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Mark Johnston, Editor-in-Chief University of Colorado-Denver Tracey DePellegrin Connelly, Editorial Board Executive Editor

CELLULAR GENETICS Thomas C. Kaufman GENOME AND SYSTEMS Wolfgang Stephan Orna Cohen-Fix Indiana University BIOLOGY University of Munich NIDDK, National Institutes of Health Kenneth J. Kemphues Jef Boeke Naoyuki Takahata Susan Dutcher Cornell University Johns Hopkins School of Medicine Graduate University for Washington University School Barbara J. Meyer Stanley Fields Advanced Studies of Medicine University of California, Berkeley University of Washington Marcy K. Uyenoyama JoAnne Engebrecht Trudi Schüpbach David Largaespada Duke University University of California, Davis Princeton University University of Minnesota Lindi M. Wahl University of Western Ontario David I. Greenstein Mariana F. Wolfner Jeffery F. Miller University of Minnesota Cornell University University of California, Los Angeles John Wakeley Michael Nonet GENE EXPRESSION Peter J. Oefner Washington University School of Stanford University H. Allen Orr (Perspectives Editor) Karen Arndt Medicine University of Pittsburgh Norbert Perrimon Adam S. Wilkins (Perspectives Editor) Mark D. Rose Harvard Medical School James A. Birchler University of Cambridge Princeton University Piali Sengupta Patricia J. Pukkila (Genetics Education Linda Siracusa Brandeis University Charles Boone Editor) University of North Carolina Jefferson Medical College of Thomas Gary Stormo University of Toronto Jasper Rine (Reviews Editor) Jefferson University Washington University School Pamela Geyer of Medicine University of California, Berkeley The University of Iowa COMPLEX TRAITS Daniel F. Voytas Michael Turelli (Reviews Editor) Justin Borevitz David Jonah Grunwald University of Minnesota University of California, Davis University of Chicago University of Utah EMPIRICAL POPULATION BOARD OF SENIOR EDITORS Edward Buckler Michael Hampsey Oliver Hobert Cornell University Robert Wood Johnson Medical GENETICS Montserrat Aguadé Columbia University Alain Charcosset School-UMDNJ Universitat de Barcelona Charles H. Langley Institut National de la Recherche Ann Hochschild David Begun University of California, Davis Agronomique Harvard Medical School University of California, Davis Terry R. Magnuson Steve Chenoweth Aaron P. Mitchell James J. Bull University of North Carolina, Chapel Hill University of Queensland Carnegie Mellon University University of Texas at Austin Lauren M. McIntyre Gary A. Churchill Craig S. Pikaard Deborah Charlesworth University of Florida The Jackson Laboratory Indiana University University of Edinburgh Rasmus Nielsen Ina Hoeschele Eric Selker Anna Di Rienzo University of Copenhagen, Centre for Virginia Polytechnic Institute and State University of Chicago Bioinformatics University John Tamkun Jeffrey Lawrence Mark Rose Corbin D. Jones University of California, Santa Cruz University of Pittsburgh Princeton University The University of North Carolina GENOME INTEGRITY AND Manyuan Long John C. Schimenti at Chapel Hill TRANSMISSION University of Chicago Cornell University Matias Kirst Eric E. Alani Michael Nachman Fred Winston University of Florida Cornell University University of Arizona Harvard Medical School Krista Nichols Sharon E. Bickel Bret Payseur Purdue University Dartmouth College University of Wisconsin-Madison GENETICS publishes high quality, Fernando F. Pardo Manuel de Villena Monica Colaiácovo Outi Savolainen original research presenting novel University of North Carolina Harvard Medical School University of Oulu findings on a range of topics bearing on heredity and variation. GENETICS Andrew H. Paterson Gregory Copenhaver Michel Veuille is a peer-edited journal—all editorial University of Georgia University of North Carolina Ecole Pratique des Hautes Etudes, David Threadgill Paris decisions are made by the authors’ Nancy Hollingsworth University of North Carolina Stephen I. Wright peers—with a tradition of rigorous Stony Brook University peer review. Publication is open Hongyu Zhao University of Toronto Andreas Houben Yale University equally to members and non-members Leibniz Institute of Plant Genetics THEORETICAL POPULATION of the Genetics Society of America. Fei Zou and Crop Plant Research GENETICS Submit manuscripts at http://submit. University of North Carolina, Chapel Hill Scott Keeney Mark Beaumont genetics.org. Address questions DEVELOPMENTAL AND Memorial Sloan-Kettering University of Bristol to the BEHAVIORAL GENETICS Cancer Center Laurent Excoffier GENETICS Editorial Office, Robert R. H. Anholt David Leach University of Bern Mellon Institute, Box I, North Carolina State University University of Edinburgh Marcus W. Feldman 4400 Fifth Avenue, Pittsburgh, PA Bonnie Bartel Jac A. Nickoloff Stanford University 15213-2683. Rice University Colorado State University Noah Rosenberg Telephone: Lynn Cooley Jeff Sekelsky Stanford University (412) 268-1812 Yale School of Medicine University of North Carolina Yun S. Song William M. Gelbart Franklin Stahl University of California, Fax: Harvard University University of Oregon Berkeley (412) 268-1813 Lawrence G. Harshman Ting (C-ting) Wu Hamish Spencer E-mail: University of Nebraska, Lincoln Harvard Medical School University of Otago [email protected]