Each video was analyzed using TrackMate Fiji, an ImageJ plugin. The cells were localized and tracked using the LAP (Linear Assignment Problem) tracker. The displacement trajectories of each of the selected cells over time along the y axis were extracted from the tracking data, and single-cell velocities were calculated by dividing the total displacements of the rolled cells by the total number of frames, during which the cell showed continuous rolling behaviors. Mean cell velocities were calculated after applying selection criteria: The cells that were not detected in more than 75 consecutive frames were considered as passing cells and were excluded from the analysis.

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