(A,B) Violin plots of pdfs of speed () across all cells, and all time points, for CR and PO. (Note the scaling of the pdfs is linear for CR, but logarithmic for PO.) (C,D) Time-averaged heatmaps of the cells’ centroid positions, for each of the four different trap sizes, overlaid with arrows showing the direction and strength of steady-state non-equilibrium fluxes computed from trajectory statistics, for CR and PO. (E) Traced cilia waveforms for CR showing the asymmetric beat pattern of the two cilia under red light. (F) The asymmetric dumbbell model used for CR with a smaller circle (radius a2) for the cell body and a larger circle (radius a1) representing the area covered by the cilia beat, which is offset from the swimming direction by angle . Comparison of the experimental mean and standard error () of the radial probability densities (G), mean-squared displacement curves (H) and mean tangential flux (I) with the simulation results for each trap size. The inset in (I) shows the time-averaged heatmap of centroid position overlaid with arrows showing the probability flux results for the simulation of the 40 μm diameter trap.