Journal of Virology

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Editor in Chief Lynn W. Enquist (2012) Princeton University Editors Karen L. Beemon (2012) Johns Hopkins University Terence S. Dermody (2009) Vanderbilt University School of Medicine Michael S. Diamond (2013) Washington University School of Medicine Daniel C. DiMaio(2011) Cover Editor Yale University School of Medicine Robert W. Doms (2010) University of Pennsylvania School of Medicine Donald E. Ganem (2012), Minireview Editor University of California, San Francisco Harry B. Greenberg (2013) Stanford University School of Medicine Beatrice H. Hahn (2009) University of Alabama Michael J. Imperiale (2011) University of Michigan Medical School Richard A. Koup (2013) National Institute of Allergy and Infectious Diseases Douglas S. Lyles (2012) Wake Forest University Grant McFadden (2012) University of Florida Jay A. Nelson (2012) Oregon Health & Science University Peter M. Palese (2010) Mount Sinai School of Medicine Stanley Perlman (2012) University of Iowa Nancy Raab-Traub (2011) University of North Carolina Susan R. Ross (2012) University of Pennsylvania Rozanne M. Sandri-Goldin (2010) University of California, Irvine Bert L. Semler (2012) University of California, Irvine Ganes C. Sen (2013) The Cleveland Clinic Foundation Anne Simon (2012) University of Maryland Ronald Swanstrom (2009) University of North Carolina School of Medicine Chairman, Publications Board Thomas E. Shenk Director, Journals Barbara M. Goldman Production Editor Judith Nedrow Assistant Production Editors Karen Esplin Julie Nachman

Editorial Board Paul Ahlquist (’08) Chris Aiken ('09) Gillian Air ('09) Lorraine Albritton ('10) Antonio Alcami ('10) David Anders (’08) Raul Andino (’08) Elliot J. Androphy (’08) Carlos F. Arias ('09) Eric Arts ('09) Ann Arvin ('09) Walter J. Atwood ('10) Steve Bachenheimer ('09) Joel Baines ('09) Susan C. Baker ('08) Dennis H. Bamford ('08) Bruce W. Banfield ('10) Lawrence Banks ('08) Ralph Baric ('09) Eric Barklis (’08) Dan Barouch ('09) Ralf Bartenschlager (’08) David J. Barton ('08) Chris Basler ('09) Jeffrey M. Bergelson ('08) Ben Berkhout ('09) Ken Berns ('09) Michael Betts ('10) Paul D. Bieniasz (’08) John A. Blaho ('10) Keril Blight ('09) Gary W. Blissard ('10) David C. Bloom ('10) James Borowiec (’08) Michele Bouloy ('09) Michel Brahic ('09) Wade Bresnahan ('08) David Brian ('09) Margo A. Brinton ('08) William Britt ('09) Chris Broder ('09) Jay Brown ('08) Michael Buchmeier ('09) Dennis R. Burton ('10) Frederic D. Bushman ('09) Andrew Byrnes ('10) Ellen D. Cahir-McFarland ('10) Craig Cameron ('09) Gabriella Campadelli-Fiume ('10) Ann Campbell ('09) Michael J. Carter (’08) Roberto Cattaneo ('09) Bruce W. Chesebro ('09) Paul Clapham ('10) John M. Coffin ('09) Jeffrey I. Cohen ('10) Anamaris Colberg-Poley ('09) Marc S. Collett ('10) Peter Collins ('09) Teresa Compton ('09) Richard C. Condit ('10) Karl-Klaus Conzelmann ('10) Susan F. Cotmore ('10) James Crowe ('09) Bryan Cullen ('10) James N. Culver ('10) James Cunningham (’08) Blossom Damania ('10) Nancy Davis ('09) Andrew J. Davison ('10) William O. Dawson ('10) Juan Carlos de la Torre ('08) Mark R. Denison ('08) Cynthia A. Derdeyn ('10) Prashant Desai ('09) Ronald Desrosiers ('09) Ulrich Desselberger ('09) Dimiter