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JCB THE JOURNAL OF CELL BIOLOGY EDITOR-IN-CHIEF EDITORIAL BOARD IRA MELLMAN NORMA ANDREWS GILLIAN GRIFFITHS RANDALL T MOON BEN BARRES ULRICH HARTL SEAN MUNRO EDITORS HUGO BELLEN REBECCA HEALD JODI NUNNARI DON W. CLEVELAND ANDREW BELMONT NOBUTAKA HIROKAWA ANDRE NUSSENZWEIG VANN BENNETT ERIKA HOLZBAUR LESLIE PARISE PIER PAOLO Dl FIORE JEFFREY BENOVIC ARTHUR HORWICH MARK PEIFER ELAINE FUCHS JUAN BONIFACINO ALAN RICK HORWITZ ANNE RIDLEY ALAN HALL TONY BRETSCHER M. ANDREW HOYT CONLY RIEDER LOUIS F. REICHARDT MARIANNE BRONNER-FRASER MARTIN HUMPHRIES DANIEL B. RIFKIN RANDY SCHEKMAN ERIC j. BROWN ED HURT DAVID RON KENNETH M. YAMADA KENNETH CLINE KENNETH JACOBSON MICHAEL RUDNICKI JUNYING YUAN PASCALE COSSART GERARD KARSENTY JOSHUA SANES CAROLINE DAMSKY HYNDA KLEINMAN TRINA SCHROER PIETRO DE CAMILLI TEYMURAS KURZCHALIA MARTIN SCHWARTZ NEWS EDITORS ELISABETTA DEJANA ARTHUR D. LANDER JEAN SCHWARZBAUER NICOLE LEBRASSEUR RAY DESHAIES JEANNE LAWRENCE RON VALE WILLIAM A. WELLS DAVID DRUBIN YURI LAZEBNIK DAVID L. VAUX RUTH WILLIAMS WILLIAM EARNSHAW VERONIQUE LEFEBVRE GRAHAM WARREN ROBERT EISENMAN LESLIE LEINWAND CLARE WATERMAN-STORER REVIEWS EDITORS SCOTT EMR RICHARD LEWIS LOIS WEISMAN WERNER FRANKE LAURA MACHESKY TIM YEN KAREN DELL HIRONORI FUNABIKI JOAN MASSAGUE MARINO ZERIAL LOUIS F. REICHARDT LARRY GERACE JACOPO MELDOLESI YIXIAN ZHENG MARK GINSBERG TOM MISTELI MANAGING EDITOR COPYEOITING MIKE ROSSNER AND PRODUCTION Phone; 212-327-8881 email: [email protected] Fax: 212-327-8576 PREFLIGHT COORDINATOR email: [email protected] .AURA SMITH COPY EDITORS EDITORIAL ASSISTANTS CAMILLE CLOWERY LINDSEY HOLLANDER SATI MOTIERAM SHAUNA O'GARRO LESLIE WALKER KARL RAMOS EMILY TAYLOR PRODUCTION EDITOR Phone: 212-327-8581 GREGORY E. KOUTROUBY Fax: 212-327-8576 NEWS PRODUCTION EDITOR email: [email protected] GREG S. BOUSTEAD COPY EDITING COORDINATOR MARY VASQUEZ PRODUCTION COORDINATOR ERINN A. GRADY PRODUCTION DIRECTOR ROBERT J. O'DONNELL ^c^feCCer]^ VOLUME 175, aoas V^niversityJ^ '^Ki90i COPYRIGHT © 2006 BY THE ROCKEfELLER UNIVERSITY PRESS NEWS In This Issue A protein long shot Not so close Brief encounter Division of labor Ring around the tubule M. Leslie Research Roundup 4 • Computing direction • The liver that caveolin built • Tether before synapse • Fixing the hole • Keeping valves clear W.A. Wells Special Report 6 • Full of sound and fury, but signifying something: XVI International AIDS Conference, Toronto, Canada. August 13-18, 2006 • The ABCDEFGHI of prevention • Science on safari • Tidbits from Toronto • Should journalists question science? • The Ultimate Hope • Too much of a good thing W.A. Wells RESEARCH ARTICLES Reports 17 Dynamic microtubules are essential for efficient chromosome capture and biorientation in S. cerevisiae Baoying Huang and Tim C. Huffaker On the cover Oligomers of the kinesin-l 3 motor form 2B Kinesin-I3s form rings around microtubules rings and spirals around microtubules. Dongyan Tan, Ana B. Asenjo, Vito Mennella, David J. Sharp, See page S5. and Hernando Sosa 33 Kidney failure in mice lacking the tetraspanin CD151 Norman Sachs, Maaike Kreft, Marius A. van den Bergh Weerman, AndyJ. Beynon, Theo A. Peters, Jan J. Weening, and Arnoud Sonnenberg Articles 4 n Mapping the assembly pathways that specify formation of the trilaminar kinetochore plates in human cells Song-Tao Liu, Jerome B. Rattner, Sandra A. Jablonski, and TimJ. Yen S5 Promyelocytic leukemia