An interference effect of observed biological movement on action.

It has been proposed that actions are intrinsically linked to perception and that imagining, observing, preparing, or in any way representing an action excites the motor programs used to execute that same action. There is neurophysiological evidence that certain brain regions involved in executing actions are activated by the mere observation of action (the so-called "mirror system;" ). However, it is unknown whether this mirror system causes interference between observed and simultaneously executed movements. In this study we test the hypothesis that, because of the overlap between action observation and execution, observed actions should interfere with incongruous executed actions. Subjects made arm movements while observing either a robot or another human making the same or qualitatively different arm movements. Variance in the executed movement was measured as an index of interference to the movement. The results demonstrate that observing another human making incongruent movements has a significant interference effect on executed movements. However, we found no evidence that this interference effect occurred when subjects observed a robotic arm making incongruent movements. These results suggest that the simultaneous activation of the overlapping neural networks that process movement observation and execution infers a measurable cost to motor control.

Is the organisation of goal-directed action modality specific? A common temporal structure.

With the help of kinematic analysis, the temporal organization of the complex daily activity 'drinking from a bottle with a glass' was described in detail. The analysis focused on the sequential action structure, the prehensile acts, and the bimanual coordination as well as on the effect of different instruction modalities on these parameters to explore the underlying representation for this complex action. Movements of the two arms were recorded in three-dimensional space with the help of an optoelectronic device in 12 normal subjects under four conditions: (1) action pantomime after verbal instruction; (2) action imitation after observation of the action performed by the experimenter without the objects; (3) action pantomime while seeing, but not touching the objects; and finally (4) action execution with objects. Despite high execution variability, the temporal structure of the action could be precisely described by the relative duration and peak velocity of action segments, by the MGA-object size-correlation, and by linear regression analysis between the onsets of functionally related action segments. A similar structure of the action as characterized by these kinematic parameters was retained across different instruction modalities. Only when the action was executed with the objects, the interval between the movement onsets of either hand and the peak velocity of the manipulative acts were reduced, while no change was observed across the other three instruction modalities. This stability of the temporal structure suggests the existence of a level in the representation of an action where all the modalities converge.

Visuo-motor control of the ipsilateral hand: evidence from right brain-damaged patients.

We investigated the extent to which the right hemisphere is involved in the control of the ipsilateral hand by analysing the kinematics of right-hand prehension in right brain-damaged (RBD) patients. We required patients to grasp one of five possible objects, equally-sized and distributed over a 40 degrees wide workspace. With the purpose of investigating the right hemisphere contribution to the on-line visuo-motor control, we also assessed patients' ability to correct their movement "in-flight", in response to a sudden change of object position. Patients' performance was compared to that of aged-matched controls. A Younger group of healthy subjects, matching the population classically tested on double-step paradigms, was also evaluated to fully assess whether patients' kinematics could be partially due to normal ageing. As a further aim, the possible influence of hemispatial neglect was evaluated by comparing the performances of right brain-damaged patients with and without neglect. In normal subjects, the results confirmed and extended the notion of (a). positional tuning of grip formation, and (b). fast reactions following a change in object position. In addition, subtle effects of ageing on visuo-motor behaviour were shown by less efficient movement correction in the Elderly group. Patients executing reach-to-grasp actions into the left contralesional hemispace were selectively affected in both temporal and spatial aspects of movements. While their performances were relatively well preserved in the right hemispace, patients did not show positional tuning of grip formation, nor fast corrections of their movements when acting in the left hemispace. Interestingly, similar deficits were found irrespective of the presence of neglect. These results show that the right hemisphere contributes to the processing of visuo-motor information that is necessary for executing actions with the ipsilateral hand in the contralateral space.

A paradoxical improvement of misreaching in optic ataxia: new evidence for two separate neural systems for visual localization.

