It is 20 years since the publication of the geometric clutch (GC) hypothesis as a mechanistic explanation of ciliary and flagellar beating (1). It was an idea that resulted from constructing wooden models of the flagellar axoneme and observing the way they distorted during bending. The key insight was that a structure composed of linear elements that experience curvature while under tension or compression experience transverse forces (t-forces) that can push the linear elements closer together or pry them apart. In the eu-karyotic axoneme of cilia and flagella, the linear elements are the nine outer doublets.
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