About the Authors

Zhiyu Liu

Contributed equally to this work with: Zhiyu Liu, Herong Shi

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Herong Shi

Contributed equally to this work with: Zhiyu Liu, Herong Shi

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Lindsey C. Szymczak

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Taner Aydin

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Sijung Yun

Affiliation Laboratory of Molecular Biology, National Institute of Diabetes and Digestive and Kidney Diseases, Bethesda, Maryland, United States of America

Katharine Constas

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Arielle Schaeffer

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Sinthu Ranjan

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Saad Kubba

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Emad Alam

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Devin E. McMahon

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Jingpeng He

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Neta Shwartz

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Chenxi Tian

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Yevgeniy Plavskin

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Amanda Lindy

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Nimra Amir Dad

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Sunny Sheth

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Nirav M. Amin

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Stephanie Zimmerman

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Dennis Liu

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Erich M. Schwarz

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Harold Smith

Affiliation Laboratory of Molecular Biology, National Institute of Diabetes and Digestive and Kidney Diseases, Bethesda, Maryland, United States of America

Michael W. Krause

Affiliation Laboratory of Molecular Biology, National Institute of Diabetes and Digestive and Kidney Diseases, Bethesda, Maryland, United States of America

Jun Liu

JL53@cornell.edu

Affiliation Department of Molecular Biology and Genetics, Cornell University, Ithaca, New York, United States of America

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Wrote the paper: ZL SY CT EMS MWK JL. Conceived and oversaw the execution of the entire project: JL. Except for those specified below, all other data were generated by: ZL HSh JL. Genetic screening and initial mapping: KC AS SR SK EA YP AL NAD SZ DL JL. Performed the genetic experiments shown in Tables 1 and 2: LCS TA DEM NMA JL. Generated data shown in Table 7, Table 8 and S3 Table: TA SS ZL JL. Performed WGS and analyzed WGS data: SY HSm MWK JL. Performed experiments shown in S1 Table: JH. Generated data included in Fig 3B: HSh CT JL. Performed the yeast two-hybrid experiments: HSh NS JL. Did the phylogenetic analysis shown in S1 Fig and S4 Table: EMS.