About the Authors

Qiaoning Guan

Contributed equally to this work with: Qiaoning Guan, Wei Zheng, Shijie Tang

Affiliation Laboratories of Genetics and Molecular Biology, University of Wisconsin, Madison, Wisconsin, United States of America

Wei Zheng

Contributed equally to this work with: Qiaoning Guan, Wei Zheng, Shijie Tang

Affiliation Laboratories of Genetics and Molecular Biology, University of Wisconsin, Madison, Wisconsin, United States of America

Shijie Tang

Contributed equally to this work with: Qiaoning Guan, Wei Zheng, Shijie Tang

Affiliation Department of Statistics, University of Wisconsin, Madison, Wisconsin, United States of America

Xiaosong Liu

Affiliation Department of Physics, University of Wisconsin, Madison, Wisconsin, United States of America

Robert A Zinkel

Affiliation Laboratories of Genetics and Molecular Biology, University of Wisconsin, Madison, Wisconsin, United States of America

Kam-Wah Tsui

Affiliation Department of Statistics, University of Wisconsin, Madison, Wisconsin, United States of America

Brian S Yandell

Affiliations Department of Statistics, University of Wisconsin, Madison, Wisconsin, United States of America , Department of Horticulture, University of Wisconsin, Madison, Wisconsin, United States of America

Michael R Culbertson

To whom correspondence should be addressed. E-mail: mrculber@wisc.edu

Affiliation Laboratories of Genetics and Molecular Biology, University of Wisconsin, Madison, Wisconsin, United States of America

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

MRC conceived and designed the experiments. QG, WZ, and RAZ performed the experiments. WZ, ST, and XL developed statistical methods, and KWT and BSY provided statistical advice and expertise. The paper was written by QG, WZ, and MRC.