Y.M. is supported by a Cancer Research Institute Dr. Keith Landesman Memorial Fellowship (CRI5247). L.S.Q. acknowledges support by National Science Foundation, National Institutes of Health and the Chau Hoi Shuen Foundation Limited. This work is supported by National Human Genome Research Institute (R21HG013133) and NIH Director’s Pioneer Award (DP1NS137219). L.S.Q. is a Chan Zuckerberg Biohub – San Francisco Investigator.