Reporting summary
Further information on research design is available in the Nature
Research Reporting Summary linked to this paper.
Data availability
Source data for Figs. 1–4 and Extended Data Figs. 1, 2, 3, 4–8 are avail-
able with the paper. All other data are available from the corresponding
authors upon reasonable request.
Acknowledgements We thank members of the Brangwynne laboratory for discussions and
comments on this manuscript. This work was supported by the Howard Hughes Medical
Institute, the St Jude Collaborative on Membraneless Organelles, and grants from the National
Institutes of Health (NIH) 4D Nucleome Program (U01 DA040601) and the Princeton Center for
Complex Materials, a Materials Research Science and Engineering Center supported by the
National Science Foundation (NSF) (DMR 1420541). L.Z. was supported by an NSF graduate
fellowship (DGE-1656466). R.W.K acknowledges support from the NIH (R01 GM115634, R35
GM131891 and P30 CA021765 (to St Jude Children’s Research Hospital)) and ALSAC. M.T.
acknowledges support from the NIH (F32 GM131524). Some images were acquired at the St
Jude Cell & Tissue Imaging Center, which is supported by St Jude Children’s Research Hospital
and the National Cancer Institute (P30 CA021765); we thank V. Frohlich and J. Peters for
technical assistance.Author contributions J.A.R., L.Z., D.M.M., R.W.K. and C.P.B. designed the research; in vivo
studies were performed and analysed by J.A.R., L.Z., D.W.S. and M.W.; in vitro studies were
performed and analysed by M.C.F., M.T. and D.M.M.; J.A.R., L.Z. and C.P.B. wrote the
manuscript, which was reviewed and edited by all authors.Competing interests R.W.K. is a consultant for, and D.M.M. has recently become employed by,
Dewpoint Therapeutics. The remaining authors declare no competing interests.
Additional information
Supplementary information is available for this paper at https://doi.org/10.1038/s41586-020-
2256-2.
Correspondence and requests for materials should be addressed to R.W.K. or C.P.B.
Peer review information Nature thanks Rohit Pappu and the other, anonymous, reviewer(s) for
their contribution to the peer review of this work.
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