重点基础研究发展计划2015CB964903;中国医学科学院医学科学创新基金2016-12M-1-003;国家自然科学基金批准号:31471116、81600083资助的课题 This work was supported by the National Basic Research Program of ChinaGrant 2015CB964903;Chinese Academy of Medical Sciences Innovation Fund for Medical SciencesGrant 2016-12M-1-003;National Natural Science Foundation of ChinaGrant 31471116,81600083