Locomotory waves of Koruga and Deltotrichonympha: flagella wag the cell

We investigated the nature of the locomotory waves of Koruga and Deltotrichonympha, flagellates living symbiotically in the hindgut of the Australian termite Mastotermes darwiniensis. The locomotory waves consist of two components: metachronal waves of flagellar beating and undulations of the cell surface, which propagate synchronously with the same wavelength, frequency, and velocity. We asked, do body waves cause flagellar waves, or vice versa? Using video microscopy and selective inhibitors and drugs, we found that (1) the amplitude of flagellar waves remains constant independent of variations in the amplitude of body waves, (2) flagellar waves can occur in the complete absence of body waves, (3) flagellar waves can induce body waves on swollen regions, (4) inhibition of flagellar beating by dynein inhibitors causes disappearance of body waves, and (5) cytochalasin D induces changes in cell shape but does not inhibit locomotory waves. Therefore, flagellar waves are not produced passively by an active contractile system in the cell cortex; instead, metachronally beating flagella exert waves of pressure that induce passive undulations of a pliant cell surface. These results support Machemer's [1974] theoretical analysis of the data of Cleveland and Cleveland [1966: Arch. Protistenk. 109:39-63], who believed the opposite.