Role of the medial part of the intraparietal sulcus in implementing movement direction.

The contribution of the posterior parietal cortex (PPC) to visually guided movements has been originally inferred from observations made in patients suffering from optic ataxia. Subsequent electrophysiological studies in monkeys and functional imaging data in humans have corroborated the key role played by the PPC in sensorimotor transformations underlying goal-directed movements, although the exact contribution of this structure remains debated. Here, we used transcranial magnetic stimulation (TMS) to interfere transiently with the function of the left or right medial part of the intraparietal sulcus (mIPS) in healthy volunteers performing visually guided movements with the right hand. We found that a "virtual lesion" of either mIPS increased the scattering in initial movement direction (DIR), leading to longer trajectory and prolonged movement time, but only when TMS was delivered 100-160 ms before movement onset and for movements directed toward contralateral targets. Control experiments showed that deficits in DIR consequent to mIPS virtual lesions resulted from an inappropriate implementation of the motor command underlying the forthcoming movement and not from an inaccurate computation of the target localization. The present study indicates that mIPS plays a causal role in implementing specifically the direction vector of visually guided movements toward objects situated in the contralateral hemifield.

Motor urgency is mediated by the contralateral cerebellum in Parkinson's disease.

BACKGROUND: In patients with Parkinson's disease (PD), motor performance may be dramatically improved in urgent and stressful situations. OBJECTIVE: The aim of this PET H(2)(15)O study was to determine the changes in brain activation pattern related to this unconscious increase in motor speed observed in the context of urgency in patients with PD. METHODS: Eight right-handed patients with PD, who had been off medication for at least 12 hours, without tremor, were enrolled. A reaching task with the right hand was performed under three conditions: self-initiated (SI), externally cued (EC) and externally cued-urgent (ECu). RESULTS: (1) Self-initiated movements (SI-EC) revealed activations in the prefrontal cortex bilaterally, the right lateral premotor cortex, anterior cingulate cortex and cerebellum, and the left primary motor cortex and thalamus; (2) Externally driven responses (EC-SI) did not involve any statistically detectable activation; (3) Urgent situations (ECu-EC) engaged the left cerebellum. Compared with a control group previously studied, the cerebellar activation was greater in patients with PD. CONCLUSIONS: This study demonstrates that the increase in movement speed in urgent situations in patients with PD is associated with the recruitment of the left (contralateral) cerebellum. This structure is a key node of the accessory motor circuitry typically recruited by patients with PD to compensate for basal ganglia dysfunction and by healthy subjects to increase movement velocity in urgent motor contexts.

An 'automatic pilot' for the hand in human posterior parietal cortex: toward reinterpreting optic ataxia.

We designed a protocol distinguishing between automatic and intentional motor reactions to changes in target location triggered at movement onset. In response to target jumps, but not to a similar change cued by a color switch, normal subjects often could not avoid automatically correcting fast aiming movements. This suggests that an 'automatic pilot' relying on spatial vision drives fast corrective arm movements that can escape intentional control. In a patient with a bilateral posterior parietal cortex (PPC) lesion, motor corrections could only be slow and deliberate. We propose that 'on-line' control is the most specific function of the PPC and that optic ataxia could result from a disruption of automatic hand guidance.

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.

Functional anatomy of saccadic adaptation in humans.

Positron emission tomography (PET) was used to investigate the neurophysiological substrate of human saccadic adaptation. Subjects made saccadic eye movements toward a visual target that was displaced during the course of the initial saccade, a time when visual perception is suppressed. In one condition, displacement was random from trial to trial, precluding any systematic modification of the initial saccade amplitude. In the second condition, the direction and magnitude of displacement were consistent, causing adaptative modification of the initial saccade amplitude. PET difference images reflecting metabolic changes attributable to the process of saccadic adaptation showed selective activation of the medioposterior cerebellar cortex. This localization is consistent with neurophysiological findings in monkeys and brain-lesioned humans.

Functional anatomy of nonvisual feedback loops during reaching: a positron emission tomography study.

