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THE JOURNAL OF BIOLOGICAL CHEMISTRY FOUNDED BY CHRISTIAN A. HERTER AND SUSTAINED IN PART BY THE CHRISTIAN A. HERTER MEMORIAL FUND

PUBLISHED BY THE AMERICAN SOCIETY FOR BIOCHEMISTRY AND MOLECULAR BIOLOGY, INC.

EDITORIAL BOARD/2005 Herbert Tabor, Editor Robert D. Simoni, Deputy Editor Norma M. Allewell, Associate Editor Judith S. Bond, Associate Editor Joan W. Conaway, Associate Editor John H. Exton, Associate Editor Peggy J. Farnham, Associate Editor Richard W. Hanson, Associate Editor

E. Dale Abel William M. Abraham Robert S. Adelstein Marcus Aebi Bharat B. Aggarwal Paul R. Albert Dario C. Altieri Jens Andersen Stefan Andersson Douglas A. Andres Toni M. Antalis Meredithe L. Applebury Suneel Apte Jonathan P. Arm Avi Ashkenazi John P. Atkinson Alan D. Attie David Auble Juan Ausio Jonathan M. Backer John Baezinger S. Paul Bajaj Sandra M. Bajjalieh Robert A. Bambara Hamid Band Philip Amos Barker Kim E. Barrett Michelle Craig Barton Robert Charles Baxter John W. Baynes Elmus G. Beale Kathleen M. Beckingham Francesco Beguinot Christoph Benning Dale John Benos Etty Benveniste John J. M. Bergeron Benjamin Berkhout Paolo Bernardi David A. Bernlohr Charles L. Bevins Ilya Bezprozvanny Narayan R. Bhat Martin Biel Paul Noble Black Carl Peter Blobel Gerd Blobel Kendall J. Blumer Paul E. Boehmer Gary Bokoch Meredith Bond Susan Bonner-Weir David R. Borchelt Mark A. Bothwell Lawrence F. Brass Patrick J. Brennan Michael Brenowitz Emery H. Bresnick H. Bryan Brewer Richard M. Breyer Barbara Brodsky Alex Brown Dennis Brown Jim Bruzik Joseph Bryan Donald A. Bryant

Vincent C. Hascall, Associate Editor Robert L. Hill, Associate Editor I. Robert Lehman, Associate Editor Jerry B Lingrel, Associate Editor Kenneth E. Neet, Associate Editor Jerrold M. Olefsky, Associate Editor John J. O’Shea, Associate Editor

Charles F. Burant Drusilla L. Burns Ashley Bush Marco Eugenio Cabrera Rafael D. Camerini-Otero Craig Eugene Cameron Marie-France T. Carlier Gordon G. Carmichael Chris Carpenter Graham Carpenter John Carpenter Nancy Carrasco Jean-Pierre Cartron Joseph R. Casey Michael Cashel Kevin Catt Elena Cattaneo Richard A. Cerione Benoit Chabot Ta-Yuan Chang Edwin R. Chapman Harry Charbonneau Maureen J. Charron Charles Chavkin Walter J. Chazin Frederic Checler Cheng-Ming Chiang Wonhwa Cho Czeslaw S. Cierniewski Gary F. Clark Robert A. Clark Henrik Clausen G. Marius Clore Vivian Cody K. Mark Coggeshall Jonathan C. Cohen Susan P. C. Cole Rosalind Coleman Karen J. Colley Marco Colonna Ronald C. Conaway Paul F. Cook John A. Corbett Robert H. Costa Rick H. Cote Benoit Coulombe Robert Craigie William A. Cramer Christine R. Cremo William Douglass Cress Richard L. Cross Rodney Croteau Rosalie K. Crouch John M. Cunningham John Cuppoletti Norman P. Curthoys Martha S. Cyert Douglas M. Cyr Michael P. Czech Luciano D’Adamio Michael E. Dahmus David Danielpour Michael David Amy L. Davidson William Sean Davidson Gideon J. Davies

David W. Russell, Associate Editor Charles E. Samuel, Associate Editor William L. Smith, Associate Editor James T. Stull, Associate Editor Thomas C. Vanaman, Associate Editor Kensal E. van Holde, Associate Editor Martha Vaughan, Associate Editor

