Claude Prablanc

Role of the posterior parietal cortex in updating reaching movements to a visual target

The exact role of posterior parietal cortex (PPC) in visually directed reaching is unknown. We propose that, by building an internal representation of instantaneous hand location, PPC computes a dynamic motor error used by motor centers to correct the ongoing trajectory. With unseen right hands, five subjects pointed to visual targets that either remained stationary or moved during saccadic eye movements. Transcranial magnetic stimulation (TMS) was applied over the left PPC during target presentation. Stimulation disrupted path corrections that normally occur in response to target jumps, but had no effect on those directed at stationary targets. Furthermore, left-hand movement corrections were not blocked, ruling out visual or oculomotor effects of stimulation.

The timing of mentally represented actions

The performance of subjects walking blindly to previously inspected visual targets (located at 5, 10 or 15 m from the subjects) was studied in 2 experiments. In Expt. 1, subjects selected as good visual imagers were instructed to build up a mental representation of the target. Then they had to either actually walk or imagine themselves walking to the target. Walking time was measured in both the actual and the mental performance. It was found that subjects took almost exactly the same time in the two conditions. Accuracy of these subjects was also measured in the actual walking task. They were found to make no direction errors and to slightly overshoot target location. Subjects from another, control, group, who received no instructions about visual imagery made much larger errors. In Expt. 2, actual and mental walking times were measured in the same subjects as in Expt. 1, while they carried a 25-kg weight on their shoulders. In this condition, actual walking time was the same as in Expt. 1, although mental walking time was found to increase systematically by about 30%. These results are discussed in terms of the neural parameters encoded in the motor program for actually executing or mentally performing an action.

Optimal response of eye and hand motor systems in pointing at a visual target

In a task requiring an optimal hand pointing (with regards to both time and accuracy) at a peripheral target, there is first a saccade of the eye within 250 ms, followed 100 ms later by the hand movement. However the latency of the hand movement is poorly correlated with that of the eye movement. When the peripheral target is cut off at the onset of the saccade, there is no correlation between the error of the gaze position and the error of the hand pointing. This suggests an early parallel processing of the two motor outputs. The duration of hand movement does not change significantly when subjects either see or not see their hand (closed or open loop). In the open loop situation, the undershoot of the hand pointing increases with target eccentricity, whatever the subjects are allowed or not to do a saccade toward the target. It suggests that the encoding of eye position by itself is a poor index for an accurately guided movement of the hand.

Visual control of reaching movements without vision of the limb

SummaryThe spatial and temporal organization of hand and eye movements were studied in normal human subjects as they pointed toward small visual targets. The experiment was designed to assess the role of information about target position in correcting the trajectory of the hand when view of the hand was not available. To accomplish this, the duration of target presentation was systematically varied across blocks of trials. The results of this experiment showed that pointing movements were about 3 times more accurate when the target was present throughout the entire pointing movement, than when the target disappeared shortly after the hand movement had begun. These data indicate that pointing movements made without view of the limb are not purely preprogrammed but instead, are corrected during their execution. These modifications to the motor program are smoothly integrated into the ongoing movement and must depend upon comparing visual information about the position of the target with non-visual information about the position of the limb. The source of this non-visual information was not directly established in the present experiment but presumably must be derived from kinesthetic reafferences and/or efference copy.

The coordination of eye, head, and arm movements during reaching at a single visual target

SummaryThe time of occurrence of eye, head, and arm movements directed at the same visual target was measured in five human subjects. The latency of activation of the corresponding neck and arm muscles was also measured. It appears that although the overt movements are sequentially ordered (starting with the eye movement, then the head and finally the arm) the EMG discharges are synchronous with respect to the eye movement onset. In addition, eye movement latency appears definitely (though weakly) correlated with either neck or arm EMG latencies. Neck and arm EMG latencies are also mutually correlated. These results indicate a clustering of segmental motor programs for target oriented actions.

From eye to hand: planning goal-directed movements.

The nature of the neural mechanisms involved in movement planning still remains widely unknown. We review in the present paper the state of our knowledge of the mechanisms whereby a visual input is transformed into a motor command. For the sake of generality, we consider the main problems that the nervous system has to solve to generate a movement, that is: target localization, definition of the initial state of the motor apparatus, and hand trajectory formation. For each of these problems three questions are addressed. First, what are the main results presented in the literature? Second, are these results compatible with each other? Third, which factors may account for the existence of incompatibilities between experimental observations or between theoritical models? This approach allows the explanation of some of the contradictions existing within the movement-generation literature. It also suggests that the search for general theories may be in vain, the central nervous system being able to use different strategies both in encoding the target location with respect to the body and in planning hand displacement. In our view, this conclusion may advance the field by both opening new lines of research and bringing some sterile controversies to an end.

A lesion of the posterior parietal cortex disrupts on-line adjustments during aiming movements

It is long known that the posterior parietal cortex (PPC) is critically involved in goal-directed movements. Nevertheless, there are still some controversies about its specific functions. Although most published studies have emphasised the role of PPC in sensorimotor planning processes, it has been recently suggested that PPC can also participate to on-line movement control. We studied kinematics of hand movements in a patient with a bilateral PPC lesion who exhibited no deficit in planning of her grasping movements in central vision. She was instructed to reach and grasp a cylinder presented at different locations and her motor performance was compared to that of four healthy control subjects. To address on-line control specifically, the cylinder was quickly and unexpectedly jumped, on a few trials, at movement onset, to a new location some 10 degrees (of apparent visual angle) from the original location. The patient could easily grasp stationary objects seen in foveal vision, exhibiting the same kinematic pattern as controls. Therefore, she could plan movements accurately. In response to the object jump, unlike the controls, the patient was unable to amend her ongoing movement. In this situation, she completed two distinct movements, a first one toward the initial object location and a second one toward the final object location. These results support the hypothesis that beyond a role in movement planning, PPC plays a major role in the on-line control of reach-to-grasp movements.

Saccadic responses evoked by presentation of visual and auditory targets

SummarySaccadic eye movements evoked by the presentation of visual and auditory targets were examined and compared. Differences were found either in the pattern of the saccadic response and in the characteristics of single saccades of the same amplitude. The longer latency and the higher percentage of multiple saccade responses in the auditory case were attributed to a more complex central processing, whereas the longer duration and the lower peak velocity of the saccades to auditory targets were attributed to reduced performances of the execution mechanism in the absence of vision.

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.

The contribution of coordinated eye and head movements in hand pointing accuracy

SummaryThe accuracy of pointing movements of the hand, directed at visual targets 10° to 40° from the midline, was measured in normal human subjects. No visual feedback from the moving hand was available to the subjects. The head could be either maintained stationary (head-fixed condition) or free to move (head-free condition) during the pointing movements. It was found that the error in pointing was reduced for all targets in the head-free condition. This reduction was more important for the more eccentric target (40°). Improvement in accuracy was observed without any significant change in either the latency or the duration of eye, head or hand movements. In the head-free condition, it was found that the head was displaced in the direction of the target by an amount representing no more than 2/3 of the target amplitude. The improvement in accuracy was not influenced by the amplitude of the head movement. A model is proposed which shows how coordinated eye and head movements could improve the encoding of target position.