Geotaxis in protozoa I. A propulsion-gravity model for Tetrahymena (Ciliata)

A propulsion-based model for negative geotaxis of ciliated protozoa is presented which views geotaxic reorientation as the unbalancing of gyrational torque by a sedimentation torque. The balanced gyrational torque results from the location of the propulsive center of effort forward of the body center of mass. When gravity is ignored, the propulsive forces generating the gyrational moments may be confined to an envelope surrounding the cell. The effect of gravity is to induce sedimentation of the body-plus-envelope system. Viscous resistance to this sedimentation at the envelope “surface” is transmitted to the beating cilia whose net constant energy output must now deal with a new source of dissipation (not “present” when gravity was ignored) which is maximal in the downswing portion of the gyration cycle. In such a manner sedimentation resistance acts as a counter torque to the downswing gyrational moment of force and an enhancing torque to the upswing moment thereby generating a net upward reorientation of the gyrational axis. Upon addition of the translational component of propulsion, the negative geotaxis behavior pattern is completed. The forward location of the center of effort which provides the basic validity indicator for the model is verified by observations from the ciliate Tetrahymena.