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Print ISSN: 0022-1767 Online ISSN: 1550-6606. All rights reserved.1451 Rockville Pike, Suite 650, Rockville, MD 20852The American Association of Immunologists, Inc.,

is published twice each month byThe Journal of Immunology

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VOL. 186 | NO. 3 | February 1, 2011 | Pages 1295–1882

IN THIS ISSUE1295

LETTERS TO THE EDITOR

1297 Comment on “Type 1 Diabetes in BioBreeding Rats Is Critically Linked to an Imbalance between Th17and Regulatory T Cells and an Altered TCR Repertoire”Jan-Luuk Hillebrands, Jan Rozing, and Jeroen T. J. Visser

1298 Response to Comment on “Type 1 Diabetes in BioBreeding Rats Is Critically Linked to an Imbalancebetween Th17 and Regulatory T Cells and an Altered TCR Repertoire”Jens van den Brandt and Holger M. Reichardt

1299 Comment on “Flagellin as an Adjuvant: Cellular Mechanisms and Potential”Janardan P. Pandey

1299 Response to Comment on “Flagellin as an Adjuvant: Cellular Mechanisms and Potential”Steven B. Mizel and John T. Bates

PILLARS OF IMMUNOLOGY

1301 Receptor Editing as a Mechanism of B Cell ToleranceSelvakumar Sukumar and Mark S. Schlissel

1303 Pillars Article: Receptor Editing: An Approach by Autoreactive B Cells to Escape Tolerance. The Journal ofExperimental Medicine. 1993. 177: 999–1008Denise Gay, Thomas Saunders, Sally Camper, and Martin Weigert

1313 Pillars Article: Receptor Editing in Self-reactive Bone Marrow B Cells. The Journal of Experimental Medicine.1993. 177: 1009–1020Susan L. Tiegs, David M. Russell, and David Nemazee

On the cover: Immunohistochemical staining for CD20 (red) and Ki-67 (brown) demonstrates that most proliferating cells ina germinal center from a human kidney with lupus nephritis are CD201 centroblasts. Original magnification 3200. Chang,A., S. G. Henderson, D. Brandt, N. Liu, R. Guttikonda, C. Hsieh, N. Kaverina, T. O. Utset, S. M. Meehan, R. J. Quigg,

E. Meffre, and M. R. Clark. 2011. In situ B cell-mediated immune responses and tubulointerstitial inflammation in humanlupus nephritis. J. Immunol. 186: 1849–1860.

The Journal of Immunology (ISSN 0022-1767) is published twice each month by The American Association of Immunologists,Inc., 9650 Rockville Pike, Bethesda, MD 20814-3994, Phone: 301-634-7197, Fax: 301-634-7829. Subscription terms: Newsubscriptions and renewals begin January 1 and expire one year later (see http://www.jimmunol.org/subscriptions for prices anddetails). A special rate is available to members of The American Association of Immunologists, Inc. (see http://www.aai.org/

membership). The American Association of Immunologists, Inc., is not responsible for undeliverable issues. Replacement issuescan be obtained, if available, at the regular price of single issues (see http://www.jimmunol.org/pdf/single_issue_purchase.pdf).Reprints of individual articles are available only from authors. Periodicals postage paid at Bethesda, MD 20814-3994, and at

additional mailing offices. Country of origin U.S.A. Printed on acid-free recyclable paper. Postmaster: Send address changes60 days in advance to The Journal of Immunology, Subscription Department, 9650 Rockville Pike, Bethesda, MD 20814-3994.Copyright � 2011 by The American Association of Immunologists, Inc.