We tested a patient (A. T.) with bilateral brain damage to the parietal lobes, whose resulting 'optic ataxia' causes her to make large pointing errors when asked to locate single light emitting diodes presented in her visual field. We report here that, unlike normal individuals, A. T.'s pointing accuracy improved when she was required to wait for 5 s before responding. This counter-intuitive result is interpreted as reflecting the very brief time-scale on which visuomotor control systems in the superior parietal lobe operate. When an immediate response was required, A. T.'s damaged visuomotor system caused her to make large errors; but when a delay was required, a different, more flexible, visuospatial coding system--presumably relatively intact in her brain--came into play, resulting in much more accurate responses. The data are consistent with a dual processing theory whereby motor responses made directly to visual stimuli are guided by a dedicated system in the superior parietal and premotor cortices, while responses to remembered stimuli depend on perceptual processing and may thus crucially involve processing within the temporal neocortex.

Grasping an object: one movement, several components.

The visuomotor transformations for producing a grasping movement imply simultaneous control of different visual mechanisms. The object size, orientation and 3D characteristics have to be encoded for the selection of the appropriate opposition space, within which the opposition forces will be applied on the object surface. These mechanisms also have to combine with those of the transport of the hand to the object location. Finally, biomechanical constraints impose categorical visuomotor decisions for positioning the opposition space according to object changes in size, orientation and spatial location. This paper examines possible interactions between the specialized structures for visuomotor transformation and the internal model that adapts prehension to its goals.

Does the type of prehension influence the kinematics of reaching?

Kinematic studies have indicated that when a subject reaches to grasp an object, the movement consists of two primary components: (a) a transport phase whereby the hand is brought towards the object and (b) a grip phase whereby the hand changes shape in anticipation of the grasp. Using a visual perturbation paradigm, we investigated the effect of different grip component strategies upon the transport phase. The distal strategy was determined by the size of the object to be grasped: for the small object (1.5 cm o.d.) subjects naturally adopted a precision grip between the index finger and thumb; for the large object (6 cm o.d.) subjects used a whole hand prehensile grip. During 20% of the reaching trials the perturbation was introduced by unexpectedly changing the object size. The results showed that corrections to the distal program in response to the perturbation were preceded by changes in the deceleration phase of the proximal component. The data supported previous findings of two visuo-motor channels for this prehensile movement but indicated that when unanticipated shifts of only the distal program are required, both channels show modifications.

Hand kinematics during reaching and grasping in the macaque monkey.

In this paper, we develop an animal model of prehension movements by examining the kinematics of reaching and grasping in monkeys and by comparing the results to published data on humans. Hand movements were recorded in three dimensions in monkeys who were trained to either point at visual targets under unperturbed and perturbed conditions, or to reach and grasp 3-D objects. The results revealed the following three similarities in the hand kinematics of monkey and man. (1) Pointing movements showed an asymmetry depending on target location relative to the hand used; in particular, movements to an ipsilateral target took longer than those to a contralateral one. (2) Perturbation of target location decreased the magnitude of the velocity peak and increased the duration of pointing movements. (3) Reaching to grasp movements displayed a bell-shaped wrist velocity profile and the maximum grip aperture was correlated with object size. These similarities indicate that the macaque monkey can be a useful model for understanding human motor control.

Temporal dissociation of motor responses and subjective awareness. A study in normal subjects.

The aim of the present study was to examine the timing of different responses given simultaneously to a single event, the sudden displacement of a visual object occurring at the onset of the grasping movement directed at that object. The subjects were requested to correct their movement in order to reach accurately for the object and to signal the time at which they became aware of its displacement by a simple vocal utterance (Tah!). The onset of the motor adjustment was measured using kinematic landmarks obtained from the hand trajectory. Movements executed during trials where the object was displaced had an earlier peak in acceleration (107 ms) than movements executed during control trials (120 ms). By contrast, the vocal signal occurred 420 ms following object displacement, that was more than 300 ms after the onset of the motor correction. Control experiments were performed in order to verify the influence of possible interferences between the two tasks. Motor corrections performed without vocal utterance had the same timing as when the vocal signal was produced. Vocal signals produced in response to object's displacements but in the absence of reaching movements had the same latency as when movements were performed. We conclude from these results that the two responses were generated independently of each other. Assuming that the vocal responses in this experiment did signal the subject's awareness, the observed delay between motor corrections and these responses suggests that neural activity must be processed during a significant and quantifiable amount of time before it can give rise to conscious experience. This dissociation between motor responses and awareness in normal subjects is discussed in the light of clinical cases where overt behaviour and conscious experience are dissociated by cerebral lesions.

