Response delay and spatial representation in pointing movements

Pointing movements decrease in accuracy when target information is removed before movement onset. This time effect was analyzed in relation with the spatial representation of the target location, which can be egocentric (i.e. in relation to the body) or exocentric (i.e. in relation to the external world) depending on the visual environment of the target. The accuracy of pointing movements performed without visual feedback was measured in two delay conditions: 0 and 5-s delay between target removal and movement onset. In each delay condition, targets were presented either in the darkness (egocentric localization) or within a structured visual background (exocentric localization). The results show that pointing was more accurate when targets were presented within a visual background than in the darkness. The time-related decrease in accuracy was observed in the darkness condition, whereas no delay effect was found in the presence of a visual background. Therefore, contextual factors applied to a simple pointing action might induce different spatial representations: a short-lived sensorimotor egocentric representation used in immediate action control, or a long-lived perceptual exocentric representation which drives perception and delayed action.     

Eye and neck proprioceptive messages contribute to the spatial coding of retinal input in visually oriented activities

Summary The egocentric localization of objects in extrapersonal space requires that the retinal and extraretinal signals specifying the gaze direction be simultaneously processed. The question as to whether the extraretinal signal is of central or peripheral origin is still a matter of controversy, however. Three experiments were carried out to investigate the following hypotheses: 1) that the proprioceptive feedback originating in eye and neck muscles might provide the CNS with some indication about the gaze direction; and 2) that the retinal and proprioceptive extraretinal inputs might be jointly processed depending on whether they are of monocular or binocular origin. Application of low amplitude mechanical vibrations to either the extraocular or neck muscles (or both) of a subject looking monocularly at a small luminous target in darkness resulted in an illusory movement of the target, the direction of which depended on which muscle was stimulated. A slow upward target displacement occurred on vibrating the eye inferior rectus or the neck sterno-cleido-mastoidus muscles, whereas a downward shift was induced when the dorsal neck muscles (trapezius and splenius) were vibrated. The extent of the perceptual effects reported by subjects was measured in an open-loop pointing task in which they were asked to point at the perceived position of the target. These results extend to visually-oriented behavior the role of extraocular and neck proprioceptive inputs previously described in the case of postural regulation, since they clearly show that these messages contribute to specifying the gaze direction. This suggests that the extraretinal signal might include a proprioceptive component. The proposition that a directional body reference frame may be based on the common processing of various proprioceptive feedbacks is discussed.     

The interaction of visual and proprioceptive inputs in pointing to actual and remembered targets

Errors in pointing to actual and remembered targets presented in three-dimensional (3D) space in a dark room were studied under various conditions of visual feedback. During their movements, subjects either had no vision of their arms or of the target, vision of the target but not of their arms, vision of a light-emitting diode (LED) on their moving index fingertip but not of the target, or vision of an LED on their moving index fingertip and of the target. Errors depended critically upon feedback condition. 3D errors were largest for movements to remembered targets without visual feedback, diminished with vision of the moving fingertip, and diminished further with vision of the target and vision of the finger and the target. Moreover, the different conditions differentially influenced the radial distance, azimuth, and elevation errors, indicating that subjects control motion along all three axes relatively independently. The pattern of errors suggest that the neural systems that mediate processing of actual versus remembered targets may have different capacities for integrating visual and proprioceptive information in order to program spatially directed arm movements.     

