JOURNAL OF VOLUME 68 * APRIL 1994 * NUMBER 4

Arnold J. Levine, Editor in Chief (1994) Vincent R. Racaniello, Editor (1997) Princeton University Columbia University Princeton, N.J. Peter M. Howley, Editor (1998) New York, N.Y Harvard Medical School John M. Coffin, Editor (1996) Boston, Mass. Thomas E. Shenk, Editor (1994) Tufts University Medical School Princeton University Boston, Mass. Michael B. A. Oldstone, Editor (1994) Princeton, N.J. Scripps Clinic & Research Foundation Ronald C. Desrosiers, Editor (1998) La Jolla, Calif. Joseph Sodroski, Editor (1998) Harvard Medical School Dana-Farber Cancer Institute Boston, Mass. Carol Prives, Editor (1996) Boston, Mass. Columbia University Mary K. Estes, Editor (1998) New N.Y Inder M. Verma, Editor (1998) Baylor College of Medicine York, The Salk Institute Houston, Tex. San Diego, Calif.

EDITORIAL BOARD Rafi Ahmed (1994) Harry B. Greenberg (1995) Elizabeth Moran (1996) Priscilla A. Schaffer (1996) Carl C. Baker (1994) Hidesaburo Hanafusa (1995) Bernard Moss (1995) Heinz Schaller (1995) Amiya K. Banerjee (1996) Ed Harlow (1996) Richard W. Moyer (1995) Sondra Schlesinger (1995) Kenneth L. Berns (1994) Gary S. Hayward (1996) James Mullins (1996) Robert J. Schneider (1994) Joseph B. Bolen (1994) Ari H. Helenius (1996) Brian R Murphy (1994) Manfred H. Schubert (1996) Thomas J.'Braciale (1994) Virginia S. Hinshaw (1996) Nicholas A. Muzyczka (1996) Bartholomew M. Sefton (1994) John Brady (1994) David D. Ho (1996) Gerald Myers (1995) Bert L. Semler (1995) Thomas R. Broker (1995) James M. Hogle (1994) Gary J. Nabel (1996) David A. Shafritz (1994) Michael J. Buchmeier (1995) John J. Holland (1996) Opendra Narayan (1994) Yosef Shaul (1995) Robert Callahan (1994) Kathryn V. Holmes (1994) Neal Nathanson (1996) Saul J. 'Silverstein (1994) Barrie J. Carter (1996) Nancy Hopkins (1995) Jay A. Nelson (1994) Daniel T. Simmons (1995) Mark Challberg (1995) Stephen H. Hughes (1994) Joseph R. Nevins (1994) Anna Marie Skalka (1996) Irvin S. Y. Chen (1996) Eric Hunter (1996) Dennis O'Callaghan (1995) Nahum Sonenberg (1996) Bruce W. Chesebro (1994) Tony Hunter (1995) Peter Palese (1994) Arne Stenlund' (1995) Donald M. Coen (1995) Nancy A. Jenkins (1995) Enzo Paoletti (1994) Mark F. Stinskj (1995) Gary H. Cohen (1-994) Richard A. Katz (1996) Takis S. Papas (1996) Jonathan P. Stoye (1996) Charles N. Cole (1995) Jack D. Keene (1994) Tristram G. Parslow (1996) James H. Strauss (1996) Peter L Collins (1994) Thomas J. Kelly (1994) J.' Thomas Parsons (1995) Bill Sugden (1996) Richard C. Condit (1995) Elliott Kief (1996) C. David Pauza (1996) Jesse Summers (1995) Neal G. Copeland (1995) Karla Kirkegaard (1996) David J. Pintel (1995) Ron Swanstrom (1996) Richard J. Courtney (1995) Geofy K Kitchingman (1995) Abraham Pinter (1994) Peter J. Tattersall (1995) Clyde S. Crumpacker II (1994) Daniel F. Klessig (1994) James M. Pipas (1996) John M. Taylor (1996) Bryan R, Cullen (1996) David M. Knipe (1994) Lewis I. Pizer (1996) Mary J. Tevethia (1996) Thomas Curran (1994) Hsing-Jien Kung (1996) Nancy