nuclear bodies behave as DNA damage sensors whose response to DNA double-strand breaks is regulated by NBSI and the kinases ATM, Chk2, and ATR Graham Dellaire, Reagan W. Ching, Kashif Ahmed, Farid Jalali, Kenneth C.K. Tse, Robert G. Bristow, and David P. Bazett-Jones Articles with related stories in the IN THIS ISSUE section hove page numbers in RED. B7 Visualization of mRNA translation in living cells Alexis J. Rodriguez, Shoilesh M. Shenoy, Robert H. Singer, and John Condeelis 77 MyoD inhibits Fstll and Utrn expression by inducing transcription ofmiR-206 Miriam I. Rosenberg, Sara A. Georges, Amy Asawachaicharn, Erwin Analau, and Stephen]. Tapscott B7 SHP-2 activates signaling of the nuclear factor of activated T cells to promote skeletal muscle growth Mara Fornaro, Peter M. Burch, Wentian Yang, Lei Zhang, Claire E. Hamilton, Jung H. Kim, Benjamin G. Neel, and Anton M. Bennett 39 Regulation of myogenic progenitor proliferation in human fetal skeletal muscle by BMP4 and its antagonist Gremlin Natasha Y. Frank, Alvin T. Kho, Tobias Schatton, George F. Murphy, Michael J. Molloy, Qian Zhan, Marco F. Ramoni, Markus H. Frank, Isaac S. Kohane, and Emanuela Gussoni n 1 BMPl controls TGF(31 activation via cleavage of latent TGFp-binding protein Gaoxiang Ge and Daniel S. Greenspan Most newly made p-actin protein (red) lan Role of FIP200 in cardiac and liver development and its regulation colocalizes with its mRNA at the of TNFct and TSC-mTOR signaling pathways leading edge. Boyi Gan, Xu Peng, Tamas Nagy, Ana Alcaraz, Hua Gu, andJun-Lin Guan See page G7. 133 Cdc42 GEF Tuba regulates the junctional configuration of simple epithelial cells Tetsuhisa Otani, Tetsuo Ichii, Shinya Aono, and Masatoshi Takeichi n47 Shootinl: a protein involved in the organization of an asymmetric signal for neuronal polarization Michinori Toriyama, Tadayuki Shimada, Ki Bum Kim, Mari Mitsuba, Eiko Nomura, Kazuhiro Katsuta, Yuichi Sakumura, Peter Roepstorff, and Naoyuki Inagaki 159 Embryonic cortical neural stem cells migrate ventrally and persist as postnatal striatal stem cells Sandrine Willaime-Morawek, Raewyn M. Seaberg, Claudia Batista, Etienne Labbe, Liliana Attisano, Jessica A. Gorski, Kevin R. Jones, Angela Kam, Cindi M. Morshead, and Derek van der Kooy 169 Transient anchorage of cross-linked glycosyl-phosphatidylinositol- anchored proteins depends on cholesterol. Src family kinases, caveolin. and phosphoinositides Yun Chen, William R. Thelin, Bing Yang, Sharon L. Milgram, and Ken Jacobson 179 Coregulation of vascular tube stabilization by endothelial cell TIMP-2 and pericyte TIMP-3 W. Brian Saunders, Brenda L. Bohnsack, Jennifer B. Faske, Nicholas]. Anthis, Kayla J. Bayless, Karen K. Hirschi, and George E. Davis Embryonic neural stem cells (green) migrate ventrally from the cortex (top) into the striatum (Str). NEWS In This Issue T 04 • Taking a Toll on neurons • Endosomes bring up the rear • Getting the embryo into shape • Membrane backtracking • Melanin's backups THE JOURNAL OF CELL BIOLOGY M. Leslie Research Roundup VOL. 175. NO. 2. OCTOBER 23. 