Energy exchanges at contact events guide sensorimotor integration.

The brain must consider the arm's inertia to predict the arm's movements elicited by commands impressed upon the muscles. Here, we present evidence suggesting that the integration of sensory information leading to the representation of the arm's inertia does not take place continuously in time but only at discrete transient events, in which kinetic energy is exchanged between the arm and the environment. We used a visuomotor delay to induce cross-modal variations in state feedback and uncovered that the difference between visual and proprioceptive velocity estimations at isolated collision events was compensated by a change in the representation of arm inertia. The compensation maintained an invariant estimate across modalities of the expected energy exchange with the environment. This invariance captures different types of dysmetria observed across individuals following prolonged exposure to a fixed intermodal temporal perturbation and provides a new interpretation for cerebellar ataxia.

State-Based Delay Representation and Its Transfer from a Game of Pong to Reaching and Tracking.

To accurately estimate the state of the body, the nervous system needs to account for delays between signals from different sensory modalities. To investigate how such delays may be represented in the sensorimotor system, we asked human participants to play a virtual pong game in which the movement of the virtual paddle was delayed with respect to their hand movement. We tested the representation of this new mapping between the hand and the delayed paddle by examining transfer of adaptation to blind reaching and blind tracking tasks. These blind tasks enabled to capture the representation in feedforward mechanisms of movement control. A Time Representation of the delay is an estimation of the actual time lag between hand and paddle movements. A State Representation is a representation of delay using current state variables: the distance between the paddle and the ball originating from the delay may be considered as a spatial shift; the low sensitivity in the response of the paddle may be interpreted as a minifying gain; and the lag may be attributed to a mechanical resistance that influences paddle's movement. We found that the effects of prolonged exposure to the delayed feedback transferred to blind reaching and tracking tasks and caused participants to exhibit hypermetric movements. These results, together with simulations of our representation models, suggest that delay is not represented based on time, but rather as a spatial gain change in visuomotor mapping.

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 "compatible pairs" 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.

How soft is that pillow? The perceptual localization of the hand and the haptic assessment of contact rigidity.

A new haptic illusion is described, in which the location of the mobile object affects the perception of its rigidity. There is theoretical and experimental support for the notion that limb position sense results from the brain combining ongoing sensory information with expectations arising from prior experience. How does this probabilistic state information affect one's tactile perception of the environment mechanics? In a simple estimation process, human subjects were asked to report the relative rigidity of two simulated virtual objects. One of the objects remained fixed in space and had various coefficients of stiffness. The other virtual object had constant stiffness but moved with respect to the subjects. Earlier work suggested that the perception of an object's rigidity is consistent with a process of regression between the contact force and the perceived amount of penetration inside the object's boundary. The amount of penetration perceived by the subject was affected by varying the position of the object. This, in turn, had a predictable effect on the perceived rigidity of the contact. Subjects' reports on the relative rigidity of the object are best accounted for by a probabilistic model in which the perceived boundary of the object is estimated based on its current location and on past observations. Therefore, the perception of contact rigidity is accounted for by a stochastic process of state estimation underlying proprioceptive localization of the hand.

Functional reorganization of upper-body movement after spinal cord injury.

Survivors of spinal cord injury need to reorganize their residual body movements for interacting with assistive devices and performing activities that used to be easy and natural. To investigate movement reorganization, we asked subjects with high-level spinal cord injury (SCI) and unimpaired subjects to control a cursor on a screen by performing upper-body motions. While this task would be normally accomplished by operating a computer mouse, here shoulder motions were mapped into the cursor position. Both the control and the SCI subjects were rapidly able to reorganize their movements and to successfully control the cursor. The majority of the subjects in both groups were successful in reducing the movements that were not effective at producing cursor motions. This is inconsistent with the hypothesis that the control system is merely concerned with the accurate acquisition of the targets and is unconcerned with motions that are not relevant to this goal. In contrast, our findings suggest that subjects can learn to reorganize coordination so as to increase the correspondence between the subspace of their upper-body motions with the plane in which the controlled cursor moves. This is effectively equivalent to constructing an inverse internal model of the map from body motions to cursor motions, established by the experiment. These results are relevant to the development of interfaces for assistive devices that optimize the use of residual voluntary control and enhance the learning process in disabled users, searching for an easily learnable map between their body motor space and control space of the device.

Adaptation to delayed force perturbations in reaching movements.

Adaptation to deterministic force perturbations during reaching movements was extensively studied in the last few decades. Here, we use this methodology to explore the ability of the brain to adapt to a delayed velocity-dependent force field. Two groups of subjects preformed a standard reaching experiment under a velocity dependent force field. The force was either immediately proportional to the current velocity (Control) or lagged it by 50 ms (Test). The results demonstrate clear adaptation to the delayed force perturbations. Deviations from a straight line during catch trials were shifted in time compared to post-adaptation to a non-delayed velocity dependent field (Control), indicating expectation to the delayed force field. Adaptation to force fields is considered to be a process in which the motor system predicts the forces to be expected based on the state that a limb will assume in response to motor commands. This study demonstrates for the first time that the temporal window of this prediction needs not to be fixed. This is relevant to the ability of the adaptive mechanisms to compensate for variability in the transmission of information across the sensory-motor system.

Perception and Action in Simulated Telesurgery.

We studied the effect of delay on perception and action in contact with a force field that emulates elastic soft tissue with a specific rigid nonlinear boundary. Such field is similar to forces exerted on a needle during teleoperated needle insertion tasks. We found that a nonlinear boundary region causes both psychometric and motor overestimation of stiffness, and that delay causes motor but not psychometric underestimation of the stiffness of this nonlinear soft tissue. In addition we show that changing the teleoperation channel gain reduces and can even cancel the motor effect of delay.

Functional reorganization of upper-body movements for wheelchair control.

In general, survivors of neuromotor disorders and injuries need to reorganize their body movements in order to achieve goals that used to be easy and natural. Often, disabled people are offered the option to control assistive devices that will facilitate the recovery of independence and capability in their daily lives. The knowledge acquired during the last few years in the motor control field can be used to study and enhance this learning process. Furthermore, this knowledge may aid in finding methods for optimizing the use of residual voluntary muscular control in disabled users and searching for an easily learnable map between body motor space and devices control space. To investigate movement reorganization we asked healthy subjects to control a cursor performing a reaching task using shoulders and upper arm movements. These movements were mapped to a lower dimensional space by principal components analysis and were used to control the cursor. We found that all subjects were able to learn to control the cursor with ease and precision while reducing the proportion of ineffective body movement components in favor of the components that mapped directly into the control space. Moreover, with practice the movements of the controlled device - the cursor - became faster, smother, more precise and with a nearly symmetric speed profile.

Somatosensory evoked potentials during a rubber-hand illusion in schizophrenia.

The rubber-hand illusion (RHI), an illusion in which tactile sensations are referred to a synthetic alien limb, is enhanced in schizophrenia patients. Somatosensory evoked responses of the illusion were compared between schizophrenia patients and normal control subjects. Schizophrenia patients had significant alterations in long latency evoked responses during the illusion. These findings support the hypothesis of alterations in associative higher-level neuronal activity in schizophrenia. The findings support previous results pointing to alterations in associative brain regions in schizophrenia.