Visual Feedback of Object Motion Direction Influences the Timing of Grip Force Modulation During Object Manipulation.

During manipulation, object slipping is prevented by modulating the grip force (GF) in synchrony with motion-related inertial forces, i.e., load force (LF). However, due to conduction delays of the sensory system, GF must be modulated in advance based on predictions of LF changes. It has been proposed that such predictive force control relies on internal representations, i.e., internal models, of the relation between the dynamic of the environment and movement kinematics. Somatosensory and visual feedback plays a primary role in building these internal representations. For instance, it has been shown that manipulation-dependent somatosensory signals contribute to building internal representations of gravity in normal and altered gravitational contexts. Furthermore, delaying the timing of visual feedback of object displacement has been shown to affect GF. Here, we explored whether and the extent to which spatial features of visual feedback movement, such as motion direction, may contribute to GF control. If this were the case, a spatial mismatch between actual (somatosensory) and visual feedback of object motion would elicit changes in GF modulation. We tested this hypothesis by asking participants to generate vertical object movements while visual feedback of object position was congruent (0° rotation) or incongruent (180° or 90°) with the actual object displacement. The role of vision on GF control was quantified by the temporal shift of GF modulation as a function of visual feedback orientation and actual object motion direction. GF control was affected by visual feedback when this was incongruent in the vertical (180°), but not horizontal dimension. Importantly, 180° visual feedback rotation delayed and anticipated GF modulation during upward and downward actual movements, respectively. Our findings suggest that during manipulation, spatial features of visual feedback motion are used to predict upcoming LF changes. Furthermore, the present study provides evidence that an internal model of gravity contributes to GF control by influencing sensory reweighting processes during object manipulation.

Motor modules account for active perception of force.

Despite longstanding evidence suggesting a relation between action and perception, the mechanisms underlying their integration are still unclear. It has been proposed that to simplify the sensorimotor integration processes underlying active perception, the central nervous system (CNS) selects patterns of movements aimed at maximizing sampling of task-related sensory input. While previous studies investigated the action-perception loop focusing on the role of higher-level features of motor behavior (e.g., kinematic invariants, effort), the present study explored and quantified the contribution of lower-level organization of motor control. We tested the hypothesis that the coordinated recruitment of group of muscles (i.e., motor modules) engaged to counteract an external force contributes to participants' perception of the same force. We found that: 1) a model describing the modulation of a subset of motor modules involved in the motor task accounted for about 70% of participants' perceptual variance; 2) an alternative model, incompatible with the motor modules hypothesis, accounted for significantly lower variance of participants' detection performance. Our results provide empirical evidence of the potential role played by muscle activation patterns in active perception of force. They also suggest that a modular organization of motor control may mediate not only coordination of multiple muscles, but also perceptual inference.

Linear Integration of Tactile and Non-tactile Inputs Mediates Estimation of Fingertip Relative Position.

While skin, joints and muscles receptors alone provide lower level information about individual variables (e.g., exerted limb force and limb displacement), the distance between limb endpoints (i.e., relative position) has to be extracted from high level integration of somatosensory and motor signals. In particular, estimation of fingertip relative position likely involves more complex sensorimotor transformations than those underlying hand or arm position sense: the brain has to estimate where each fingertip is relative to the hand and where fingertips are relative to each other. It has been demonstrated that during grasping, feedback of digit position drives rapid adjustments of fingers force control. However, it has been shown that estimation of fingertips' relative position can be biased by digit forces. These findings raise the question of how the brain combines concurrent tactile (i.e., cutaneous mechanoreceptors afferents induced by skin pressure and stretch) and non-tactile (i.e., both descending motor command and joint/muscle receptors signals associated to muscle contraction) digit force-related inputs for fingertip distance estimation. Here we addressed this question by quantifying the contribution of tactile and non-tactile force-related inputs for the estimation of fingertip relative position. We asked subjects to match fingertip vertical distance relying only on either tactile or non-tactile inputs from the thumb and index fingertip, and compared their performance with the condition where both types of inputs were combined. We found that (a) the bias in the estimation of fingertip distance persisted when tactile inputs and non-tactile force-related signals were presented in isolation; (b) tactile signals contributed the most to the estimation of fingertip distance; (c) linear summation of the matching errors relying only on either tactile or non-tactile inputs was comparable to the matching error when both inputs were simultaneously available. These findings reveal a greater role of tactile signals for sensing fingertip distance and suggest a linear integration mechanism with non-tactile inputs for the estimation of fingertip relative position.

