  Cilia and flagella in biological systems often show large scale cooperative
behaviors such as the synchronization of their beats in "metachronal waves".
These are beautiful examples of emergent dynamics in biology, and are essential
for life, allowing diverse processes from the motility of eukaryotic
microorganisms, to nutrient transport and clearance of pathogens from mammalian
airways. How these collective states arise is not fully understood, but it is
clear that individual cilia interact mechanically,and that a strong and long
ranged component of the coupling is mediated by the viscous fluid. We review
here the work by ourselves and others aimed at understanding the behavior of
hydrodynamically coupled systems, and particularly a set of results that have
been obtained both experimentally and theoretically by studying actively driven
colloidal systems. In these controlled scenarios, it is possible to selectively
test aspects of the living motile cilia, such as the geometrical arrangement,
the effects of the driving profile and the distance to no-slip boundaries. We
outline and give examples of how it is possible to link model systems to
observations on living systems, which can be made on microorganisms, on cell
cultures or on tissue sections. This area of research has clear clinical
application in the long term, as severe pathologies are associated with
compromised cilia function in humans.
