国家重点基础研究发展计划973计划2015CB964903;中国医学科学院医学科学创新基金2016-12M-1-003,2017-I2M-1-015;国家自然科学基金31471116,81600083,31700783;"协和青年基金"和"中央高校基本科研业务费专项资金"2017310023,2017PT31033National Basic Research Program of China973 Program2015CB964903;Chi-nese Academy of Medical SciencesCAMSInnovation Fund for Medical Sciences2016-12M-1-003,2017-I2M-1-015;National Natural Science Foundation of China31471116,81600083,31700783;Pe-king Union Medical CollegePUMCYouth Fund and Fundamental Research Funds for the Central Univer-sities2017310023,2017PT31033