Reaching movements performed without vision of the moving limb are continuously monitored, during their execution, by feedback loops (designated nonvisual). In this study, we investigated the functional anatomy of these nonvisual loops using positron emission tomography (PET). Seven subjects had to "look at" (eye) or "look and point to" (eye-arm) visual targets whose location either remained stationary or changed undetectably during the ocular saccade (when vision is suppressed). Slightly changing the target location during gaze shift causes an increase in the amount of correction to be generated. Functional anatomy of nonvisual feedback loops was identified by comparing the reaching condition involving large corrections (jump) with the reaching condition involving small corrections (stationary), after subtracting the activations associated with saccadic movements and hand movement planning [(eye-arm-jumping minus eye-jumping) minus (eye-arm-stationary minus eye-stationary)]. Behavioral data confirmed that the subjects were both accurate at reaching to the stationary targets and able to update their movement smoothly and early in response to the target jump. PET difference images showed that these corrections were mediated by a restricted network involving the left posterior parietal cortex, the right anterior intermediate cerebellum, and the left primary motor cortex. These results are consistent with our knowledge of the functional properties of these areas and more generally with models emphasizing parietal-cerebellar circuits for processing a dynamic motor error signal.

On-line motor control in patients with Parkinson's disease.

Recent models based, in part on a study of Huntington's disease, suggest that the basal ganglia are involved in on-line movement guidance. Two experiments were conducted to investigate this idea. First, we studied advanced Parkinson's disease patients performing a reaching task known to depend on on-line guidance. The task was to 'look and point' in the dark at visual targets displayed in the peripheral visual field. In some trials, the target location was slightly modified during saccadic gaze displacement (when vision is suppressed). In both patient and control groups, the target jump induced a gradual modification of the movement which diverged smoothly from its original path to reach the new target location. No deficit was found in the patients, except for an increased latency to respond to the target jump (Parkinson's disease: 243 ms; controls: 166 ms). A computational simulation indicated that this response slowing was likely to be a by-product of bradykinesia. The unexpected inconsistency between this result and previous reports was investigated in a second experiment. We hypothesized that the relevant factor was the characteristics of the corrections to be performed. To test this prediction, we investigated a task requiring corrections of the same type as investigated in Huntington's disease, namely large, consciously detected errors induced by large target jumps at hand movement onset. In contrast with the smooth adjustments observed in the first experiment, the subjects responded to the target jump by generating a discrete corrective sub-movement. While this iterative response was relatively rapid in the control subjects (220 ms), Parkinson's disease patients exhibited either dramatically late (>730 ms) or totally absent on-line corrections. When on-line corrections were absent, the initial motor response was completed before a second corrective response was initiated (the latency of the corrective response was the same as the latency of the initial response). Considered together, these results suggest that basal ganglia dependent circuits are not critical for feedback loops involving a smooth modulation of the ongoing command. These circuits may rather contribute to the generation of discrete corrective sub-movements. This deficit is in line with the general impairment of sequential and simultaneous actions in patients with basal ganglia disorders.

Sensorimotor mapping of the human cerebellum during pineal region surgery.

BACKGROUND: The cerebellum is a fundamental structure of the central nervous system. However, in humans, its anatomo-functional organization and the processes through which this organization adapts in response to injuries remain widely unknown. METHODS: Motor and somatosensory evoked potentials were used to map functional representations in the posterior cerebellum of patients with extra- and intracellebellar injuries. Extracerebellar patients had injuries outside the cerebellum (e.g. pineal region, quadrigeminal plate) while intracerebellar patients had injuries within the cerebellum. Data were collected in 20 extracerebellar patients for motor representations. Only preliminary data were gathered for somatosensory representations and intracerebellar patients. RESULTS: In extracerebellar patients, electrical stimulation induced muscle contractions in the ipsilateral hemibody. These representations were somatotopically organized with large overlaps between the face and upper limb in the superior posterior cerebellum and the upper and lower limb in the inferior posterior cerebellum. Neck muscles were represented in the oculomotor vermis. In intracerebellar patients, preliminary data seem to indicate that motor plasticity is achieved by recruiting the contralesional (healthy) cerebellar hemisphere. CONCLUSIONS: Although still ongoing, this project could eventually lead to an improvement of the surgical treatment of patients with lesions of the posterior fossa, by improving our knowledge of cerebellar organization and the process of post-lesional plasticity.

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.

[Effects on children's cognitive development of chronic exposure to screens]. / Effets de l'exposition chronique aux écrans sur le développement cognitif de l'enfant.

During the last few years, the time spent in front of various screens, including TV sets, video games, smartphones and computers, has dramatically increased. Numerous studies show, with a remarkable consistency, that this trend has a strong negative influence on the cognitive development of children and teenagers. The affected fields include, in particular, scholastic achievement, language, attention, sleep and aggression. We believe that this often disregarded - not to say denied - problem should now be considered a major public health issue. Primary care physicians should inform parents and children about this issue to support efficient prevention.

