Key Laboratory for Neuroinformation of Ministry of Education,University of Electronic Sciences and Technology of China,Chengdu,China;Center for Life Sciences,Shanghai Institutes of Biological Sciences,Chinese Academy of Sciences,Shanghai,China;,Key Laboratory for Neuroinformation of Ministry of Education,University of Electronic Sciences and Technology of China,Chengdu,China;Key Laboratory for Neuroinformation of Ministry of Education,University of Electronic Sciences and Technology of China,Chengdu,China,Center for Life Sciences, Shanghai Institutes of Biological Sciences, Chinese Academy of Sciences, Shanghai, China

This work was supported by grants from National Basic Research Program of China (2013CB329401), China Postdoctoral Science Foundation (2012M521681, 2013T60844), and The National Natural Science Foundation of China (31300912, 31100797, 90820301, 31000492)