JOURNAL OF VOLUME 67 * AUGUST 1993 * NUMBER 8 Arnold J. Levine, Editor in Chief (1994) Peter M. Howley, Editor (1998) Vincent R. Racaniello, Editor (1997) Princeton University National Cancer Institute Columbia University Princeton, N.J. Bethesda, Md. New York, N.Y. Robert A. Lamb, Editor (1997) John M. Coffin, Editor (1996) Northwestern University Thomas E. Shenk, Editor (1994) Tufts University Medical School Evanston, Ill. Princeton University Boston, Mass. Princeton, N.J. Michael B. A. Oldstone, Editor (1994) Ronald C. Desrosiers, Editor (1998) Scripps Clinic & Research Foundation Inder M. Verma, Editor (1998) Harvard Medical School La Jolla, Calif. The Salk Institute Boston, Mass. Carol Prives, Editor (1996) San Diego, Calif. Columbia University New York, N. Y. EDITORIAL BOARD Rafi Ahmed (1994) Paul D. Friesen (1994) Edward S. Mocarski (1993) Manfred H. Schubert (1993) James Alwine (1994) Denise A. Galloway (1995) Peter Model (1995) Bartholomew M. Sefton (1994) Carl C. Baker (1994) Don Ganem (1994) Bernard Moss (1995) Bert L. Semler (1995) Amiya K. Banerjee (1993) Costa P. Georgopoulos (1995) Richard W. Moyer (1995) David A. Shafritz (1994) Kenneth I. Berns (1994) Joseph C. Glorioso (1995) James Mullins (1993) Yosef Shaul (1995) Joseph B. Bolen (1994) Harry B. Greenberg (1995) Brian R. Murphy (1994) Saul Thomas J. Braciale (1994) Hidesaburo Hanafusa (1995) Nicholas A. Muzyczka (1993) J. Silverstein (1994) John Brady (1994) Ed Harlow (1993) Gerald Myers (1995) Daniel T. Simmons (1995) Thomas R. Broker (1995) An H. Helenius (1993) Opendra Narayan (1994) Anna Marie Skalka (1993) Michael J. Buchmeier (1995) Virginia S. Hinshaw (1993) Neal Nathanson (1993) Joseph Sodraski (1995) Robert Callahan (1994) James M. Hogle (1994) Jay A. Nelson (1994) Nahum Sonenberg (1993) Barrie J. Carter (1993) John J. Holland (1993) Joseph R. Nevins (1994) Arne Stenlund (1995) Mark Challberg (1995) Kathryn V. Holmes (1994) Dennis O'Caliaghan (1995) Mark F. Stinski (1995) Irvin S. Y. Chen (1993) Nancy Hopkdns (1995) Peter Palese (1994) James Strauss (1993) Bruce W. Chesebro (1994) Alice S. Huang (1993) Enzo Paoletti (1994) Bill Sugden (1993) Donald M. Coen (1995) Stephen H. Hughes (1994) Takis Papas (1993) Jesse Summers (1995) Gary H. Cohen (1994) Eric Hunter (1993) J. Thomas Parsons (1995) Ron Swanstrom (1993) Charles N. Cole (1995) Tony Hunter (1995) David J. Pintel (1995) Peter J. Tattersall (1995) Peter L. Collins (1994) Nancy A. Jenkins (1995) Abraham Pinter (1994) John M. Taylor (1993) Richard W. Compans (1993) Richard A. Katz (1993) James Pipas (1993) Howard M. Temin (1994) Richard C. Condit (1995) Jack D. Keene (1994) Lewis I. Pizer (1993) Mary J. Tevethia (1993) Neal G. Copeland (1995) Thomas J. Kelly (1994) Nancy Raab-Traub (1993) Satvir S. Tevethia (1994) Richard J. Courtney (1995) Elliott Kieff (1993) Arnold B. Rabson (1993) David A. Thorley-Lawson Clyde S. Crumpacker 11(1994) Karla Kirkegaard (1993) R. Frank Ramig (1994) (1994) Bryan R. Cullen (1993) Geoffrey R. Kitchingman (1995) Dan S. Ray (1995) Paula Traktman (1995) Thomas Curran (1994) Daniel F. Klessig (1994) Alan Rein (1993) Philip N. Tsichlis (1995) Samuel Dales (1994) David M. Knipe (1994) Solon L. Rhode (1995) Kenneth L. Tyler (1994) Daniel DiMaio (1994) Hsing-Jien Kung (1993) William S. Robinson (1995) Luis