EDITOR IN CHIEF (2016)
Harold L. Drake
University of Bayreuth
EDITORS
Johanna Björkroth, Editor(2018)
University of Helsinki
Axel A. Brakhage, Editor(2016)
HKI and University of Jena
Daniel Cullen, Editor (2017)
USDA Forest Products Laboratory
Christopher A. Elkins, Editor (2018)
U.S. FDA Center for Food Safety and Applied Nutrition
Heidi Goodrich-Blair, Editor(2017)
University of Wisconsin—Madison
Mansel W. Griffiths, Editor (2015)
University of Guelph
Robert M. Kelly, Editor(2017)
North Carolina State University
Maia Kivisaar, Editor (2017)
Tartu University
Joel E. Kostka, Editor (2016)
Georgia Institute of Technology
Shuang-Jiang Liu, Editor (2016)
Chinese Academy of Sciences
Frank E. Löffler, Editor (2018)
University of Tennessee
Charles R. Lovell, Editor (2015)
University of South Carolina
George T. Macfarlane, Editor (2016)
University of Dundee
Volker Müller, Minireview Editor(2014)
Goethe University Frankfurt am Main
Hideaki Nojiri, Editor (2017)
The University of Tokyo
Rebecca E. Parales, Editor (2016)
University of California—Davis
Matthew R. Parsek, Cover Editor (2014)
University of Washington
M. Julia Pettinari, Editor (2016)
University of Buenos Aires
Donald W. Schaffner, Editor (2015)
Rutgers, The State University of New Jersey
Janet L. Schottel, Editor (2018)
University of Minnesota
Alfred M. Spormann, Editor (2016)
Stanford University
Gerrit Voordouw, Editor (2017)
University of Calgary
K. Eric Wommack, Editor (2017)
University of Delaware
Thomas K. Wood, Minireview Editor (2017)
Pennsylvania State University
Marylynn V. Yates, Editor (2014)
University of California—Riverside
Thomas E. Shenk, Chairman, Journals Board
Barbara M. Goldman, Director, Journals
Barbara S. Slinker, Production Editor
Stephanie L. D'Agostino, Assistant Production Editor
Michael E. Lerman, Assistant Production Editor
EDITORIAL BOARD
Adly Abd-Alla (2015)
Luis A. Actis (2014)
Gladys Alexandre (2015)
Kevin J. Allen (2015)
Eric Altermann (2015)
Rustam Aminov (2013)
Gary L. Andersen (2013)
Matthew J. Arduino (2013)
Arthur I. Aronson (2015)
Shota Atsumi (2015)
Brian Austin (2013)
Beate Averhoff (2013)
Abdu F. Azad (2013)
M. Andrea Azcarate-Peril (2015)
Reha O. Azizoglu (2015)
Charles W. Bacon (2015)
Michael Bagdasarian (2015)
Alan T. Bakalinsky (2013)
Petr Baldrian (2013)
Jeffrey Banas (2015)
Ravi D. Barabote (2015)
József Baranyi (2013)
Florian F. Bauer (2015)
Gwyn A. Beattie (2014)
Teresa M. Bergholz (2015)
Stefan Bertilsson (2015)
Thomas Besser (2013)
Arvind A. Bhagwat (2015)
Valeria Bianciotto (2015)
Linda F. Bisson (2013)
Christopher Blackwood (2013)
Rafael Blasco (2013)
Melanie Blokesch (2013)
Paul L. Bodelier (2015)
Eckhard Boles (2015)
Matthias Boll (2015)
Daniel R. Bond (2015)
Violaine Bonnefoy (2013)
Robert A. Bonomo (2013)
Kathryn J. Boor (2013)
James Borneman (2014)
