The authors thank Prof. Lei Liu of Capital Medical University for his comments and Prof. Christopher J. Chang of University of California for kindly providing the RFAP-1 probe. This work was supported by the Beijing Brain Initiative of Beijing Municipal Science & Technology Commission Z181100001518104; the NSFC 61625102, 61827810, and 81571044 and HTRDC 2016YFC1305903 and 2016YFC1306302; 973 Program, 2015CB856402; the Beijing Natural Science Foundation of China 7172022; the Scientific Research Common Program of Beijing Municipal Commission of Education KM201510025014; and the Major Projects Fund of Beijing Institute for Brain Disorders ZD2015-08, China. W.S. was the holder of the Tier 1 Canada Research Chair in Alzheimer's Disease. Y.Z. is the recipient of Michael Smith Foundation for Health Research Post-Doctoral Fellowship Award. Author contributions: H.H. and Z.T. conceived the study. W.S. H.H. and Z.T. designed the study and supervised all analyses. XY, YZ, WS, and ZT wrote the manuscript. R.H. and B.S. contributed to study design and experiments. X.Y. Y.M. H.L. L.A. Y.L. C.D. G.G. H.Y. B.S. and R.L. performed experiments. H.H. and Z.T. performed the data analyses. All authors reviewed the manuscript. The authors declare that they have no competing financial interests.