The beat pattern of rabbit tracheal cilia has been investigated using high-speed cine photography and scanning electron microscopy, on cultured epithelia of known orientation.

The cilia normally rest in the position reached at the end of the effective stroke, the ciliary tips pointing towards the oropharynx. Each beat begins with a recovery (or preparative) stroke in which a bend is propagated up the cilium causing the cilium to rotate backwards in a clockwise sweep, as viewed from above. At the end of its recovery stroke the cilium progresses immediately into the effective (or power) stroke, which is almost planar and in a cephalad direction. The active cilium describes an arc of almost 110° before reaching the rest stage. This beat pattern is not significantly altered over an increase in frequency from 13 –29 Hz; the relative durations of the 2 active phases of the beat remain similar over this range.

Metachronal waves exist in the form of short erratic areas of coordinated beating which travel only short distances. Within each area, the non-planar recovery strokes initiate an antilaeoplectic wave of activity which recruits inactive cilia to extend the wave. As cilia perform their effective strokes, adjacent cilia in the plane of beating move in an antiplectic sequence. This pattern of coordination is related to the pattern of beat of the cilia and their distribution on the epithelium.