Coordination between equilibrium and hand trajectories during whole body pointing movements

We examined the coordination between equilibrium and voluntary pointing movements executed from the standing position, using the whole body. It has previously been shown that trunk movement has little effect upon kinematic characteristics of hand pointing when movements are executed in the sitting position. The present study asked if elements of hand trajectory are modified by requirements of large trunk displacements and fine equilibrium control when pointing movements are executed from the standing position. To achieve this, center of pressure (CoP) and center of mass (CoM) displacements were analyzed along with the kinematics of the pointing hand. Results showed that the CoM was not stabilized (it displaced between 23% and 61+/-21% of the foot's length), confirming that instead of a compensation of mechanical perturbations due to arm and trunk movements, the present equilibrium strategy consisted of controlling CoM acceleration towards the target. Hand paths were curved and were not distance or speed invariant. Rather than simple inefficiencies in programming or execution, path curvature suggested that different hand movement strategies were chosen as a function of equilibrium constraints. In light of these results, we hypothesize that postural stability may play a role in the generation of hand trajectory for complex, whole-body pointing movements, in addition to constraints placed upon end-effector kinematics or the dynamic optimization of upper-limb movements. A dependent regulation of equilibrium and spatial components of the movement is proposed.     

Gaze anchoring to a pointing target is present during the entire pointing movement and is driven by a non-visual signal

A well-coordinated pattern of eye and hand movements can be observed during goal-directed arm movements. Typically, a saccadic eye movement precedes the arm movement, and its occurrence is temporally correlated with the start of the arm movement. Furthermore, the coupling of gaze and aiming movements is also observable after pointing initiation. It has recently been observed that saccades cannot be directed to new target stimuli, away from a pointing target stimulus. Saccades directed to targets presented during the final phase of a pointing movement were delayed until after pointing movement offset ("gaze anchoring"). The present study investigated whether ocular gaze is anchored to a pointing target during the entire pointing movement. In experiment 1, new targets were presented at various times during the duration of a pointing movement, triggered by the kinematics arm moment itself (movement onset, peak acceleration/velocity/deceleration, and offset). Subjects had to make a saccade to the new target as fast as possible while maintaining the pointing movement to the initial target. Saccadic latencies were increased by an amount of time that approximately equaled the remaining pointing time after saccadic target presentation, with the majority of saccades executed after pointing movement offset. The nature of the signal driving gaze stabilization during pointing was investigated in experiment 2. In previous experiments where ocular gaze was anchored to a pointing target, subjects could always see their moving arm, thus it was unknown whether a visual image of the moving arm, an afferent (proprioceptive) signal or an efferent (motor control related) signal produced gaze anchoring. In experiment 2 subjects had to point with or without vision of the moving arm to test whether a visual signal is used to anchor gaze to a pointing target. Results indicate that gaze anchoring was also observed without vision of the moving arm. The findings support the existence of a mechanism enforcing ocular gaze anchoring during the entire duration of a pointing movement. Moreover, such a mechanism uses an internally generated, or proprioceptive, nonvisual signal. Possible neural substrates underlying these processes are discussed, as well as the role of selective attention.     

Amplitude and direction errors in kinesthetic pointing

We investigated the accuracy with which, in the absence of vision, one can reach again a 2D target location that had been previously identified by a guided movement. A robotic arm guided the participants hand to a target (locating motion) and away from it (homing motion). Then, the participant pointed freely toward the remembered target position. Two experiments manipulated separately the kinematics of the locating and homing motions. Some robot motions followed a straight path with the bell-shaped velocity profile that is typical of natural movements. Other motions followed curved paths, or had strong acceleration and deceleration peaks. Current motor theories of perception suggest that pointing should be more accurate when the homing and locating motion mimics natural movements. This expectation was not borne out by the results, because amplitude and direction errors were almost independent of the kinematics of the locating and homing phases. In both experiments, participants tended to overshoot the target positions along the lateral directions. In addition, pointing movements towards oblique targets were attracted by the closest diagonal (oblique effect). This error pattern was robust not only with respect to the manner in which participants located the target position (perceptual equivalence), but also with respect to the manner in which they executed the pointing movements (motor equivalence). Because of the similarity of the results with those of previous studies on visual pointing, it is argued that the observed error pattern is basically determined by the idiosyncratic properties of the mechanisms whereby space is represented internally.     

