley, 1592 Kalff, Martha, 1716 Peterse, Johannes, 2019 Craig, Elizabeth A., 1906 Kaplan, Paul L., 1830 Piatigorsky, Joram, 1807 van de Bersselaar, Rene, 2019 Kaul, K., 1629 Propst, F., 1629 Van De Putte, Piet, 2024 Davidson, Jeffrey N., 1961 Kawai, Sadaaki, 1776 van de Vijver, Marc, 2019 de Lange, Titia, 1697 Kawakita, Masao, 1999 Quanrud, Myra, 2031 Vande Woude, G. F., 1629 Denaro, M., 1947 Kaziro, Yoshito, 1999 Quatannens, B., 1995 Van Zeeland, Albert A., 2024 Denhez, F., 1995 Kishi, Koichiro, 1873 Varmus, Harold, 1697 Derman, E., 1938, 1947 Kucherlapati, Raju, 1656 Ramachandran, Venetianer, Aniko, 1856 De Rose, Margaret L., 1740 Kueng, Valentin, 1933 Kuzhalmannam L., 1865 Villeponteau, Bryant, 1917 Devilee, Peter, 2019 Kung, Hsing-Jien, 1978 Ramsay, Gary, 1697 Viola, Michael V., 1984 Dick, Robert, 1865 Ranchalis, Jane E., 1585 Von Hoff, Daniel D., 1740 Dodgson, Jerry B., 1751 Labhart, Paul, 1900 Rao, Gadiparthi N., 1961 Dozier, C., 1995 Lacal, Juan Carlos, 1984 Reddel, Roger R., 2031 Dreyfuss, Gideon, 1731 Laffier, Thomas G., 1933 Reeder, Ronald H., 1900 Wahl, Geoffrey M., 1740 Duboule, Denis, 2003 Lane, D. P., 1782 Reynolds, Fred H., Jr., 1955 Wheeler, Esther F., 1673 Larkin, John C., 1764 Roberts, Robert, 1973 Wilkie, Thomas M., 1646 Eckhart, Walter, 1830 Larsen, Anne S., 1967 Robins, Diane M., 1716 Wolgemuth, Debra J., 1791 Emery, John F., 1740 Lazarow, Paul B., 1848 Rosenberg, M. P., 1629 Woolford, John L., Jr., 1764 Engel, James Douglas, 1751 Lee, David C., 1585 Rowekamp, Walter G., 1823 Lee, William, 1697 Yamaguchi, Masamitsu, 2012 Farrell, Michael P., 2031 LeVan, Kay, 1731 Sala-Trepat, Jose Maria, 1856 Yamamoto, Tadashi, 1776 Flavell, Richard A., 1865 Levens, David, 1815 Satoh, Takaya, 1999 Yang, Ke, 2031 Folk, William R., 1681 Lim, G. Jonathan, 1585 Saule, S., 1995 Young, Julie E., 1710 Fu, Ying-Hui, 1691 Linz, John E., 1925 Scafe, Charles, 1602 Young, Richard, 1602 Fujita, Donald J., 1978 Loreni, Fabrizio, 1716 Schneider, Michael D., 1973 Fukui, Mikiharu, 1776 Luskey, Kenneth L., 1881 Schwab, Manfred, 1638 Fuscoe, James C., 2007 Lyon, Sue Brockett, 1759 Sen, Ranjan, 1989 Zakeri, Zahra F., 1791

i If you are working in the area ofeucaryotic cell biology, then you need EDITOR IN CHIEF: Aaron J. Shatkin EDITORS: Harvey F. Lodish. David J. L. Luck. Louis Siminovitch. Joan A. Steitz. Jack Szostak. Robert Tjian. Harold E. Varmus

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PUBLISHER: Ameria for Microbiology El^1v1 1913 I Streeto N.W. Society Washington, DC 20006 When the focus is on basic microbial research, readers turn to the followingASMjournals: .l _~~~~~~ _. _~~~~~~~~~~~~~~~~~~~~~~%. we_ Molecular and Cellular Biology EDITOR IN CHIEF. Aaron J. Shatkin MCB is devoted to the advancement and dissemination of fundamental knowledge concerning the molecular biology of eucaryotic cells. of both microbial and I~~~ *S OS higher organisms. The journal publishes papers on cellular morphology and function. genome organization. regulation of genetic expression. morphogenesis, and somatic cell genetics. In addition. it contains articles concerning plas- mid vectors and virus-infected cells in which emphasis is clearly on the cell. Monthly. ISSN: 0270-7306. 4.500 pages in 1987. Nonmember Member (US) Member (Foreign) $280.00 $43.00 $57.00 Add $145.00 for foreign airmail service. Journal of Bacteriology N S _ EDITOR IN CHIEF: Simon Silver The oldest of the ASM journals. JB is the leading periodical. worldwide, in its field. It is devoted to the advancement and dissemination of fundamental Knowledge concerning bacteria and other microorganisms. including fungi and other unicellular, eucaryotic organisms. Regular features include articles on structure and function. plant microbiology membranes. eucaryotic cells. genetics and molecular biology. plasmids and transposons. physiology and metabolism, and enzymology. Monthly. ISSN: 0021-9193. 4.700 pages in 1987. I Nonmember Member (US) Member (Foreign) $340.00 $41.00 $56.00 Add $1 70.00 for foreign airmail service. Joura of Virology EDITOR IN CHIEF: Arnold J. Levine This leading research journal is devoted to the dissemination of fundamental knowledge concerning viruses of bacteria. plants. and animals. JVI publishes reports of original research in all areas of basic virology. including biochemistr>. rs ~~ 55 biophysics. genetics, immunology. morphology. physiology. and pathogenesis and immunity. Monthly. ISSN: 0022-538X. 4.200 pages in 1987. Nonmember Member (US) Member (Foreign) i S w $350.00 $41.00 $56.00 Add $1 70.00 for foreign airmail service.

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