We thank Drs. A. Thomas Look, David M. Langenau, Anurag Singh, Neil J. Ganem, and Herbert Cohen for helpful discussion and suggestions; as well as Dr. Michael T. Kirber from the Cellular Imaging Core facility at Boston University School of Medicine for his expert help with imaging acquisitions. This study was supported by grants from the NIH (R00CA134743, R56CA215059 and Boston University [BU] pilot grants through 1UL1TR001430 to H.F., R01CA096899 to M.A.K., predoctoral training grant through T32GM008541 to L.N.H.) and fellowship grants from the Rally Foundation and the Alex Lemonade Stand Foundation (N.M.A.). H.F. also acknowledges funding support through a Karin Grunebaum Faculty Fellowship, a BU Ralph Edwards Career Development Professorship, a Young Investigator Award from the Leukemia Research Foundation, a St. Baldrick Scholar grant, and the American Cancer Society (IRG –72-001-36-IRG and RSG-17-204-01-TBG). Y.S., B.L., and J.W.C received the Undergraduate Research Opportunity Program Award at BU. The content of this research is solely the responsibility of the authors and does not necessarily represent the official views of the NIH.

Author contributions:

H.F. conceived and supervised the project. The experiments were designed by H.F., L.N.H., N.M.A, F.J.F.L., J.E.R., and C.T.L; and performed by L.N.H., Y.S., B.L., Y.W.W., J.W.C., G.L.Z., L.W., C.T.L., and J.E.R.. Data analyses were performed by H.F., L.N.H., L.W., C.T.L., and G.L.Z.. Manuscripts were written by H.F. and L.N.H., and revised by N.M.A., Y.S., B.L., and M.A.K.