Oculo-manual tracking of visual targets: control learning,coordination control and coordination model

Summary The processes which develop to coordinate eye and hand movements in response to motion of a visual target were studied in young children and adults. We have shown that functional maturation of the coordination control between eye and hand takes place as a result of training. We observed, in the trained child and in the adult, that when the hand is used either as a target or to track a visual target, the dynamic characteristics of the smooth pursuit system are markedly improved: the eye to target delay is decreased from 150 ms in eye alone tracking to 30 ms, and smooth pursuit maximum velocity is increased by 100%. Coordination signals between arm and eye motor systems may be responsible for smooth pursuit eye movements which occur during self-tracking of hand or finger in darkness. These signals may also account for the higher velocity smooth pursuit eye movements and the shortened tracking delay when the hand is used as a target, as well as for the synkinetic eye-arm motions observed at the early stage of oculo-manual tracking training in children. We propose a model to describe the interaction which develops between two systems involved in the execution of a common sensorimotor task. The model applies to the visuo-oculo-manual tracking system, but it may be generalized to other coordinated systems. According to our definition, coordination control results from the reciprocal transfer of sensory and motor information between two or more systems involved in the execution of single, goal-directed or conjugate actions. This control, originating in one or more highly specialized structures of the central nervous system, combines with the control processes normally operating in each system. Our model relies on two essential notions which describe the dynamic and static aspects of coordination control: timing and mutual coupling.     

The coordination of eye, head, and arm movements during rapid gaze orienting and arm pointing

This study aimed to investigate the coordination of multiple control actions involved in human horizontal gaze orienting or arm pointing to a common visual target. The subjects performed a visually triggered reaction time task in three conditions: (1) gaze orienting with a combined eye saccade and head rotation (EH), (2) arm pointing with gaze orienting by an eye saccade without head rotation (EA), and (3) arm pointing with gaze orienting by a combined eye saccade and head rotation (EHA). The subjects initiated eye movement first with nearly constant latencies across all tasks, followed by head movement in the EH task, by arm movement in the EA task, and by head and then arm movements in the EHA task. The differences of onset times between eye and head movements in the EH task, and between eye and arm movements in the EA task, were both preserved in the EHA task, leading to an eye-to-head-to-arm sequence. The onset latencies of eye and head in the EH task, eye and arm in the EA task, and eye, head and arm in the EHA task, were all positively correlated on a trial-by-trial basis. In the EHA task, however, the correlation coefficients of eye–head coupling and of eye–arm coupling were reduced and increased, respectively, compared to those estimated in the two-effector conditions (EH, EA). These results suggest that motor commands for different motor effectors are linked differently to achieve coordination in a task-dependent manner.     

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

Summary The 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.Supported by INSERM (Unité 94)     

Motor control prior to movement onset: preparatory mechanisms for pointing at visual targets

Summary The present study investigated the mechanisms involved in the preparation of pointing movements in humans. We provided visual precues on the location of the upcoming target, and registered the effect of these precues on the reaction time (RT = interval between target appearance and movement onset). Generally, precues were found to reduce RT, suggesting that some aspects of the preparatory process have been advanced in time. In Exp. 1, precues fully specified the direction required for the upcoming movement while indicating only a range of movement amplitudes; in Exp. 2, precues fully specified the amplitude and indicated a range of directions. In both experiments, RT was shorter than in control trials without precues, and gradually increased with the size of the precued amplitude or direction range. This result suggests that the preparation of either parameter is possible without knowing the precise value of the other, i.e. amplitude and direction are not prepared in a fixed order. Furthermore, our results are consistent with the view that movement preparation includes a progressive contraction of the precued range towards the final value. The speed of this process can be estimated as 0.31 cm/ms for amplitude, and 1.7 deg/ms for direction ranges. In Exp. 3 and 4, precues indicated both amplitude and direction as ranges only. The size of the amplitude range was held constant while the size of the direction range was varied (Exp. 3), or vice versa (Exp. 4). Under these conditions, RT increased with the size of the varied range. For all range sizes tested, RT when precuing both amplitude and direction as ranges corresponded to the longer of the two RTs obtained in control trials where only one parameter was precued as range (like in Exp. 1 and 2), This outcome supports the hypothesis that amplitude and direction are prepared in parallel. The contraction speeds of amplitude and direction ranges estimated from Exp. 3 and 4 were comparable to those estimated from Exp. 1 and 2, indicating that processing speed is not reduced if both parameters rather than just one have to be prepared. In Exp. 5, the precued amplitude and direction range was held constant while precue area, and thus the range of possible final arm positions, was varied. RT was independent of precue area, which argues against a major contribution of position control mechanisms. Taken together, the present data support the hypothesis that amplitude and direction are the two most predominant parameters of movement preparation.     

