Does the cerebellum learn strategies for the optimal time-varying control of joint stiffness ?

Although there is increasing agreement that the cerebellum plays an important role in motor learning, the basic substance of what constitutes motor learning has been difficult to define. Unless motor learning is somehow radically different from other forms of learning, it must involve relatively simple stimulus-stimulus and stimulus-response associations. All forms of learning, including purely sensory associations and cognitive learning as well as motor learning, effect changes in behavior. However, a singular characteristic of motor learning is that it adjusts joint and limb mechanics by altering the neural input to muscles through practice and mental rehearsal. The hypothesis proposed here is that the cerebellum plays an important role in motor learning by forming and storing associated muscle activation patterns or the time-varying control oflimb mechanics. By modulating the cocontraction of agonist-antagonist muscles through adjustments in the timing and amplitude of muscle activity, the viscoelastic properties of joints can be appropriately regulated throughout movement and adapted for transitions between postures and movements. Optimal control of joint viscoelastic properties cannot be achieved by online corrections initiated by reflex feedback because of the delays and consequent instabilities incurred. Instead, strategies for optimizing muscle activation patterns or synergies must be learned from the temporal association of proprioceptive stimuli signaling muscle lengths and forces and the rates of changes in these parameters, with reinforcement occurring when the movement achieves its objective. Such strategies would involve varying degrees of cocontraction or reciprocal inhibition of agonist-antagonist muscles that ultimately contribute to joint and limb stiffness. Evidence from neural recordings and clinical and experimental lesion studies are presented, suggesting that the cerebellum uses teleceptive and proprioceptive feedback as feedforward conditioned stimuli for specific muscle activation patterns contributing to joint stiffness (i.e., agonist-antagonist muscle synergies) for particular tasks and postures. A wide variety of observations are thought to be consistent with such a role for the cerebellum, but ultimately additional experiments could confirm or disconfirm this hypothesis.