Energetics of synchronization for model flagella and cilia.

Synchronization is often observed in the swimming of flagellated cells, either for multiple appendages on the same organism or between the flagella of nearby cells. Beating cilia are also seen to synchronize their dynamics. In 1951, Taylor showed that the observed in-phase beating of the flagella of coswimming spermatozoa was consistent with minimization of the energy dissipated in the surrounding fluid. Here we revisit Taylor's hypothesis for three models of flagella and cilia: (1) Taylor's waving sheets with both longitudinal and transverse modes, as relevant for flexible flagella, (2) spheres orbiting above a no-slip surface to model interacting flexible cilia, and (3) whirling rods above a no-slip surface to address the interaction of nodal cilia. By calculating the flow fields explicitly, we show that the rate of working of the model flagella or cilia is minimized in our three models for (1) a phase difference depending on the separation of the sheets and precise waving kinematics, (2) in-phase or opposite-phase motion depending on the relative position and orientation of the spheres, and (3) in-phase whirling of the rods. These results will be useful in future models probing the dynamics of synchronization in these setups.