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MOLECULAR and CELLULAR BIOLOGY VOLUME 10 * MAY 1990 * NUMBER 5 Aaron J MOLECULAR AND CELLULAR BIOLOGY VOLUME 10 * MAY 1990 * NUMBER 5 Aaron J. Shatkin, Editor in Chief (1990) Louis Siminovitch, Editor (1990) Center for Advanced Mount Sinai Hospital Biotechnology and Medicine David J. L. Luck, Editor (1992) Toronto, Canada Piscataway, N.J. Rockefeller University Shirley M. Tilghman, Editor Don W. Cleveland, Editor (1994) New York, N. Y. (1993) The Johns Hopkins University Steven L. McKnight, Editor (1992) Princeton University Baltimore, Md. Carnegie Institution ofWashington Princeton, N.J. Anita K. Hopper, Editor (1994) Baltimore, Md. Alan M. Weiner, Editor (1993) The Milton S. Hershey Medical Center Randy W. Schekman, Editor Yale University Pennsylvania State University (1992) New Haven, Conn. Hershey, Pa. University of California Owen N. Witte, Editor (1993) Tony Hunter, Editor (1993) Berkeley Molecular Biology Institute The Salk Institute University of California San Diego, Calif. Los Angeles EDITORIAL BOARD Frederick W. Alt (1990) Mary P. Edmonds (1990) Monty Krieger (1992) Naomi E. Rosenberg (1991) Susan Berget (1990) Robert N. Eisenman (1991) Elizabeth Lacy (1990) Rodney Rothstein (1990) Arnold J. Berk (1991) Sarah C. R. Elgin (1991) Alan Lambowitz (1990) Norman P. Salzman (1991) Alan Bernstein (1990) Scott Emr (1991) Lester F. Lau (1992) Paul Schedl (1990) Barbara K. Birshtein (1990) Walton L. Fangman (1991) Michael Lenardo (1991) Milton J. Schlesinger (1992) J. Michael Bishop (1990) Thomas D. Fox (1990) Arthur D. Levinson (1990) Matthew P. Scott (1992) Elizabeth H. Blackburn (1991) Mary-Jane Gething (1990) Susan Lindquist (1990) Charles J. Sherr (1991) Jef D. Boeke (1991) Stephen P. Goff (1992) Douglas Lowy (1990) Arthur Skoultchi (1991) Michael R. Botchan (1990) Jack F. Greenblatt (1991) Paul T. Magee (1991) Frank Solomon (1991) David Botstein (1990) Rudolf Grosschedl (1992) James Manley (1992) Michael B. Sporn (1991) Bruce P. Brandhorst (1990) Leonard P. Guarente (1991) Kunihiro Matsumoto (1991) Allan C. Spradling (1991) Marjorie C. Brandriss (1991) Christine Guthrie (1992) Frank McCormick (1992) Karen U. Sprague (1992) Rodrigo Bravo (1992) James E. Haber (1990) Janet E. Mertz (1990) Pamela Stanley (1991) Kathryn Calame (1991) Hidesaburo Hanafusa (1992) N. Ronald Morris (1992) Nat L. Sternberg (1992) Judith Campbell (1992) Leland D. Hartwell (1990) Bernardo Nadal-Ginard (1990) Bruce Stillman (1991) Mario R. Capecchi (1990) Nathaniel Heintz (1991) Paul Neiman (1992) Kevin Struhl (1992) John A. Carbon (1990) Ari Helenius (1990) Joseph R. Nevins (1990) Kenneth D. Stuart (1991) Marian Carlson (1992) Steven Henikoff (1991) Carol Newlon (1991) Bill Sugden (1991) C. Thomas Caskey (1990) Ira Herskowitz (1990) Mary Ann Osley (1990) Lawrence H. Thompson (1991) Lawrence A. Chasin (1991) Alan Hinnebusch (1991) Harvey L. Ozer (1991) Geoffrey Wahl (1992) Nam-Hai Chua (1991) Michael J. Holland (1990) Mary Lou Pardue (1991) Randolph Wall (1992) Jonathan A. Cooper (1992) Greg Hollis (1990) David Patterson (1991) Peter Walter (1990) Nicholas Cowan (1990) Peter M. Howley (1991) Gianni Piperno (1991) Jonathan R. Warner (1990) Elizabeth A. Craig (1991) Stephen H. Hughes (1992) John R. Pringle (1991) Reed B. Wickner (1991) Thomas Curran (1991) Mark Johnston (1991) Randall R. Reed (1992) Lewis T. Williams (1991) Mark Davis (1990) Katherine A. Jones (1992) Steven I. Reed (1991) Fred Winston (1991) Thomas Donahue (1992) Daniel Klessig (1992) Jean-Paul Revel (1991) Elton T. Young (1990) Gideon Dreyfuss (1990) Marilyn Kozak (1991) Jasper D. Rine (1991) Michael Young (1991) Helen R. Whiteley, Chairman, Publications Board Linda M. Ihig, Managing Editor, Journals Catherine E. Blickendorfer, Production Editor Molecular and Cellular Biology (ISSN 0270-7306), a publication of the American Society for Microbiology, 1325 Massachu- setts Ave., N.W., Washington, DC 20005, 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 Department. The journal is published monthly, one volume per year. The nonmember subscription prices are $340 (U.S. and Canada) and $390 (foreign; air drop shipping) per year; single copies are $40. The member subscription prices are $43 (U.S. and Canada) and $93 (foreign; air drop shipping) per year; single copies are $10. Correspondence relating to subscriptions, reprints, defective copies, availability of back issues, lost or late proofs, disposition of submitted manuscripts, and general editorial matters should be directed to the ASM Publications Department, 1325 Massachusetts Ave., N.W., Washington, DC 20005 (area 202 737-3600). Claims for missing issues from residents of the United States, Canada, and Mexico must be submitted within 3 months after publication of the issues; residents of all other countries must submit claims within 6 months of publication of the issues. Claims for issues missing because of failure to report an address change or for issues "missing from files" will not be allowed. Second-class postage paid at Washington, DC 20005, and at additional mailing offices. POSTMASTER: Send address changes to Molecular and Cellular Biology, ASM, 1325 Massachusetts Ave., N.W., Washington, DC 20005. Made in the United States of America. Printed on acid-free paper. Copyright © 1990, American Society for Microbiology. 1:liL±$¶I 1wl E %Jif.1#i ( 4I] All Rights Reserved. The code at the top of the first page of an article in thisjournal indicates the copyright owner's consent that copies ofthe 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 Akeson, Richard, 2012 Franke, Jakob, 1921 Linial, Maxine L., 1891 Sawai, Shoji, 2017 Alcorta, David A., 2413 French, Randall P., 2081 Linsley, Peter S., 1882 Sax, Christina M., 2349 Alt, Frederick W., 2096 Frontali, C., 2423 Lipsick, Joseph S., 2285 Saxe, Stephen A., 2367 Altanchimeg, Rencendorj, London, Lucille, 1901 Schmidt, Ann, 2176 2432 Gaffney, Robert, 1931 Schuster, Wolfgang, 2428 Amrein, Kurt E., 1853 Gallego, Maria E., 