Raab-Traub (1996) Satvir S. Tevethia (1994) Samuel Dales (1994) Ching-Juh Lai (1996) Arnold B. Rabson (1996) David A. Thorley-Lawson (1994) Daniel DiMaio (1994) Michael Lai (1994) R Frank Ramig (1994) Paula Traktman (1995) Walter Doerfler (1995) Jonathan Leis (1994) Dan S. Ray (1995) Didier Trono (1996) Elvera Ehrenfeld (1995) Jack Lenz (1996) Alan R. Rein (1996) Philip N. Tsichlis (1995) Robert N. Eisenman (1994) Norman L Letvin (1995) Carol S. Reiss (1996) Kenneth L. Tyler (1994) John Elder (1996) Maxine Linial (1994) Solon L Rhode (1995) Luis V. Villarreal (1994) Michael Emerman (1995) David M. Livingston (1994) William S. Robinson (1995) Peter K. Vogt (1996) Suzanne U. Emerson (1995) Douglas R Lowy (1995) Bernard Roizman (1994) Edward K. Wagner (1995) Lynn W. Enquist (1994) Paul J. Maddon (1996) John K. Rose (1994) Robert G. Webster (1996) Hung Fan (1994) James L Manley (1995) Craig Rosen (1996) Raymond M. Welsh (1994) Ellen Fanning (1994) Malcolm A. Martin (1995) Naomi Rosenberg (1995) Gail Wertz (1994) Bernard Fleckenstein (1996) William S. Mason (1994) Barry T. Rouse (1996) J. Lindsay Whitton (1995) Jane Flint (1996) Michael B. Mathews (1994) Roland R Rueckert (1994) Reed B. Wickner (1994) William R Folk (1994) Alan McLachlan (1996) Norman P. Salzman (1996) John W. Wills (1995) Paul D. Friesen (1994) A. Dusty Miller (1994) Charles E. Samuel (1995) Eckard Wimmer (1994) Denise A. Galloway (1995) Lois I. Miller (1994) Richard Jude Samulski (1995) Flossie Wong-Staal (1996) Don Ganem (1994) Edward S. Mocarski (1996) Peter Sarnow (1996) Moshe Yaniv (1995) Costa P. Georgopoulos (1995) Peter Model (1995) Nava Sarver (1996) Charles Hamish Young (1995) Joseph C. Glorioso (1995) John P. Moore (1996) Dorothea L Sawicki (1994) Harald zur Hausen (1995)

Barbara H. Iglewski, Chairman, Publications Board Linda M. Illig, Director, Journals Judith Nedrow, Production Editor Arthur Gelmis, Assistant Production Editor

The Journal of Virology, a publication of the American Society for (ASM), 1325 Massachusetts Ave., N.W., Washington, DC 20005-4171, is devoted to the dissemination of fundamental knowledge concerning viruses of bacteria, plants, and animals. Investigators are invited to submit reports of original research in all areas of basic virology, including , biophysics, , , morphology, physiology, and pathogenesis and immunity. Instructions to authors are published in the January issue each year; reprints are available from the editors and the Journals Division. The Journal is issued monthly, one volume per year. The nonmember print subscription prices are $380 (U.S.) (Canadians add 7% GST) and $454 (other countries) per year; single copies are $43 (Canadians add 7% GST). The member print subscription prices are $81 (U.S.) (Canadians add 7% GST) and $106 (other countries); single copies are $11 (Canadians add 7% GST). For prices of CD-ROM versions, contact the Subscriptions Unit, ASM. Correspondence relating to subscriptions, defective copies, missing issues, and availability of back issues should be directed