2006 106 • Rosettes for elongation • Lipid microscopy • Innate tumorigenesis • Mopping up chemokines • Spinning septins W.A. Wells REVIEWS Comments *100 Darkness descends with two Rabs Michael S. Marks Mini-Reviews SO'l Membranes of the world unite! Leonid V. Chernomordik, Joshua Zimmerberg, and Michael M. Kozlov RESEARCH ARTICLES Reports SO0 Toll-like receptor 8 functions as a negative regulator of neurite outgrowth and inducer of neuronal apoptosis Yinghua Ma, Jianxue Li, Isaac Chiu, Yawen Wang, Jacob A. Sloone, Jining LLi, Bela Kasaras, Richard L. Sidman, Joseph J. Voipe, and Timothy Vartanian On the cover Cell adhesion sites in the rear of a 217 Growth control by EGF repeats of the C. elegam Fibulin-1C isoform migrating cell are loosened with help Daniel Hesselson and Judith Kimble from endosomes carrying EndolSO (red), which leads to myosin light chain (blue) activation. Articles See page 33S. 225 Defective mitochondrial peroxiredoxin-3 results in sensitivity to oxidative stress in Fanconi anemia Sudit S. Mukhopadhyay, Kathryn S. Leung, M. John Hicks, PhilipJ. Hastings, Hagop Youssoufian, and Sharon E. Plon 237 Dynamic subcompartmentalization of the mitochondrial inner membrane Frank Vogel, Carsten Bornhovd, Walter Neupert, and Andreas S. Reichert 240 Reconstitution of protein targeting to the inner envelope membrane of chloroplasts Ming Li and DannyJ. Schnell Articles with related stories in the IN THIS ISSUE section have page numbers in RED; articles with COMMENTS have page numbers in BLUE. S61 SELIL. the homologue of yeast Hrd3p, is involved in protein dislocation from the mammalian ER Britta Mueller, Brendan N. Lilley, and Hidde L. Pbegh S7'\ Rab38 and Rab32 control post-Golgi trafficking of melanogenic enzymes Christina Wasmeier, Maryse Romao, Lynn Plowright, Dorothy C. Bennett, Graga Raposo, and Miguel C. Seabra SB3 MyoD-positive epiblast cells regulate skeletal muscle differentiation in the embryo Jacquelyn Gerhart, Justin Elder, Christine Neely, Jared Schure, Tage Kvist, Karen Knudsen, and Mindy George-Weinstein SS3 Protein tyrosine phosphatase TbPTPl: a molecular switch controlling life cycle differentiation in trypanosomes Baidzs Szoor, Jude Wilson, Helen McElhinney, Lydia Tabernero, and Keith R. Matthews 4 303 Phosphorylation and activity of the tumor suppressor Merlin and the ERM protein Moesin are coordinately regulated by the Slik kinase Sarah C. Hughes and Richard G. Fehon Proteins of the mitochondrial inner 315 Rapid actin monomer-insensitive depolymerization of Listeria actin membrane are unevenly distributed. comet tails by cofilin, coronin, and Aipl Corel (labeled) is part of the cristae William M. Brieher, Hao Yuan Kueh, Bryan A. Ballif, and TimothyJ. Mitchison membrane, whereas others belong to the inner boundary membrane. A model membrane section, used for plotting the 3S5 Spectrin functions upstream of ankyrin in a spectrin cytoskeleton location of gold particles, is shown. assembly pathway See page S37. Amlan Das, Christine Base, Srilakshmi Dhulipala, and Ronald R. Dubreuil 337 Endosomes generate localized Rho-ROCK-MLC2-based contractile signals via EndolSO to promote adhesion disassembly Justin Sturge, Dirk Wienke, and Clare M. Isacke 349 How the headpiece hinge angle is opened: new insights into the dynamics of integrin activation Eileen Puklin-Faucher, Mu Gao, Klaus Schulten, and Viola Vogel 361 Correction NEWS In This Issue 3G4 • Lipids be gone eIF4E exports proliferation mRNAs Histone exchange program An integrin’s gut instincts Cortex construction N. LeBrasseur Research Roundup 366 Centrosomes deliver the death blow Cosmc converter tRNA transport to the nucleus Ubp6 delays destruction Point of no return for senescence R. Williams RESEARCH ARTICLES Reports 366 A new method reveals microtubule minus ends throughout the meiotic spindle Kendro