About the Authors

John A. Mengshol

Affiliations Department of Medicine, Division of Gastroenterology & Hepatology, University of Colorado School of Medicine, Aurora, Colorado, United States of America, Denver Veterans Affairs Medical Center, Denver, Colorado, United States of America

Lucy Golden-Mason

Affiliations Department of Medicine, Division of Gastroenterology & Hepatology, University of Colorado School of Medicine, Aurora, Colorado, United States of America, Division of Clinical Immunology, University of Colorado School of Medicine and National Jewish Hospital, Denver, Colorado, United States of America

Tomohiro Arikawa

Affiliation Department of Immunology and Immunopathology, Kagawa Medical University, Kagawa, Japan

Maxwell Smith

Affiliation Department of Pathology, University of Colorado School of Medicine, Denver, Colorado, United States of America

Toshiro Niki

Affiliation GalPharma, Kagawa, Japan

Ryan McWilliams

Affiliation Department of Medicine, Division of Gastroenterology & Hepatology, University of Colorado School of Medicine, Aurora, Colorado, United States of America

Jessica A. Randall

Affiliation Department of Medicine, Division of Gastroenterology & Hepatology, University of Colorado School of Medicine, Aurora, Colorado, United States of America

Rachel McMahan

Affiliation Department of Medicine, Division of Gastroenterology & Hepatology, University of Colorado School of Medicine, Aurora, Colorado, United States of America

Michael A. Zimmerman

Affiliation Division of Transplant Surgery, University of Colorado School of Medicine, Aurora, Colorado, United States of America

Manu Rangachari

Affiliation Center for Neurologic Diseases, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts, United States of America

Evgenia Dobrinskikh

Affiliation Department of Medicine, Division of Gastroenterology & Hepatology, University of Colorado School of Medicine, Aurora, Colorado, United States of America

Pierre Busson

Affiliation Institut de Cancérologie Gustave Roussy, Villejuif, France

Stephen J. Polyak

Affiliation Division of Laboratory Medicine, University of Washington, Seattle, Washington, United States of America

Mitsuomi Hirashima

Affiliation Department of Immunology and Immunopathology, Kagawa Medical University, Kagawa, Japan

Hugo R. Rosen

Hugo.Rosen@ucdenver.edu

Affiliations Department of Medicine, Division of Gastroenterology & Hepatology, University of Colorado School of Medicine, Aurora, Colorado, United States of America, Denver Veterans Affairs Medical Center, Denver, Colorado, United States of America, Division of Clinical Immunology, University of Colorado School of Medicine and National Jewish Hospital, Denver, Colorado, United States of America

Competing Interests

Dr. Niki works for GalPharma and supplied antibodies for this study. This does not alter adherence to all the PLoS ONE policies on sharing data and materials, as detailed online.

Author Contributions

Conceived and designed the experiments: JAM LGM TA MS RM MZ SJP MH HRR. Performed the experiments: JAM LGM TA MS TN RM JAR RM ED SJP MH. Analyzed the data: JAM LGM TA MS RM JAR RM MR ED PB SJP MH HRR. Contributed reagents/materials/analysis tools: JAM LGM TA MS TN RM MZ MR ED PB SJP MH. Wrote the paper: JAM LGM RM HRR.