Testing basal ganglia motor functions through reversible inactivations in the posterior internal globus pallidus.

To test current hypotheses on the contribution of the basal ganglia (BG) to motor control, we examined the effects of muscimol-induced inactivations in the skeletomotor region of the internal globus pallidus (sGPi) on visually directed reaching. Injections were made in two monkeys trained to perform four out-and-back reaching movements in quick succession toward four randomly selected target locations. Following sGPi inactivations the following occurred. 1) Peak velocity and acceleration were decreased in nearly all sessions, whereas movement duration lengthened inconsistently. 2) Reaction times were unaffected on average, although minor changes were observed in several individual sessions. 3) Outward reaches showed a substantial hypometria that correlated closely with bradykinesia, but directional accuracy was unaffected. 4) Endpoint accuracy was preserved for the slow visually guided return movements. 5) No impairments were found in the rapid chaining of out-and-back movements, in the selection or initiation of four independent reaches in quick succession or in the quick on-line correction of initially misdirected reaches. 6) Inactivation-induced reductions in the magnitude of movement-related muscle activity (EMG) correlated with the severity of slowing and hypometria. There was no evidence for inactivation-induced alterations in the relative timing of EMG bursts, excessive cocontraction, or impaired suppression of antagonist EMG. Therefore disconnecting the BG motor pathway consistently produced bradykinesia and hypometria, but seldom affected movement initiation time, feedback-mediated guidance, the capacity to produce iterative reaches, or the ability to abruptly reverse movement direction. These results are discussed with reference to the idea that the BG motor loop may regulate energetic expenditures during movement (i.e., movement "vigor").

Perceptual factors contribute to akinesia in Parkinson's disease.

Parkinson's disease (PD) patients have longer reaction time (RT) than age-matched control subjects. During the last decades, conflicting results have been reported regarding the source of this deficit. Here, we addressed the possibility that experimental inconsistencies originated in the composite nature of RT responses. To investigate this idea, we examined the effect of PD on different processes that compose RT responses. Three variables were manipulated: the signal quality, the stimulus-response compatibility and the foreperiod duration. These variables have been shown to affect, respectively, the ability to extract the relevant features of the stimulus (perceptual stage), the intentional selection of the motor response (cognitive stage) and the implementation of the muscle command (motor stage). Sixteen PD patients were tested on and off-medication and compared with an age and gender-matched control group. Results indicated that degrading the legibility of the response stimulus affected the latency of simple key-press movements more dramatically in the off-medication PD group than in the control population. The stimulus-response compatibility and the foreperiod duration had similar effects in the two groups. Interestingly, the response slowing associated with the degradation of the stimulus was the same whether the patients were on or off dopaminergic medication. This suggests that the high-level perceptual deficits observed in the present study do not have a dopaminergic origin.

Normalizing motor-related brain activity: subthalamic nucleus stimulation in Parkinson disease.

OBJECTIVE: To test whether therapeutic unilateral deep brain stimulation (DBS) of the subthalamic nucleus (STN) in patients with Parkinson disease (PD) leads to normalization in the pattern of brain activation during movement execution and control of movement extent. METHODS: Six patients with PD were imaged off medication by PET during performance of a visually guided tracking task with the DBS voltage programmed for therapeutic (effective) or subtherapeutic (ineffective) stimulation. Data from patients with PD during ineffective stimulation were compared with a group of 13 age-matched control subjects to identify sites with abnormal patterns of activation. Conjunction analysis was used to identify those areas in patients with PD where activity normalized when they were treated with effective stimulation. RESULTS: For movement execution, effective DBS caused an increase of activation in the supplementary motor area (SMA), superior parietal cortex, and cerebellum toward a more normal pattern. At rest, effective stimulation reduced overactivity of SMA. Therapeutic stimulation also induced reductions of movement related "overactivity" compared with healthy subjects in prefrontal, temporal lobe, and basal ganglia circuits, consistent with the notion that many areas are recruited to compensate for ineffective motor initiation. Normalization of activity related to the control of movement extent was associated with reductions of activity in primary motor cortex, SMA, and basal ganglia. CONCLUSIONS: Effective subthalamic nucleus stimulation leads to task-specific modifications with appropriate recruitment of motor areas as well as widespread, nonspecific reductions of compensatory or competing cortical activity.