Peter J. A. Davies John H. Dawson Ted Dawson Anthony John Day Dennis R. Dean Michael Dean Neta Dean Louis J. De Felice Anne Dell Paul Dent Sharon Y. R. Dent John M. Denu Thomas Dever Lakshmi A. Devi David L. DeWitt Enrico Di Cera Fabio Di Lisa Ivan Dikic Rick T. Dobrowsky Julie Donaldson Gerald W. Dorn Kristina Downing Boris Draznin Lester R. Drewes Arnold J. M. Driessen Raymond N. Dubois Ronald R. Dubreuil Colin S. Duckett Maria Dufau Stanley Dunn Anindya Dutta Douglas C. Eaton Dale E. Edmondson Michael D. Ehlers David J. Eide Lee Eiden Betty A. Eipper Edward Eisenstein Wafik S. El-Deiry Per Erik Elias Lisa M. Ellerby Timothy John Elliott John F. Engelhardt Caroline Enns James Ervasti Mary E. Fabry Mike Fainzilber Ellen Fanning Joshua Marion Farber Stephen R. Farmer Eric R. Fearon Martha J. Fedor Michael A. J. Ferguson Pascal Ferre Alan P. Fields Gregg B. Fields Janet S. Finer-Moore Anthony L. Fink Edward A. Fisher David J. Fitzgerald Veit Flockerzi Thomas Force Michael D. Forgac Mike Forte Amanda J. Fosang Joan Elizabeth Brothwell Fox

Paul L. Fox Claire M. Fraser Paul Fraser Robert S. Freeman Lloyd Fricker Judith L. Fridovich-Keil Errol C. Friedberg Raymond A. Frizzell Robert P. P. Fuchs Robert Stuart Fuller James L. Funderburgh Masamitsu Futai Daniel Gallie Maria Garcia Narasimhan Gautam Michael Gelb Craig Gerard Thomas A. Gerken Michael Scott German Jorge A. Ghiso Sankar Ghosh Harry J. Gilbert Charles G. Glabe Marc Glucksman Daniel Goldberg Douglas Golenbock F. Xavier Gomis-Ruth Frank Gonzalez Jeffrey P. Gorski Joel Gottesfeld Channe Gowda Robert M. Graham Gregory A. Grant Lee M. Graves Michael Green Daniel S. Greenspan Jack Griffith Michael Jeffrey Grusby F. Peter Guengerich Michel Guertin Richard I. Gumport Claes M. Gustafsson Arthur Gutierrez-Hartmann Jesper Haeggstrom Tsonwin Hai Kasturi Halder Christine Hall Otto Albrecht Haller Susan Hamilton Thomas Alan Hamilton Marie-Louise Hammarskjold Jiahuai Han Fumio Hanaoka T. Kendall Harden Daniel M. Hardy David Harris Robert A. Harris Robert Adron Harris Marietta L. Harrison Gerald W. Hart David J. Hartshorne Maria Hatzoglou Craig A. Hauser Edward Hawrot Jeffrey J. Hayes Stanley L. Hazen