Mapping Muscles Activation to Force Perception during Unloading.

It has been largely proved that while judging a force humans mainly rely on the motor commands produced to interact with that force (i.e., sense of effort). Despite of a large bulk of previous investigations interested in understanding the contributions of the descending and ascending signals in force perception, very few attempts have been made to link a measure of neural output (i.e., EMG) to the psychophysical performance. Indeed, the amount of correlation between EMG activity and perceptual decisions can be interpreted as an estimate of the contribution of central signals involved in the sensation of force. In this study we investigated this correlation by measuring the muscular activity of eight arm muscles while participants performed a quasi-isometric force detection task. Here we showed a method to quantitatively describe muscular activity ("muscle-metric function") that was directly comparable to the description of the participants' psychophysical decisions about the stimulus force. We observed that under our experimental conditions, muscle-metric absolute thresholds and the shape of the muscle-metric curves were closely related to those provided by the psychophysics. In fact a global measure of the muscles considered was able to predict approximately 60% of the perceptual decisions total variance. Moreover the inter-subjects differences in psychophysical sensitivity showed high correlation with both participants' muscles sensitivity and participants' joint torques. Overall, our findings gave insights into both the role played by the corticospinal motor commands while performing a force detection task and the influence of the gravitational muscular torque on the estimation of vertical forces.

Visuomotor adaptation to a visual rotation is gravity dependent.

Humans perform vertical and horizontal arm motions with different temporal patterns. The specific velocity profiles are chosen by the central nervous system by integrating the gravitational force field to minimize energy expenditure. However, what happens when a visuomotor rotation is applied, so that a motion performed in the horizontal plane is perceived as vertical? We investigated the dynamic of the adaptation of the spatial and temporal properties of a pointing motion during prolonged exposure to a 90° visuomotor rotation, where a horizontal movement was associated with a vertical visual feedback. We found that participants immediately adapted the spatial parameters of motion to the conflicting visual scene in order to keep their arm trajectory straight. In contrast, the initial symmetric velocity profiles specific for a horizontal motion were progressively modified during the conflict exposure, becoming more asymmetric and similar to those appropriate for a vertical motion. Importantly, this visual effect that increased with repetitions was not followed by a consistent aftereffect when the conflicting visual feedback was absent (catch and washout trials). In a control experiment we demonstrated that an intrinsic representation of the temporal structure of perceived vertical motions could provide the error signal allowing for this progressive adaptation of motion timing. These findings suggest that gravity strongly constrains motor learning and the reweighting process between visual and proprioceptive sensory inputs, leading to the selection of a motor plan that is suboptimal in terms of energy expenditure.

Visual gravity influences arm movement planning.

When submitted to a visuomotor rotation, subjects show rapid adaptation of visually guided arm reaching movements, indicated by a progressive reduction in reaching errors. In this study, we wanted to make a step forward by investigating to what extent this adaptation also implies changes into the motor plan. Up to now, classical visuomotor rotation paradigms have been performed on the horizontal plane, where the reaching motor plan in general requires the same kinematics (i.e., straight path and symmetric velocity profile). To overcome this limitation, we considered vertical and horizontal movement directions requiring specific velocity profiles. This way, a change in the motor plan due to the visuomotor conflict would be measurable in terms of a modification in the velocity profile of the reaching movement. Ten subjects performed horizontal and vertical reaching movements while observing a rotated visual feedback of their motion. We found that adaptation to a visuomotor rotation produces a significant change in the motor plan, i.e., changes to the symmetry of velocity profiles. This suggests that the central nervous system takes into account the visual information to plan a future motion, even if this causes the adoption of nonoptimal motor plans in terms of energy consumption. However, the influence of vision on arm movement planning is not fixed, but rather changes as a function of the visual orientation of the movement. Indeed, a clear influence on motion planning can be observed only when the movement is visually presented as oriented along the vertical direction. Thus vision contributes differently to the planning of arm pointing movements depending on motion orientation in space.