Postural control of three-dimensional prehension movements.

This experiment was carried out to test the hypothesis that three-dimensional upper limb movements could be initiated and controlled in the joint space via a mechanism comparing an estimate of the current postural state of the upper arm with a target value determined by one specific inverse static transform converting the coordinates of the object into a set of arm, forearm, and wrist angles. This hypothesis involves two main predictions: 1) despite joint redundancy, the posture reached by the upper limb should be invariant for a given context; and 2) a movement programmed in joint space should exhibit invariant characteristics of the joint covariation pattern as well as a corresponding variable hand path curvature in the task space. To test these predictions, we examined prehension movements toward a cylindrical object presented at a fixed spatial location and at various orientations without vision of the moving limb. Once presented, the object orientation was either kept constant (unperturbed trials) or suddenly modified at movement onset (perturbed trials). Three-dimensional movement trajectories were analyzed in both joint and task spaces. For the unperturbed trials, the task space analysis showed a variable hand path curvature depending on object orientation. The joint space analysis showed that the seven final angles characterizing the upper limb posture at hand-to-object contact varied monotonically with object orientation. At a dynamic level, movement onset and end were nearly identical for all joints. Moreover, for all joints having a monotonic variation, maximum velocity occurred almost simultaneously. For the elbow, the only joint presenting a reversal, the reversal was synchronized with the time to peak velocity of the other joint angles. For the perturbed trials, a smooth and complete compensation of the movement trajectory was observed in the task space. At a static level the upper limb final posture was identical to that obtained when the object was initially presented at the orientation following the perturbation. This result was particularly remarkable considering the large set of comfortable postures allowed by joint redundancy. At a dynamic level, the joints' covariation pattern was updated to reach the new target posture. The initial synergies were not disrupted by the perturbation, but smoothly modified, the different joints' movements ending nearly at the same time. Taken together, these results support the hypothesis that prehension movements are initiated and controlled in the joint space on the basis of a joint angular error vector rather than a spatial error vector.

Final posture of the upper limb depends on the initial position of the hand during prehension movements.

The question of knowing how the nervous system transforms a desired position and orientation of the hand into a set of arm and forearm angles has been widely addressed during the last few decades. Despite this fact, it still remains unclear as to whether a unique posture of the arm is associated with every location and orientation of the hand in space. The main objective of the present study a was to address this question. To this end, we studied a prehension task requiring human subjects to reach and grasp a cylindrical object presented at different locations, along variable orientations. In contrast to previous investigations, we considered the influence of the initial position of the hand. Results showed that the posture of the arm: (1) varied systematically as a function of the movement starting point; (2) was stereotyped for a particular subject given a configuration of the object and a movement starting location; (3) was altered at both the distal and proximal levels when the orientation of the object was changed; (4) was similarly influenced by the experimental factors in all the subjects, except one. When considered together, the previous results support three main conclusions: First, the nervous system solves the joint redundancy problem using fixed strategies. Second, these fixed strategies do not provide a single correspondence between hand configuration and arm posture. Third, the position and orientation of the hand in space are unlikely to be controlled through separate independent neural pathways.

Postural invariance in three-dimensional reaching and grasping movements.

The question of whether the final arm posture to be reached is determined in advance during prehension movements remains widely debated. To address this issue, we designed a psychophysical experiment in which human subjects were instructed to reach and grasp, with their right arm, a small sphere presented at various locations. In some trials the sphere remained stationary, while in others (the perturbed trials) it suddenly jumped, at movement onset, to a new unpredictable position. Our data indicate that the final configuration of the upper limb is highly predictable for a given location of the sphere. For movements directed at stationary objects, the variability of the final arm posture was very small in relation to the variability allowed by joint redundancy. For movements directed at "jumping" objects, the initial motor response was quickly amended, allowing an accurate grasp. The final arm posture reached at the end of the perturbed trials was neither different from nor more variable than the final arm posture reached at the end of the corresponding stationary trials (i.e. the trials sharing the same final object location). This latter result is not trivial, considering both joint redundancy and the motor reorganization imposed by the change in sphere location. In contrast to earlier observations, our data cannot be accounted for by biomechanical or functional factors. Indeed, the spherical object used in the present study did not constrain the final arm configuration or the hand trajectory. When considered together, our data support the idea that the final posture to be reached is planned in advance and used as a control variable by the central nervous system.

Vectorial coding of movement: vision, proprioception, or both?

