About the Authors

Jamie J. Arnold

jja5@psu.edu (JJA); cec9@psu.edu (CEC)

Affiliation Department of Biochemistry and Molecular Biology, The Pennsylvania State University, University Park, Pennsylvania, United States of America

Suresh D. Sharma

Affiliation Department of Biochemistry and Molecular Biology, The Pennsylvania State University, University Park, Pennsylvania, United States of America

Joy Y. Feng

Affiliation Gilead Sciences, Inc., Foster City, California, United States of America

Adrian S. Ray

Affiliation Gilead Sciences, Inc., Foster City, California, United States of America

Eric D. Smidansky

Affiliation Department of Biochemistry and Molecular Biology, The Pennsylvania State University, University Park, Pennsylvania, United States of America

Maria L. Kireeva

Affiliation Frederick National Laboratory for Cancer Research, NCI, Frederick, Maryland, United States of America

Aesop Cho

Affiliation Gilead Sciences, Inc., Foster City, California, United States of America

Jason Perry

Affiliation Gilead Sciences, Inc., Foster City, California, United States of America

Jennifer E. Vela

Affiliation Gilead Sciences, Inc., Foster City, California, United States of America

Yeojin Park

Affiliation Gilead Sciences, Inc., Foster City, California, United States of America

Yili Xu

Affiliation Gilead Sciences, Inc., Foster City, California, United States of America

Yang Tian

Affiliation Gilead Sciences, Inc., Foster City, California, United States of America

Darius Babusis

Affiliation Gilead Sciences, Inc., Foster City, California, United States of America

Ona Barauskus

Affiliation Gilead Sciences, Inc., Foster City, California, United States of America

Blake R. Peterson

Affiliation Department of Medicinal Chemistry, The University of Kansas, Lawrence, Kansas, United States of America

Averell Gnatt

Affiliation Department of Pharmacology, University of Maryland School of Medicine, Baltimore, Maryland, United States of America

Mikhail Kashlev

Affiliation Frederick National Laboratory for Cancer Research, NCI, Frederick, Maryland, United States of America

Weidong Zhong

Address Current address: Novartis Institutes for BioMedical Research, Emeryville, California, United States of America.

Affiliation Gilead Sciences, Inc., Foster City, California, United States of America

Craig E. Cameron

jja5@psu.edu (JJA); cec9@psu.edu (CEC)

Affiliation Department of Biochemistry and Molecular Biology, The Pennsylvania State University, University Park, Pennsylvania, United States of America

Competing Interests

JYF, ASR, AC, JP, JEV, YP, YX, YT, DB, OB and WZ are current or former employees and shareholders of Gilead Sciences, Inc. that is responsible for the active clinical development of GS-7977 and has a partnership with Roche related to the development of mericitabine. JJA and CEC are collaborating with INDIGO Biosciences, Inc. for the development and distribution of mitochondrial toxicity assays. This does not alter our adherence to all PLOS Pathogens policies on sharing data and materials.

Author Contributions

Conceived and designed the experiments: JJA SDS JYF ASR EDS MLK MK WZ CEC. Performed the experiments: JJA SDS JYF ASR EDS MLK AC JP JEV YP YX YT DB OB. Analyzed the data: JJA SDS JYF ASR EDS MLK MK CEC. Contributed reagents/materials/analysis tools: JYF ASR BRP AG MK WZ CEC. Wrote the paper: JJA JYF ASR EDS MLK CEC.