to the Subscriptions Unit, ASM; correspondence relating to reprint orders should be directed to the Reprint Order Unit, ASM; and correspondence relating to disposition of submitted manuscripts, proofs, and general editorial matters should be directed to the Journals Division, American Society for Microbiology, 1325 Massachusetts Ave., N.W., Washington, DC 20005-4171. Phone: (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 Journal of Virology, ASM, 1325 Massachusetts Ave., N.W., Washington, DC 20005-4171. Made in the U.S.A. Printed on acid-free paper. ISSN 0022-538X CODEN: JOVIAM Copyright © 1994, 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 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, 222 Rosewood Drive, Danvers, MA 01923, 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

Aasted, Bent, 2720 Dow, Steven W., 2230 Jansen-Diirr, Pidder, 2206 Marion, Patricia L., 2091 Adelmant, Guillaume, 2097 Dubin, Gary, 2478 Javaherian, Kashi, 2624 Markham, Phillip, 2693 Akahane, Yoshihiro, 2671 Jenkins, Nancy A., 2320 Martin, Jennifer E., 2649 Albrecht, Julia, 2142 Eddleston, Michael, 2383 Jeong, Yong Seok, 2615 Martin, Malcolm A., 2503 Alexandersen, Soren, 2720 Eilers, Martin, 2206 Jiang, Min, 2065 Mason, William S., 2286 Alford, Patricia, 2693 Einfeld, David A., 2513 Johansen, Bjarne, 2398 Mayumi, Makoto, 2671 Allan, Jonathan, 2693 Eisenberg, Roselyn J., 2529 Johansen, Tere, 2398 McDougall, Ann S., 2151 Aloni, Yosef, 2741 Elkins, William R., 2649 Johnson, Marion D., III, 2331 McGregor, Martha W., 2051 Alonso-Caplen, Firelli, 2433 Engelke, Ute, 2752 Johnson, Philip R., 2649 McKnight, Kevin L., 2683 Angliker, Herbert, 2772 Ensoli, Barbara, 2677 Johnston, Robert E., 2683 Mehdi, Haider, 2415 Anlar, Fehim Yasar, 2415 Erickson, Ann L., 2756 Joswig, Silvia, 2206 Menendez-Arias, Luis, 2313 Menzo, Stefano, 2495 Bachmann, Barbara, 2765 Fauci, Anthony S., 2598 Kabat, David, 2570 Miller, Lois K., 2521 Baer, Margaret L., 2194 Fink, D. J., 2239 Kaiwar, Ravi, 2118 Mindich, Leonard, 2331 Bagnarelli, Patrizia, 2495 Folks, Thomas M., 2726 Kalyanaraman, Vaniambadi, Miyakawa, Yuzo, 2671 Balfe, Peter, 2161 Franchini, Genoveffa, 2693 2693 Mizushima, Hiroto, 2731 Bargmanti, William, 2073 Freed, Eric O., 2503 Kaplan, Michael J., 2415 Moens, Ugo, 2398 Basu, Mitali, 2478 Friedman, H. M., 2478 Katzenstein, Gil E., 2135 Monastyrskaya, Katherine, Basu, Saswata, 2478 Friedrich, Thomas D., 2355 Kaushal, S., 2468 2169 Bayer, Robert, 2415 Fullen, Jake, 2693 Kemper, Phillip, 2383 Montefiori, David C., 2362, Bear, Susan E., 2320 Kent, Karen A., 2624 2624 Bellacosa, Alfonso, 2320 Gallo, Robert C., 2693 Kim, Carol H., 2777 Morin, Nathalie, 2589 Ben-Asher, Edna, 2741 Ganem, Don, 2091 Kimmel, Paul L., 2224 Morita, Elaine, 2662 Birnbaum, Mark J., 2521 Garten, Wolfgang, 2772 Kimura, Koichi, 2731 Morrison, Mary E., 2578 Biswas, Priscilla, 2598 Gehrke, Lee, 2194 Kinchington, P. R., 2468 Mosca, J. D., 2468 Boeri, Enzo, 2693 Gessain, Antoine, 2693 Kitchell, Barbara E., 2230 Muflins, James I., 2161, 2230 Bolognesi, Dani P., 2624 