About the Authors

Ziying Han

Affiliation Department of Pathobiology, School of Veterinary Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, United States of America

Jonathan J. Madara

Affiliation Department of Pathobiology, School of Veterinary Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, United States of America

Andrew Herbert

Affiliation United States Army Medical Research Institute of Infectious Diseases, Fort Detrick, Maryland, United States of America

Laura I. Prugar

Affiliation United States Army Medical Research Institute of Infectious Diseases, Fort Detrick, Maryland, United States of America

Gordon Ruthel

Affiliation Department of Pathobiology, School of Veterinary Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, United States of America

Jianhong Lu

Current address: Cancer Research Institute, Central South University, Changsha, Hunan, People's Republic of China

Affiliation Department of Pathobiology, School of Veterinary Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, United States of America

Yuliang Liu

Current address: Poultry Disease Laboratory, National Veterinary Diagnostic Center (NVDC), China Animal Disease Control Center (CADC), Beijing, China

Affiliation Department of Pathobiology, School of Veterinary Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, United States of America

Wenbo Liu

Current address: Animal Infectious Disease Laboratory, School of Veterinary Medicine, Yangzhou University, Yangzhou, Jiangsu, China

Affiliation Department of Pathobiology, School of Veterinary Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, United States of America

Xiaohong Liu

Affiliation Department of Pathobiology, School of Veterinary Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, United States of America

Jay E. Wrobel

Affiliation Fox Chase Chemical Diversity Center, Inc., Doylestown, Pennsylvania, United States of America

Allen B. Reitz

Affiliation Fox Chase Chemical Diversity Center, Inc., Doylestown, Pennsylvania, United States of America

John M. Dye

Affiliation United States Army Medical Research Institute of Infectious Diseases, Fort Detrick, Maryland, United States of America

Ronald N. Harty

rharty@vet.upenn.edu (RNH); bruce@vet.upenn.edu (BDF)

Affiliation Department of Pathobiology, School of Veterinary Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, United States of America

Bruce D. Freedman

rharty@vet.upenn.edu (RNH); bruce@vet.upenn.edu (BDF)

Affiliation Department of Pathobiology, School of Veterinary Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, United States of America

Competing Interests

ABR and JEW are employed by Fox Chase Chemical Diversity, Inc. a privately held biopharmaceutical company. The company, along with RNH and BDF at the University of Pennsylvania, have a pending patent application in the area of this publication. This does not alter our adherence to all PLOS Pathogens policies on sharing data and materials.

Author Contributions

Conceived and designed the experiments: ZH JJM GR JL YL WL XL JEW ABR AH LIP JMD RNH BDF. Performed the experiments: ZH JJM GR JL YL WL XL AH LIP. Analyzed the data: ZH JJM GR JL YL WL XL JEW ABR AH LIP JMD RNH BDF. Contributed reagents/materials/analysis tools: ZH JJM GR JL YL WL XL JEW ABR AH LIP JMD RNH BDF. Wrote the paper: ZH JJM GR JL YL WL XL JEW ABR AH LIP JMD RNH BDF.