Correspondence to Masahide Kikkawa: mkikkawa@m.u-tokyo.ac.jp Abbreviations used in this paper: BCCP, biotin carboxyl carrier protein; CCD, charge-coupled device; CP, central pair apparatus; DMT, doublet microtubule; fps, frames per second; IC 50 , half-maximal inhibitory concentration; IDA, inner dynein arm; ODA, outer dynein arm; RS, radial spoke; RSP, radial spoke protein.