Dimitrov ('09) Dirk Dittmer ('10) Valerian Dolja (’08) Esteban Domingo ('09) Philip R. Dormitzer ('10) Tanya Dragic ('08) Theo Dreher (’08) Jean D. Dubuisson ('08) Jackie Dudley ('09) Rebecca E.Dutch ('09) Ellie Ehrenfeld ('10) Maribeth Eiden ('09) Roselyn J. Eisenberg ('09) John H. Elder (’08) Richard M. Elliott ('10) Michael Emerman ('09) Alan Engelman ('09) Francis Ennis (’08) Mariano Esteban ('10) Roger Everett ('10) Erik Falck-Pedersen ('10) Paul J. Farrell ('10) Michael Farzan ('09) Heinrich U. Feldmann ('08) Ana Fernandez-Sesma ('09) Jane Flint (’08) Ron Fouchier ('10) Genveffa Franchini ('09) Manuel Franco ('09) Nigel Fraser ('09) Eric O. Freed (’08) David H. Fremont ('10) Teryl Frey ('09) Ilya Frolov ('10) Klaus Frueh ('10) Robert S. Fujinami (’08) Dana H. Gabuzda (’08) Michael Gale ('10) Thomas Gallagher (’08) Robert Garcea ('09) J. Victor Garcia-Martinez ('10) Adolfo Garcia-Sastre ('10) Richard Gaynor ('09) Adam P. Geballe ('10) Chou-Zen Giam ('09) Donald Gilden (’08) Stephen P. Goff ('09) Francisco Gonzalez-Scarano (’08) Felicia Goodrum ('10) Heinrich Gottlinger (’08) Patrick Green ('10) Charles Grose ('09) Linda Guarino ('09) Nancy Haigwood ('09 Otto A. Haller ('08) Linda Hanley-Bowdoin ('10) J. Marie Hardwick (’08) Reuben Harris ('10) Ronald Harty ('09) Kim Hasenkrug ('09) S. Diane Hayward ('07 Patrick Hearing ('09) Franz Heinz ('09) Mark Heise ('08) Ann B. Hill ('08) Vanessa Hirsch ('09) James M. Hogle ('09) Brenda Hogue ('09) Thomas Hohn (’08) Eddie Holmes ('09) Fred Homa ('09) Edward A. Hoover ('09) Thomas Hope ('09) Curt M. Horvath ('10) Jim Hoxie ('09) Jianming Hu ('10) Stephen Hughes ('09) Lindsey Hutt-Fletcher (’08) Bertram Jacobs ('08) Michael A. Jarvis ('10) Ronald T.  Javier ('10) Kuan-Teh Jeang ('09) David C. Johnson (’08) R. Paul Johnson ('10) Welkin Johnson ('09) Paul Jolicoeur (’08) Jae Jung ('09) David Kabat (’08) Cheng Kao ('09) Michael Katze (’08) Yoshihiro Kawaoka ('10) Kenneth Kaye ('09) Vineet N. KewalRamani ('09) Alexander Khromykh ('10) Elliott Kieff (’08) Margaret Kielian ('10) Karla Kirkegaard (’08) Paul R. Kinchington ('08) Frank Kirchhoff ('10) David Knipe ('08) Georg Kochs ('10) Dan Kolakofsky (’08) Sarah Kopecky-Bromberg ('10) Ulrich Koszinowski ('09) Mark Krystal (’08) Richard Kuhn ('10) Laimonis Laimins ('09 Michael Lairmore ('09)

Robert A. Lamb ('09) Paul F. Lambert ('10) Nathaniel Landau ('09) Benhur Lee ('09) David A. Leib (’08) Andrew Leigh Brown ('09) Stanley M. Lemon ('10) Laura Levy ('10) Jeffrey Lifson ('10) Brett Lindenbach ('10) Maxine Linial ('10) Howard Lipton ('09) Richard Lloyd ('09) Dan Loeb ('10) Volker Lohmann ('10) Richard Longnecker ('10) Susana Lopez Charreton ('09) Anice Lowen ('10) Jeremy Luban (’08) Guangxiang G. Luo ('10) Margaret MacDonald ('10) Erich Mackow ('09) Shinji Makino ('10) Malcolm A. Martin (’08) John Mascola ('10) William S. Mason ('09) Paul S. Masters ('10) Thomas Mettenleiter ('08) A. Dusty Miller ('09) Chris Miller ('09) Edward S. Mocarski (’08) David Montefiori ('09) Ronald C. Montelaro (’08) Darius Moradpour ('09) Elizabeth Moran (’08) Thomas Moran ('10) Casey Morrow ('09) Anne Moscona ('08) Ashley B. Moses ('08) Don Mosier ('09) Bernard Moss ('10) Karen L. Mossman ('08) Karl Munger ('09) Brian R. Murphy ('08) Nicholas A. Muzyczka (’08) Peter Nagy ('10) Glen R. Nemerow ('09) Max L. Nibert (’08) Stuart T. Nichol ('10) Janko Nikolich-Zugich ('10) Edward G. Niles ('10) Douglas Nixon ('09) Isabel Suarez Novella ('10) Don Nuss ('09) Jennifer Nyborg ('09) Peter O'Hare ('09) David A. Ornelles (’08) Klaus Osterrieder ('10) James Ou ('09) Leslie Parent ('10) Greg Pari ('09) Griffith D. Parks ('10) Colin R. Parrish ('10) John Patton ('09) Mark E. Peeples ('10) Andrew Pekosz ('09) Philip E. Pellett ('10) Michael Perdue ('09) Lenore Pereira ('09) Daniel Perez ('09) Matija Peterlin (’08) Louis Picker ('09) Theodore C. Pierson ('10) David J. Pintel ('10) James M. Pipas (’08) Paula Pitha-Rowe ('10) B.V. Venkataram Prasad ('10) Vinayaka R. Prasad ('10) Chris Preston ('09) Vincent R. Racaniello ('08) Kathryn L. Radke (’08) Glenn Rall ('10) Robert F. Ramig ('10) Lee Ratner ('09) Stuart C. Ray ('10) Matthias Reddehase ('09) Barbara Rehermann ('10) Alan R. Rein (’08) Rolf Renne ('10) Marilyn Resh ('10) Felix A. Rey ('10) Andrew Rice ('09) Alan Rickinson (’08) Bert K. Rima ('10) Charles R. Rinaldo (’08) Erle S. Robertson ('10) harriet L. Robinson ('08) George Rohrman ('10) Bernard Roizman ('09) Raymond P. Roos ('10) John K. Rose ('10) Monica Roth ('10) Rob Ruigrok ('09) Bill Ruyechan ('09) Charles E. Samuel ('08) Peter Sarnow (’08) Quentin Sattentau ('09) Dorothea L. Sawicki ('09) Priscilla A. Schaffer (’08) Brian Schaffhausen ('10) John Schiller ('10) Connie S. Schmaljohn ('10) Matthias S. Schnell (’08) Paul Sharp ('09) Megan Shaw ('10) Kunitada Shimotohno ('10) Stuart Siddell ('09) Aleem Siddiqui ('09) Robert Siliciano ('09) Robert Silverman ('09) Guido Silvestri ('09) Tony Simmons ('09) John Sinclair ('10) John Sixbey ('10) Mark Slifka ('10) James Smiley ('09) Greg Smith ('09) Eric Snijder ('09) Joseph Sodroski ('09) Willy J. M. Spaan ('09) Richard R. Spaete ('10) Patricia G. Spear (’08) Paul Spearman ('09) Samuel Speck ('09) Deborah Spector ('09) Katherine R. Spindler ('08) Arun Srivastava ('09) Mario Stevenson (’08) Mark F. Stinski ('10) Leonidas Stamatatos ('09) Victor Stollar ('09) Jonathan P. Stoye (’08) Klaus Strebel ('09) Daniel Streblow ('10) William Sugden (’08) Ren Sun ('08) Wesley I. Sundquist (’08) Richard Sutton ('09) Yuri Svitkin ('09) Sankar Swaminathan ('10) John M. Taylor ('10) Alice Telesnitsky (’08) Timothy Tellinghuisen ('10) Mary J. Tevethia (’08) Heinz-Juergen Thiel ('09) David A. Thorley-Lawson ('10) Steven Tracy ('09) Paula Traktman ('10) Didier Trono (’08) Terrance Tumpey ('09) Kenneth L. Tyler ('10) Luis P. Villarreal ('09) Herbert Virgin IV (’08) Peter K. Vogt (’08) Volker M. Vogt (’08) Bruce Walker ('09) Christopher Walker ('09) Fred C. Wang ('09) Richard Ward ('09) David Watkins (’08) Scott Weaver ('10) Susan Weiss ('09) Sandra Weller (’08) Judith White ('09) J. Lindsay Whitton ('10) Reed B. Wickner ('09) Luc L. Willems (’08) Bryan Williams ('10) Carolyn Williamson ('10) Scott W. Wong ('10) David L. Woodland ('10) Richard Wyatt ('08) Otto Yang ('09) T.S. Benedict Yen ('10) Jonathan Yewdell (’08) Jerome Zack ('10)

The Journal of Virology, a publication of the American Society for Microbiology (ASM), 1752 N Street, N.W., Washington, DC 20036-2904, is devoted to the dissemination of fundamental knowledge concerning viruses of 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.