Biomechanics of a muscular hydrostat: a model of lapping by a reptilian tongue

Abstract

We have developed a quantitative model of an example of a muscular hydrostat, a reptilian tongue, and have used this model to study a functional movement, protrusion and retrusion, a form of lapping. The model tongue consists of a longitudinal muscle that shortens the tongue when it contracts, and a circumferential muscle wrapped around the longitudinal muscle that lengthens the tongue when it contracts. The anatomy of the model tongue and the pattern of activation of its muscles are based on studies of the tongue of the lizard Tupinambis nigropunctatus (Smith 1984). The mechanics of pressure vessels were used to derive a relationship between the forces in the two muscles. Muscle force production was modelled as the product of length/tension properties, force/velocity properties, and activation due to neural inputs (incorporating both recruitment and firing period). Passive forces were modeled as a force in parallel with the longitudinal muscle. Muscle activation dynamics were modeled as a first order low pass filter. When the model tongue is short, the two muscles can lengthen or shorten it with comparable forces, but as it lengthens, the force that the circumferential muscle can exert drops precipitously. When the tongue is long, it can neither be very stiff, nor can it generate much force. The model also reproduces the kinematics of lapping movements actually observed in Tupinambis.