S. Burbank, Aaron C. Groen, Zachary E. Perlman, Daniel S. Fisher, and Timothy J. Mitchison ^77 Rapidly inducible changes in phosphatidylinositol 4,5-bisphosphate levels influence multiple regulatory functions of the lipid in intact living cells Peter Varnai, Baskaran Thyagarajan, Tiber Rohacs, and Tomas Balia 383 Activation of p38a/p MAPK in myogenesis via binding of the scaffold protein JLP to the cell surface protein Cdo Giichi Takaesu, Jong-Sun Kang, Gyu-Un Bae, Min-Jeong Yi, Clement M. Lee, E. Premkumar Reddy, and Robert S. Krauss Articles 383 A novel histone exchange factor, protein phosphatase 2Cy, mediates the exchange and dephosphorylation of H2A-H2B Hiroshi Kimura, Nanako Takizawa, Eric Allemand, Tetsuya Hori, Francisco J. Iborra, Naohito Nozaki, AAichiko AAuraki, Masatoshi Hagiwara, Adrian R. Krainer, Tatsuo Fukagawa, and Kafsuya Okawa 40*1 UV-induced fragmentation of Cajal bodies Mario Cioce, Severine Boulon, A. Gregory Matera, and Angus 1. Lomond 4 T 5 eIF4E is a central node of an RNA regulon that governs cellular proliferation Biljana Culjkovic, Ivan Topisirovic, Lucy Skrabanek, Melisa Ruiz-Gutierrez, and Katherine L.B. Borden 427 Spatial codes in dendritic BCl RNA Ilham A. Muslimov, Anna lacoangeli, Jurgen Brosius, and Henri Tiedge 441 PGEi stimulation of HEK293 cells generates multiple contiguous domains with different [cAMP]: role of compartmentalized phosphodiesterases Anna Terrin, Giulietta Di Benedetto, Vanessa Pertegato, York-Fong Cheung, George Baillie, Martin J. Lynch, Nicola Elvassore, Anke Prinz, Friedrich W. Herberg, Miles D. Houslay, and Manuela Zaccolo Articles with related stories in the IN THIS ISSUE section hove page numbers in RED. 453 Differential activation and function of Rho GTPases during Salmonella-)\os,[ cell interactions Jayesh C. Patel and Jorge E. Galan 465 Shigella applies molecular mimicry to subvert vinculin and invade host cells Tina Izard, Guy Tran Van Nhieu, and Philippe RJ. Bois 47'7 Reassembly of contractile actin cortex in cell blebs Guillaume T, Charras, Chi-Kuo Hu, Margaret Coughlin, and Timothy J. Mitchison 491 A postsynaptic Spectrin scaffold defines active zone size, spacing, and efficacy at the Drosophila neuromuscular junction Jan Pielage, Richard D. Fetter, and Graeme W. Davis 505 Conditional deletion of pi integrins in the intestinal epithelium causes a loss of Hedgehog expression, intestinal hyperplasia, and early postnatal lethality Robert G. Jones, Xiufen Li, Phillip D. Gray, Jinqiu Kuang, Frederic Clayton, Wade S. Samowitz, Blair B. Madison, Deborah L. Gumucio, and Scott K. Kuwada The normal (top) size and spacing of postsynaptic active and peri-active (green) zones, surrounding glutamate receptors (magenta), are perturbed in neuromuscular junctions lacking a-Spectrin (bottom). See page 4S*1. NEWS In This Issue 516 • Togetherness, molecular style • Karyopherins play the field • Untangling a deadly conspiracy • Lining up microtubules • Molecular mothballs for RNA THE JOURNAL OF CELL BIOLOGY M. Leslie Research Roundup VOL. 175, IMO. 4, NOVEMBER 20, 2006 SIB • Spitz on growth control • Fight infection with oxidation • A minus end checkpoint • Yeast’s primitive urea • Myosin’s need for speed N. LeBrasseur REVIEWS Mini-Reviews 52*1 Why yeast cells can undergo apoptosis: death in times of peace, love, and war Sobrina Buftner, Tobias Eisenberg, Eva Herker, Didac Carmona-Gutierrez, Guido Kroemer, and Frank Madeo RESEARCH ARTICLES Reports 52'7 ZBPl