cAMP-dependent phosphorylation of a 29-kDa axonemal polypeptide (p29) increases the swimming speed of permeabilized Paramecium and in vitro translocation velocity of bovine brain microtubules over 22S dynein extracted from Paramecium axonemes. A quantitative relationship between microtubule translocation velocity and beat frequency is developed. We conclude that p29 acts as a regulatory light chain of outer arm dynein in the control of ciliary beat frequency.