While combining this tool with other methodologies has already made cryo-electron tomography more biologically significant, we need to continue this cooperation using recently developed biotechnology and cell biology approaches. We expect further development of this technique for specimen preparation, data acquisition, and analysis. Now, after nearly 10 years of application of this technique, we are turning a corner and are at the stage to discuss the next steps. Since the mid 2000s, cryo-electron tomography has provided us with new knowledge, concepts, and questions in the area of cilia research. In particular, we have identified the locations and conformations of all the regular inner and outer dyneins, as well as various regulators such as radial spokes. It opened a way to reconstruct 3D conformations of proteins in cilia at 3-nm resolution, revealed networks of a number of component proteins in cilia, and has even allowed the study of component dynamics. Cryo-electron tomography has been a valuable tool in the analysis of 3D structures of cilia at molecular and cellular levels.
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