Role of the cerebellum in visuomotor coordination

The initiation of coupled eye and arm movements was studied in six patients with mild cerebellar dysfunction and in six age-matched control subjects. The experimental paradigm consisted of 40 deg step-tracking elbow movements made under different feedback conditions. During tracking with the eyes only, saccadic latencies in patients were within normal limits. When patients were required to make coordinated eye and arm movements, however, eye movement onset was significantly delayed. In addition, removal of visual information about arm versus target position had a pronounced differential effect on movement latencies. When the target was extinguished for 3 s immediately following a step change in target position, both eye and arm onset times were further prolonged compared to movements made to continuously visible targets. When visual information concerning arm position was removed, onset times were reduced. Eye and arm latencies in control subjects were unaffected by changes in visual feedback. The results of this study clearly demonstrate that, in contrast to earlier reports of normal saccadic latencies associated with cerebellar dysfunction, initiation of both eye and arm movements is prolonged during coordinated visuomotor tracking thus supporting a coordinative role for the cerebellum during oculo-manual tracking tasks.     

Effect of target size on spatial and temporal characteristics of a pointing movement in man

Summary The effects of constraints related to movement accuracy on the spatial and temporal characteristics of pointing movements of the arm to a target were investigated. It was found that movement time increased, even at slow speeds, when target size decreased. Spatial variability of the trajectory of the index finger was also reduced, but only in proximity to the target, when higher accuracy was demanded while variability of motion at the wrist showed little change. The effect of varying the angular orientation of the target on the trajectories of the wrist and finger was also investigated. The data support the hypothesis that motion at the shoulder and elbow joints, which is closely linked, is determined primarily by target position while motion at the wrist joint, which is only loosely coupled to the motion at the more proximal joints, is related principally to the angular orientation of the target in space. The data also suggest that wrist motion is controlled separately from motion at the more proximal joints.     

Memory pointing in children and adults: dissociations in the maturation of spatial and temporal movement parameters

In previous studies a systematic directional error (the “motor oblique effect”) was found in 2D memory pointing movements of healthy adults. In this study we extend these observations to observe that healthy children displayed the same motor oblique effect. In contrast other spatial and temporal movement parameters (mean amplitude error, square directional and amplitude error, latency and the time to maximum velocity) changed with increasing age. Memory delay increased the square directional and amplitude error independent of age. Finally failure of movement inhibition during the delay was more frequent in children compared to adults. These results favor the hypothesis that the motor oblique effect related to perceptual processing biases is constant from childhood while other movement parameters are modulated by age reflecting the continuing optimization of motor control from childhood to adulthood. The dissociation of memory and age effects suggests that motor working memory is already mature in young children.     

Spatial and temporal aspects of eye-hand coordination across different tasks

The way in which saccadic eye movements are elicited influences their latency and accuracy. Accordingly, different tasks elicit different types of saccades. Using the tasks steps, gap, memory, scanning and antisaccade, we analyzed combined eye and hand movements to determine whether both motor systems share control strategies. Errors and latencies were measured to examine whether changes in eye motor behavior are reflected in hand motor behavior. Directional and variable errors of eye and hand changed differently according to the tasks. Moreover, errors of the two systems did not correlate for any of the tasks investigated. Contrary to errors, mean latencies of eye movements were organized in the same pattern as hand movements. A correlation of latencies indicates that both motor systems rely on common information to initiate movement. Temporal coupling was stronger for intentional tasks than for reflexive tasks.     

