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Microtubules driven by dense kinesin motility assay : from individual motion to large-scale collective emergent structures

谷田, 桜子 東京大学 DOI:10.15083/0002004336

2022.06.22

概要

Microtubules (MTs) are functional intracellular scaffolds. They do not just support the cell body. They also produce abundant cell motion emerging in essential periods of the cell cycle, for example, cell division. During the cell motion, MTs are moving, growing, shrinking, and colliding, and then finally create cooperative structure and motion. Such cooperative structure and motion seem to have similarities with the dynamics of a collection composed of motile elements. For example, MTs confined between the endoplasmic reticulum and the cell membrane align themselves parallel. Similarly, rod-shaped bacteria confined in a thin chamber also spontaneously align their direction of the long axis parallel. However, it is necessary to carefully examine whether they are the same in properties or just similar on the impression. In other words, we must discuss similarities and differences by expressing the structure and motion as appropriate order parameters and examining their control-parameter dependencies. The cooperative structure and motion of MTs also need to be investigated for their order parameters. We assume that the cooperative structure and motion of MTs are comparable with the dynamics of another collection of motile elements. Then, what we should focus on is the dynamics and the interaction of MTs rather than on various reactions of MTs with multiple molecules. However, many molecules are involved in a living cell. So, it is hard to investigate the cooperative structure and motion which are purely produced by the dynamics and interaction of MTs. Thus, a bottom-up approach by a simple experiment is needed.

In order to study the cooperative structure and motion of MTs in a bottom-up approach based on the motivation described above, we investigated the collective motion of MTs in a reconstructed system called “motility assay”. The motility assay is composed of stabilized MTs and kinesin motors. The kinesins are immobilized on a glass surface. They kick MTs along the long axis of the MTs by using the energy of ATP hydrolysis. We found that the several types of collective motion emerge by changing the kinesin density. At a high kinesin density [(6.2 ± 0.6) × 103 kinesins per µm2 ], MTs form clusters that move in a random direction when MT density was above 0.04∼0.08 MTs per µm2 . On the other hand, at a lower kinesin density [(2.5 ± 0.7) × 103 kinesins per µm2 ], MTs align their orientation globally when MT density was above 0.05 MTs per µm2 . To understand the mechanisms of those pattern formations, the interaction between MTs was examined. When two MTs collided with each other, MTs aligned their orientation perfectly at a certain probability. Additionally, an MT was able to cross over the other MT at a probability only at a low kinesin density. At 2.5 × 103 kinesins per µm2 , about 5% of colliding pairs showed cross over. At 6.2 × 103 kinesins per µm2 , no pair of crossing was observed. Whether MTs can cross or not can be interpreted as the difference in the strength of the steric interaction. These results indicate that too much strong steric interaction breaks the global alignment even though MTs have alignment interaction. In addition to those collective emergent structures, rotational phenomena have been observed. At a high kinesin density, loops of 1-µm radius are formed. At a low kinesin density, the orientation of global alignment rotates in a counterclockwise direction.

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