On Primitives in Motor Control

The concept of primitives has been used in motor control both as a theoretical construct and as a means of describing the results of experimental studies involving multiple moving elements. This concept is close to Bernstein’s notion of engrams and level of synergies. Performance primitives have been explored in spaces of peripheral variables but interpreted in terms of neural control primitives. Performance primitives reflect a variety of mechanisms ranging from body mechanics to spinal mechanisms and to supraspinal circuitry. This review suggests that primitives originate at the task level as preferred time functions of spatial referent coordinates or at mappings from higher level referent coordinates to lower level, frequently abundant, referent coordinate sets. Different patterns of performance primitives can emerge depending, in particular, on the external force field.

Stability of hand force production. II. Ascending and descending synergies

We combined the theory of neural control of movement with referent coordinates and the uncontrolled manifold hypothesis to investigate multifinger coordination. We tested hypotheses related to stabilization of performance by covarying control variables, translated into apparent stiffness and referent coordinate, at different levels of an assumed hierarchy of control. Subjects produced an accurate combination of total force and total moment of force with the four fingers under visual feedback on both variables and after feedback was partly or completely removed. The “inverse piano” device was used to estimate control variables. We observed strong synergies in the space of hypothetical control variables that stabilized total force and moment of force, as well as weaker synergies stabilizing individual finger forces; whereas the former were attenuated by alteration of visual feedback, the latter were much less affected. In addition, we investigated the organization of “ascending synergies” stabilizing task-level control variables by covaried adjustments of finger-level control variables. We observed intertrial covariation of individual fingers’ referent coordinates that stabilized hand-level referent coordinate, but we observed no such covariation for apparent stiffness. The observations suggest the existence of both descending and ascending synergies in a hierarchical control system. They confirm a trade-off between synergies at different levels of control and corroborate the hypothesis on specialization of different fingers for the control of force and moment. The results provide strong evidence for the importance of central back-coupling loops in ensuring stability of action.NEW & NOTEWORTHY We expand analysis of action in the space of hypothetical control variables to hierarchically organized multieffector systems. We also introduce the novel concept of ascending synergies, which reflect covariation of control variables to individual effectors (fingers) that stabilize task-specific control variables at a hierarchically higher, task-specific level (hand).

Force illusions and drifts observed during muscle vibration

We explored predictions of a scheme that views position and force perception as a result of measuring proprioceptive signals within a reference frame set by ongoing efferent process. In particular, this hypothesis predicts force illusions caused by muscle vibration and mediated via changes in both afferent and efferent components of kinesthesia. Healthy subjects performed accurate steady force production tasks by pressing with the four fingers of one hand (the task hand) on individual force sensors with and without visual feedback. At various times during the trials, subjects matched the perceived force using the other hand. High-frequency vibration was applied to one or both of the forearms (over the hand and finger extensors). Without visual feedback, subjects showed a drop in the task hand force, which was significantly smaller under the vibration of that forearm. Force production by the matching hand was consistently higher than that of the task hand. Vibrating one of the forearms affected the matching hand in a manner consistent with the perception of higher magnitude of force produced by the vibrated hand. The findings were consistent between the dominant and nondominant hands. The effects of vibration on both force drift and force mismatching suggest that vibration led to shifts in both signals from proprioceptors and the efferent component of perception, the referent coordinate and/or coactivation command. The observations fit the hypothesis on combined perception of kinematic-kinetic variables with little specificity of different groups of peripheral receptors that all contribute to perception of forces and coordinates. NEW & NOTEWORTHY We show that vibration of hand/finger extensors produces consistent errors in finger force perception. Without visual feedback, finger force drifted to lower values without a drift in the matching force produced by the other hand; hand extensor vibration led to smaller finger force drift. The findings fit the scheme with combined perception of kinematic-kinetic variables and suggest that vibration leads to consistent shifts of the referent coordinate and, possibly, of coactivation command to the effector.

On the nature of unintentional action: a study of force/moment drifts during multifinger tasks

We explored the origins of unintentional changes in performance during accurate force production in isometric conditions seen after turning visual feedback off. The idea of control with referent spatial coordinates suggests that these phenomena could result from drifts of the referent coordinate for the effector. Subjects performed accurate force/moment production tasks by pressing with the fingers of a hand on force sensors. Turning the visual feedback off resulted in slow drifts of both total force and total moment to lower magnitudes of these variables; these drifts were more pronounced in the right hand of the right-handed subjects. Drifts in individual finger forces could be in different direction; in particular, fingers that produced moments of force against the required total moment showed an increase in their forces. The force/moment drift was associated with a drop in the index of synergy stabilizing performance under visual feedback. The drifts in directions that changed performance (non-motor equivalent) and in directions that did not (motor equivalent) were of about the same magnitude. The results suggest that control with referent coordinates is associated with drifts of those referent coordinates toward the corresponding actual coordinates of the hand, a reflection of the natural tendency of physical systems to move toward a minimum of potential energy. The interaction between drifts of the hand referent coordinate and referent orientation leads to counterdirectional drifts in individual finger forces. The results also demonstrate that the sensory information used to create multifinger synergies is necessary for their presence over the task duration.