About the Authors

Sarah E. Belisle

Contributed equally to this work with: Sarah E. Belisle, Jiangmei Yin

Affiliation Department of Microbiology, University of Washington, Seattle, Washington, United States of America

Jiangmei Yin

Contributed equally to this work with: Sarah E. Belisle, Jiangmei Yin

Affiliation Department of Pathology and Laboratory Medicine, University of Pennsylvania School of Medicine, Philadelphia, Pennsylvania, United States of America

Devon J. Shedlock

Affiliation Department of Pathology and Laboratory Medicine, University of Pennsylvania School of Medicine, Philadelphia, Pennsylvania, United States of America

Anlan Dai

Affiliation Department of Pathology and Laboratory Medicine, University of Pennsylvania School of Medicine, Philadelphia, Pennsylvania, United States of America

Jian Yan

Affiliation Department of Pathology and Laboratory Medicine, University of Pennsylvania School of Medicine, Philadelphia, Pennsylvania, United States of America

Lauren Hirao

Affiliation Department of Pathology and Laboratory Medicine, University of Pennsylvania School of Medicine, Philadelphia, Pennsylvania, United States of America

Michele A. Kutzler

Affiliation Department of Medicine, Drexel University College of Medicine, Philadelphia, Pennsylvania, United States of America

Mark G. Lewis

Affiliation Research Section, Bioqual, Rockville, Maryland, United States of America

Hanne Andersen

Affiliation Research Section, Bioqual, Rockville, Maryland, United States of America

Simon M. Lank

Affiliation Wisconsin National Primate Research Center, University of Wisconsin-Madison, Madison, Wisconsin, United States of America

Julie A. Karl

Affiliation Wisconsin National Primate Research Center, University of Wisconsin-Madison, Madison, Wisconsin, United States of America

David H. O'Connor

Affiliation Wisconsin National Primate Research Center, University of Wisconsin-Madison, Madison, Wisconsin, United States of America

Amir Khan

Affiliation Inovio Pharmaceuticals, Blue Bell, Pennsylvania, United States of America

Niranjan Sardesai

Affiliation Inovio Pharmaceuticals, Blue Bell, Pennsylvania, United States of America

Jean Chang

Affiliation Department of Microbiology, University of Washington, Seattle, Washington, United States of America

Lauri Aicher

Affiliation Department of Microbiology, University of Washington, Seattle, Washington, United States of America

Robert E. Palermo

Affiliation Department of Microbiology, University of Washington, Seattle, Washington, United States of America

David B. Weiner

Affiliation Department of Pathology and Laboratory Medicine, University of Pennsylvania School of Medicine, Philadelphia, Pennsylvania, United States of America

Michael G. Katze

Affiliation Department of Microbiology, University of Washington, Seattle, Washington, United States of America

Jean Boyer

boyerj@mail.med.upenn.edu

Affiliation Department of Pathology and Laboratory Medicine, University of Pennsylvania School of Medicine, Philadelphia, Pennsylvania, United States of America

Competing Interests

Mark G. Lewis and Hanne Anderson are employed by Bioqual, and Amir Khan and Niranjan Sardesai are employed by Inovio. Inovio provided DNA for the immunization. Primate studies were completed at Bioqual. There are no patents, products in development or marketed products to declare. This does not alter the authors' adherence to all the PLoS ONE policies on sharing data and materials.

Author Contributions

Conceived and designed the experiments: JB DBW. Performed the experiments: SEB J. Yin DS AD J. Yan LH MGL LA HA. Analyzed the data: REP MGK JB. Contributed reagents/materials/analysis tools: DBW AK NS MAK. Wrote the paper: JB SEB DS AK NS. Provided guidance: DHO. Completed analysis: JAK JC SML. Completed animal work: HA.