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Brief Reviews

1325 Recent Developments in Cancer VaccinesKarolina Palucka, Hideki Ueno, and Jacques Banchereau

Cutting Edge

1333 Cutting Edge: NLRC5-Dependent Activation of the InflammasomeBeckley K. Davis, Reid A. Roberts, Max T. Huang, Stephen B. Willingham, Brian J. Conti, W. June Brickey,

Brianne R. Barker, Mildred Kwan, Debra J. Taxman, Mary-Ann Accavitti-Loper, Joseph A. Duncan,

and Jenny P.-Y. Ting

1338 Cutting Edge: TIGIT Has T Cell-Intrinsic Inhibitory FunctionsNicole Joller, Jason P. Hafler, Boel Brynedal, Nasim Kassam, Silvia Spoerl, Steven D. Levin, Arlene H. Sharpe,

and Vijay K. Kuchroo

CELLULAR IMMUNOLOGY AND IMMUNE REGULATION

1343 Postselection Thymocyte Maturation and Emigration Are Independent of IL-7 and ERK5Michael A. Weinreich, Stephen C. Jameson, and Kristin A. Hogquist

1348 Diverse Endogenous Antigens for Mouse NKT Cells: Self-Antigens That Are Not GlycosphingolipidsBo Pei, Anneliese O. Speak, Dawn Shepherd, Terry Butters, Vincenzo Cerundolo, Frances M. Platt, and Mitchell Kronenberg

1361 Concurrent Allorecognition Has a Limited Impact on Posttransplant VaccinationTeresa Manzo, Rodrigo Hess Michelini, Veronica Basso, Alessia Ricupito, Jian-Guo Chai, Elizabeth Simpson,

Matteo Bellone, and Anna Mondino

1369 Dragon (Repulsive Guidance Molecule b) Inhibits IL-6 Expression in MacrophagesYin Xia, Virna Cortez-Retamozo, Vera Niederkofler, Rishard Salie, Shanzhuo Chen, Tarek A. Samad, Charles C. Hong,

Silvia Arber, Jatin M. Vyas, Ralph Weissleder, Mikael J. Pittet, and Herbert Y. Lin

1377 Langerin+ Dermal Dendritic Cells Are Critical for CD8+ T Cell Activation and IgH g-1 Class Switching inResponse to Gene Gun VaccinesAngelika Stoecklinger, Tekalign D. Eticha, Mehrnaz Mesdaghi, Adrien Kissenpfennig, Bernard Malissen, Josef Thalhamer,

and Peter Hammerl

1384 Epigenetic Reduction in Invariant NKT Cells following In Utero Vitamin D Deficiency in MiceSanhong Yu and Margherita T. Cantorna

1391 Glycogen Synthase Kinase-3 Is an Early Determinant in the Differentiation of Pathogenic Th17 CellsEl�eonore Beurel, Wen-I Yeh, Suzanne M. Michalek, Laurie E. Harrington, and Richard S. Jope

1399 A Unique Role of the Cholera Toxin A1-DD Adjuvant for Long-Term Plasma and Memory B CellDevelopmentMats Bemark, Peter Bergqvist, Anneli Stensson, Anna Holmberg, Johan Mattsson, and Nils Y. Lycke

1411 CCL20 and b-Defensin-2 Induce Arrest of Human Th17 Cells on Inflamed Endothelium In Vitro underFlow ConditionsSoufiane Ghannam, C�ecile Dejou, Nathalie Pedretti, Jean-Philipe Giot, Karim Dorgham, Hassan Boukhaddaoui,

Virginie Deleuze, Francois-Xavier Bernard, Christian Jorgensen, Hans Yssel, and J�erome P�ene

1421 Functional Redundancy between Thymic CD8a+ and Sirpa+ Conventional Dendritic Cells in Presentationof Blood-Derived Lysozyme by MHC Class II ProteinsDanielle F. Atibalentja, Kenneth M. Murphy, and Emil R. Unanue

1432 CD1d-Restricted IFN-g–Secreting NKT Cells Promote Immune Complex-Induced Acute Lung Injury byRegulating Macrophage-Inflammatory Protein-1a Production and Activation of Macrophages andDendritic CellsJi Hyung Kim and Doo Hyun Chung