V. V'illarreal (1994) Walter Doerfler (1995) Ching-Juh Lai (1993) Bernard Roizman (1994) Peter K. Vogt (1993) Elvera Ehrenfeld (1995) Michael Lai (1994) John K. Rose (1994) Edward K. Wagner (1995) Robert N. Eisenman (1994) Jonathan Leis (1994) Craig Rosen (1993) Robert G. Webster (1993) John Elder (1993) Norman L. Letvin (1995) Naomi Rosenberg (1995) Raymond M. Welsh (1994) Michael Emerman (1995) Maxine Linial (1994) Roland R. Rueckert (1994) Gail Wertz (1994) Suzanne U. Emerson (1995) David M. Livingston (1994) Norman P. Salzman (1993) J. Lindsay Whitton (1995) Lynn W. Enquist (1994) Douglas R. Lowy (1995) Charles E. Samuel (1995) Reed B. Wickner (1994) Mary K. Estes (1995) James L. Manley (1995) Richard Jude Samulski (1995) John W. Wills (1995) Hung Fan (1994) Malcolm A. Martin (1995) Dorothea L. Sawicki (1994) Eckard Wimmer (1994) Ellen Fanning (1994) William S. Mason (1994) Priscilla A. Schaffer (1993) Flossie Wong-Staal (1993) Bernard Fleckenstein (1993) Michael B. Mathews (1994) Heinz Schaller (1995) Moshe Yaniv (1995) S. Jane Flint (1993) A. Dusty Miller (1994) Sondra Schlesinger (1995) Charles Hamish Young (1995) William RI Folk (1994) Lois K. Miller (1994) Robert J. Schneider (1994) Harald zur Hausen (1995)

Barbara H. Iglewski, Chairman, Publications Board Linda M. Illig, Director, Journals Judith Nedrow, Production Editor The Journal of Vrology, 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 $362 (U.S.) (Canadians add 7% GST) and $432 (other countries) per year; single copies are $40 (Canadians add 7% GST). The member print subscription prices are $81 (U.S.) (Canadians add 7% GST) and $106 (other countries); single copies are $10 (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 Copyright © 1993, American Society for Microbiology. CODEN: JOVIAM 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, 27 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

Aiken, Christopher, 4945 Garcia, J. Victor, 4923 La Rosa, Corinna, 4742 Pavlovic, Jovan, 4760 Akagi, Tsuyoshi, 4665 Garzelli, Carlo, 4742 Lausch, Robert N., 4777 Pensaert, Maurice B., 4492 Akerblom, Lennart, 4785 Ghosh, Hara P., 4856 Lee, Stephen C., 4566 Perera, L. P., 4474 Albright, Allen G., 4842 Ghosh, Kakoli, 4856 Lenches, Edith, 5003 Perez, Luis, 4543 Anderson, Michelle M., 4964 Gilbert, Ilka, 4992 Lewis, George K., 4785 Pfeiffer, Brian, 4611 Anderson, Stephen, 4923 Gine, Nuria R., 5045 Li, Guangpu, 5062 Pinter, A., 4932 Anderson, W. French, 4712 Glueck, Deborah H., 4639 Liang, Xiaoping, 4896 Pletnev, Alexander G., 4956 Apt, Doris, 4455 Gombold, James L., 4504 Liu, Yichun, 4455 Poma, Eric, 4981 Arnheiter, Heinz, 4484 Gottlieb, Paul, 4914 Lombardi, Stefania, 4742 Poon, Alice P. W., 4497 Arthur, Avril K., 4992 Graham, S., 4932 Longnecker, Richard, 5068 Poon, Shu-Wing, 4785 Green, Maurice, 4804 Lu, Meng-Ji, 4905 Posner, Marshall, 4557 Babiuk, Lorne A., 4896 Green, William R., 5045 Luckow, Verne A., 4566 Prives, Carol, 4750 Balfe, P., 4932 Griffin, Diane E., 4605 Lyles, Douglas S., 4814 Bamford, Dennis H., 4696 Grose, Charles, 4464 Qiao, Xueying, 4914 Banham, Alison H., 4549 Gu, Baohua, 4676 Maldarelli, Frank, 5056 Barry, Gerard F., 4566 Malvaldi, Gino, 4742 Ramsingh, Arlene, 4797 Bastin, Marcel, 5025 Hadlock, Kenneth G., 4659 Manfredi, James J., 4750 Ramsperger, Uwe, 4992 