Albert Bosch (2015)
Peter Bottomley (2015)
Kostas Bourtzis (2015)
Byron F. Brehm-Stecher (2013)
Frederick Breidt, Jr. (2013)
Harry Brumer (2014)
Alison Buchan (2014)
Robert L. Buchanan (2015)
Daniel H. Buckley (2014)
William Burkhardt (2015)
H. J. Busscher (2013)
Douglas R. Call (2013)
Amy Callaghan (2015)
Terri A. Camesano (2015)
Isaac K. O. Cann (2015)
Michelle Q. Carter (2015)
Shane M. Ceraul (2015)
Woo-Suk Chang (2015)
Jinru Chen (2014)
Shicheng Chen (2014)
Wilfred Chen (2015)
Yi Chen (2014)
Yuhuan Chen (2015)
Michael L. Chikindas (2013)
Ludmila Chistoserdova (2015)
Kuang-Ren Chung (2014)
Jeffrey D. Cirillo (2014)
Luca Cocolin (2013)
Michael T. Collins (2013)
Doug Cossar (2015)
Don A. Cowan (2015)
Patricia Cruz (2013)
Holger Daims (2015)
Rolf Daniel (2015)
Steven L. Daniel (2014)
Michelle Danyluk (2013)
Atin R. Datta (2013)
Svetlana Dedysh (2015)
Zixin Deng (2013)
Angelo DePaola, Jr. (2015)
Mickaël Desvaux (2015)
Ronald P. de Vries (2014)
Thomas J. DiChristina (2015)
Francisco Diez-Gonzalez (2013)
Alan A. DiSpirito (2014)
Jan Dolfing (2015)
James S. Dooley (2015)
Irina S. Druzhinina (2015)
Doris d'Souza (2014)
Nicole Dubilier (2013)
Edward Dudley (2015)
Siobain Duffy (2013)
Jonathan Dworkin (2014)
Daniel C. Eastwood (2014)
Leo Eberl (2014)
Elizabeth A. Edwards (2013)
Henry L. Ehrlich (2015)
Patrick Eichenberger (2015)
Marie Elliot (2014)
Mostafa Elshahed (2015)
David Emerson (2015)
K.-D. Entian (2013)
Slava Epstein (2013)
Danilo Ercolini (2015)
Joseph O. Falkinham III (2013)
Seamus Fanning (2015)
Matthew W. Fields (2014)
Martin Filion (2015)
Steven E. Finkel (2013)
Steven Foley (2014)
Philipp Franken (2015)
Herbert L. Fredrickson (2013)
Michael W. Friedrich (2015)
Cormac Gahan (2015)
Rebecca Gast (2014)
Beilei Ge (2013)
Elke Genersch (2015)
Terry Joe Gentry (2014)
Johannes Gescher (2015)
Emanuel Goldman (2013)
Mark Gomelsky (2014)
Ramon Gonzalez (2013)
Lawrence Goodridge (2015)
Lisa Gorski (2015)
Joerg Graf (2014)
Jeffrey Gralnick (2015)
Irene R. Grant (2013)
Stefan J. Green (2014)
Leslie Gregg-Jolly (2014)
Richard L. Gregory (2015)
Jean Guard (2013)
Jan O. Haerter (2015)
Thomas E. Hanson (2015)
Eiji Haramoto (2015)
Hauke Harms (2013)
Steven Harris (2014)
Valerie J. Harwood (2015)
Paul B. Hatzinger (2015)
Terry C. Hazen (2013)
Ji-Zheng He (2014)
Qiang He (2013)
Zhili He (2013)
Brian Hedlund (2014)
Henry S. Heine (2015)
Hermann J. Heipieper (2014)
Ute Hentschel (2015)
Niels B. Hendriksen (2015)
Laura Hendrix (2015)
Salvador Herrero (2015)
Paul S. Hoffman (2015)
Dirk Hoffmeister (2015)
Andrew J. Holmes (2013)
Allen L. Honeyman (2015)
Dallas G. Hoover (2015)
David J. Hopper (2013)
Marcus A. Horn (2014)
Hong-Ying Hu (2013)
John Andrew Hudson (2014)
Jeroen Hugenholtz (2015)
Hor-Gil Hur (2015)
Michele M. Igo (2015)
Fumio Inagaki (2013)
