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In conditions in which ciliated cortical sheets prepared from detergent-extracted Paramecium multimicronucleatum cells adhered to glass coverslips on a microscope stage, perfusion of a reactivation medium containing ATP plus cyclic AMP or cyclic GMP generated metachronal waves. An analysis of the ciliary movements that generate these metachronal waves yielded the following results. During the generation of metachronal waves, there were phase differences in the ciliary orientation of adjacent cilia in the direction of wave propagation. Addition of cyclic AMP or cyclic GMP increased the rotational angular velocities during the effective stroke of ciliary beating, but did not increase the rotational angular velocity of the recovery stroke. When the ATP concentration in the cyclic GMP reactivation medium was increased, the rotational angular velocity during the effective stroke rose steeply and saturated at 0.8 mmol l-1 ATP, whereas that during the recovery stroke rose gradually. Addition of cyclic nucleotides caused a single cilium isolated from neighbouring cilia on the cortical sheet to incline almost parallel to the cortical surface during the recovery stroke. Addition of cyclic GMP increased the amplitude of bending of cilia detached from the cortical sheet. From these results, it was concluded that increases in the asymmetrical movement of individual cilia, caused by the addition of cyclic nucleotides, create the ciliary interaction that generates the metachronal waves.

Reconstitution of Metachronal Waves in Ciliated Cortical Sheets of Paramecium: II. Asymmetry of the Ciliary Movements