Bates, Patricia, 4676 Haenggi, Markus, 5041 Mangold, Constance M. T., Randall, Sandra K., 4992 Baylis, Sally A., 4549 Hannibal, Mark C., 5035 4588 Rasile, Len, 4856 Bendinelli, Mauro, 4742 Hanssens, Filip P., 4492 Mansell, Eric A., 4848 Rassart, Eric, 4722 Bernard, Hans-Ulrich, 4455 Hart, Clyde E., 5020 Markovitz, David M., 5035 Raviprakash, Kanakatte, 4856 Berneman, Zwi N., 4611 Hay, J., 4474 Martin, Annette, 4630 Renne, Rolf, 4598 Black, Brian L., 4814 Hayward, G. S., 4474 Masuda, Michiaki, 4580 Renshaw-Gegg, Lisa W., 4598 Blair, D. G., 4704 Hayward, S. Diane, 4875 Masuda, Tohru, 4533 Repke, Heinrich, 4557 Blair, Wade S., 5030 Hellen, Christopher U. T., Matsubayashi, Yuji, 4533 Rhodes, Richard B., 4814 Blondel, Bruno, 4630 4688 McCance, Dennis J., 4521 Ricciardi, Robert P., 4804 Bogerd, Hal P., 5030 Henning, Ulf, 4905 McKeating, J. A., 4932 Rice, Charles M., 5062 Bolmstedt, A., 4932 Herchenroeder, Ottmar, 4598 McKenzie, Margie, 4814 Rivera-Gonzalez, Ramon, Bray, Michael, 4956 Hijikata, Makoto, 4665 Mehigh, Christine S., 4964 4676 Bridger, J. C., 5012 Hingley, Susan T., 4504 Mergia, Ayalew, 4598 Robertson, Michael N., 4533 Hinkle, Patricia M., 4521 Miao, X-Qian, 5068 Robinson, James, 4557 Caggana, Michele, 4797 Ho, David D., 4557, 4785 Middeldorp, Jaap M., 4875 Roehl, Holger H., 4964 Calain, Philippe, 4822 Hoffman, Paul M., 4580 Miller, Cheryl L., 5068 Rogove, Andrew, 5075 Carrasco, Luis, 4543 Hogle, James, 5075 Mindich, Leonard, 4914 Roizman, Bernard, 4497 Chan, Perry, 4797 Hopfner, Matthias, 4992 Mitchell, William J., 4484 Rossignol, Jean-Michel, 4886 Chandran, Bala, 4611 Horaud, Florian, 4630 Miyazawa, Masaaki, 4533 Roth, Michael G., 4831 Chang, Cheow K., 4611 Horisberger, Michel A., 4760 Mizushima, Hiroto, 4665 Roux, Laurent, 4822 Charles, MacArthur, 4785, Howard, C. J., 5012 Moarefi, Ismail F., 4992 Ru, Ming, 4722 4932 Hua, Jian, 4848 Molla, Akhteruzzaman, 4688 Ruscetti, Sandra K., 4580 Chen, Mao-Yuan, 5056 Huang, Eng-Shang, 4981 Monteiro, Carlos A., 4777 Russo, Alicia A. R., 4992 Chen, Mei-Ru, 4875 Huong, Shu-mei, 4981 Moore, John P., 4785 Ruyechan, W. T., 4474 Chesebro, Bruce, 4533 Moran, Elizabeth, 4804 Chong, Terence, 4455 Sattentau, Quentin J., 4785 Isaacs, Stuart N., 4732 Morgan, Richard A., 4712 Harri, 4696 Chow, Marie, 5075 Ishimoto, Akinori, 4533 Mori, Shigehisa, 4665 Savilahti, Clark, Nina, 5035 Schlicht, Hans-Jurgen, 4867 Iwashiro, Michihiro, 4533 Morrow, Casey D., 4621 Christine, 4992 Conrad, Susan E., 4964 Mosca, J. D., 4474 Schneider, Cordell, J., 4932 Jackson, Wallen, 4464 Schneider-Schaulies, Sibylle, Moscufo, Nicola, 5075 4760 Couture, Larry, 4712 Jameson, Bradford A., 4785 Moss, Bernard, 4732 Cubitt, Christopher L., 4777 Janetzko, Alfred, 4972 Schnorr, Jens-Jorg, 4760 Mous, Jan, 5041 Gerald, 5020 Cullen, Bryan R., 5030 Jean-Jean, Olivier, 4886 Muenchau, Daryl D., 4712 Schochetman, Jenkins, Frank J., 4842 Seifer, Maria, 4513 Daikoku, Tohru, 4651 Jewers, Richard J., 4521 Nabel, Gary J., 5035 Seillier-Moiseiwitsch, Dean, C., 4932 Naim, Hussein Y., 4831 Francoise, 4639 Delpeyroux, Francis, 4630 Kakiuchi, Nobuko, 4665 Nantermet, Pascale V., 4639 Sharp, Phillip A., 4769 DeLuca, Neal A., 4676 Kaplan, Jonathan E., 4659 Nauwynck, Hans J., 4492 Shimizu, Takeyuki, 4533 De Santo, Ronald J., 4484 Kato, Nobuyuki, 4665 Neipel, Frank, 4611 Shimotohno, Kunitada, 