The control of multi-muscle systems: human jaw and hyoid movements

A model is presented of sagittal plane jaw and hyoid motion based on the lambda model of motor control. The model, which is implemented as a computer simulation, includes central neural control signals, position- and velocity-dependent reflexes, reflex delays, and muscle properties such as the dependence of force on muscle length and velocity. The model has seven muscles (or muscle groups) attached to the jaw and hyoid as well as separate jaw and hyoid bone dynamics. According to the model, movements result from changes in neurophysiological control variables which shift the equilibrium state of the motor system. One such control variable is an independent change in the membrane potential of alpha-motoneurons (MNs); this variable establishes a threshold muscle length (lambda) at which MN recruitment begins. Motor functions may be specified by various combinations of lambda s. One combination of lambda s is associated with the level of coactivation of muscles. Others are associated with motions in specific degrees of freedom. Using the model, we study the mapping between control variables specified at the level of degrees of freedom and control variables corresponding to individual muscles. We demonstrate that commands can be defined involving linear combinations of lambda change which produce essentially independent movements in each of the four kinematic degrees of freedom represented in the model (jaw orientation, jaw position, vertical and horizontal hyoid position). These linear combinations are represented by vectors in lambda space which may be scaled in magnitude. The vector directions are constant over the jaw/hyoid workspace and result in essentially the same motion from any workspace position. The demonstration that it is not necessary to adjust control signals to produce the same movements in different parts of the workspace supports the idea that the nervous system need not take explicit account of musculo-skeletal geometry in planning movements.