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ii Editorial Board continued Peter C. Heinrich Martin E. Hemler Barbara L. Hempstead Susan A. Henry Brian Herman Louis B. Hersh John W. B. Hershey Paul J. Hertzog John D. Hildebrandt Russ Hille Douglas James Hilton Michael Hippler John Hiscott Timothy Hla Jan Hoek P. Mark Hogarth Hazel M. Holden Richard Horuk Curt Horvath Robert L. Houtz Ara G. Hovanessian O. M. Zack Howard Philip H. Howe Tao Hsieh Xin-Yun Huang Billy G. Hudson David Y. Hui David J. S. Hulmes Martin James Humphries John Hunt William Hunter James B. Hurley Daniel H. Hwang Carlos Fernando Ibanez Anthony Imbalzano Giuseppe Inesì Renato V. Iozzo Nancy Ip David G. Jackson Steven J. Jacobs David Ernest James Mariusz Jaskolski Kuan-Teh Jeang Robert L. Jernigan Tae H. Ji Eric F. Johnson Kenneth A. Johnson Randall S. Johnson Jim Johnston Larry R. Jones Teresa L. Z. Jones Frank Jordan Suresh Joseph Karl Kadler Barbara Barken Kahn Satish C. Kalhan Patricia M. Kane Neil Kaplowitz Kenneth G. Keegstra Jack Keene John Kehrl Michael Kilberg Donghee Kim George Liang King David Charles Klein William L. Klein Dwight John Klemm Rachel E. Klevit Thomas R. Kleyman Amira Klip James R. Knox Erik Knudsen Brian Kobilka Walter J. Koch Murray Korc Sally A. Kornbluth Antonis E. Koromilas Stephen C. Kowalczykowski Reinhard Kraemer Ken Kreuzer Martin Kro¨nke Nicholas T. Ktistakis Robert Kuchta Thomas B. Kuhn Rakesh Kumar Lawrence C. Kuo

John M. Kyriakis Dale W. Laird Ratnesh Lal Paul Lampe Stephen M. Lanier Martin R. Larsen Thomas M. Laue James D. Lechleiter Hon-Cheung Lee Stephen Lee Todd Leff Sylvain Lehmann Nancy J. Leidenheimer Vance P. Lemmon Thomas L. Leto John J. Letterio Ellis R. Levin Stewart J. Levine Alexander Levitzki Xiao-Jiang Li Jane B. Lian James Kuang-Jan Liao Bob Liddington Thomas M. Lincoln Darrell T. Liu Leroy F. Liu Richard A. Lockshin Eric Long Carlos Lopez-Otin Paul E. Love Richard F. Luduena Louis M. Luttrell Jianjie Ma Ian G. Macara Ormond A. MacDougald Steven James Madore Mark Magnuson Philip W. Majerus George I. Makhatadze James J. Manfredi David R. Manning Thomas F. J. Martin Grant Mauk Bernd Mayer Neil McDonald David McKay Patrick A. McKee James H. McKerrow Chris R. McMaster Elizabeth M. McNally Linda C. McPhail Susan O. Meakin Gerhard Meissner Ronald L. Mellgren Ralf R. Mendel Alfred H. Merrill Joe M. Miano Anthony J. Michael Gregory A. Mignery Edith W. Miles Audrey Minden Henry M. Miziorko Edward Mocarski Vincent M. Monnier Kevin L. Moore Kelley Moremen Simon J. Morley James H. Morrissey Jason D. Morrow Randall Morse Deane F. Mosher David M. Mosser Dick D. Mosser Shmuel Muallem Kathleen M. Mulder Gerd Multhaup Philip M. Murphy George E. O. Muscat Tomas M. Mustelin Toshi Nakayama Joseph L. Napoli Linda O. Narhi Mohandas Narla William M. Nauseef Gary L. Nelsestuen Michael E. Nesheim

Eisuke Nishida Thomas Nowak Richard Mark O’Brien Luke O’Neil John A. Oates Stefan Offermanns Rolf I. Ohlsson Richard W. Olsen Steven T. Olson Santa Jeremy Ono Peter A. B. Orlean Daniel S. Ory Philip Osdoby Neil Osheroff Michael Ostrowski Keiko Ozato Radhakrishnan Padmanabhan Himadri Pakrasi Zhen-Qiang Pan David S. Park Armando J. Parodi Henry Paulson Edward J. Pearce Trevor Penning John J. Perona Fred W. Perrino Richard G. Pestell Martha L. Peterson Lawrence Marc Pfeffer Douglas R. Pfeiffer Kenneth D. Philipson Caroline Philpott Susan Pierce Linda Pike Daniel Pipeleers Leonidas C. Platanias Edward F. Plow Bert Poolman James D. Potter Jacques Pouyssegur Susan Powers-Lee Leonard Presta Glenn Prestwich S. Russ Price Suzette Priola Richard L. Proia James P. Quigley Mark T. Quinn Michael J. Quon Daniel M. Raben Rick Rachubinski Steve Ragsdale Francesco Ramirez Paul Randazzo Rajini Rao Basil Rapoport Alan C. Rapraeger Ivan Rayment John R. Raymond Reinhart Reithmeier Marilyn D. Resh Robert E. Rhoads Christopher J. Rhodes David Richardson John P. Richardson Kevin D. Ridge Charles T. Roberts David D. Roberts Mary Roberts Gail Robertson Phillip J. Robinson Paul A. Roche William J. Roesler Dina Ron Ze’ev A. Ronai David S. Roos David W. Rose Barry P. Rosen Paul R. Rosevear Marsha R. Rosner Alonzo H. Ross Gregory M. Ross Bryan L. Roth Michael G. Roth Lawrence I. Rothblum Peter Rotwein