Are arm trajectories planned in kinematic or dynamic coordinates? An adaptation study

There are several invariant features of pointto-point human arm movements: trajectories tend to be straight, smooth, and have bell-shaped velocity profiles. One approach to accounting for these data is via optimization theory; a movement is specified implicitly as the optimum of a cost function, e.g., integrated jerk or torque change. Optimization models of trajectory planning, as well as models not phrased in the optimization framework, generally fall into two main groups-those specified in kinematic coordinates and those specified in dynamic coordinates. To distinguish between these two possibilities we have studied the effects of artificial visual feedback on planar two-joint arm movements. During self-paced point-to-point arm movements the visual feedback of hand position was altered so as to increase the perceived curvature of the movement. The perturbation was zero at both ends of the movement and reached a maximum at the midpoint of the movement. Cost functions specified by hand coordinate kinematics predict adaptation to increased curvature so as to reduce the visual curvature, while dynamically specified cost functions predict no adaptation in the underlying trajectory planner, provided the final goal of the movement can still be achieved. We also studied the effects of reducing the perceived curvature in transverse movements, which are normally slightly curved. Adaptation should be seen in this condition only if the desired trajectory is both specified in kinematic coordinates and actually curved. Increasing the perceived curvature of normally straight sagittal movements led to significant (P<0.001) corrective adaptation in the curvature of the actual hand movement; the hand movement became curved, thereby reducing the visually perceived curvature. Increasing the curvature of the normally curved transverse movements produced a significant (P<0.01) corrective adaptation; the hand movement became straighter, thereby again reducing the visually perceived curvature. When the curvature of naturally curved transverse movements was reduced, there was no significant adaptation (P>0.05). The results of the curvature-increasing study suggest that trajectories are planned in visually based kinematic coordinates. The results of the curvature-reducing study suggest that the desired trajectory is straight in visual space. These results are incompatible with purely dynamicbased models such as the minimum torque change model. We suggest that spatial perception-as mediated by vision-plays a fundamental role in trajectory planning.     

Reaching during virtual rotation: context specific compensations for expected coriolis forces

Subjects who are in an enclosed chamber rotating at constant velocity feel physically stationary but make errors when pointing to targets. Reaching paths and endpoints are deviated in the direction of the transient inertial Coriolis forces generated by their arm movements. By contrast, reaching movements made during natural, voluntary torso rotation seem to be accurate, and subjects are unaware of the Coriolis forces generated by their movements. This pattern suggests that the motor plan for reaching movements uses a representation of body motion to prepare compensations for impending self-generated accelerative loads on the arm. If so, stationary subjects who are experiencing illusory self-rotation should make reaching errors when pointing to a target. These errors should be in the direction opposite the Coriolis accelerations their arm movements would generate if they were actually rotating. To determine whether such compensations exist, we had subjects in four experiments make visually open-loop reaches to targets while they were experiencing compelling illusory self-rotation and displacement induced by rotation of a complex, natural visual scene. The paths and endpoints of their initial reaching movements were significantly displaced leftward during counterclockwise illusory rotary displacement and rightward during clockwise illusory self-displacement. Subjects reached in a curvilinear path to the wrong place. These reaching errors were opposite in direction to the Coriolis forces that would have been generated by their arm movements during actual torso rotation. The magnitude of path curvature and endpoint errors increased as the speed of illusory self-rotation increased. In successive reaches, movement paths became straighter and endpoints more accurate despite the absence of visual error feedback or tactile feedback about target location. When subjects were again presented a stationary scene, their initial reaches were indistinguishable from pre-exposure baseline, indicating a total absence of aftereffects. These experiments demonstrate that the nervous system automatically compensates in a context-specific fashion for the Coriolis forces associated with reaching movements.     

