About the Authors

Khalid S. Mohammad

Affiliation Department of Internal Medicine, Division of Endocrinology, University of Virginia, Charlottesville, Virginia, United States of America

Carol G. Chen

Affiliations Graduate Program in Oral and Craniofacial Sciences, University of California San Francisco, San Francisco, California, United States of America, Department of Orthopaedic Surgery, University of California San Francisco, San Francisco, California, United States of America

Guive Balooch

Affiliation Department of Materials Science and Engineering, University of California, Berkeley and Materials Science Division, Lawrence Berkeley National Laboratories, Berkeley, California, United States of America

Elizabeth Stebbins

Affiliation Scios, Inc, Fremont, California, United States of America

C. Ryan McKenna

Affiliation Department of Internal Medicine, Division of Endocrinology, University of Virginia, Charlottesville, Virginia, United States of America

Holly Davis

Affiliation Department of Internal Medicine, Division of Endocrinology, University of Virginia, Charlottesville, Virginia, United States of America

Maria Niewolna

Affiliation Department of Internal Medicine, Division of Endocrinology, University of Virginia, Charlottesville, Virginia, United States of America

Xiang Hong Peng

Affiliation Department of Internal Medicine, Division of Endocrinology, University of Virginia, Charlottesville, Virginia, United States of America

Daniel H. N. Nguyen

Affiliation Department of Orthopaedic Surgery, University of California San Francisco, San Francisco, California, United States of America

Sophi S. Ionova-Martin

Affiliation Scios, Inc, Fremont, California, United States of America

John W. Bracey

Affiliation Department of Orthopaedic Surgery, Center for Orthopaedic Research, Barton Research Institute, University of Arkansas for Medical Sciences, Little Rock, Arkansas, United States of America

William R. Hogue

Affiliation Department of Orthopaedic Surgery, Center for Orthopaedic Research, Barton Research Institute, University of Arkansas for Medical Sciences, Little Rock, Arkansas, United States of America

Darren H. Wong

Affiliations Scios, Inc, Fremont, California, United States of America, Pfizer RTC, Cambridge, Massachusetts, United States of America

Robert O. Ritchie

Affiliation Department of Materials Science and Engineering, University of California, Berkeley and Materials Science Division, Lawrence Berkeley National Laboratories, Berkeley, California, United States of America

Larry J. Suva

Affiliation Department of Orthopaedic Surgery, Center for Orthopaedic Research, Barton Research Institute, University of Arkansas for Medical Sciences, Little Rock, Arkansas, United States of America

Rik Derynck

Affiliations Graduate Program in Oral and Craniofacial Sciences, University of California San Francisco, San Francisco, California, United States of America, Department of Cell and Tissue Biology, University of California San Francisco, San Francisco, California, United States of America, Institute of Regeneration Medicine, University of California San Francisco, San Francisco, California, United States of America

Theresa A. Guise

Affiliation Department of Internal Medicine, Division of Endocrinology, University of Virginia, Charlottesville, Virginia, United States of America

Tamara Alliston

tamara.alliston@ucsf.edu

Affiliations Graduate Program in Oral and Craniofacial Sciences, University of California San Francisco, San Francisco, California, United States of America, Department of Orthopaedic Surgery, University of California San Francisco, San Francisco, California, United States of America, Institute of Regeneration Medicine, University of California San Francisco, San Francisco, California, United States of America

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: KM RD. Performed the experiments: KM SIM. Analyzed the data: KM. Contributed reagents/materials/analysis tools: KM. Wrote the paper: KM. Wrote the manuscript with critical input from all authors: TA. Planned and performed most of the experiments with assistance from CRM, HD, MN, XHP: KSM. Planned and performed nanoindentation analyses: CC. Performed XTM: GB. Participated in study design: ES. Administered drugs and did DXA measurements: HD RM. Performed CFU-F & CFU-OB: MN. Did the histology: XHP. Planned and performed gene expression studies: DN TA. Performed microCT: WH JB. Participated in study design: DW. Planned fracture toughness tests and X-ray tomography, which were performed by SSI-M and GB respectively: RR. Planned micro-tomography and macromechanical tests, which were performed by JWB and WRH: LS. Designed and coordinated the study and supervised all experiments: TAG TA.