From Parallel Sequence Representations to Calligraphic Control: A Conspiracy of Neural Circuits

Calligraphic writing presents a rich set of challenges to the human movement control system. These challenges include: initial learning, and recall from memory, of prescribed stroke sequences; critical timing of stroke onsets and durations; fine control of grip and contact forces; and letter-form invariance under voluntary size scaling, which entails fine control of stroke direction and amplitude during recruitment and derecruitment of musculoskeletal degrees of freedom. Experimental and computational studies in behavioral neuroscience have made rapid progress toward explaining the learning, planning and contTOl exercised in tasks that share features with calligraphic writing and drawing. This article summarizes computational neuroscience models and related neurobiological data that reveal critical operations spanning from parallel sequence representations to fine force control. Part one addresses stroke sequencing. It treats competitive queuing (CQ) models of sequence representation, performance, learning, and recall. Part two addresses letter size scaling and motor equivalence. It treats cursive handwriting models together with models in which sensory-motor tmnsformations are performed by circuits that learn inverse differential kinematic mappings. Part three addresses fine-grained control of timing and transient forces, by treating circuit models that learn to solve inverse dynamics problems.National Institutes of Health (R01 DC02852

Gamma Band Oscillation Response to Somatosensory Feedback Stimulation Schemes Constructed on Basis of Biphasic Neural Touch Representation

abstract: Prosthetic users abandon devices due to difficulties performing tasks without proper graded or interpretable feedback. The inability to adequately detect and correct error of the device leads to failure and frustration. In advanced prostheses, peripheral nerve stimulation can be used to deliver sensations, but standard schemes used in sensorized prosthetic systems induce percepts inconsistent with natural sensations, providing limited benefit. Recent uses of time varying stimulation strategies appear to produce more practical sensations, but without a clear path to pursue improvements. This dissertation examines the use of physiologically based stimulation strategies to elicit sensations that are more readily interpretable. A psychophysical experiment designed to investigate sensitivities to the discrimination of perturbation direction within precision grip suggests that perception is biomechanically referenced: increased sensitivities along the ulnar-radial axis align with potential anisotropic deformation of the finger pad, indicating somatosensation uses internal information rather than environmental. Contact-site and direction dependent deformation of the finger pad activates complimentary fast adapting and slow adapting mechanoreceptors, exhibiting parallel activity of the two associate temporal patterns: static and dynamic. The spectrum of temporal activity seen in somatosensory cortex can be explained by a combined representation of these distinct response dynamics, a phenomenon referred in this dissertation to “biphasic representation.” In a reach-to-precision-grasp task, neurons in somatosensory cortex were found to possess biphasic firing patterns in their responses to texture, orientation, and movement. Sensitivities seem to align with variable deformation and mechanoreceptor activity: movement and smooth texture responses align with potential fast adapting activation, non-movement and coarse texture responses align with potential increased slow adapting activation, and responses to orientation are conceptually consistent with coding of tangential load. Using evidence of biphasic representations’ association with perceptual priorities, gamma band phase locking is used to compare responses to peripheral nerve stimulation patterns and mechanical stimulation. Vibrotactile and punctate mechanical stimuli are used to represent the practical and impractical percepts commonly observed in peripheral nerve stimulation feedback. Standard patterns of constant parameters closely mimic impractical vibrotactile stimulation while biphasic patterns better mimic punctate stimulation and provide a platform to investigate intragrip dynamics representing contextual activation.Dissertation/ThesisDoctoral Dissertation Biomedical Engineering 201

Cortical Networks for Control of Voluntary Arm Movements Under Variable Force Conditions

A neural model of voluntary movement and proprioception functionally interprets and simulates cell types in movement related areas of primate cortex. The model circuit maintains accurate proprioception while controlling voluntary reaches to spatial targets, exertion of force against obstacles, posture maintenance despite perturbations, compliance with an imposed movement, and static and inertial load compensations. Computer simulations show that model cell properties mimic cell properties in areas 4 and 5. These include delay period activation, response profiles during movement, kinematic and kinetic sensitivities, and latency of activity onset. Model area 4 phasic and tonic cells compute velocity and position commands which activate alpha and gamma motor neurons, thereby shifting the mechanical equilibrium point. Anterior area 5 cells compute limb position using corollary discharges from area 4 and muscle spindle feedback. Posterior area 5 cells use the perceived position and target position signals to compute a desired movement vector. The cortical loop is closed by a volition-gated projection of this movement vector to area 4 phasic cells. Phasic-tonic cells in area 4 incorporate force command components to compensate for static and inertial loads. Predictions are made for both motor and parietal cell types under novel experimental protocols.Office of Naval Research (N00014-92-J-1309, N00014-93-1-1364, N00014-95-l-0409, N00014-92-J-4015); National Science Foundation (IRI-90-24877, IRI-90-00530