Dissociating visual and kinesthetic coordinates during pointing movements

Goal-directed movements imply that the visual coordinates in which the localisation of the goal is coded are transformed into proprioceptive coordinates in which the arm movement is coded. The two systems of coordinates are normally superimposed. Using a virtual reality device attached to the subject's head, we have created a situation where these systems were dissociated from each other. The virtual environment involved virtual visual targets and an image of the subject's hand reconstructed from the output of a data glove wore by the subject's right hand. When the subject's head was rotated, the visual targets and the image of the hand rotated by the same amount. Movements of the real hand were thus in conflict with those of the reconstructed hand, which appeared to err in the direction of head rotation. Pointing movements directed at five targets (0°, 26° and 52° on each side) were studied for five different head positions (0°, 45° and 80° to the right and to the left). The results showed a significant pointing bias towards head position, except for the left-most targets in the right head rotations. Constant errors in azimuth were proportional to the amount of head rotation. When the head was rotated to the right, constant errors in azimuth were greater during pointing towards right than left targets. Similarly, they were greater for left than for right stimuli when the head was rotated to the left. Errors in amplitude were not influenced by the direction nor the amount of head rotation. Finally, a decrease in the directional bias took place within blocks of trials. These results indicate that head position signals are used during the process of transforming motor coordinates from the visual to the pro-prioceptive system of reference.     

Multimodal reference frame for the planning of vertical arms movements

In this study we investigated the reference frames used to plan arm movements. Specifically, we asked whether the body axis, visual cues and graviception can each play a role in defining "up" and "down" in the planning and execution of movements along the vertical axis. Horizontal and vertical pointing movements were tested in two postures (upright and reclined) and two visual conditions (with and without vision) to identify possible effects of each of these cues on kinematics of movement. Movements were recorded using an optical 3D tracking system and analysis was conducted on velocity profiles of the hand. Despite a major effect of gravity, our analysis shows an effect of the movement direction with respect to the body axis when subjects were reclined with eyes closed. These results suggest that our CNS takes into account multimodal information about vertical in order to compute an optimal motor command that anticipates the effects of gravity.     

Eye-head-hand coordination in pointing at visual targets: spatial and temporal analysis

This study investigated whether the execution of an accurate pointing response depends on a prior saccade orientation towards the target, independent of the vision of the limb. A comparison was made between the accuracy of sequential responses (in which the starting position of the hand is known and the eye centred on the target prior to the onset of the hand pointing movement) and synergetic responses (where both hand and gaze motions are simultaneously initiated on the basis of unique peripheral retinal information). The experiments were conducted in visual closed-loop (hand visible during the pointing movement) and in visual openloop conditions (vision of hand interrupted as the hand started to move). The latter condition eliminated the possibility of a direct visual evaluation of the error between hand and target during pointing. Three main observations were derived from the present work: (a) the timing of coordinated eye-head-hand pointing at visual targets can be modified, depending on the executed task, without a deterioration in the accuracy of hand pointing; (b) mechanical constraints or instructions such as preventing eye, head or trunk motion, which limit the redundancy of degrees of freedom, lead to a decrease in accuracy; (c) the synergetic movement of eye, head and hand for pointing at a visible target is not trivially the superposition of eye and head shifts added to hand pointing. Indeed, the strategy of such a coordinated action can modify the kinematics of the head in order to make the movements of both head and hand terminate at approximately the same time. The main conclusion is that eye-head coordination is carried out optimally by a parallel processing in which both gaze and hand motor responses are initiated on the basis of a poorly defined retinal signal. The accuracy in hand pointing is not conditioned by head movement per se and does not depend on the relative timing of eye, head and hand movements (synergetic vs sequential responses). However, a decrease in the accuracy of hand pointing was observed in the synergetic condition, when target fixation was not stabilised before the target was extinguished. This suggests that when the orienting saccade reaches the target before hand movement onset, visual updating of the hand motor control signal may occur. A rapid processing of this final input allows a sharper redefinition of the hand landing point.     

Sensorimotor adaptation of point-to-point arm movements after spaceflight: the role of internal representation of gravity force in trajectory planning

