A model for the micro-structure in ciliated organisms

Improved models for the movement of fluid by cilia are presented. A theory which models the cilia of an organism by an array of flexible long slender bodies distributed over and attached at one end to a plane surface is developed. The slender bodies are constrained to move in similar patterns to the cilia of the microorganisms Opalina, Paramecium and Pleurobrachia . The velocity field is represented by a distribution of force singularities (Stokes flow) along the centre-line of each slender body. Contributions to the velocity field from all the cilia distributed over the plane are summed, to give a streaming effect which in turn implies propulsion of the organism. From this we have been able to model the mean velocity field through the cilia sublayer for the three organisms. We find that, in a frame of reference situated in the organism, the velocity near the surface of the organism is very small – up to one half the length of the cilium – but it increases rapidly to near the velocity of propulsion from then on. This is because of the beating pattern of the cilia; they beat in a near rigid-body rotation during the effective (‘power’) stroke, but during the recovery stroke move close to the wall. Backflow (‘reflux’) is found to occur in the organisms exhibiting antiplectic metachronism (i.e. Paramecium and Pleurobrachia ). The occurrence of gradient reversal, but not backflow, has recently been confirmed experimentally (Sleigh & Aiello 1971). Other important physical values that are obtained from this analysis are the force, bending moment about the base of a cilium and the rate of working. It is found, for antiplectic metachronism, that the force exerted by a cilium in the direction of propulsion is large and positive during the effective stroke whereas it is small and negative during the recovery stroke. However, the duration of the recovery stroke is longer than the effective stroke so the force exerted over one cycle of a ciliary beat is very small. The bending moment follows a similar pattern to the component of force in the direction of propulsion, being larger in the effective stroke for antiplectic metachronism. In symplectic metachronism (i.e. Opalina ) the force and bending moment are largest in magnitude when the bending wave is propagated along the cilium. The rate of working indicates that more energy is consumed in the effective stroke for Paramecium and Pleurobrachia than in the recovery stroke, whereas in Opalina it is found to be large during the propagation of the bending wave.