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MOLECULAR AND CELLULAR VOLUME 8 SEPTEMBER 1988 NUMBER 9

Aaron J. Shatkin, Editor in Chief(1990) Randy W. Schekman, Editor Center for Advanced (1992) Robert Tjian, Editor (1991) Biotechnology and Medicine University of California University of California Piscataway, N.J. Berkeley Berkeley David J. L. Luck, Editor (1992) Louis Siminovitch, Editor (1990) Alan M. Weiner, Editor (1993) Mount Sinai Hospital Yale University New York, N. Y. Toronto, Canada New Haven, Conn. Owen N. Witte, Editor (1993) Steven L. McKnight, Editor (1992) Joan A. Steitz, Editor (1990) Institute Carnegie Institution of Washington Yale University University of California Baltimore, Md. New Haven, Conn. Los Angeles EDITORIAL BOARD Frederick W. Alt (1990) Steve Goff (1989) Susan Lindquist (1990) Norman P. Salzman (1988) Susan Berget (1990) Michael Green (1988) Stuart M. Linn (1989) Paul Schedl (1990) Arnold J. Berk (1988) Jack F. Greenblatt (1988) Douglas Lowy (1990) Milton J. Schlesinger (1989) Alan Bernstein (1990) Leonard P. Guarente (1988) Paul T. Magee (1988) Matthew P. Scott (1989) Barbara K. Birshtein (1990) Christine Guthrie (1989) James Manley (1989) Fred Sherman (1988) J. Michael Bishop (1990) James E. Haber (1990) Janet E. Mertz (1990) Arthur Skoultchi (1988) Michael R. Botchan (1990) Hidesaburo Hanafusa (1989) Robert L. Metzenberg (1988) Barbara Sollner-Webb (1989) (1990) Leland D. Hartwell (1990) Robert K. Mortimer (1988) Frank Solomon (1988) Bruce P. Brandhorst (1990) Ari Helenius (1990) Bernardo Nadal-Ginard (1990) Karen Sprague (1989) James R. Broach (1988) (1990) Paul Neiman (1989) Pamela Stanley (1988) Joan Brugge (1988) James B. Hicks (1989) Joseph R. Nevins (1990) Nat Sternberg (1989) Mario R. Capecchi (1990) Alan Hinnebusch (1988) Carol Newlon (1988) Bruce Stillman (1988) John A. Carbon (1990) Michael J. Holland (1990) Mary Ann Osley (1990) Kevin Struhl (1989) Marian Carlson (1989) Greg Hoilis (1990) Brad Ozanne (1989) Bill Sugden (1988) C. Thomas Caskey (1990) Anita K. Hopper (1988) Harvey L. Ozer (1988) Lawrence H. Thompson (1988) Lawrence A. Chasin (1988) Peter M. Howley (1988) Mary Lou Pardue (1988) Shirley M. Tilghman (1990) Nam-Hai Chua (1988) Tony Hunter (1989) Ira H. Pastan (1988) Geoffrey Wahl (1989) Don W. Cleveland (1990) Larry Kedes (1988) David Patterson (1988) (1990) Nicholas Cowan (1990) Robert S. Kerbel (1988) Gianni Piperno (1988) Jonathan R. Warner (1990) Elizabeth A. Craig (1988) Daniel Klessig (1989) John R. Pringle (1988) Harold Weintraub (1988) James E. Darnell, Jr. (1988) Barbara Knowles (1989) Jean-Paul Revel (1988) Reed B. Wickner (1988) Mark Davis (1990) Marilyn Kozak (1988) Daniel B. Rifkin (1988) Fred Winston (1988) Gideon Dreyfuss (1990) Monty Krieger (1989) G. S. Roeder (1988) Elton T. Young (1990) Mary P. Edmonds (1990) Elizabeth Lacy (1990) Robert G. Roeder (1988) Michael Young (1988) Thomas D. Fox (1990) Alan Lambowitz (1990) Naomi E. Rosenberg (1988) Edward Ziff (1988) Mary-Jane Gething (1990) Arthur D. Levinson (1990) Rodney Rothstein (1990) Helen R. Whiteley, Chairman, Publications Board Kirk Jensen, Director of Publications Linda M. Ihig, Managing Editor, Journals Linda M. Illig, Production Editor Molecular and Cellular Biology (ISSN 0270-7306), a publication of the American Society for Microbiology, 1913 I St., N.W., Washington, DC 20006-5107, 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 price is $340 per year; single copies are $30. The member subscription price is $43 (foreign, $63 [air drop shipping]) 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., N.W., Washington, DC 20006-5107 (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., N.W., Washington, DC 20006- 5107. Made in the United States of America. a A M;1 9 JAL i rTqf*. (I'J c Copyright ©) 1988, American Society for Microbiology. 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 arti- cle 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