After an exposure to weightlessness, the central nervous system operates under new dynamic and sensory contexts. To find optimal solutions for rapid adaptation, cosmonauts have to decide whether parameters from the world or their body have changed and to estimate their properties. Here, we investigated sensorimotor adaptation after a spaceflight of 10 days. Five cosmonauts performed forward point-to-point arm movements in the sagittal plane 40 days before and 24 and 72 h after the spaceflight. We found that, whereas the shape of hand velocity profiles remained unaffected after the spaceflight, hand path curvature significantly increased 1 day after landing and returned to the preflight level on the third day. Control experiments, carried out by 10 subjects under normal gravity conditions, showed that loading the arm with varying loads (from 0.3 to 1.350 kg) did not affect path curvature. Therefore, changes in path curvature after spaceflight cannot be the outcome of a control process based on the subjective feeling that arm inertia was increased. By performing optimal control simulations, we found that arm kinematics after exposure to microgravity corresponded to a planning process that overestimated the gravity level and optimized movements in a hypergravity environment (～1.4 g). With time and practice, the sensorimotor system was recalibrated to Earth's gravity conditions, and cosmonauts progressively generated accurate estimations of the body state, gravity level, and sensory consequences of the motor commands (72 h). These observations provide novel insights into how the central nervous system evaluates body (inertia) and environmental (gravity) states during sensorimotor adaptation of point-to-point arm movements after an exposure to weightlessness.     

Postural configuration does not alter unperturbed or perturbed reach movement kinematics

This study investigated whether postural configuration has a significant effect upon the kinematics of arm movements when humans performed unconstrained reach movements to visual targets. Eight subjects were required to reach to static visual targets (unperturbed REACH movements) or correct reach movements when the position of a target unexpectedly changed during the execution of a planned movement (perturbed reaches, or online corrections, OC). Subjects were required to execute REACH and OC movements in sitting and standing (STAND) positions. The height of the targets, distance from the right shoulder (acromion) and eccentricity in terms of the body midline were standardized between the two postural conditions before movements begun. Unperturbed REACH movements were executed to a central target placed at 130 % of outstretched arm length, along the midline (0°). Perturbed (OC) movements involved subjects initiating an arm movement to the 0° target upon its illumination. Two hundred milliseconds after the onset of the hand movement, the 0° target was extinguished and the target at 60° to the right of the midline (still at 130 % outstretched arm distance) illuminated. Subjects had to correct their reach movements online to the new target. Results demonstrated that, despite evident differences in postural kinematics between the four experimental conditions (e.g. pelvis obliquity and trunk/pelvis rotation), postural configuration had little or no effect upon the endpoint kinematics of the finger. Most importantly, the STAND position, with its greater postural constraints, did not affect the time taken to initiate an OC, nor did it lengthen the time taken to complete the REACH or OC movements. Our results suggest, therefore, that postural constraints are accounted for by the central nervous system when executing complex arm reaching movements.     

Postural adjustments for online corrections of arm movements in standing humans

The aim of this study was to investigate how humans correct ongoing arm movements while standing. Specifically, we sought to understand whether the postural adjustments in the legs required for online corrections of arm movements are predictive or rely on feedback from the moving limb. To answer this question we measured online corrections in arm and leg muscles during pointing movements while standing. Nine healthy right-handed subjects reached with their dominant arm to a visual target in front of them and aligned with their midline. In some trials, the position of the target would switch from the central target to one of the other targets located 15°, 30°, or 45° to the right of the central (midline) target. For each target correction, we measured the time at which arm kinematics, ground reaction forces, and arm and leg muscle electromyogram significantly changed in response to the target displacement. Results show that postural adjustments in the left leg preceded kinematic corrections in the limb. The corrective postural muscle activity in the left leg consistently preceded the corrective reaching muscle activity in the right arm. Our results demonstrate that corrections of arm movements in response to target displacement during stance are preceded by postural adjustments in the leg contralateral to the direction of target shift. Furthermore, postural adjustments preceded both the hand trajectory correction and the arm-muscle activity responsible for it, which suggests that the central nervous system does not depend on feedback from the moving arm to modify body posture during voluntary movement. Instead, postural adjustments lead the online correction in the arm the same way they lead the initiation of voluntary arm movements. This suggests that forward models for voluntary movements executed during stance incorporate commands for posture that are produced on the basis of the required task demands.     