Ralph W. Jack (2015)
Paul J. Jackson (2015)
Mohamed Jamal (2015)
Janet K. Jansson (2013)
Edward L. Jarroll (2015)
Thomas W. Jeffries (2014)
Dieter Jendrossek (2015)
Paul R. Jensen (2014)
Ian Jenson (2014)
Nianzhi Jiao (2014)
Zackary I. Johnson (2015)
Yves Jouanneau (2015)
Juan Luis Jurat-Fuentes (2015)
Matthias Kaestner (2013)
Boran Kartal (2014)
Daniel Kearns (2013)
James E. Keen (2013)
John J. Kelly (2015)
William Kenealy (2015)
Angela D. Kent (2013)
Philip J. Kersten (2013)
Eungbin Kim (2015)
James E. Kirby (2013)
Mary Jo Kirisits (2013)
Martin G. Klotz (2014)
Kalmia E. Kniel (2014)
Rob Knight (2013)
Steffen Kolb (2014)
Konstantinos T. Konstantinidis (2015)
Shige Koseki (2015)
Balasubrahmanyam Kottapalli (2014)
George Kowalchuk (2015)
Sven Krappmann (2015)
Lee R. Krumholz (2015)
Indira T. Kudva (2013)
Ursula Kües (2013)
Kirsten Kuesel (2013)
Timothy M. LaPara (2015)
Gisèle LaPointe (2013)
Robert Alan LaRossa (2015)
Luis Larrondo (2014)
Peter C. K. Lau (2015)
Jared R. Leadbetter (2015)
Brian Lefebvre (2013)
Jeffrey T. LeJeune (2015)
Paulo Alexandre da Costa Lemos (2015)
Tine Rask Licht (2014)
Nic D. Lindley (2013)
John T. Lisle (2015)
Wen-Tso Liu (2013)
Xueduan Liu (2013)
Karen Lloyd (2015)
Martin J. Loessner (2015)
John C. Londesborough (2015)
Alexander Loy (2014)
Yahai Lu (2013)
Petra Luber (2014)
Tillmann Lueders (2015)
Yanhe Ma (2013)
Sandra Macfarlane (2014)
Robert L. Mach (2015)
Eugene L. Madsen (2015)
Jon K. Magnuson (2013)
Mike Manefield (2014)
Maria L. Marco (2015)
Helene Marquis (2015)
Phyllis A. W. Martin (2014)
Stephen Martin (2015)
Jan Martinussen (2014)
Emma Master (2015)
Timothy E. Mattes (2014)
Ann G. Matthysse (2015)
Harold D. May (2013)
Andrew J. McBain (2015)
Timothy R. McDermott (2015)
James McKinlay (2015)
Robert J. McLean (2015)
Brian B. McSpadden-Gardener (2013)
Rainer U. Meckenstock (2015)
Rick J. Meinersmann (2015)
Aubrey F. Mendonca (2015)
John Scott Meschke (2014)
Chris H. Miller (2015)
Patricia D. Millner (2015)
Kiwamu Minamisawa (2015)
Melanie R. Mormile (2013)
Jochen A. Mueller (2015)
Yuji Nagata (2015)
István Nagy (2013)
Cindy H. Nakatsu (2013)
Josh D. Neufeld (2014)
Nancy Nichols (2015)
Walter G. Niehaus, Jr. (2013)
Brendan A. Niemira (2015)
R. Sean Norman (2015)
Kevin P. O'Connell (2014)
Julie B. Olson (2015)
Huub Op den Camp (2015)
Ronald S. Oremland (2015)
Mark Osborn (2013)
Konstantinos Papadimitriou (2013)
Ilenys M. Perez-Diaz (2015)
Gregg S. Pettis (2014)
Jennifer Pett-Ridge (2015)
Laurent Philippot (2014)
Allen T. Phillips (2015)
Flynn W. Picardal (2013)
Carmen Pin (2015)
James L. Pinckney (2014)
Antonio G. Pisabarro (2014)
Mircea Podar (2014)
David L. Popham (2015)
Rolf Prade (2015)
M. Auxiliadora Prieto (2015)
Jim I. Prosser (2014)
Cheryl L. Quinn (2015)
Ralf Rabus (2014)
Andreja Rajkovic (2015)
Gemma Reguera (2015)