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1442 Transplantation Tolerance to a Single Noninherited MHC Class I Maternal Alloantigen Studied in aTCR-Transgenic Mouse ModelYoshinobu Akiyama, St�ephane M. Caucheteux, C�ecile Vernochet, Yoshiko Iwamoto, Katsunori Tanaka,

Colette Kanellopoulos-Langevin, and Gilles Benichou

1450 Runx1/Cbfb2 Complexes Are Required for Lymphoid Tissue Inducer Cell Differentiation at TwoDevelopmental StagesMasashi Tachibana, Mari Tenno, Chieko Tezuka, Machiko Sugiyama, Hisahiro Yoshida, and Ichiro Taniuchi

1458 IFN Regulatory Factor 8 Restricts the Size of the Marginal Zone and Follicular B Cell PoolsJianxun Feng, Hongsheng Wang, Dong-Mi Shin, Marek Masiuk, Chen-Feng Qi, and Herbert C. Morse, III

1467 P-Rex1 and Vav1 Cooperate in the Regulation of Formyl-Methionyl-Leucyl-Phenylalanine–DependentNeutrophil ResponsesCampbell D. Lawson, Sarah Donald, Karen E. Anderson, Daniel T. Patton, and Heidi C. E. Welch

1477 Plasmacytoid Dendritic Cell Dichotomy: Identification of IFN-a Producing Cells as a Phenotypicallyand Functionally Distinct SubsetPia Bjorck, H. Xianne Leong, and Edgar G. Engleman

1486 TNF Receptor-1 Is Required for the Formation of Splenic Compartments during Adult, but Not EmbryonicLifeNovica M. Mili�cevi�c , Karola Klaperski, Klaus Nohroudi, �Zivana Mili�cevi�c , Katja Bieber, Babett Baraniec,

Maike Blessenohl, Kathrin Kalies, Carl F. Ware, and Jurgen Westermann

1495 IL-23–Mediated Psoriasis-Like Epidermal Hyperplasia Is Dependent on IL-17AHeather L. Rizzo, Shinji Kagami, Kevin G. Phillips, Stephen E. Kurtz, Steven L. Jacques, and Andrew Blauvelt

1503 CpG Promotes Cross-Presentation of Dead Cell-Associated Antigens by Pre-CD8a+ Dendritic CellsChristelle de Brito, Martine Tomkowiak, Raffaella Ghittoni, Christophe Caux, Yann Leverrier, and Jacqueline Marvel

1512 Invariant NKT Cells Inhibit Autoreactive B Cells in a Contact- and CD1d-Dependent MannerJun-Qi Yang, Xiangshu Wen, Peter J. Kim, and Ram Raj Singh

1521 Lung Cancer-Derived Galectin-1 Mediates Dendritic Cell Anergy through Inhibitor of DNA Binding3/IL-10 Signaling PathwayPo-Lin Kuo, Jen-Yu Hung, Shau-Ku Huang, Shah-Hwa Chou, Da-En Cheng, Yuh-Jyh Jong, Chih-Hsing Hung,

Chih-Jen Yang, Ying-Ming Tsai, Ya-Ling Hsu, and Ming-Shyan Huang

1531 Restricted Microbiota and Absence of Cognate TCR Antigen Leads to an Unbalanced Generation of Th17CellsMatthias Lochner, Marion B�erard, Shinichiro Sawa, Siona Hauer, Val�erie Gaboriau-Routhiau, Tahia Diana Fernandez,

Johannes Snel, Philippe Bousso, Nadine Cerf-Bensussan, and G�erard Eberl

1538 A Human NK Cell Activation/Inhibition Threshold Allows Small Changes in the Target Cell SurfacePhenotype To Dramatically Alter Susceptibility to NK CellsTim D. Holmes, Yasser M. El-Sherbiny, Adam Davison, Sally L. Clough, G. Eric Blair, and Graham P. Cook

