The Mechanism of Tongue Projection in Chameleons: II. Role of Shape Change in a Muscular Hydrostat

In this paper we investigate the interaction between the accelerator muscle (the muscle that powers tongue projection) and the entoglossal process (the tongue's skeletal support) that occurs during tongue projection in chamaeleonid lizards. Previous work has shown that there is a delay of about 185 ms between the onset of accelerator muscle activity and the onset of tongue projection. In conjunction with anatomical observations, in vitro preparations of the accelerator muscle mounted on isolated entoglossal and surrogate processes were stimulated tetanically, and the resulting movements were recorded on video at 200 fields s−1. Three results indicate that morphological features of the entoglossus and the accelerator muscle delay the onset of tongue projection following the onset of accelerator contraction: (1) the entoglossus is parallel-sided along the posterior 90% of its shaft, only tapering at the very tip, (2) the sphincter-like portion of the accelerator muscle, which effects tongue projection, makes up the posterior 63% of the muscle and does not contact the tapered region of the entoglossus at rest, and (3) accelerator muscles mounted on the entoglossus undergo longitudinal extension and lateral constriction for 83 ms following the onset of electrical stimulation, before projecting off the entoglossus. It is proposed that, during elongation of the accelerator muscle, the sphincter-like region ultimately comes into contact with the tapered region of the entoglossus, causing the onset of projection. This conclusion is supported by the observation that the time between the onset of stimulation and the onset of projection was longer in preparations with surrogate entoglossal processes that had no tapered tip and shorter with surrogate processes that had a tapered tip about four times as long as the natural entoglossus. Tetanically stimulated accelerator muscles reached 90% of peak force 110 ms after the onset of stimulation, indicating that the 185 ms delay between the onset of accelerator activity and the onset of projection seen in vivo allows the accelerator to achieve peak force prior to the onset of projection. Thus, the delay in projection may be crucial in maximizing the acceleration and velocity achieved by the projected chameleon tongue.