The curvature of human arm movements in the absence of visual experience

It has been suggested that the spatial path of the hand is an important controlled feature of normal human arm movements and that the desired path is a straight line through external space. Recent experiments have suggested that distortions in visual perception of external space may lead to errors in its representation and thus influence the curvature of movements. The movements of blind and normal blind-folded subjects were therefore compared in a task requiring point-to-point hand movements in six directions across a horizontal worktop. Movement curvature varied with direction in both groups but was significantly higher for the blindfolded control subjects. Thus, the normals' distorted visual experience of straight lines in some orientations may lead them to make curved movement paths. The perception of curvature was also tested in the two groups in a task in which they traced the curved edge of a ruler. The blind group were slightly better at this task, although the difference was not significant. We conclude that visual experience influences point-to-point hand movements, leading to higher curvature for movements made in the fronto-parallel plane by sighted subjects due to visual distortions. These data therefore support the hypothesis that the spatial path followed by the hand is influenced by sensory inputs and is a controlled feature of human reaching movements. The data argue against the hypothesis that movement curvature is a result of optimising only the dynamics of the limb control.     

Effects of roll visual motion on online control of arm movement: reaching within a dynamic virtual environment

Reaching toward a visual target involves the transformation of visual information into appropriate motor commands. Complex movements often occur either while we are moving or when objects in the world move around us, thus changing the spatial relationship between our hand and the space in which we plan to reach. This study investigated whether rotation of a wide field-of-view immersive scene produced by a virtual environment affected online visuomotor control during a double-step reaching task. A total of 20 seated healthy subjects reached for a visual target that remained stationary in space or unpredictably shifted to a second position (either to the right or left of its initial position) with different inter-stimulus intervals. Eleven subjects completed two experiments which were similar except for the duration of the target’s appearance. The final target was either visible throughout the entire trial or only for a period of 200 ms. Movements were performed under two visual field conditions: the virtual scene was matched to the subject’s head motion or rolled about the line of sight counterclockwise at 130°/s. Nine additional subjects completed a third experiment in which the direction of the rolling scene was manipulated (i.e., clockwise and counterclockwise). Our results showed that while all subjects were able to modify their hand trajectory in response to the target shift with both visual scenes, some of the double-step movements contained a pause prior to modifying trajectory direction. Furthermore, our findings indicated that both the timing and kinematic adjustments of the reach were affected by roll motion of the scene. Both planning and execution of the reach were affected by roll motion. Changes in proportion of trajectory types, and significantly longer pauses that occurred during the reach in the presence of roll motion suggest that background roll motion mainly interfered with the ability to update the visuomotor response to the target displacement. Furthermore, the reaching movement was affected differentially by the direction of roll motion. Subjects demonstrated a stronger effect of visual motion on movements taking place in the direction of visual roll (e.g., leftward movements during counterclockwise roll). Further investigation of the hand path revealed significant changes during roll motion for both the area and shape of the 95% tolerance ellipses that were constructed from the hand position following the main movement termination. These changes corresponded with a hand drift that would suggest that subjects were relying more on proprioceptive information to estimate the arm position in space during roll motion of the visual field. We conclude that both the spatial and temporal kinematics of the reach movement were affected by the motion of the visual field, suggesting interference with the ability to simultaneously process two consecutive stimuli.     

Coordination of multi-joint arm movements in cerebellar ataxia: Analysis of hand and angular kinematics

Kinematic abnormalities of fast multijoint movements in cerebellar ataxia include abnormally increased curvature of hand trajectories and an increased hand path and are thought to originate from an impairment in generating appropriate levels of muscle torques to support normal coordination between shoulder and elbow joints. Such a mechanism predicts that kinematic abnormalities are pronounced when fast movements are performed and large muscular torques are required. Experimental evidence that systematically explores the effects of increasing movement velocities on movement kinematics in cerebellar multijoint movements is limited and to some extent contradictory. We, therefore, investigated angular and hand kinematics of natural multijoint pointing movements in patients with cerebellar degenerative disorders and healthy controls. Subjects performed self-paced vertical pointing movements with their right arms at three different target velocities. Limb movements were recorded in three-dimensional space using a two-camera infrared tracking system. Differences between patients and healthy subjects were most prominent when the subjects performed fast movements. Peak hand acceleration and deceleration were similar to normals during slow and moderate velocity movements but were smaller for fast movements. While altering movement velocities had little or no effect on the length of the hand path and angular motion of elbow and shoulder joints in normal subjects, the patients exhibited overshooting motions (hypermetria) of the hand and at both joints as movement velocity increased. Hypermetria at one joint always accompanied hypermetria at the neighboring joint. Peak elbow angular deceleration was markedly delayed in patients compared with normals. Other temporal movement variables such as the relative timing of shoulder and elbow joint motion onsets were normal in patients. Kinematic abnormalities of multijoint arm movements in cerebellar ataxia include hypermetria at both the elbow and the shoulder joint and, as a consequence, irregular and enlarged paths of the hand, and they are marked with fast but not with slow movements. Our findings suggest that kinematic movement abnormalities that characterize cerebellar limb ataxia are related to an impairment in scaling movement variables such as joint acceleration and deceleration normally with movement speed. Most likely, increased hand paths and decomposition of movement during slow movements, as described earlier, result from compensatory mechanisms the patients may employ if maximum movement accuracy is required.     

