About the Authors

Izumi C Mori

Contributed equally to this work with: Izumi C Mori, Yoshiyuki Murata, Yingzhen Yang

Current address: Research Institute for Bioresources, Okayama University, Chuo, Kurashiki, Japan

Affiliation Cell and Developmental Biology Section, Division of Biological Sciences and Center for Molecular Genetics, University of California San Diego, La Jolla, California, United States of America

Yoshiyuki Murata

Contributed equally to this work with: Izumi C Mori, Yoshiyuki Murata, Yingzhen Yang

Affiliations Cell and Developmental Biology Section, Division of Biological Sciences and Center for Molecular Genetics, University of California San Diego, La Jolla, California, United States of America , Department of Agriculture, Okayama University, Okayama, Japan

Yingzhen Yang

Contributed equally to this work with: Izumi C Mori, Yoshiyuki Murata, Yingzhen Yang

Affiliation Cell and Developmental Biology Section, Division of Biological Sciences and Center for Molecular Genetics, University of California San Diego, La Jolla, California, United States of America

Shintaro Munemasa

Affiliations Cell and Developmental Biology Section, Division of Biological Sciences and Center for Molecular Genetics, University of California San Diego, La Jolla, California, United States of America , Department of Agriculture, Okayama University, Okayama, Japan

Yong-Fei Wang

Affiliation Cell and Developmental Biology Section, Division of Biological Sciences and Center for Molecular Genetics, University of California San Diego, La Jolla, California, United States of America

Shannon Andreoli

Affiliation Cell and Developmental Biology Section, Division of Biological Sciences and Center for Molecular Genetics, University of California San Diego, La Jolla, California, United States of America

Hervé Tiriac

Affiliation Cell and Developmental Biology Section, Division of Biological Sciences and Center for Molecular Genetics, University of California San Diego, La Jolla, California, United States of America

Jose M Alonso

Current address: Department of Genetics, North Carolina State University, Raleigh, North Carolina, United States of America

Affiliation The Salk Institute of Biological Studies, La Jolla, California, United States of America

Jeffery F Harper

Affiliation Biochemistry Department, University of Nevada, Reno, Nevada, United States of America

Joseph R Ecker

Affiliation The Salk Institute of Biological Studies, La Jolla, California, United States of America

June M Kwak

Current address: Department of Cell Biology and Molecular Genetics, University of Maryland, College Park, Maryland, United States of America

Affiliation Cell and Developmental Biology Section, Division of Biological Sciences and Center for Molecular Genetics, University of California San Diego, La Jolla, California, United States of America

Julian I Schroeder

To whom correspondence should be addressed. E-mail: julian@biomail.ucsd.edu

Affiliation Cell and Developmental Biology Section, Division of Biological Sciences and Center for Molecular Genetics, University of California San Diego, La Jolla, California, United States of America

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

JIS conceived and designed the experiments. ICM, YM, YY, SM, YFW, SA, HT, and JMK performed the experiments. ICM, YM, YY, JMK, and JIS analyzed the data. JMA, JFH, and JRE contributed reagents/materials/analysis tools. ICM and JIS wrote the paper with contributions from JMK.