Orientation of the opposition axis in mentally simulated grasping.

Five normal subjects were tested in a simulated grasping task. A cylindrical container filled with water was placed on the center of a horizontal monitor screen. Subjects used a precision grip formed by the thumb and index finger of their right hand. After a preliminary run during which the container was present, it was replaced by an image of the upper surface of the cylinder appearing on the horizontal computer screen on which the real cylinder was placed during the preliminary run. In each trial the image was marked with two contact points which defined an opposition axis in various orientations with respect to the frontal plane. The subjects' task consisted, once shown a stimulus, of judging as quickly as possible whether the previously experienced action of grasping the container full of water and pouring the water out would be easy, difficult or impossible with the fingers placed according to the opposition axis indicated on the circle. Response times were found to be longer for the grasps judged to be more difficult due to the orientation and position of the opposition axis. In a control experiment, three subjects actually performed the grasps with different orientations and positions of the opposition axis. The effects of these parameters on response time followed the same trends as during simulated movements. This result shows that simulated hand movements take into account the same biomechanical limitations as actually performed movements.

Influence of object position and size on human prehension movements.

Prehension movements of the right hand were recorded in normal subjects using a computerized motion analyzer. The kinematics and the spatial paths of markers placed at the wrist and at the tips of the index finger and thumb were measured. Cylindrical objects of different diameters (3, 6, 9 cm) were used as targets. They were placed at six different positions in the workspace along a circle centered on subject's head axis. The positions were spaced by 10 degrees starting from 10 degrees on the left of the sagittal axis, up to 40 degrees on the right. Both the transport and the grasp components of prehension were influenced by the distance between the resting hand position and the object position. Movement time, time to peak velocity of the wrist and time to maximum grip aperture varied as a function of distance from the object, irrespective of its size. The variability of the spatial paths of wrist and fingers sharply decreased during the phase of the movement prior to contact with the object. This indicates that the final position of the thumb and the index finger is a controlled parameter of visuomotor transformation during prehension. The orientation of the opposition axis (defined as the line connecting the tips of the thumb and the index finger at the end of the movement) was measured. Several different frames of reference were used. When an object-centered frame was used, the orientation of the opposition axis was found to change by about 10 degrees from one object position to the next. By contrast, when a body-centered frame was used (with the head or the forearm as a reference), this orientation was found to remain relatively invariant for different object positions and sizes. The degree of wrist flexion was little affected by the position of the object. This result, together with the invariant orientation of the opposition axis, shows that prehension movements aimed at cylindrical objects are organized so as to minimize changes in posture of the lower arm.

The coupling of arm and finger movements during prehension.

The experiments reported here were aimed at testing the degree of coupling of motor components during the act of prehension. Hand movements were recorded bidimensionnally by a Selspot system which monitored the displacement of IREDS placed at the thumb and index finger tips, at the metacarpophalangeal joint of the index and at the radial styloid. Targets were three-dimensional translucent dowels placed concentrically at 30 cm from the subject. The dowels were 10 degrees apart from each other. In blocked and control trials, one dowel was illuminated and served as a target for the movement. In the perturbed trials (20% of cases) one dowel was illuminated first and the light was unexpectedly shifted to another dowel at the onset of the subject's movements. Kinematic analysis of the movement revealed the following: 1. In blocked and control trials, the wrist moved with a single acceleration to the target dowel. Meanwhile, the finger grip (computed as the distance between thumb and index IREDS) increased up to a maximum size, located in time at about 60% of movement time and then decreased until contact with the dowel. 2. In perturbed trials the initial wrist acceleration was aborted. A new acceleration started about 180 ms after the first, in order to reorient the hand to the new target. Similarly, the initial grip aperture also aborted and reincreased in synchrony with the second wrist acceleration. 3. Perturbations increased movement time by only 95 ms on average. The first peak in acceleration indicating abortion of the initial movement occurred 100 ms after the movement onset, i.e., 30 ms earlier than in non perturbed trials.(ABSTRACT TRUNCATED AT 250 WORDS)

Selective perturbation of visual input during prehension movements. 2. The effects of changing object size.

