Tongues, tentacles and trunks: the biomechanics of movement in muscular-hydrostats

Muscular-hydrostats, muscular organs which lack typical systems of skeletal support, include the tongues of mammals and lizards, the arms and tentacles of cephalopod molluscs and the trunks of elephants. In this paper the means by which such organs produce elongation, shortening, bending and torsion are discussed. The most important biomechanical feature of muscular-hydrostats is that their volume is constant, so that any decrease in one dimension will cause a compensatory increase in at least one other dimension. Elongation of a muscular-hydrostat is produced by contraction of transverse, circular or radial muscles which decrease the cross-section. Shortening is produced by contraction of longitudinal muscles. The relation between length and width of a constant volume structure allows amplification of muscle force or displacement in muscular-hydrostats and other hydrostatic systems. Bending requires simultaneous contraction of longitudinal and antagonistic circular, transverse or radial muscles. In bending, one muscle mass acts as an effector of movement while the alternate muscle mass provides support for that movement. Torsion is produced by contraction of muscles which wrap the muscular-hydrostat in a helical fashion.