Superposition of independent units of coordination during pointing movements involving the trunk with and without visual feedback

Previous studies addressing the problem of the control of multiple degrees of freedom have examined the influence of trunk movement on pointing movements within the arm's reach. Such movements may be controlled by two functionally independent units of coordination (synergies): one involving only arm joints and producing the hand trajectory to the target (the transport synergy), and the other coordinating trunk and arm movements leaving the hand trajectory unchanged (the compensatory synergy). The question of whether or not this functional subdivision depends on visual feedback was addressed in the present study. We also tested whether or not the motor effects of different synergies are summated as independent components, a control strategy called "superposition." Finally, we investigated whether or not the relationship between different degrees of freedom within each synergy could be considered linear resulting in proportional changes in different joint angles. Seated subjects produced fast, uncorrected arm movements to an ipsi- or a contralateral target in the direction of +/-45 degrees to the sagittal midline of the trunk. Targets could be reached using the arm alone (control trials) or by combining the arm motion with a forward or backward trunk motion produced by hip flexion or extension (test trials), with and without visual feedback. The shape of the hand trajectory, its direction and tangential velocity, movement precision, joint angles and the sequence of the trunk and hand recruitment and de-recruitment were measured. In both visual conditions, the direction of the hand trajectory observed in control trials was generally preserved in test trials. In terms of sequencing, even in the absence of vision, the trunk movement was initiated before the onset of and outlasted the hand shift, indicating that the potential influence of the trunk on the hand movement was compensated by rotations in the elbow and shoulder joint. The analysis of other variables also implied that the effects of trunk recruitment on the hand trajectory were minor compared to those which could be observed if these effects were not compensated by appropriate changes in the arm joint angles. It was concluded that an arm-trunk compensatory synergy is present in pointing movements regardless of visual feedback. Principal component analysis showed that the relationship between elbow, shoulder and hip joint angles in individual arm and combined arm-trunk movements cannot be considered linear, implying that this relationship is adjusted according to the changing arm geometry. The changes in each arm joint angle (elbow, shoulder) elicited by a forward trunk bending in one block of trials were compared with those elicited by a backward bending in another block, whereas the hand moved to the same target in both blocks. These changes were opposite but of similar magnitude. As a result, for each moment of movement, the mean joint angle obtained by averaging across two directions of trunk motion was practically identical to that in control trials in which the trunk was motionless. It is concluded that the transport and arm-trunk compensatory synergies are combined as independent units, according to the principle of superposition. This principle may simplify the control of the coordination of a redundant number of degrees of freedom.     

Effects of a tilted visual background on human sensory-motor coordination

 The present study investigated the effects of a tilted visual background on perceived hand orientation, and on the execution of aimed arm movements. Subjects were seated in a room tilted about their mid-sagittal axis to the left or right. They were asked to indicate the gravitational vertical or the body midline by rotating their supported or free, unseen hand about the longitudinal forearm axis. They were further asked to draw vertical lines with their unseen arm, and to point with the hand at visual targets. Our results indicate that if the hand is stationary, tilted environments induce an illusory hand and body tilt in the opposite direction; the effects on the hand is substantially smaller than that on the body. We found no evidence for illusory hand tilt with line drawing, and pointing movements were not noticeably modified by background tilt. We concluded that the latter two tasks provide dynamic cues about hand orientation, which remain veridical in tilted environments, and can be utilized for fast corrections of motor commands. Received: 24 April 1995 / Accepted: 17 January 1997     

Contribution of retinal versus extraretinal signals towards visual localization in goal-directed movements