The development of consistency and flexibility in manual pointing during middle childhood

Goal‐directed actions become truly functional and skilled when they are consistent yet flexible. In manual pointing, end‐effector consistency is characterized by the end position of the index fingertip, whereas flexibility in movement execution is captured by the use of abundant arm‐joint configurations not affecting the index finger end position. Because adults have been shown to exploit their system's flexibility in challenging conditions, we wondered whether during middle childhood children are already able to exploit motor flexibility when demanded by the situation. We had children aged 5–10 years and adults perform pointing movements in a nonchallenging and challenging condition. Results showed that end‐effector errors and flexibility in movement execution decreased with age. Importantly, only the 9‐10‐year‐olds and adults showed increased flexibility in the challenging condition. Thus, while consistency increases and flexibility decreases during mid‐childhood development, from the age of nine children appear able to employ more flexibility with increasing task demands.     

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.     

Combined eye-head gaze shifts to visual and auditory targets in humans

We studied the characteristics of combined eye-head gaze shifts in human subjects to determine whether they used similar strategies when looking at visual (V), auditory (A), and combined (V+A) targets located at several target eccentricities along the horizontal meridian. Subjects displayed considerable variability in the combinations of eye and head movement used to orient to the targets, ranging from those who always aligned their head close to the target, to those who relied predominantly on eye movements and only moved their head when the target was located beyond the limits of ocular motility. For a given subject, there was almost no variability in the amount of eye and head movement in the three target conditions (V, A, V+A). The time to initiate a gaze shift was influenced by stimulus modality and eccentricity. Auditory targets produced the longest latencies when located centrally (less than 20° eccentricity), whereas visual targets evoked the longest latencies when located peripherally (greater than 40° eccentricity). Combined targets (V+A) elicited the shortest latency reaction times at all eccentricities. The peak velocity of gaze shifts was also affected by target modality. At eccentricities between 10 and 30°, peak gaze velocity was greater for movements to visual targets than for movements to auditory targets. Movements to the combined target were of comparable speed with movements to visual targets. Despite the modality-specific differences in reaction latency and peak gaze velocity, the consistency of combinations of eye and head movement within subjects suggests that visual and auditory signals are remapped into a common reference frame for controlling orienting gaze shifts. A likely candidate is the deeper layers of the superior colliculus, because visual and auditory signals converge directly onto the neurons projecting to the eye and head premotor centers.     

Dynamics of learning and transfer of muscular and spatial relative phase in bimanual coordination: evidence for abstract directional codes

The present study addressed whether the timing of muscle activation and the relative direction of limb movements are dissociable constraints that may affect learning and transfer of bimanual coordination patterns, either independently or in combination. Subjects were assigned to two experimental groups in which the to-be-learned muscular phasing (135°) was either practiced with 45° (i.e., predominantly isodirectional) or 135° (i.e., predominantly nonisodirectional) of spatial relative phase (RP) across 2 days of practice. Prior to, during, and following practice, probe tests were held in which various relative phasing patterns were administered to assess transfer of learning. Converging evidence was obtained that the relative direction of moving limbs prominently constrained transfer of learning rather than muscular relationships. Acquisition of a specific pattern resulted in spontaneous positive transfer of learning to a new coordination pattern having the same spatial RP but not to a pattern with a different spatial RP, irrespective of muscular phasing relationships. In summary, the present results suggest that learning and transfer of coordination patterns is mediated by abstract directional codes that become part of the memory representation for bimanual coordination.     

