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Eukaryotes swim with coordinated flagellar (ciliary) beating and steer byfine-tuning the coordination. The model organism for studying flagellatemotility, C. reinhardtii (CR), employs synchronous, breast-stroke-likeflagellar beating to swim, and it modulates the beating amplitudesdifferentially to steer. This strategy hinges on both inherent flagellarasymmetries (e.g. different response to chemical messengers) and suchasymmetries being effectively coordinated in the synchronous beating. In CR,the synchrony of beating is known to be supported by a mechanical connectionbetween flagella, however, how flagellar asymmetries persist in the synchronyremains elusive. For example, it has been speculated for decades that oneflagellum leads the beating, as its dynamic properties (i.e. frequency,waveform, etc.) appear to be copied by the other one. In this study, we combineexperiments, computations, and modeling efforts to elucidate the roles playedby each flagellum in synchronous beating. With a non-invasive technique toselectively load each flagellum, we show that the coordinated beatingessentially responds to only load exerted on the cis flagellum; and that suchasymmetry in response derives from a unilateral coupling between the twoflagella. Our results highlight a distinct role for each flagellum incoordination and have implication for biflagellates tactic behaviors.

The younger flagellum coordinates the beating in C. reinhardtii