Ghiasi, Homayon, 2118, 2347 Kleiman, Lawrence, 2065 Murphey-Corb, Michael, 2624 Booth, Timothy, 2169 Ghosh, Hara P., 2186, 2272 Kleine, Kerstin, 2142 Murthy, Krishna, 2693 Bose, Henry R., Jr., 2073 Ghosh, Kakoli, 2272 Klenk, Hans-Dieter, 2772 Bremer, Christine, 2179 Ghysdael, Jacques, 2632 Koito, Atsushi, 2253 Nehyba, Jiri, 2039, 2371 Brooks, Diane M., 2059 Gilbert, Renald, 2272 Kbppe, Britta, 2313 Neil, James C., 2151 Brown, Katherine, 2135 Ginsberg, Harold S., 2127 Koralnik, Igor J., 2693 Nel, Louis, 2169 Broyles, Steven S., 2605 Giraldo, Gaetano, 2677 Kozak, Susan L., 2570 Nesburn, Anthony B., 2118, Bruggeman, Cathrien A., Glorioso, J. C., 2239 Kralova, Jarmila, 2073 2347 2297, 2305 Goins, W. F., 2239 Krause, P. R., 2239 Nishio, Jane, 2059 Buonaguro, Franco M., 2677 Goldstein, Simoy, 2649 Krauskopf, Anat, 2741 Buonaguro, Luigi, 2677 Grez, Manuel, 2161 Krug, Robert M., 2425, 2433 O'Brien, Stephen J., 2693 Burch, John B. E., 2286 Guo, Hong-Guang, 2693 Kumar, Ajit, 2224 Ohresser, Marc, 2589 Bumy, Arsene, 2632 Kuroki, Kazuyuki, 2091 Ohuchi, Masanobu, 2772 Bushman, Frederic D., 2215 Haagmans, Bart L., 2297, Okamoto, Hiroaki, 2671 Butera, Salvatore T., 2726 2305 Laffin, Judith A., 2355 Oldstone, Michael B. A., 2383 Hammarskjbld, Marie-Louise, Lagrou, Christian, 2097 Orenstein, Jan M., 2598 Cai, Jun-Ming, 2194 2065 Lam, Lee, 2726 Oroszlan, Stephen, 2313 Calvert, Jacqueline, 2084 Harrowe, Greg, 2253 Langlois, Alphonse J., 2624 Ortmann, D6rte, 2772 Campbell, lain L., 2383 Hasenkrug, Kim J., 2059 LaRosa, Gregory J., 2624 Overbaugh, Julie, 2458 Capone, John P., 2339 Hayward, G. S., 2468 Laudet, Vincent, 2097 Carrington, James C., 2388 He, Yuan-Jing, 2578 Lazo, Pedro A., 2320 Pagano, Michele, 2206 Casey, James W., 2777 Heidner, Hans W., 2683 Lee, Chang Woo, 2708 Pan, Reyes, 2746 Caughey, Byron, 2135 Hendricks, R. L., 2239 Lee, Ki Jeong, 2708 Paras, Peter, Jr., 2632 Cerutti, Martine, 2589 Hijikata, Makoto, 2731 Lees, Emma, 2206 Parent, Leslie J., 2556 Chang, Jun, 2708 Hinshaw, Virginia S., 2051 Lehman, John M., 2355 Patriotis, Christos, 2320 Cheney, Carolyn, 2151 Hirscb, Vanessa M., 2649, Lengert, Maike, 2142 Peeples, Mark E., 2415 Cheng-Mayer, Cecilia, 2253 2693 Letvin, Norman L., 2362 Pennington, Michelle J., 2605 Chesebro, Bruce, 2059, 2570 Hobbs, Monte V., 2383 Levine, M., 2239 Perera, L. P., 2468 Cheung, Ramsey, 2091 Hodge, Thomas, 2726 Levy, Jay A., 2253 Petry, Harald, 2765 Chiang, Hsien-Yuan, 2529 Hogle, James M., 2578 Li, Jing, 2605 Picard, Didier, 2206 Claassen, Eric, 2305 Homa, Fred L., 2442 Liem, Karel F., Jr., 2194 Poli, Guido, 2598 Clem, Rollie J., 2521 Hoover, Edward A., 2230, Lin, Wenyu, 2362 Popova, Bilyana, 2339 Clementi, Massimo, 2495 2458 Linenberger, Michael L., 2458 Pugh, John C., 2487 Cohen, Gary H., 2529 Horikita, Minoru, 2671 Lipsick, Joseph S., 2752 Connolly, Lynn, 2760 Horzinek, Marian C., 2297, Liu, Chen, 2286 Qian, Xiao-Yan, 2433 Cooley, A. James, 2051 2305 Lo Monico, Anita, 2693 Qiu, Yun, 2425 Copeland, Neal G., 2320 Houser-Scott, Felicia, 2194 London, William T., 2649 Quatannens, Brigitte, 