Summary In human subjects, we investigated the accuracy of goal-directed arm movements performed without sight of the arm; errors of target localization and of motor control thus remained uncorrected by visual feedback, and became manifest as pointing errors. Target position was provided either as retinal eccentricity or as eye position. By comparing the results to those obtained previously with combined retinal plus extraretinal position cues, the relative contribution of the two signals towards visual localization could be studied. When target position was provided by retinal signals, pointing responses revealed an over-estimation of retinal eccentricity which was of similar size for all eccentricities tested, and was independent of gaze direction. These findings were interpreted as a magnification effect of perifoveal retinal areas. When target position was provided as eye position, pointing was characterized by a substantial inter-, and intra-subject variability, suggesting that the accuracy of localization by extraretinal signals is rather limited. In light of these two qualitatively different deficits, possible mechanisms are discussed how the two signals may interact towards a more veridical visual localization.     

Integration of target and hand position signals in the posterior parietal cortex: effects of workspace and hand vision

Previous findings suggest the posterior parietal cortex (PPC) contributes to arm movement planning by transforming target and limb position signals into a desired reach vector. However, the neural mechanisms underlying this transformation remain unclear. In the present study we examined the responses of 109 PPC neurons as movements were planned and executed to visual targets presented over a large portion of the reaching workspace. In contrast to previous studies, movements were made without concurrent visual and somatic cues about the starting position of the hand. For comparison, a subset of neurons was also examined with concurrent visual and somatic hand position cues. We found that single cells integrated target and limb position information in a very consistent manner across the reaching workspace. Approximately two-thirds of the neurons with significantly tuned activity (42/61 and 30/46 for left and right workspaces, respectively) coded targets and initial hand positions separably, indicating no hand-centered encoding, whereas the remaining one-third coded targets and hand positions inseparably, in a manner more consistent with the influence of hand-centered coordinates. The responses of both types of neurons were largely invariant with respect to the presence or absence of visual hand position cues, suggesting their corresponding coordinate frames and gain effects were unaffected by cue integration. The results suggest that the PPC uses a consistent scheme for computing reach vectors in different parts of the workspace that is robust to changes in the availability of somatic and visual cues about hand position.     

Errors in pointing are due to approximations in sensorimotor transformations

1. We define an extrinsic frame of reference to represent the location of a point in extrapersonal space relative to a human subject's shoulder, and we define an intrinsic frame of reference to represent the orientation of the arm and forearm. 2. We examined the relations between coordinates in the extrinsic and intrinsic frames of reference under two experimental conditions: when subjects made inaccurate movements by pointing to virtual targets in the dark and when they made accurate movements by pointing to actual targets in the light. 3. When subjects made inaccurate movements, there was a close-to-linear relationship between the orientation angles of the arm (intrinsic coordinates) at its final position and the extrinsic coordinates of the target. When they made accurate movements, these relationships were more nonlinear. 4. Specifically, arm and forearm elevations depended principally on target distance and elevation, whereas the two yaw angles depended mainly on the target's azimuth. 5. We propose that errors in pointing occur because subjects implement a linear approximation to the transformation from extrinsic to intrinsic coordinates and that this transformation is one step in the process of transforming a visually derived representation of target location into an appropriate pattern of muscle activity.     

Eye–hand coupling is not the cause of manual return movements when searching

When searching for a target with eye movements, saccades are planned and initiated while the visual information is still being processed, so that subjects often make saccades away from the target and then have to make an additional return saccade. Presumably, the cost of the additional saccades is outweighed by the advantage of short fixations. We previously showed that when the cost of passing the target was increased, by having subjects manually move a window through which they could see the visual scene, subjects still passed the target and made return movements (with their hand). When moving a window in this manner, the eyes and hand follow the same path. To find out whether the hand still passes the target and then returns when eye and hand movements are uncoupled, we here compared moving a window across a scene with moving a scene behind a stationary window. We ensured that the required movement of the hand was identical in both conditions. Subjects found the target faster when moving the window across the scene than when moving the scene behind the window, but at the expense of making larger return movements. The relationship between the return movements and movement speed when comparing the two conditions was the same as the relationship between these two when comparing different window sizes. We conclude that the hand passing the target and then returning is not directly related to the eyes doing so, but rather that moving on before the information has been fully processed is a general principle of visuomotor control.