TSN, MKK and SYC conceived and designed the experiments. SHL, KHK and JHK performed the experiments. WY analysed MR images. TSN, SHL, JHK, WY, SHH, MJK, BAS, MDP, HEC, JJ, MKK and SYC analysed and interpreted the data. TSN, JJ, MKK and SYC wrote the manuscript. All authors read and approved the final manuscript.