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Axonemal dyneins are the molecular motors responsible for the beating ofcilia and flagella. These motors generate sliding forces between adjacentmicrotubule doublets within the axoneme, the motile cytoskeletal structureinside the flagellum. To create regular, oscillatory beating patterns, theactivities of the axonemal dyneins must be coordinated both spatially andtemporally. It is thought that coordination is mediated by stresses or strainsthat build up within the moving axoneme, but it is not known which componentsof stress or strain are involved, nor how they feed back on the dyneins. Toanswer this question, we used isolated, reactivate axonemes of the unicellularalga Chlamydomonas as a model system. We derived a theory for beat regulationin a two-dimensional model of the axoneme. We then tested the theory bymeasuring the beat waveforms of wild type axonemes, which have asymmetricbeats, and mutant axonemes, in which the beat is nearly symmetric, usinghigh-precision spatial and temporal imaging. We found that regulation bysliding forces fails to account for the measured beat, due to the short lengthsof Chlamydomonas cilia. We found that regulation by normal forces (which tendto separate adjacent doublets) cannot satisfactorily account for the symmetricwaveforms of the mbo2 mutants. This is due to the model's failure to producereciprocal inhibition across the axes of the symmetrically beating axonemes.Finally, we show that regulation by curvature accords with the measurements.Unexpectedly, we found that the phase of the curvature feedback indicates thatthe dyneins are regulated by the dynamic (i.e. time-varying) component ofaxonemal curvature, but not by the static one. We conclude that a high-passfiltered curvature signal is a good candidate for the signal that feeds back tocoordinate motor activity in the axoneme.

Dynamic curvature regulation accounts for the symmetric and asymmetric beats of Chlamydomonas flagella