MGR performed all the analyses and wrote the paper. DBJ conceived of many of the analyses: directed the project: and extensively revised the paper. CR and CN contributed analytic ideas and helped craft and edit the manuscript. MGR: CR: CN and DBJ jointly devised the experimental design. MC: AH: NJL: and RH wrote and edited portions of the text: designed and developed lab protocols: and provided intellectual input. All authors read and approved the final manuscript.