Bernd H. Rehm (2015)
Gregor Reid (2015)
Christian Riedel (2015)
Kirsti M. Ritalahti (2015)
Jorge Rodrigues (2013)
David Rodrígues-Lázaro (2015)
Margie F. Romine (2014)
Michael J. Sadowsky (2013)
Daâd A. Saffarini (2014)
Badal C. Saha (2013)
Maria Sandkvist (2015)
Robert A. Sanford (2015)
Anderson S. Sant'Ana (2015)
Sima Sariaslani (2015)
Mahfuzur R. Sarker (2013)
Brian D. Sauders (2013)
Bernhard Schink (2015)
Patrick D. Schloss (2015)
Monika Schmoll (2014)
Dirk Schüler (2015)
Lynne Sehulster (2013)
Bernhard Seiboth (2014)
Verena Seidl (2015)
Jeremy D. Semrau (2014)
Peter Setlow (2015)
Gordon Shephard (2013)
Joshua D. Shrout (2015)
Mark W. Silby (2015)
Simon D. Silver (2014)
Mitchell H. Singer (2014)
Steven W. Singer (2015)
Brajesh K. Singh (2013)
David Singleton (2015)
Kaarina Sivonen (2014)
Panos N. Skandamis (2013)
Evan Skowronski (2015)
Joan L. Slonczewski (2014)
Helena Solo-Gabriele (2015)
John R. Spear (2013)
Andrew J. Spiers (2013)
Eric V. Stabb (2013)
Alfons J. M. Stams (2013)
Thaddeus B. Stanton (2015)
Lisa Y. Stein (2013)
Alexander Steinbüchel (2014)
Roger Stephan (2014)
Brian Stevenson (2013)
George C. Stewart (2015)
John F. Stolz (2013)
Wolfgang R. Streit (2015)
Joseph M. Sturino (2013)
Marcelino Suzuki (2013)
Ken Takai (2013)
Ralph Tanner (2014)
Gerald W. Tannock (2014)
Sam Telford (2015)
Andreas P. Teske (2015)
Michael G. Thomas (2015)
Kai Martin Thormann (2015)
Thomas Thurnheer (2015)
Nicolas Toro Garcia (2014)
Effie Tsakalidou (2013)
Masataka Tsuda (2015)
Wouter R. van der Star (2014)
Paul W. van der Wielen (2014)
Tina Kangas Van Dyk (2015)
Marco Ventura (2015)
Alain A. Vertes (2013)
Claire Vieille (2015)
Adrián A. Vojnov (2013)
Dmitriy V. Volokhov (2015)
Julia A. Vorholt (2015)
Stéphane Vuilleumier (2015)
Carmen Wacher (2013)
Irene Wagner-Döbler (2014)
Aljoscha Wahl (2015)
Judy D. Wall (2015)
Aijie Wang (2013)
Todd J. Ward (2013)
Keith Warriner (2014)
J. Scott Weese (2015)
Bart C. Weimer (2013)
Helge Weingart (2015)
Alan Welman (2015)
Tingyi Wen (2015)
Volker F. Wendisch (2015)
Marvin Whiteley (2013)
Giovanni Widmer (2013)
Steve Wilhelm (2015)
Charles F. Wimpee (2014)
Regina-Michaela Wittich (2014)
Randy W. Worobo (2013)
Hong-sheng Wu (2015)
Qinglong Wu (2015)
Chuanwu Xi (2014)
Hua Xiang (2015)
Lihua Xiao (2015)
Luying Xun (2015)
Jagjit S. Yadav (2014)
S. Steve Yan (2013)
X. Frank Yang (2015)
Ming Yang (2013)
Yunfeng Yang (2015)
Chris Yeager (2013)
Suleyman Yildirim (2014)
Daniel R. Zeigler (2015)
Jill Zeilstra-Ryalls (2014)
Karsten Zengler (2014)
Chuanlun Zhang (2015)
Qijing Zhang (2014)
Wei Zhang (2015)
Yu-Zhong Zhang (2014)
Ning-Yi Zhou (2015)
Yong-Guan Zhu (2013)
Erik R. Zinser (2013)
Erwin G. Zoetendal (2015)
Angeles Zorreguieta (2013)
Sergey B. Zotchev (2015)
Peter Alan Zuber (2014)
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