1546 Foxp3+ Regulatory T Cells Control Humoral Autoimmunity by Suppressing the Development ofLong-Lived Plasma CellsEunkyeong Jang, Wang Sik Cho, Mi-La Cho, Hyun-Joo Park, Hye-Joa Oh, Sang Mee Kang, Doo-Jin Paik, and Jeehee Youn

1554 CD22 Is a Recycling Receptor That Can Shuttle Cargo between the Cell Surface and EndosomalCompartments of B CellsMary K. O’Reilly, Hua Tian, and James C. Paulson

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MOLECULAR AND STRUCTURAL IMMUNOLOGY

1564 Autophagy Regulates Endoplasmic Reticulum Homeostasis and Calcium Mobilization in T LymphocytesWei Jia, Heather H. Pua, Qi-Jing Li, and You-Wen He

IMMUNOGENETICS

1575 Although Divergent in Residues of the Peptide Binding Site, Conserved Chimpanzee Patr-ALand Polymorphic Human HLA-A*02 Have Overlapping Peptide-Binding RepertoiresMichael Gleimer, Angela R. Wahl, Heather D. Hickman, Laurent Abi-Rached, Paul J. Norman, Lisbeth A. Guethlein,

John A. Hammond, Monia Draghi, Erin J. Adams, Sean Juo, Roxana Jalili, Baback Gharizadeh, Mostafa Ronaghi,

K. Christopher Garcia, William H. Hildebrand, and Peter Parham

HOST DEFENSE

1589 Persistence of Gut Mucosal Innate Immune Defenses by Enteric a-Defensin Expression in the SimianImmunodeficiency Virus Model of AIDSMelinda M. Zaragoza, Sumathi Sankaran-Walters, Don R. Canfield, Jason K. S. Hung, Enrique Martinez,

Andr�e J. Ouellette, and Satya Dandekar

1598 CD4 T Cells Promote Rather than Control Tuberculosis in the Absence of PD-1–Mediated InhibitionDaniel L. Barber, Katrin D. Mayer-Barber, Carl G. Feng, Arlene H. Sharpe, and Alan Sher

1608 TLR 2 and CD14 Mediate Innate Immunity and Lung Inflammation to Staphylococcal Panton–ValentineLeukocidin In VivoAna Zivkovic, Omar Sharif, Karin Stich, Bianca Doninger, Mario Biaggio, Jacques Colinge, Martin Bilban, Ildiko Mesteri,

Parastoo Hazemi, Rosa Lemmens-Gruber, and Sylvia Knapp

1618 RIG-I/MDA5/MAVS Are Required To Signal a Protective IFN Response in Rotavirus-Infected IntestinalEpitheliumAlexis H. Broquet, Yoshihiro Hirata, Christopher S. McAllister, and Martin F. Kagnoff

1627 Mycobacterium tuberculosis Directs Immunofocusing of CD8+ T Cell Responses Despite VaccinationJoshua S. Woodworth, Daniel Shin, Mattijs Volman, Cl�audio Nunes-Alves, Sarah M. Fortune, and Samuel M. Behar

1638 c-Maf–Dependent Growth of Mycobacterium tuberculosis in a CD14hi Subpopulation ofMonocyte-Derived MacrophagesRohan Dhiman, Anuradha Bandaru, Peter F. Barnes, Sudipto Saha, Amy Tvinnereim, Ramesh C. Nayak,

Padmaja Paidipally, Vijaya Lakshmi Valluri, L. Vijaya Mohan Rao, and Ramakrishna Vankayalapati

1646 NLRP4 Negatively Regulates Autophagic Processes through an Association with Beclin1Nao Jounai, Kouji Kobiyama, Masaaki Shiina, Kazuhiro Ogata, Ken J. Ishii, and Fumihiko Takeshita

1656 Pyk2 Is Required for Neutrophil Degranulation and Host Defense Responses to Bacterial InfectionLynn A. Kamen, Joseph Schlessinger, and Clifford A. Lowell

