Helical Nature of the Ciliary Beat of Paramecium multimicronucleatum*

Abstract

SYNOPSIS. Phase and interference cinemicrographs of cilia of Paramecium multimicronucleatum, immersed 3–24 hours in 1.0% methyl cellulose, revealed that 1) in swimming Paramecium the cilia beat with a traveling helical wave from base to tip rather than with the back and forth movement usually assumed, 2) during ciliary reversal the cilia merely change direction, but continue to beat with a traveling helical wave, and 3) in stationary Paramecium the beat is conicoidal. The traveling wave appears as an undulatory wave about 1 1/4 wave lengths long in both surface and profile views, and therefore must be helical. Envelope of the wave is cylindrical except near the base. Observations were confirmed in media without methyl cellulose by means of high speed cinemicrography, up to 4000 frames/sec.The back and forth movement, as described in all textbooks and monographs, is based mostly on 1) analogy to the abfrontal cilia (cirri) of Mytilus, which do beat with a back and forth movement, and 2) conclusions drawn from fixed preparations which do not represent what actually happens in a living animal.In a stationary Paramecium the envelope of the beat is conicoidal as seen in profile, but probably is a spiral wave, i.e., similar to a helix but increasing in diameter from base to tip. This change in wave form could be caused by the increase in resistance of the water in a stationary organism over one that is moving.Cilia and flagella (also ciliates and flagellates) are usually distinguished on the basis of wave form, but the present observations, together with previous data on flagella, show that such distinctions are untenable.