2097 Cox, Nancy J., 2051 Hrdlitkovh, Radmila, 2039, Lukas, Jiri, 2206 Croen, K. D., 2239 2371 Luke, Wolfgang, 2765 Racaniello, Vincent R., 2578 Huang, Liping, 2108 Racz, Paul, 2362 Dapolito, George, 2649 Humphries, Eric H., 2039, Mahambre, Gurudas, 2161 Ransil, Bernard J., 2362 Datta, Ketaki, 2320 2371 Mak, Johnson, 2065 Rasile, Len, 2272 Davis, Nancy L., 2683 Hunsmann, Gerhard, 2765 Makino, Shinji, 2615 Raymond, Gregory J., 2135 DeGasperi, Rita, 2662 Hunter, Eric, 2513 Mallory, Diana L. P., 2478 Regnery, Helen L., 2051 Delsert, Claude, 2589 Hutchison, Michele R., 2662 Malone, Cheryl L., 2108 Reimann, Keith A., 2362 Delwart, Eric L., 2161 Maniar, Janak K., 2161 Rekosh, David, 2065 Dietrich, Ursula, 2161 Itoh, Keiichi, 2671 Manzin, Aldo, 2495 Resh, Marilyn D., 2556 i ii AUTHOR INDEX J. VIROL.

Restrepo-Hartwig, Maria A., Shimotohno, Kunitada, 2731 Tenner-Racz, Klara, 2362 Wechsler, Steven L., 2118, 2388 Shpaer, Eugene G., 2230 Terry, Anne, 2151 2347 Roberts, Beverly D., 2726 Shukla, Ram R., 2224 Thomas, Claire, 2179 Wehrly, Kathy, 2570 Rodriguez, Javier M., 2746 Simmons, Heidi, 2487 Thompson, Betsy L., 2051 Wentworth, David E., 2051 Rodriguez, Jose F., 2746 Slanina, Susan, 2118, 2347 Thomsen, Darrell R., 2442 Wernert, Nicolas, 2097 Rohn, Jennifer L., 2458 Slattery, Jill Pecon, 2693 Thresher, Wayne, 2135 Wien, Michelle W., 2578 Rojko, Jennifer, 2151 Smibert, Craig A., 2339 Tolle, Tanja, 2765 Wills, John W., 2556 Roof, Lori L., 2442 Smiley, James R., 2339 Tommasino, Massimo, 2206 Wong, Paul K. Y., 2735 Rosas, Diana H., 2662 Sodora, Donald L., 2230 Torpier, Gerard, 2097 Wyman, Peggy D., 2624 Rosl, Frank, 2142 Sosa, Miguel A. Gama, 2662 Traavik, Terje, 2398 Roy, Polly, 2169, 2179 Spangrude, Gerald J., 2059 Tsichlis, Philip N., 2320 Xiao, Peter, 2339 Rubsamen-Waigmann, Helga, Spitkovsky, Dimitry, 2206 Tsuda, Fumio, 2671 Xu, Xiyan, 2051 2161 Stals, Frans S., 2297, 2305 Tzavaras, Theodore, 2151 Ruprecht, Ruth M., 2662 Stehelin, Dominique, 2097 Ryan, Kelly A., 2624 Steiner, Philipp, 2206 Yamamoto, Kayoko, 2671 Sternberg, L. R., 2239 Valenza, Anna, 2495 Yaniez, Rafael J., 2746 Saha, Kunal, 2735 Stinski, Mark F., 2108 van den Eertwegh, Alfons J. Yang, Xu, 2415 Salas, Maria L., 2746 Stott, James, 2624 M., 2305 Yasutomi, Yasuhiro, 2362 Saule, Simon, 2097 Straus, S. E., 2239, 2468 van der Meide, Peter H., Yotsumoto, Shigeru, 2671 Saxinger, W. Carl, 2693 Strebel, Klaus, 2260 2297, 2305 Young, John A. T., 2760 Scalise, Giorgio, 2495 Subramaniam, Nanthakumar, Van Lint, Carine, 2632 Yuen, P. H., 2735 Scarano, Frank J., 2355 2398 Varaldo, Pietro E., 2495 Schatzle, John D., 2073 Summers, Jesse, 2084 Verdin, Eric, 2632 Zawatzky, Rainer, 2142 Schijns, Virgil E. C. J., 2297, Sung, Young Chul, 2708 Viniuela, Eladio, 2746 Zhang, Jiayou, 2409 2305 Sutherland, Karen, 2415 Viuff, Birgitte, 2720 Zhang, Li, 2186 Schneider, Robert J., 2544 von Briesen, Hagen, 2161 Zhang, Yan, 2544 Schraven, Burkhart, 2142 Tal-Singer, Ruth, 2478 Zhao, Yuan, 2179 Schubert, Ulrich, 2260 Tang, Qingquan, 2127 Wainberg, Mark A., 2065 Zhou, Wenjun, 2556 Schulze, Almut, 2206 Tanji, Yasunori, 2731 Walker, Christopher M., 2756 Zingler, Kurt, 2760 Shaw, Elliott, 2772 Temin, Howard M., 2409 Wang, Beibei, 2215 zur Hausen, Harald, 2142 f