Hand-eye coordination for rapid pointing movements. Arm movement direction and distance are specified prior to saccade onset

Visually guided arm movements such as reaching or pointing are accompanied by saccadic eye movements that typically begin prior to motion of the arm. In the past, some degree of coupling between the oculomotor and limb motor systems has been demonstrated by assessing the relative onset times of eye and arm movement, and by the demonstration of a gap effect for arm movement reaction times. However, measures of limb movement onset time based on kinematics are affected by factors such as the relatively high inertia of the limb and neuromechanical delays. The goal of the present study was thus to assess the relative timing of rapid eye and arm movements made to visual targets by examining electromyographic (EMG) activity of limb muscles in conjunction with eye and arm position measures. The observation of a positive correlation between eye and limb EMG onset latencies, and the presence of a gap effect for limb EMG onset times (a reduction in reaction time when a temporal gap is introduced between the disappearance of a central fixation point and the appearance of a new target) both support the idea that eye and arm movement initiation are linked. However, limb EMG onset in most cases precedes saccade onset, and the magnitude of EMG activity prior to eye movement is correlated with both the direction and amplitude of the upcoming arm movement. This suggests that, for the rapid movements studied here, arm movement direction and distance are specified prior to the onset of saccades.     

Location and features of instructive spatial cues do not influence the time course of covert shifts of visual spatial attention

The time course of shifting visual spatial attention to flickering stimuli in the left and right visual hemifield was investigated. The goal was to test whether an instructive peripheral salient cue located close to the newly to-be-attended location triggers faster shifts per se compared to a central cue. Besides behavioural data an objective electrophysiological measure, the steady-state visual evoked potential (SSVEP) was used to measure the time course of visual pathway facilitation in the human brain for centrally and peripherally cued shifts of spatial attention. Results revealed that both spatial cues resulted in identical time courses of shifts of covert spatial attention. This was true with respect to behavioural data and SSVEP amplitude. Results support the notion that a salient peripheral spatial cue does not automatically produce faster shifts of spatial attention to the to-be-attended location when this cue is informative and embedded in an ongoing stimulation.     

Oculo-manual coordination control: respective role of visual and non-visual information in ocular tracking of self-moved targets

We evaluated the role of visual and non-visual information in the control of smooth pursuit movements during tracking of a self-moved target. Previous works have shown that self-moved target tracking is characterised by shorter smooth pursuit latency and higher maximal velocity than eye-alone tracking. In fact, when a subject tracks a visual target controlled by his own arm, eye movement and arm movement are closely synchronised. In the present study, we showed that, in a condition where the direction of motion of a self-moved visual target was opposite to that of the arm (same amplitude, same velocity, but opposite direction of movement), the resulting smooth pursuit eye movements occurred with low latency, and continued for about 140 ms in the direction of the arm movement rather than in the direction of the actual visual target movement. After 140 ms, the eye movement direction reversed through a combination of smooth pursuit and saccades. Subsequently, while arm and visual target still moved in opposite directions, smooth pursuit occurred in pace with the visual target motion. Subjects were also submitted to a series of 60 tracking trials, for which the arm-to-target motion relationship was systematically reversed. Under these conditions subjects were able to initiate early smooth pursuit in the actual direction of the visual target. Overall, these results confirm that non-visual information produced by the arm motor system can trigger and control smooth pursuit. They also demonstrate the plasticity of the neuronal network handling eye-arm coordination control.     

The role of predictive visual temporal information in the coordination of muscle activity in catching

Summary This study addresses the question as to the nature of the information on which the preactivation of the appropriate muscles in the grasping of the ball in a onehanded catching task is initiated and coordinated. High speed film and electromyograms were recorded while experiences subjects (N = 4) caught balls — projected towards them by a ball-machine at different speeds (11.9, 13.9 and 16.2 m/s — resulting in significantly different flight times of 508, 443 and 355 ms, respectively). Tau-margins (times to contact) values were calculated at the time of the initiation of the grasp movement for each subject at each speed. No significant differences were found between taumargins at different speeds. Further, the onset of the muscle activity for the initiation of the grasp movement was shown to be independent of ball speed. These findings lend support to the contention that the initiation of the grasp movement in catching is controlled and coordinated by the optical variable tau which specifies (directly) this time-to-contact. Given that the muscle group selected includes both flexors and extensors, co-activation on the basis of tau information is evidenced.