JOURNAL of VIROLOGY VOLUME 63 * DECEMBER 1989 NUMBER 12 Arnold J
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JOURNAL OF VIROLOGY VOLUME 63 * DECEMBER 1989 NUMBER 12 Arnold J. Levine, Editor in Chief Robert A. Lamb Editor (1992) (1994) Northwestern University Princeton University Stephen P. Goff, Editor (1994) Evanston, Ill. Princeton, N.J. Columbia University Michael B. A. Oldstone, Editor (1993) Joan S. Brugge, Editor (1994) New York, N. Y. Scripps Clinic & Research University ofPennsylvania Peter M. Howley, Editor (1993) Foundation Philadelphia, Pa. National Cancer Institute La Jolla, Calif. Bernard N. Fields, Editor (1993) Bethesda, Md. Thomas E. Shenk, Editor (1994) Harvard Medical School Princeton University Boston, Mass. Princeton, N.J. EDITORIAL BOARD Rafi Ahmed (1991) Emanuel A. Faust (1990) Robert A. Lazzarini (1990) William S. Robinson (1989) James Alwine (1991) S. Jane Flint (1990) Jonathan Leis (1991) Bernard Roizman (1991) David Baltimore (1990) William R. Folk (1991) Myron Levine (1991) John K. Rose (1991) Amiya K. Banerjee (1990) Donald Ganem (1991) Arthur D. Levinson (1991) Naomi Rosenberg (1989) Tamar Ben-Porat (1990) Costa Georgopolous (1989) Maxine Linial (1991) Roland R. Rueckert (1991) Kenneth I. Berns (1991) Walter Gerhard (1989) David M. Livingston (1991) Norman P. Salzman (1990) Joseph B. Bolen (1991) Mary-Jane Gething (1990) Douglas R. Lowy (1989) Charles E. Samuel (1989) Michael Botchan (1989) Joseph C. Glorioso (1989) Malcohm Martin (1989) Priscilla A. Schaffer (1990) Thomas J. Braciale (1991) Larry M. Gold (1991) Robert Martin (1990) Sondra Schlesinger (1989) Dalius J. Briedis (1991) Hidesaburo Hanafusa (1989) Warren Masker (1990) Robert J. Schneider (1991) Michael J. Buchmeier (1989) John Hassell (1989) James McDougall (1990) Bart Sefton (1991) Robert Callahan (1991) William S. Hayward (1990) Thomas Merigan (1989) Bert L. Semler (1989) Barrie J. Carter (1990) Ari H. Helenius (1990) Lois K. Miller (1991) Charles J. Sherr (1990) Sherwood Casjens (1990) Roger Hendrix (1990) Peter Model (1989) Saul J. Silverstein (1991) Bruce Chesebro (1991) James M. Hogle (1991) Bernard Moss (1989) Patricia G. Spear (1990) John M. Coffin (1989) John J. Holland (1990) Brian R. Murphy (1991) Mark F. Stinski (1989) Gary H. Cohen (1991) Kathryn V. Holmes (1991) Opendra Narayan (1991) James Strauss (1990) Charles N. Cole (1989) Nancy Hopkins (1989) Joseph R. Nevins (1991) F. William Studier (1990) Peter L. Collins (1991) Alice S. Huang (1990) Erling Norrby (1989) Bfi Sugden (1990) Neal Copeland (1989) Steve Hughes (1991) Nancy G. Nossal (1990) John M. Taylor (1990) Donald Court (1990) Eric Hunter (1990) Dennis O'Callaghan (1989) Howard M. Temin (1991) Richard Courtney (1989) Tony Hunter (1989) Joseph Pagano (1990) David A. Thorley-Lawson (1991) Clyde S. Crumpacker II Richard Hyman (1989) Peter Palese (1991) Inder Verma (1989) (1991) Nancy Jenkins (1989) Enzo Paoletti (1991) Luis V. Villarreal (1991) Bryan R. Cullen (1990) Richard A. Katz (1990) Takis Papas (1990) Volker M. Vogt (1990) Thomas Curran (1991) Jack D. Keene (1991) J. Thomas Parsons (1989) Edward K. Wagner (1989) Samuel Dales (1991) Thomas J. Kelly (1991) Ulf G. Pettersson (1989) Gail Wertz (1991) Daniel DiMaio (1991) Eli Keshet (1990) Lennart Philipson (1990) Reed B. Wickner (1991) Walter Doerfler (1989) Elliott Kieff (1990) Lewis I. Pizer (1990) Eckard Wimmer (1991) Elvera Ehrenfeld (1989) Daniel F. Klessig (1991) Carol Prives (1989) Owen Witte (1989) Robert N. Eisenman (1991) David Knipe (1991) Vincent Racaniello (1990) Charles Hamish Young (1989) Suzanne U. Emerson (1989) Daniel Kolakofsky (1989) R. Frank Ramig (1991) Harald zur Hausen (1989) Lynn W. Enquist (1991) Hsing-Jien Kung (1990) Dan S. Ray (1989) Mary K. Estes (1989) Ching-Juh Lai (1990) Solon L. Rhode (1989) Ellen Fanning (1991) Michael Lai (1991) Harriet Robinson (1991) Helen R. Whiteley, Chairman, Publications Board Linda M. Illig, Managing Editor, Journals Judith Nedrow, Production Editor The Journal of Virology (ISSN 0022-538X), a publication of the American Society for Microbiology, 1325 Massachusetts Ave., N.W., Washington, DC 20005, 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 biochemistry, biophysics, genetics, immunology, 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 Publications Department. The Journal is issued monthly, one volume per year. The nonmember subscription price is $350 per year; single copies are $30. The member subscription price is $41 (foreign, $71 [air drop shipping]) per year; single copies are $8.00. 