About the Authors

Cheng Cui

Affiliations Department of Developmental Biology, University of Pittsburgh School of Medicine, Pittsburgh, Pennsylvania, United States of America, Laboratory of Developmental Biology, National Heart Lung and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America

Bishwanath Chatterjee

Affiliations Department of Developmental Biology, University of Pittsburgh School of Medicine, Pittsburgh, Pennsylvania, United States of America, Laboratory of Developmental Biology, National Heart Lung and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America

Thomas P. Lozito

Affiliation Center for Cellular and Molecular Engineering, Department of Orthopedic Surgery, University of Pittsburgh School of Medicine, Pittsburgh, Pennsylvania, United States of America

Zhen Zhang

Affiliation Department of Developmental Biology, University of Pittsburgh School of Medicine, Pittsburgh, Pennsylvania, United States of America

Richard J. Francis

Affiliations Department of Developmental Biology, University of Pittsburgh School of Medicine, Pittsburgh, Pennsylvania, United States of America, Laboratory of Developmental Biology, National Heart Lung and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America

Hisato Yagi

Affiliation Department of Developmental Biology, University of Pittsburgh School of Medicine, Pittsburgh, Pennsylvania, United States of America

Lisa M. Swanhart

Current address: Department of Biology, Canisius College, Buffalo, New York, United States of America.

Affiliation Department of Developmental Biology, University of Pittsburgh School of Medicine, Pittsburgh, Pennsylvania, United States of America

Subramaniam Sanker

Affiliation Department of Developmental Biology, University of Pittsburgh School of Medicine, Pittsburgh, Pennsylvania, United States of America

Deanne Francis

Affiliation Laboratory of Developmental Biology, National Heart Lung and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America

Qing Yu

Affiliation Laboratory of Developmental Biology, National Heart Lung and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America

Jovenal T. San Agustin

Affiliation Program in Molecular Medicine, University of Massachusetts Medical Center, Worcester, Massachusetts, United States of America

Chandrakala Puligilla

Current address: Department of Pathology and Laboratory Medicine, Medical University of South Carolina, Charleston, South Carolina, United States of America.

Affiliation Section on Developmental Neuroscience, National Institute on Deafness and Other Communication Disorders, National Institutes of Health, Bethesda, Maryland, United States of America

Tania Chatterjee

Affiliation Laboratory of Developmental Biology, National Heart Lung and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America

Terry Tansey

Affiliation Laboratory of Developmental Biology, National Heart Lung and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America

Xiaoqin Liu

Affiliation Department of Developmental Biology, University of Pittsburgh School of Medicine, Pittsburgh, Pennsylvania, United States of America

Matthew W. Kelley

Affiliation Section on Developmental Neuroscience, National Institute on Deafness and Other Communication Disorders, National Institutes of Health, Bethesda, Maryland, United States of America

Elias T. Spiliotis

Affiliation Department of Biology, Drexel University, Philadelphia, Pennsylvania, United States of America

Adam V. Kwiatkowski

Affiliation Department of Cell Biology, University of Pittsburgh School of Medicine, Pittsburgh, Pennsylvania, United States of America

Rocky Tuan

Affiliation Center for Cellular and Molecular Engineering, Department of Orthopedic Surgery, University of Pittsburgh School of Medicine, Pittsburgh, Pennsylvania, United States of America

Gregory J. Pazour

Affiliation Program in Molecular Medicine, University of Massachusetts Medical Center, Worcester, Massachusetts, United States of America

Neil A. Hukriede

Affiliation Department of Developmental Biology, University of Pittsburgh School of Medicine, Pittsburgh, Pennsylvania, United States of America

Cecilia W. Lo

cel36@pitt.edu

Affiliations Department of Developmental Biology, University of Pittsburgh School of Medicine, Pittsburgh, Pennsylvania, United States of America, Laboratory of Developmental Biology, National Heart Lung and Blood Institute, National Institutes of Health, Bethesda, Maryland, United States of America

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

The author(s) have made the following declarations about their contributions: Conceived and designed the experiments: CC CWL. Performed the experiments: CC BC TPL ZZ RJF HY LMA SS DF QY JTS CP TC TT XL. Analyzed the data: CC BC TPL ZZ RJF HY LMA SS MWK ETS AVK GJP NAH CWL. Contributed reagents/materials/analysis tools: MWK ETS AVK RT GJP NAH CWL. Wrote the paper: CC CWL.