About the Authors

Eli S Groban

Contributed equally to this work with: Eli S Groban, Arjun Narayanan

Affiliations Department of Pharmaceutical Chemistry, University of California San Francisco, San Francisco, California, United States of America , Graduate Group in Biophysics, University of California San Francisco, San Francisco, California, United States of America

Arjun Narayanan

Contributed equally to this work with: Eli S Groban, Arjun Narayanan

Affiliations Department of Pharmaceutical Chemistry, University of California San Francisco, San Francisco, California, United States of America , Graduate Group in Biophysics, University of California San Francisco, San Francisco, California, United States of America

Matthew P Jacobson

To whom correspondence should be addressed. E-mail: matt@cgl.ucsf.edu

Affiliations Department of Pharmaceutical Chemistry, University of California San Francisco, San Francisco, California, United States of America , Graduate Group in Biophysics, University of California San Francisco, San Francisco, California, United States of America

Competing Interests

MPJ is a member of the Scientific Advisory Board of Schrodinger, Inc., a company that distributes scientific software, including software that he has written. However, MPJ states that he believes that the work described in this manuscript has no significant commercial value, since the study is concerned with a fundamental problem in biophysics (i.e., it is unrelated to drug discovery, which is the company's focus).

Author Contributions

MPJ conceived and designed the experiments. ESG, AN, and MPJ performed the experiments. ESG, AN, and MPJ analyzed the data. ESG, AN, and MPJ wrote the paper.