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First published online 5 May 2004
doi: 10.1242/jcs.01082

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Analysis of microtubule sliding patterns in Chlamydomonas flagellar axonemes reveals dynein activity on specific doublet microtubules

Department of Biological Sciences, Dartmouth College, Hanover, NH 03755, USA

^* Author for correspondence (e-mail: elizabeth.f.smith{at}dartmouth.edu)

Accepted 5 January 2004

Generating the complex waveforms characteristic of beating eukaryotic cilia and flagella requires spatial regulation of dynein-driven microtubule sliding. To generate bending, one prediction is that dynein arms alternate between active and inactive forms on specific subsets of doublet microtubules. Using an in vitro microtubule sliding assay combined with a structural approach, we determined that ATP induces sliding between specific subsets of doublet microtubules, apparently capturing one phase of the beat cycle. These studies were also conducted using high Ca²⁺ conditions. In Chlamydomonas, high Ca²⁺ induces changes in waveform which are predicted to result from regulating dynein activity on specific microtubules. Our results demonstrate that microtubule sliding in high Ca²⁺ buffer is also induced by dynein arms on specific doublets. However, the pattern of microtubule sliding in high Ca²⁺ buffer significantly differs from that in low Ca²⁺. These results are consistent with a `switching hypothesis' of axonemal bending and provide evidence to indicate that Ca²⁺ control of waveform includes modulation of the pattern of microtubule sliding between specific doublets. In addition, analysis of microtubule sliding in mutant axonemes reveals that the control mechanism is disrupted in some mutants.

Key words: Flagella, Microtubule, Dynein, Chlamydomonas

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