We thank J. K. Kim for discussing single-cell sequencing. This work was supported by grants from Suh Kyungbae Foundation and IBS-R002-D1 to J.H.L. (last author), the Korean Health Technology R&D Project, Ministry of Health & Welfare, South Korea (H15C3143 and H16C0415 to J.H.L. (last author), HI14C1324 and HI17C2586 to S.G.K.), the Global PhD Fellowship Program through the National Research Foundation (NRF) of Korea funded by the Ministry of Education, Republic of Korea (NRF-2014H1A2A1021321 to J.H.L. (first author)), the Basic Science Research Program through the NRF of Korea (NRF-2016R1D1A1A09916521 to S.G.K.) funded by the Ministry of Education, the NRF of Korea grant (NRF-2017M2A2A7A01071036 to S.G.K.) funded by the Korean Ministry of Science, ICT and Future Planning, the Basic Science Research Program through the NRF of Korea (NRF-2017R1A2B2006526 to S.P.) funded by the Ministry of Science, ICT and Future Planning, and Korea Health Technology R&D Project through the Korea Health Industry Development Institute (HI16C2387 to Y.S.J.) funded by the Ministry of Health and Welfare. Non-cancer brain tissues were provided by the Netherlands Brain Bank (project number Lee-835) to J.H.L. (last author).

Reviewer information

Nature thanks M. Taylor and the other anonymous reviewer(s) for their contribution to the peer review of this work.