4665 Dixon, Linda K., 4549 Kayman, S. C., 4932 Nishiyama, Yukihiro, 4651 Shoeman, Robert L., 4972 Drelich, Meera, 5041 Kelly, Thomas J., 4992 Nussbaum, Ofer, 4712 Shotton, C., 4932 Dunn, K. J., 4704 Kieff, Elliott, 5068 Shu, Liming, 4712 Durda, Paul J., 4785 Kimura, Koichi, 4665 Oakes, John E., 4777 Shugars, Diane C., 4639, 4923 4722 4742 Kjems, J0rgen, 4769 Odenwald, Ward F., 4484 Shustik, Chaim, Esposito, Fulvio, Kobayashi, Akio, 4651 Oldham, G., 5012 Simon-Jodicke, Angela, 4760 Facke, Michael, 4972 Kock, Josef, 4867 Olins, Peter O., 4566 Small, Don, 4992 Fanning, Ellen, 4992 Kondo, Tokukazu, 4533 Olofsson, S., 4932 Smith, Colton A., 4676 Folks, Thomas M., 4659 Kowalik, Timothy F., 4981 Onodera, Shiroh, 4914 Smith, Geoffrey L., 4549 Fouillot, Nathalie, 4886 Kowalski, Jacek, 4896 Otaka, Akira, 4533 Smith, Marilyn S., 4639 Fridell, Robert A., 5030 Krausslich, Hans-Georg, 4972 Sodroski, Joseph, 4557, 4785, Frilander, Mikko, 4914 Kuhn, Richard J., 4605 Page, M., 4932 4932 Fujii, Nobutaka, 4533 Kurachi, Ryutaro, 4651 Pal-Ghosh, Ruma, 4621 Song, Jinping, 4945 Fung, Michael S., 4785 Kuribayashi, Kagemasa, 4533 Pardi, Diane, 4659 Specter, Steven, 4742 Furnari, Beth A., 4981 Pardigon, Nathalie, 5003 Stahl, Hans, 4992 Lai, Ching-Juh, 4956 Parsons, K. R., 5012 Standring, David N., 4513 Galphin, Judith C., 5020 Lal, Renu B., 4659 Patton, John T., 4848 Stierhof, York-Dieter, 4905 i ii AUTHOR INDEX J. VIROL.

St-Onge, Luc, 5025 Takahashi, Katsutoshi, 4533 VanDerMaaten, Martin J., Wolffe, Elizabeth J., 4732 Stout, Michael W., 4598 Tamai, Katsuyuki, 4651 5051 Wood, Charles, 5051 Straight, Samuel W., 4521 Tanaka, Torahiko, 4665 Vignaux, Francoise, 4785 Wu, Z., 4932 Strassman, Jeffrey, 4914 Tang, Min, 4896 von Schwedler, Uta, 4945 Wyatt, Richard, 4557 Straus, S. E., 4474 Tardy-Panit, Maryse, 4630 Vydelingum, Soopayah, 4549 Tekaia, Fredj, 4630 Strauss, Ellen G., 4605 Yaciuk, Peter, 4804 Strauss, James H., 4605, 5003 ter Meulen, Volker, 4760 Thali, Markus, 4557, Yafal, Alicia Gomez, 5075 Strebel, Klaus, 5056 4785 Wahren, Britta, 4785 Yamagishi, Hideo, 4533 Streeck, Rolf E., 4588 Tirwatnapong, Suranan, 4611 Wang, Hwei-Gene Heidi, 4804 Tlouzeau, Sylvie, 4886 Weiss, R. A., 4932 Yao, Zhengbin, 4464 Suarez, David L., 5051 Tomkinson, Blake, 5068 Weiss, Susan R., 4504 Yokoyama, Kazushige, 4804 Sullivan, Nancy, 4557, 4932 Tozzini, Franco, 4742 Westhafer, Mark A., 5020 Yuan, C. C., 4704 Sun, Bill, 4785 Trono, Didier, 4945 Whetstone, Cecelia A., 5051 Swanstrom, Ronald, 4639, Tsurumi, Tatsuya, 4651 White, Hillary D., 5045 Zaccaro, Lucia, 4742 4923 Tucker, Pamela C., 4605 Willey, Ronald L., 5056 Zamb, Tim J., 4896 Switzer, William M., 4659 Tykocinski, Mark L., 4896 Wimmer, Eckard, 4688 Zhang, Shang-Ding, 4484 Discover the Latest From eucaryotes to viruses... depend in Virology. on these two ASM journals for The Viruses cutting-edge science Series Editors: Heinz Fraenkel-Conrat Molecular and Cellular and Robert R. Wagner THE RETROVIRIDAE Biology Volume 1 Editor in Chief: Alan M. Weiner edited by Jay A. Levy This volume offers definitive studies of the IDesearchers find MCB to be an authoritative source of biology, pathology, immunology, and molecular fundamental knowledge and new developments in all features of retroviruses. 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