Assembly, Functions and Evolution of Archaella, Flagella and Cilia

Cells from all three domains of life on Earth utilize motile macromolecular devices that protrude from the cell surface to generate forces that allow them to swim through fluid media. Research carried out on archaea during the past decade or so has led to the recognition that, despite their common function, the motility devices of the three domains display fundamental differences in their properties and ancestry, reflecting a striking example of convergent evolution. Thus, the flagella of bacteria and the archaella of archaea employ rotary filaments that assemble from distinct subunits that do not share a common ancestor and generate torque using energy derived from distinct fuel sources, namely chemiosmotic ion gradients and FlaI motor-catalyzed ATP hydrolysis, respectively. The cilia of eukaryotes, however, assemble via kinesin-2-driven intraflagellar transport and utilize microtubules and ATP-hydrolyzing dynein motors to beat in a variety of waveforms via a sliding filament mechanism. Here, with reference to current structural and mechanistic information about these organelles, we briefly compare the evolutionary origins, assembly and tactic motility of archaella, flagella and cilia.