Addison, Christine, 3740 Giguere, Vincent, 3847 LaRosa, Gregory J., 3906 Sakiyama, Shigeru, 3929, 3934 Altman, Ronni, 3717 Gottesdiener, Keith M., 3809 Latt, S. A., 3611 Samuelson, Linda C., 3857 Amin, Jahanshah, 3761 Grosveld, Frank, 3847 Legrain, P., 3755 Sanders, Nancy J., 3571 Ananthan, Jayakumar, 3761 Groudine, Mark, 3710 Leiden, Jeffrey M., 3809, 3820 Sariban, Eric, 3951 Anderson, Philip, 3748 Grummt, Ingrid, 3891 Levine, Arthur S., 3943 Scher, Charles D., 3696 Arai, Ken-ichi, 3777 Gudas, Lorraine J., 3906 Lindsten, Tullia, 3809, 3820 Schiestl, Robert H., 3619 Austerberry, Charles F., 3947 Linn, Stuart, 3964 Schoneberg, Christa, 3891 Haber, James E., 3918 Lyons, Sue, 3734 Scott, Gary K., 3670 Bartsch, Ingrid, 3891 Hanke, Paul D., 3591 Seraphin, B., 3755 Basson, Michael E., 3797 Harrich, David, 3717 Majumdar, Subrata, 36% Smith, Deborah H., 3882 Bayley, Stanley T., 3955 Hayes, Pliny H., 3670 Manley, James L., 3582 Solski, Patricia A., 3960 Beier, David R., 3864 Heartlein, M. W., 3611 Martin, G. Steven, 3603 Spanopoulou, Eugenia, 3847 Bishayee, Subal, 3696 Heindl, Lisa M., 3676 Maru, Yoshiro, 3770 Stern, David F., 3969 Boothroyd, John C., 3837 Hinnebusch, Alan G., 3827 Mason, Thomas L., 3636, Stimac, Eva, 3734 Branton, Philip E., 3955 Hirai, Hisamaru, 3770 3647 Storti, Robert V., 3591 Buss, Janice E., 3960 Hodgkin, Jonathan, 3627 Matsumoto, Kunihiro, 3777 Strominger, Jack L., 3809 Holloman, William K., 3703 Mehta, V. B., 3661 Stroud, Robert M., 3797 Cao, Huining, 3969 Horiguchi, Junko, 3951 Meisler, Miriam H., 3857 Sturzbecher, H.-W., 3740 Carleton, Stephen, 3703 Howe, John A., 3955 Meyer-Leon, Leslie, 3784 Chao, Janna, 3717 Huang, Li-Chun, 3784 Mueller, Peter P., 3827 Takaku, Fumimaro, 3770 Colgan, John D., 3582 Huxley, Clare, 3898 Michael L., 3837 Cone, Roger D., 3864 Muhich, Takeda, Kyoko, 3929 Cox, Michael M., 3784 Inman, Ross B., 3784 Muller, M. T., 3661 Takenaga, Keizo, 3929, 3934 Curcio, M. Joan, 3571 Munson, Peter J., 3943 Thompson, Craig B., 3809, Jelsma, Tony N., 3955 3820 Darlington, Gretchen J., 3857 Jenkins, John R., 3740 Nakamura, Yohko, 3934 Thorsness, Mary, 3797 Davies, Peter L., 3670 Jensen, Lee, 3964 Nakayama, Naoki, 3777 Tokunaga, Katsuo, 3929 Der, Channing J., 3960 Jones, Nancy H., 3809 Tsukuda, Toyoko, 3703 Dickson, Robert C., 3726 June, Carl H., 3820 Ostrowski, Michael C., 3872 Dixon, Kathleen, 3943 Umlauf, Scott W., 3784 Kageyama, Hajime, 3929 Partaledis, Judith A., 3647 Egan, Cay, 3955 Kamiyama, Kiyofumi, 3929 Perry, Michael, 3676 Voellmy, Richard, 3761 Kamps, Mark P., 3969 Pious, Donald, 3734 Fearon, Kathleen, 3636 Karpinski, Beverly A., 3809 Prakash, Satya, 3619 Waterston, Robert H., 3627 Ferguson, Blair, 3955 Kayne, Paul, 3717 Prochownik, Edward V., 3683 Watson, Roger J., 3938 Ferrell, James E., Jr., 3603 Kaziro, Yoshito, 3777 Pulak, Rock A., 3748 Weil, Thomas S., 3676 Finer-Moore, Janet, 3797 Keller, Paul R., 3857 Williams, David A., 3864 Fletcher, Garth L., 3670 Khan, Shabbir, 36% Rienhoff, Hugh Young, Jr., Williams, Norma P., 3827 Fotheringham, Scott, 3703 Knoll, J. H. M., 3611 3710 Williams, Trevor, 3898 Fried, Mike, 3898 Kondo, Kazunori, 3627 Rine, Jasper, 3797 Witte, Michael M., 3726 Fu, Xin-Yuan, 3582 Kornuc, Masayo, 3717 Rodgers, Claire, 3683 Kufe, Donald, 3951 Roilides, Emmanuel, 3943 Yao, Meng-Chao, 3947 Garcia, Joseph, 3717 Kukowska, Jolanta, 3683 Rosbash, M., 3755 Yoshida, Michihiro C., 3770 Garfinkel, David J., 3571 Rosenblatt, Margery F., 3864 Gaynor, Richard, 3717 Langer, Stephen J., 3872 Rudin, Norah, 3918 Ziff, Edward B., 3882 When the focus is on basic microbial research, readers turn to the followingASMjournals: Molecular and Cellular Biology EDITOR IN CHIEF: Aaron J. Shatkin MCB is devoted to the advancement and dissemination of fundamental knowl- edge concerning the molecular biology of eucaryotic cells, of both microbial and '11-.., higher organisms. The journal publishes papers on cellular morphology and function, genome organization, regulation of genetic expression, morphogen- esis, 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. 4,900 pages in 1988. Nonmember Member (US) Member (Foreign) $340.00 $43.00 $63.00 Add $165.00 for foreign airmail service. Journal o Bacteriology EDITOR IN CHIEF: Samuel Kaplan 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. 5,700 pages in 1988. Nonmember Member (US) Member (Foreign) $340.00 $41.00 $63.00 Add $210.00 for foreign airmail service. Journal 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 , biophysics, genetics, immunology, morphology, physiology, and pathogenesis and immunity. Monthly. ISSN: 0022-538X. 4,400 pages in 1988. Nonmember Member (US) Member (Foreign) $350.00 $41.00 $63.00 Add $170.00 for foreign airmail service.