1. Subjects were asked to point toward visual targets without visual reafference from the moving hand in two conditions. In both conditions the pointing fingertip was viewed only before movement onset. 2. In one condition, the pointing fingertip was viewed through prisms that created a visual displacement without altering the view of the target. In another experimental condition, vision of the fingertip was not displaced. Comparison of these two conditions showed that virtually shifting finger position before movement through prisms induced a pointing bias in the direction opposite to the shift. The extent of this pointing bias was about one third of the prismatic shift applied to the fingertip. 3. Analysis of movement initial direction demonstrated that it was also less deviated than predicted from the prismatic shift. In addition, the reaction time and movement time of the reaching movement were increased. 4. This result is interpreted in the framework of the vectorial coding of reaching movement. Proprioception and vision provide two possible sources of information about initial hand position, i.e., the origin of the movement vector. The question remains as to how these two sources of information interact in specifying initial hand position when they are simultaneously available. 5. Our results are thus discussed with respect to a visual-to-visual movement vector hypothesis and a proprioceptive-to-visual vector hypothesis. It is argued that the origin of the putative movement vector is encoded by weighted fusion of the visual and the proprioceptive information about hand initial position.

Constrained and unconstrained movements involve different control strategies.

This experiment was carried out to test whether or not the rules governing the execution of compliant and unconstrained movements are different (a compliant motion is defined as a motion constrained by external contact). To answer this question we examined the characteristics of visually directed movements performed with either the index fingertip (unconstrained) or a hand-held cursor (compliant). For each of these categories of movements, two experimental conditions were investigated: no instruction about hand path, and instruction to move the fingertip along a straight-line path. The results of the experiment were as follows: 1) The spatiotemporal characteristics of the compliant and unconstrained movements were fundamentally different when the subjects were not required to follow a specific hand path. 2) The instruction to perform straight movements modified the characteristics of the unconstrained movements, but not those of the compliant movements. 3) The target eccentricity influenced selectively the curvature of the "unconstrained-no path instruction" movements. Taken together, these results suggest that compliant and unconstrained movements involve different control strategies. Our data support the hypothesis that unconstrained motions are, unlike compliant motions, not programmed to follow a straight-line path in the task space. These observations provide a theoretical reference frame within which some apparently contradictory results reported in the movement generation literature may be explained.

Pointing errors reflect biases in the perception of the initial hand position.

By comparing the visuomotor performance of 10 adult, normal subjects in three tasks, we investigated whether errors in pointing movements reflect biased estimations of the hand starting position. In a manual pointing task with no visual feedback, subjects aimed at 48 targets spaced regularly around two starting positions. Nine subjects exhibited a similar pattern of systematic errors across targets, i.e., a parallel shift of the end points that accounted, on average, for 49% of the total variability. The direction of the shift depended on the starting location. Systematic errors decreased dramatically in the second condition where subjects were allowed to see their hand before movement onset. The third task was to use a joystick held by the left hand to estimate the location of their (unseen) right hand. The systematic perceptual errors in this condition were found to be highly correlated with the motor errors in the first condition. The results support the following conclusions. 1) Kinesthetic estimation of hand position may be consistently biased. Some of the mechanisms responsible for these biases are always active, irrespective of whether position is estimated overtly (e.g., with a matching paradigm), or covertly as part of the motor planning for aimed movements. 2) Pointing errors reflect to a significant extent the erroneous estimation of initial hand position. This suggests that aimed hand movements are planned vectorially, i.e., in terms of distance and direction, rather than in terms of absolute position in space.

Forward modeling allows feedback control for fast reaching movements.

Delays in sensorimotor loops have led to the proposal that reaching movements are primarily under pre-programmed control and that sensory feedback loops exert an influence only at the very end of a trajectory. The present review challenges this view. Although behavioral data suggest that a motor plan is assembled prior to the onset of movement, more recent studies have indicated that this initial plan does not unfold unaltered, but is updated continuously by internal feedback loops. These loops rely on a forward model that integrates the sensory inflow and motor outflow to evaluate the consequence of the motor commands sent to a limb, such as the arm. In such a model, the probable position and velocity of an effector can be estimated with negligible delays and even predicted in advance, thus making feedback strategies possible for fast reaching movements. The parietal lobe and cerebellum appear to play a crucial role in this process. The ability of the motor system to estimate the future state of the limb might be an evolutionary substrate for mental operations that require an estimate of sequelae in the immediate future.