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Work was supported by NIH as R01 AG028730-01 and P01 AG027916 to D.S. D.S. is supported by the Glenn Foundation for Medical Research and a Senior Fellowship from the Ellison Medical Foundation Senior Scholar in Aging, A.R. gratefully acknowledges support from the NIH, a Leducq Foundation Network of Research Excellence, Judith and David Ganz, and the Maxwell Hurston Charitable Foundation. A.R. is a principal faculty member of the Harvard Stem Cell Institute