Gravitational and Dynamic Components of Muscle Torque Underlie Tonic and Phasic Muscle Activity during Goal-Directed Reaching

Human reaching movements require complex muscle activations to produce the forces necessary to move the limb in a controlled manner. How gravity and the complex kinetic properties of the limb contribute to the generation of the muscle activation pattern by the central nervous system (CNS) is a long-standing and controversial question in neuroscience. To tackle this issue, muscle activity is often subdivided into static and phasic components. The former corresponds to posture maintenance and transitions between postures. The latter corresponds to active movement production and the compensation for the kinetic properties of the limb. In the present study, we improved the methodology for this subdivision of muscle activity into static and phasic components by relating them to joint torques. Ten healthy subjects pointed in virtual reality to visual targets arranged to create a standard center-out reaching task in three dimensions. Muscle activity and motion capture data were synchronously collected during the movements. The motion capture data were used to calculate postural and dynamic components of active muscle torques using a dynamic model of the arm with 5 degrees of freedom. Principal Component Analysis (PCA) was then applied to muscle activity and the torque components, separately, to reduce the dimensionality of the data. Muscle activity was also reconstructed from gravitational and dynamic torque components. Results show that the postural and dynamic components of muscle torque represent a significant amount of variance in muscle activity. This method could be used to define static and phasic components of muscle activity using muscle torques

The role of noise in sensorimotor control

Goal-directed arm movements show stereotypical trajectories, despite the infinite possible ways to reach a given end point. This thesis examines the hypothesis that this stereotypy arises because movements are optimised to reduce the consequences of signal-dependent noise on the motor command. Both experimental and modelling studies demonstrate that signal-dependent noise arises from the normal behaviour of the muscle and motor neuron pool, and has a particular distribution across muscles of different sizes. Specifically, noise decreases in a systematic fashion with increasing muscle strength and motor unit number. Simulations of obstacle avoidance performance in the presence of signal-dependent noise demonstrate that the optimal trajectory for reaching the target accurately and without collision matches the observed trajectories. Isometric force generation is also shown to have systematic changes in variability with posture, which can be explained by the presence of signal-dependent noise in the muscles of the arm. These results confirm the tested hypothesis and imply that consideration of the statistics of action is crucial to human movement planning. To investigate the importance of feedback in the motor system, the impact of static position on motor excitability was examined using transcranial magnetic stimulation and systematic changes in motor evoked potentials were observed. Force generated at the wrist following stimulation was analysed in terms of different possible movement representations, and the differences between force fields arising from stimulation over the cervical spinal cord and from stimulation over primary motor cortex are determined. These results demonstrate the structured influence of proprioceptive feedback on the human motor system. All the experiments are discussed in relation to current theories describing the control of human movements and the impact of noise in the motor system

Ubiquitous haptic feedback in human-computer interaction through electrical muscle stimulation

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Learning to push and learning to move: The adaptive control of contact forces

To be successful at manipulating objects one needs to apply simultaneously well controlled movements and contact forces. We present a computational theory of how the brain may successfully generate a vast spectrum of interactive behaviors by combining two independent processes. One process is competent to control movements in free space and the other is competent to control contact forces against rigid constraints. Free space and rigid constraints are singularities at the boundaries of a continuum of mechanical impedance. Within this continuum, forces and motions occur in \u201ccompatible pairs\u201d connected by the equations of Newtonian dynamics. The force applied to an object determines its motion. Conversely, inverse dynamics determine a unique force trajectory from a movement trajectory. In this perspective, we describe motor learning as a process leading to the discovery of compatible force/motion pairs. The learned compatible pairs constitute a local representation of the environment's mechanics. Experiments on force field adaptation have already provided us with evidence that the brain is able to predict and compensate the forces encountered when one is attempting to generate a motion. Here, we tested the theory in the dual case, i.e., when one attempts at applying a desired contact force against a simulated rigid surface. If the surface becomes unexpectedly compliant, the contact point moves as a function of the applied force and this causes the applied force to deviate from its desired value. We found that, through repeated attempts at generating the desired contact force, subjects discovered the unique compatible hand motion. When, after learning, the rigid contact was unexpectedly restored, subjects displayed after effects of learning, consistent with the concurrent operation of a motion control system and a force control system. Together, theory and experiment support a new and broader view of modularity in the coordinated control of forces and motions