2133 Marck, Christian, 2437 Sefton, Bartholomew M., 1853 Anraku, Yasuhiro, 2214 Gashler, Andrea, 1931 Martin, Christopher, 1908 Sentenac, Andrd, 2437 Aronoff, Rachel, 1891 Gaugler, Luc, 1959 Martin, Debra J., 1950 Shago, Mary, 2335 Azizkhan, Jane C., 2111 Giguere, Vincent, 2335 Matsushime, Hitoshi, 2261 Shaw, Andrey S., 1853 Glaser, Scott M., 1873 McDonald, Vicki L., 1969 Shay, Jerry W., 2027 Babiss, Lee E., 2027 Glass, Charles A., 2164 McDonnell, Timothy J., 1901 Sherr, Charles J., 2407 Ballesta, Juan P. G., 2182 Goridis, Christo, 1959 McKearn, John P., 1901 Shibuya, Masabumi, 2261 Bally-Cuif, Laure, 1959 Grant, Barbara J., 2164 McKiernan, Martin, 1931 Shortle, David, 2308 Baltimore, David, 2327 Green, Janell M., 1969 Memet, Sylvie, 2049 Shoyab, Mohammed, 1969 Bandyopadhyay, Ram, 2060 Miller, Brian R., 1873 Simpson, Robert T., 2247 Barriocanal, Javier G., 2294 Hall, Anne L., 1921 Mitchell, Aaron P., 2104 Smith, Harold E., 2104 Barta, Endre, 2432 Hammond, Craig, 1853 Morris, Charles F., 1931 Spear, David H., 2315 Baum, Mary P., 1863 Hartwell, Leland H., 2202 Morrison, Deborah K., 2359 Steiner, Bart, 1891 Beltrame, Monica, 2341 Hatfield, Dolph, 1940 Mueller, LeMoyne, 2154 Stewart, Valerie, 2096 Bennett, Jeffrey L., 2191 Hay, Rick V., 1931 Stratling, Wolf H., 2302 Benvenisty, Nissim, 2418 Herrick, David, 2269 Nadal-Ginard, Bernardo, 2133 Strommer, Judith N., 2090 Bernards, Rene, 2012 Hiebert, Scott W., 1989 Nagata, Shigekazu, 2002 Sukhatme, Vikas P., 1931 Bianchi, Marco E., 2341 Hiesel, Rudolf, 2428 Neubauer, Michael G., 1882, Sullivan, Kevin F., 2164 Biemann, Hans-Peter N., 2122 Hinnebusch, Alan G., 2294 1969 Sweet, Laurel J., 2413 Blackburn, Elizabeth H., 2070 Hirsch, Marie-Rose, 1959 Ng, Davis T. W., 1989 Szab6, Piroska, 2432 Blasband, Andrew J., 2111 Hirschfeld, Steven, 2041 Nishizawa, Mikio, 2002 Bolen, Joseph B., 2035 Hockenberry, David, 1901 Nomura, Masayasu, 2049 Bosze, Zsuzsa, 2432 Nunez, Gabriel, 1901 Bowen-Pope, Daniel F., 2237 Ibanez, Carlos E., 2285 Takagi, Nobuo, 2261 Bozzoni, Irene, 2437 Iman, Deborah S., 2027 Ochs, Vincent, 1882 Tan, Eng M., 2164 Brawerman, George, 2060 Ogawa, Nobuo, 2224 Jackson, Catherine L., 2202 Tanaka, Kenji, 2214 Brennicke, Axel, 2428 Okamura, Sara, 1908 Tate, Peri, 2335 Buhler, Jean-Marie, 2049 Jacobson, Allan, 2269 Ortiz, Daniel F., 2090 Jacobson, K. Bruce, 1940 Teruya, Julie H., 2315 Oshima, Yasuji, 2224 Todaro, George J., 1969 Cao, Xinmin, 1931 Jinno, Atsushi, 2261 Ostertag, Wolfram, 2302 Johnson, Jeffrey J., 2081 Transy, Catherine, 2407 Chalkley, Roger, 2390 Ozato, Keiko, 2041 Trus, Michael, 2418 Chalupny, Jan, 1853 Johnson, Kyle, 1891 Chan, Edward K. L., 2164 Jones, Steven W., 2413 Pace, T., 2423 Chen, Xiaorong, 2111 Park, Eun-Chung, 2081 Ciafre, Silvia-Anna, 2437 Kallestad, Jeffery, 1882 Parker, Roy, 2269 Kanbe, Toshio, 2214 Paulson, K. Eric, 1841 Van Ness, Brian G., 1950 Clarke, Catherine F., 2315 Varmuza, Sue, 2335 Clarke, Louise, 1863 Kaplan, David R., 2359 Phi-Van,
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