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 (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 offailure 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. Made in the U.S.A. Printed on acid-free paper. Copyright ©3 1989, American Society for Microbiology. i9W,& r4{U (I1'4I,t 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, 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 Abe, Shinobu, 5354 Frey, Alan B., 5440 Lahn, Sharon, 5194 Rice, Nancy R., 5194 Ahlquist, Paul, 5302 Fu, Jianlin, 5492 Lai, Ching-Juh, 5086 Richards, Douglas, 5302 Ahn, Jeong K., 5425 Lai, Michael M. C., 5285 Righthand, V. Fay, 5268 Anderson, Mary E., 5175 Gallo, Michael L., 5023 Langdon, Wallace Y., 5420 Robey, W. Gerard, 5046 Arias, Carlos F., 5393 Gallo, Robert C., 5046 Langlois, Alphonse J., 5046 Roos, Raymond P., 5344, 5492 Arita, Mineo, 5354 Gao, Min, 5258 Lawson, T. Glen, 5013 Rose, John K., 5328 Arthur, Larry O., 5046 Garber, Christa, 5119 Le, Shu-Yun, 5006 Rosenblum, Betsy, 5364 Ayata, Minoru, 5464 Garcia, Gabriela, 5393 Leis, Jonathan, 5319 Rossignol, Jean-Michel, 5451, Garoff, Henrik, 4991, 5111 Lequarre, Anne-Sophie, 5194 5497 Baier, Michael, 5119 Garsky, Victor M., 5037 Levine, Arnold J., 5440 Ruusala, Aino, 5328 Bennett, Ellen R., 5371 Gavin, Brian J., 5425 Levis, Robin, 5216 Ben-Porat, Tamar, 5475 Gazzolo, Louis, 5413 Linial, Maxine L., 5092 Salhi, Samia, 5497 Berkhout, Ben, 5501 Ghattas, Ingrid, 5310 Lippe, Roger, 5133 Schaller, Heinz, 5399 Bernard, Alain, 5413 Graham, Frank L., 5133 Liu, Qingyun, 5062 Schlesinger, Sondra, 5310 Bess, Julian W., Jr., 5046 Grakoui, Arash, 5216 Lopez, Susana, 5393 Schlicht, Hans-Jurgen, 5399 Biegalke, Bonita J., 5092 Gruffat, Henri, 5276 L6pez-Turiso, Jose Angel, Schulze, Thomas, 5489 Blackburn, Robert V., 5268 Grumont, Raelene J., 5420 5445 Semler, Bert L., 5344, 5492 Boere, Wil A. M., 4991 Grundy, Frank J., 5101 Lowy, Douglas R., 5469 Bolen, Joseph B., 5076 Grunhaus, Avraham, 5455 Sergeant, Alain, 5276 Bray, Michael, 5086 Maizel, Jacob, 5006 Sexton, Connie J., 5505 Brunner, Dieter, 5483 Hacker, David, 5166 Makino, Shinji, 5285 Shaw, Andrey, 5328 Buisson, Monique, 5276 Haddad, Ruwaida S., 4998 Malim, Michael H., 5006 Siddiqui, Aleem, 5293 Bukowski, Jack F., 5086 Haenni, Anne-Lise, 5153 Manet, Evelyne, 5276 Skalka, Anna Marie, 5319 Hagino-Yamagishi, Kimiko, Martinez, Concepci6n, 5445 Smith, Lisa L., 5013 Callahan, Pia L., 5037 5354, 5386 McBride, Alison A., 5076 Sodora, Donald L., 5184 Capone, John P., 5509 Hartle, Heather T., 5054 Metsikko, Kalervo, 5111 Spector, Deborah H., 5334 Carlier, Damien, 5497 Hartung, Susanne, 5119 Mettenleiter, Thomas C., 5475 Srinivasan, Ashok, 5459 Casey, James W., 5194 Harty, Ronald N., 5101 Moelling, Karin, 5489 Stacy, Terryl, 5208 Chamberlain, Michele, 5208 Hassell, John A., 5371 Mondesert, G., 5124 Stein, Steven, 5492 Chen, Chu, 5092 Hauber, Joachim, 5006 Morch, Marie-Dominique, Steinhauer, David A., 5030 Cichutek, Klaus, 5119 Hirano, Akiko, 5464 5153 Stephens, Robert M., 5194 Clarke, David, 5030 Holland, John J., 5030 Morin, Nathalie, 5228 Sugg, Nancy, 5475 Cohen, Gary H., 5184 Honda, Ayae, 5142 Morse, Herbert C., III, 5420 Sugimura, Haruhiko, 5405 Cohen, Jeffrey I., 5364 Horwitz, Marshall S., 5244, Muller, Carolin, 5119 Summers, William C., 5062 Cole, Charles N., 5208 5455 Murdin, Andrew D., 5251 Szafranski, Przemyslaw, 5153 Colle, Clarence F., 5101 Howley, Peter M., 5076 Colonno, Richard J., 5037 Huang, Henry V., 5216 Nara, Peter L., 5046 Tago, Katsuhiko, 5354 Cordingley, Michael G., 5037 Hutt-Fletcher, Lindsey M., Naujokas, Monica,