THE JOURNAL OF CELL BIOLOGY

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JCB THE JOURNAL OF CELL BIOLOGY

EDITOR-IN-CHIEF JODI NUNNARI EXECUTIVE EDITOR REBECCA ALVANIA email: [email protected]

EDITORS PIER PAOLO DI FIORE ELAINE FUCHS REBECCA HEALD IAN MACARA IRA MELLMAN LOUIS F. REICHARDT KENNETH M. YAMADA JUNYING YUAN SCIENTIFIC EDITORS MELINA CASADIO email:

[email protected]

SHAWN N. JORDAN email:

[email protected]

MARIE ANNE O'DONNELL email: [email protected]

NEWS EDITOR BENJAMIN SHORT

EDITORIAL BOARD JOHN AITCHISON LAURA ATTARDI JOHAN AUWERX BEN BARRES HUGO BELLEN BILL BEMENT VANN BENNETT JEFFREY BENOVIC MONICA BETTENCOURT-DIAS SUE BIGGINS CÉDRIC BLANPAIN JULIUS BRENNECKE TONY BRETSCHER MARIANNE BRONNER DON W. CLEVELAND DANIEL COLÓN-RAMOS NIKA DANIAL TITIA DE LANGE ARSHAD DESAI RAY DESHAIES IVAN DIKIC DANIEL DUROCHER WILLIAM EARNSHAW JAN ELLENBERG SCOTT EMR JEFFREY ESKO MARC FREEMAN HIRONORI FUNABIKI MARGARET GARDEL MELISSA GARDNER LARRY GERACE FRANK GERTLER DAVID GILBERT

SERGIO GRINSTEIN ULRICH HARTL RAMANUJAN HEGDE MARTIN HETZER NOBUTAKA HIROKAWA ERIKA HOLZBAUR ARTHUR HORWICH MARTIN HUMPHRIES ANNA HUTTENLOCHER LUISA IRUELA-ARISPE NANCY IP JOHANNA IVASKA SUSAN KAECH GERARD KARSENTY SCOTT KEENEY ALEXEY KHODJAKOV JÜRGEN KNOBLICH AKIHIRO KUSUMI THOMAS LANGER JEANNE LAWRENCE DANNY LEW JIRI LUKAS JENS LYKKE-ANDERSEN VIVEK MALHOTRA JOAN MASSAGUE JACOPO MELDOLESI ALEX MOGILNER RANDALL T. MOON SEAN MUNRO EVA NOGALES KAREN OEGEMA ROY PARKER MARK PEIFER

ELIOR PELES ANA POMBO THOMAS RANDO JEREMY REITER DANIEL B. RIFKIN DAVID RON MICHAEL ROUT CRAIG ROY MICHAEL RUDNICKI ERIK SAHAI DANNY SCHNELL TRINA SCHROER MARTIN SCHWARTZ JEAN SCHWARZBAUER ANDREY SHAW ZU-HANG SHENG MICHAEL SIXT TIM STEARNS JOAN STEITZ HARALD STENMARK AARON STRAIGHT RON VALE PATRIK VERSTREKEN AMY WAGERS ERWIN WAGNER TOBIAS WALTHER LOIS WEISMAN MATT WELCH TIM YEN TAMOTSU YOSHIMORI RICHARD YOULE MARINO ZERIAL YIXIAN ZHENG

EDITORIAL OFFICE

PRODUCTION OFFICE

Phone: 212-327-8581 Fax: 212-319-1082 email: [email protected]

email: [email protected]

MANAGING EDITOR DEMANTIE (SATI) MOTIERAM MANUSCRIPT COORDINATORS CAROLIENA CABADA LINDSEY HOLLANDER PREFLIGHT COORDINATOR LAURA SMITH

COPY EDITOR DIEDRA HOWSON-BARKER PRODUCTION EDITOR MARY VASQUEZ PRODUCTION MANAGER CAMILLE CLOWERY PRODUCTION DESIGNER ERINN A. GRADY

COPYRIGHT TO ARTICLES PUBLISHED IN THIS JOURNAL IS HELD BY THE AUTHORS. ARTICLES ARE PUBLISHED BY THE ROCKEFELLER UNIVERSITY PRESS UNDER LICENSE FROM THE AUTHORS. CONDITIONS FOR REUSE OF THE ARTICLES BY THIRD PARTIES ARE LISTED AT HTTP://WWW.RUPRESS.ORG/TERMS

PRINT ISSN: 0021-9525 ONLINE ISSN: 1540-8140

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EXECUTIVE DIRECTOR SUSAN KING FINANCE DIRECTOR RAYMOND T. FASTIGGI BUSINESS DEVELOPMENT DIRECTOR GREGORY FLYNN MALAR DIRECTOR OF PUBLISHING TECHNOLOGIES ROBERT J. O'DONNELL ADVERTISING SALES DIRECTOR LORNA PETERSEN OFFICE ADMINISTRATOR JOANN GREENE ASSISTANTS TO THE FINANCE DIRECTOR SARAH S. KRAFT ANN TRAVERS BUSINESS DEVELOPMENT SPECIALIST COLLEEN M. SMITH MARKETING ASSOCIATE LARAINE KARL

THE JOURNAL OF CELL BIOLOGY (ISSN 0021-9525) IS PUBLISHED BIWEEKLY BY THE ROCKEFELLER UNIVERSITY PRESS, 950 THIRD AVENUE, NEW YORK, NY 10022. PERIODICAL POSTAGE PAID AT NEW YORK, NY AND ADDITIONAL MAILING OFFICES. 2016 SUBSCRIPTION RATES: INSTITUTIONAL RATES: ONLINE PRINT+ONLINE

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PERSONAL RATES: PRINT ONLY:$500 FOR MORE INFORMATION, PLEASE CONTACT THE CIRCULATION DEPARTMENT: Phone: 212-327-8572 Fax: 212-319-1080 email: [email protected] ADVERTISING REQUESTS: Phone: 212-327-8880 Fax: 212-319-1080 email: [email protected] PERMISSION REQUESTS: email: [email protected] MEDIA REQUESTS: email: [email protected] POSTMASTER: SEND ADDRESS CHANGES TO THE JOURNAL OF CELL BIOLOGY, CIRCULATION DEPARTMENT, THE ROCKEFELLER UNIVERSITY PRESS, 950 THIRD AVENUE, FLOOR 2, NEW YORK, NY 10022-2705.