1666 Influenza A Inhibits Th17-Mediated Host Defense against Bacterial Pneumonia in MiceAnupa Kudva, Erich V. Scheller, Keven M. Robinson, Chris R. Crowe, Sun Mi Choi, Samantha R. Slight,

Shabaana A. Khader, Patricia J. Dubin, Richard I. Enelow, Jay K. Kolls, and John F. Alcorn

1675 IL-17 Contributes to Cell-Mediated Defense against Pulmonary Yersinia pestis InfectionJr-Shiuan Lin, Lawrence W. Kummer, Frank M. Szaba, and Stephen T. Smiley

1685 A Novel Role for IFN-Stimulated Gene Factor 3II in IFN-g Signaling and Induction of Antiviral Activity inHuman CellsAngel N. Morrow, Hana Schmeisser, Takaya Tsuno, and Kathryn C. Zoon

1694 EBV Lytic-Phase Protein BGLF5 Contributes to TLR9 Downregulation during Productive InfectionMichiel van Gent, Bryan D. Griffin, Eufemia G. Berkhoff, Daphne van Leeuwen, Ingrid G. J. Boer, Marlyse Buisson,

Franca C. Hartgers, Wim P. Burmeister, Emmanuel J. Wiertz, and Maaike E. Ressing

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1703 T Cell-Intrinsic Factors Contribute to the Differential Ability of CD8+ T Cells To Rapidly Secrete IFN-g inthe Absence of AntigenElsa N. Bou Ghanem, Christina C. Nelson, and Sarah E. F. D’Orazio

1713 Functional Consequences of Natural Sequence Variation of Murine Cytomegalovirus m157 for Ly49Receptor Specificity and NK Cell ActivationAlexandra J. Corbett, Jerome D. Coudert, Catherine A. Forbes, and Anthony A. Scalzo

INFLAMMATION

1723 Mechanistic Role of MicroRNA-146a in Endotoxin-Induced Differential Cross-Regulation of TLRSignalingMd A. Nahid, Minoru Satoh, and Edward K. L. Chan

1735 Controlling Herpes Simplex Virus-Induced Ocular Inflammatory Lesions with the Lipid-Derived MediatorResolvin E1Naveen K. Rajasagi, Pradeep B. J. Reddy, Amol Suryawanshi, Sachin Mulik, Per Gjorstrup, and Barry T. Rouse

1747 IL-1a Modulates Neutrophil Recruitment in Chronic Inflammation Induced by Hydrocarbon OilPui Y. Lee, Yutaro Kumagai, Yuan Xu, Yi Li, Tolga Barker, Chao Liu, Eric S. Sobel, Osamu Takeuchi, Shizuo Akira,

Minoru Satoh, and Westley H. Reeves

1755 CR2+ Marginal Zone B Cell Production of Pathogenic Natural Antibodies Is C3 IndependentKeith M. Woods, Michael R. Pope, Sara M. Hoffman, and Sherry D. Fleming

1763 Identification of Endothelial Cell Junctional Proteins and Lymphocyte Receptors Involved inTransendothelial Migration of Human Effector Memory CD4+ T CellsThomas D. Manes and Jordan S. Pober

1769 A Virus-Like Particle-Based Anti-Nerve Growth Factor Vaccine Reduces Inflammatory Hyperalgesia:Potential Long-Term Therapy for Chronic PainTill A. Rohn, William T. Ralvenius, Jolly Paul, Petra Borter, Marcela Hernandez, Robert Witschi, Paula Grest,

Hanns Ulrich Zeilhofer, Martin F. Bachmann, and Gary T. Jennings

1781 Tumor-Associated a2 Vacuolar ATPase Acts As a Key Mediator of Cancer-Related Inflammation byInducing Pro-Tumorigenic Properties in MonocytesChristina Kwong, Alice Gilman-Sachs, and Kenneth Beaman