= 1-3 Infection and 1 or Immunity ff:/i- U Editor in Chief: Graham C Walker Editorin Chief: rincentA. Fischetti JB sovers a broad area of IAI continues to publish a broad range of subjects, including structure topics important to immunologists, epi- and function-, plant microbi- demiologists, pathologists, and clini- ology, cell surfaces, eukary- cians. Subjects include infections otic cells, genetics and mo- caused by pathogenic bacteria, fungi, lecular biology, population and unicellular parasites; ecology, epi- genetics and evolution, plasmids and demiology, and evolution of pathogenic microbes; --i transposons, bacteriophages, physiology and me- mechanisms of pathogenicity and virulence factors such as tabolism, and enzymes and proteins. MinQeviews are a regular toxins and microbial surface structures; nonspecific factors in feature. host resistance-and susceptibility to infection; immunology of microbial infection; and development and evaluation of vac- cines against pathogens. ISSN 0021-9193. 8,400 pages/year. Twice monthly. Members of ASM may subscribe at $79.00 (U.S.A.) (Canadians add 7%O GST), $104.00 (other countries). ISSN 0019-9567. 5,550 pages/year. Monthly. Members of ASM may subscribe at $51.00 (U.S.A.) (Canadians add 7% GST), $89.00 (other countries). Journal of Molecular and Virology Cellular JVI maintains its premier position as a source of broad-based, high-qual- Editorin Chief:Alan M. Weiner ity, original research con- Researchers find MCB to be an author- cerning viruses of bacte- itative source of fundamental knowl- ria, plants, and animals: edge and new developments in all The journal provides aspects of the of fundamental new informa- eukaryotic cells, of both microbial and higher organisms. The tion using cross-disciplinary approaches scope includes regulation of gene expression, transcription, of biochernistry, biophysics, cell biology, genetics, immunol- cell growth and development, oncogenesis, cell and.organelle ogy, molecular biology, morphology, physiology, and patho- structure and assembly, DNA replication and recombination, genesis and immunity. Specific sections include: Viral and chromosome structure, and somatic cell genetics. The journal Cellular Oncogenes, Viral Pathogenesis and Immunity, Virus- publishes work on microbial as well as higher organisms and Cell Interactions, Animal Viruses, Bacterial Viruses, Plant on viral systems where the emphasis is clearly on the cell. Viruses, and Protozoan and Fungal Viruses. Also included are minireviews. ISSN 0270-7306. 8,200 pages/year. Monthly. Members of ASM may subscribe at $80.00 (U.S.A.) (Canadians add 7% GST), $105.00 (other countries). ISSN 0022-538X. 8,100 pages/year. Monthly. Members of ASM may subscribe at $81.00 (U.S.A.) (Canadians add 7o GST), $106.00 (ofer countries).

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