David R. Rowley Ananda Roy Peter Rubenstein Neil B. Ruderman Gary Rudnick Tommaso Russo Robert O. Ryan David B. Sacks David Samols Linda Sandell Konrad Sandhoff John D. Sandy Massimo Santoro H. Uri Saragovi Kevin D. Sarge Rashmin C. Savani Thomas S. Scanlan Suzanne F. Scarlata Martin Scheffner Philipp E. Scherer Ann Marie Schmidt Jonathan Scholey Debra A. Schwinn Miguel Seabra Bartholomew M. Sefton Deborah L. Segaloff James Sellers Charles N. Serhan James Alan Shayman Shirish Shenolikar Yang Shi Ali Shilatifard Takao Shimizu Toru Shimizu G. Graham Shipley Joseph Shlomai Gary E. Shull Gerald I. Shulman Ulrich Siebenlist Maya Sieber-Blum James N. Siedow Richard Siegel Roberto Sitia Bonnie Fiedorek Sloane Robert John Smith Mark Smyth Jaro Sodek R. John Solaro Dieter G. Soll Avril V. Somlyo Linda L. Spremulli M. Sharon Stack Michael R. Stallcup Bruce A. Stanton Donald F. Steiner Jacqueline Stephens F. Anne Stephenson Jeffry B. Stock Stanko S. Stojilkovic Judith Storch Philip J. Stork Christopher D. Stubbs Thomas C. Sudhof Hei Sook Sul Witold Surewicz Mark A. Sussman John Svaren Lawrence Tabak Robert Tampe Wei-Jen Tang Naoyuki Taniguchi Martin Teichman David B. Teplow Steven M. Theg Gopal Thinakaran Andrew P. Thomas Andrew Thorburn Gabor Tigyi Alex Toker Peter J. Tonge Bryan P. Toole James Travis Steve Triezenberg Karl Tryggvason Yuk-Ching Tse-Dinh Yoshihide Tsujimoto

iii Editorial Board continued Jeremy E. Turnbull Christopher Turner R. James Turner Bik-Kwoon Tye Michael D. Uhler Raul Urrutia Bennett van Houten Gerrit F. B. P. van Meer William E. van Nostrand Elizabeth Vierling Richard D. Vierstra Steven B. Vik Giuseppe Villani Nicolas Vitale Chris J. Vlahos

Brian E. Wadzinski Elmar Wahle David M. Waisman Warren W. Wakarchuk Susan Wallace David Wallach Ulrich Walter Xiao-Fan Wang Paul A. Watkins Patricia C. Weber Nancy Weigel Peter Anthony Weil Michael Welsh David E. Wemmer Robert J. Wenthold

David Westaway Sidney W. Whiteheart Chris Whitfield Thomas M. Wilkie Keith D. Wilkinson Bart Williams Carol L. Williams Ian M. Willis Dennis R. Winge Malcolm E. Winkler Beatrice A. Wittenberg Alexander Wlodawer Michael S. Wolfe Benjamin Wolozin Marie W. Wooten

Jo Rae Wright Kenji Yamamoto Honggao Yan Alpha S. Yap Philip L. Yeagle Timothy J. Yen John D. York Howard A. Young Stephen G. Young David Yule Peter D. Yurchenco Jillian Zhang Zhizhuang Joe Zhao Yehiel Zick

Barbara A. Gordon, Executive Officer Nancy J. Rodnan, Director of Publications

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