1790 IFN-g Attenuates Hypoxia-Inducible Factor (HIF) Activity in Intestinal Epithelial Cells throughTranscriptional Repression of HIF-1bLouise E. Glover, Karina Irizarry, Melanie Scully, Eric L. Campbell, Brittelle E. Bowers, Carol M. Aherne,

Douglas J. Kominsky, Christopher F. MacManus, and Sean P. Colgan

1799 Spontaneous Pregnancy Loss Mediated by Abnormal Maternal Inflammation in Rats Is Linked to DeficientUteroplacental PerfusionStephen J. Renaud, Tiziana Cotechini, Jill S. Quirt, Shannyn K. Macdonald-Goodfellow, Maha Othman,

and Charles H. Graham

1809 Senescent CD14+CD16+ Monocytes Exhibit Proinflammatory and Proatherosclerotic ActivityAna Merino, Paula Buendia, Alejandro Martin-Malo, Pedro Aljama, Rafael Ramirez, and Julia Carracedo

1816 Endothelial ICAM-1 Protein Induction Is Regulated by Cytosolic Phospholipase A2a via Both NF-kBand CREB Transcription FactorsNurit Hadad, Liron Tuval, Vered Elgazar-Carmom, Ron Levy, and Rachel Levy

CLINICAL IMMUNOLOGY

1828 Provision of Continuous Maturation Signaling to Dendritic Cells by RIG-I–Stimulating Cytosolic RNASynthesis of Sendai VirusShinji Okano, Yoshikazu Yonemitsu, Ken Shirabe, Yoshihiro Kakeji, Yoshihiko Maehara, Mamoru Harada,

Yasunobu Yoshikai, Makoto Inoue, Mamoru Hasegawa, and Katsuo Sueishi

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1840 Daratumumab, a Novel Therapeutic Human CD38 Monoclonal Antibody, Induces Killing of MultipleMyeloma and Other Hematological TumorsMichel de Weers, Yu-Tzu Tai, Michael S. van der Veer, Joost M. Bakker, Tom Vink, Dani€elle C. H. Jacobs,

Lukas A. Oomen, Matthias Peipp, Thomas Valerius, Jerry W. Slootstra, Tuna Mutis, Wim K. Bleeker,

Kenneth C. Anderson, Henk M. Lokhorst, Jan G. J. van de Winkel, and Paul W. H. I. Parren

1849 In Situ B Cell-Mediated Immune Responses and Tubulointerstitial Inflammation in Human LupusNephritisAnthony Chang, Scott G. Henderson, Daniel Brandt, Ni Liu, Riteesha Guttikonda, Christine Hsieh, Natasha Kaverina,

Tammy O. Utset, Shane M. Meehan, Richard J. Quigg, Eric Meffre, and Marcus R. Clark

1861 Gene Expression Patterns of Th2 Inflammation and Intercellular Communication in Asthmatic AirwaysDavid F. Choy, Barmak Modrek, Alexander R. Abbas, Sarah Kummerfeld, Hilary F. Clark, Lawren C. Wu,

Grazyna Fedorowicz, Zora Modrusan, John V. Fahy, Prescott G. Woodruff, and Joseph R. Arron

1870 IL-1b Is Overexpressed and Aberrantly Regulated in Corticosteroid Nonresponders with AutoimmuneInner Ear DiseaseShresh Pathak, Elliot Goldofsky, Esther X. Vivas, Vincent R. Bonagura, and Andrea Vambutas

CORRECTIONS

1880 Correction: An Inflammasome-Independent Role for Epithelial-Expressed Nlrp3 in RenalIschemia-Reperfusion InjuryAlana A. Shigeoka, James L. Mueller, Amanpreet Kambo, John C. Mathison, Andrew J. King, Wesley F. Hall,

Jean da Silva Correia, Richard J. Ulevitch, Hal M. Hoffman, and Dianne B. McKay

1881 AUTHOR INDEX

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