regulates mRNA stability during cellular stress Nadine Sfbhr, Marcell Lederer, Claudia Reinke, Sylke Meyer, Mechfhild Hafzfeld, Robert H. Singer, and Stefan Hiittelmaier S35 High levels of Notch signaling down-regulate Numb and Numblike Gavin Chapman, Lining Liu, Cecilia Sahlgren, Camilla Dahlqvist, and Urban Lendahl On the cover 541 Tau-dependent microtubule disassembly initiated The main limiting factor for nuclear by prefibrillar (3-amyloid import is the time it takes for o transport Michelle E. King, Ho-Man Kan, Peter W. Baas, Alev Erisir, Charles G. Glabe, factor (red cells) to find its cargo (green) and George S. Bloom in o sea of nonspecific competition. See page 57S. The von Hippel-Lindau tumor suppressor protein controls ciliogenesis by orienting microtubule growth Bernhard Schermer, Cristina Ghenoiu, Malte Bartram, Roman Ulrich Muller, Fruzsina Katsis, Martin Hohne, Wolfgang Kuhn, Manuela Rapka, Roland Nitschke, FHanswalter Zentgraf, Manfred Fliegauf, Heymut Omran, Gerd Walz, and Thomas Benzing 555 Filopodia formation mediated by receptor tyrosine kinase Ror2 is required for Wnt5a-induced cell migration Michiru Nishita, Sa Kan Yoo, Akira Nomachi, Shuichi Kani, Nagako Sougawa, Yasutaka Ohta, Shinji Takada, Akira Kikuchi, and Yasuhiro Minami 563 CCN3 controls 3D spatial localization of melanocytes in the human skin through DDR I Mizuho Fukunaga-Kalabis, Gabriela Martinez, Zhao-Jun Liu, Jiri Kalabis, Paul Mrass, Wolfgang Weninger, Sue M. Firth, Nathalie Planque, Bernard Perbal, and Meenhard F4erlyn Articles with related stories in the IN THIS ISSUE section have page numbers in RED. 57'I Comparative proteomics of clathrin-coated vesicles Georg H.H. Borner, Michael Harbour, Svenja Hester, Kathryn S. Lilley, and Margaret S. Robinson Articles 575 Simple kinetic relationships and nonspecific competition govern nuclear import rates in vivo Benjamin L. Timney, Jaclyn Tetenbaum-Novatt, Diana S. Agate, Rosemary Williams, Wenzhu Zhang, Brian T. Chait, and Michael P. Rout 595 Calpain is required for macroautophagy in mammalian cells Francesca Demarchi, Cosetta Bertoli, Tamara Copetti, Isei Tanida, Claudio Brancolini, Eeva-Liisa Eskelinen, and Claudio Schneider 607 IKKp programs to turn on the GADD45a-MKK4-JNK apoptotic cascade specifically via p50 NF-kB in arsenite response Proteomic analysis reveals genuine Lun Song, Jingxia Li, Dongyun Zhang, Zheng-gang Liu, Jianping Ye, Qimin Zhon, clathrin-coated vesicle proteins as red Han-Ming Shen, Matt Whiteman, and Chuanshu Huang spots and contaminants as green-to-yellow. See page 571. 619 The major human and mouse granzymes are structurally and functionally divergent Dion Koisermon, Catherina H. Bird, Jiuru Sun, Antony Motthews, Kheng Ung, James C. Whisstock, Philip E. Thompson, Joseph A. Trapani, and Phillip I. Bird 631 Transport of LAPTM5 to lysosomes requires association with the ubiquitin ligase Nedd4, but not LAPTM5 ubiquitination Youngshil Pak, Wioletta K. Glowacka, M. Christine Bruce, Nam Pham, and Daniela Rotin 6*17 Specific inhibition of GPI-anchored protein function by homing and self-association of specific GPI anchors Thomas B. Nicholson and Clifford P. Stanners 661 Lipid defect underlies selective skin barrier impairment of an epidermal-specific deletion of Gata-3 Cristino de Guzman Strong, Philip W. Wertz, Chenwei Wong, Fan Yang, Paul S. Meltzer, Thomas Andl, Sarah E. Millar, IGheng Ho, Sung-Yun Pai, and Julia A. Segre 671 Correction 673 Correction