Infection and Immunity EDITOR IN CHIEF: Joseph W. Shands, Jr.

IAI publishes articles on: infections caused by pathogenic bacteria, fungi, and parasites; ecology and epidemiology of pathogenic microbes; virulence factors, such as toxins and microbial surface structures; nonspecific factors in host resistance and susceptibility to infection; and immunology of microbial infection. Oral microbiology is given extensive coverage. Monthly. ISSN: 0019-9567. 3,700 pages in 1988. Nonmember Member (US) Member (Foreign) $340.00 $41.00 $63.00 Add $140.00 for foreign airmail service. I PFP' -DIl

Moleclular Biologyof *~' Micmobial Dffeierntiaton Editors: James A. Hoch Peter Setlow Proceedings ofthe Ninth International Spores Conference, Asiloma, California, 3-6 September 1984 The Ninth Intemational Spores Conference held in September 1984 focused on one exciting and rapidly progressing area of bacterial development: the molecular biology of the sporulation and germination processes. Symposium presentations from this important meeting and review articles have been compiled and edited for ready reference. Keep pace with the progress in spore research Molecular Biology ofMicrobial Differentiation presents the latest conclusions in the molecular biology ofdifferentiation in higher procaryotes, particularly Bacillussubtilis. Thirty-eight excellent articles cover these major areas: * Molecular Cloning and Genetics of Sporulation and Germination * Characterization of Cloned Genes and Their Regulatory Signals * Control of Developmental Transcription For more information, complete the coupon below * Metabolic Control of Expression and mail to: and Sporulation Dr. Robert K. Nauman * Endotoxin Formation in Sporulating Chairman, Membership Committee Box ASMN 1083- 1913 Street N.W. M Organisms Washington, DC 20006 * Microbial Differentiation Systems ______Order this valuable new reference tool for I'm interested in membership in the American Society for or soon. Microbiology. Please send information and a Membership the laboratory library Application to: Publication date: April 1985 280 pages, illustrated, index Name Hardcover (ISBN 0-914826-75-1) Address ASM member, $39.00; nonmember, $49.00 City American Society for Microbiology Publication Sales State Zip Code 1913 I Street, N.W, Washington, DC 20006 USA Country Student: El Yes O No Date