1. Subjects were instructed to reach and grasp cylindrical objects, using a precision grip. The objects were two concentric dowels made of translucent material placed at 35 cm from the subject. The inner ("small") dowel was 10 cm high and 1.5 cm in diameter. The outer ("large") dowel was 6 cm high and 6 cm in diameter. Prehension movements were monitored using a Selspot system. The displacement of a marker placed at the wrist level was used as an index for the transport of the hand at the location of the object. Markers placed at the tips of the thumb and the index finger were used for measuring the size of aperture of the finger grip. 2. Kinematics of transport and grasp components were computed from the filtered displacement signals. Movement time (MT), time to peak velocity (TPV) and time to peak deceleration (TPD) of the wrist, time to peak velocity of grip aperture (TGV), time to maximum grip aperture (TGA) were the main parameters used for comparing the movements in different conditions. Spatial paths of the wrist, thumb and index markers were reconstructed in two dimensions. Variability of the spatial paths over repeated trials was computed as the surface of the ellipses defined by X and Y standard deviations from the mean path. 3. Computer controlled illumination of one of the dowels was the signal for reaching toward that dowel. Blocks of trials were made to the small dowel and to the large dowel. Mean MT during blocked trials was 550 ms. The acceleration phase of the movements (measured by parameter TPV) represented 33% of MT. About half of MT (52%) was spent after TPD in a low velocity phase while the hand was approaching the object. This kinematic pattern was not influenced by whether movements were directed at small or large dowels. 4. Grip aperture progressively increased during transport of the hand. TGA corresponded to about 60% of MT, that is, maximum grip aperture was reached during the low velocity phase of transport. Following TGA, fingers closed around the object until contact was made. This pattern of grip formation differed whether the movement was directed at the large or the small dowel: TGA occurred often earlier for the small dowel, and the size of the maximum grip aperture was larger for the large dowel. Variability of both the wrist and finger spatial paths was larger during the first half of MT, and tended to become very low as the hand approached the dowels.(ABSTRACT TRUNCATED AT 400 WORDS)

Selective perturbation of visual input during prehension movements. 1. The effects of changing object position.

Prehension involves processing information in two hypothesized visuomotor channels: one for extrinsic object properties (e.g., the spatial location of objects) and one for intrinsic objects properties (e.g., shape and size). The present study asked how the two motor components that correspond to these channels (transport and grasp, respectively) are related. One way to address this question is to create a situation where unexpected changes occur at the input level of one of the visuomotor channels, and to observe how the movement reorganizes. If transport and grasp are independent components, then changing the object location, for example, should affect only the transport, not the grasp component. Subjects were requested to reach, grasp and lift as accurately as possible one of three dowels using the distal pads of the thumb and index finger. On certain trials, upon movement initiation towards the middle dowel, the dowel was made to instantaneously change its location to one of the two other positions, requiring the subject to reorient the hand to the new dowel location. Results consisted of comparing the movement characteristics of the transport and grasp components of these perturbed movements with appropriate control movements. Kinematics of the wrist trajectory showed fast adjustments, within 100 ms, to the change of dowel position. This duration seems to correspond to the minimum delay required within the visuomotor system for visual and/or proprioceptive reafferents to influence the ongoing movement. In addition, these delays are much shorter than has been found for conditions where object location changes before movement initiation (approximately 300 ms). The faster times may relate to the dynamic character of the deviant limb position signals, with the only constraint being the physiological delays for visual and kinaesthetic signals to influence the movement. A spatiotemporal variability analysis of the movement trajectories for non-perturbed trials showed variability to be greatest during the acceleration part of the movement, interpreted as due to control by a relatively inaccurate directional coding mechanism. Control during the deceleration phase, marked by low trajectory variability, was seen to be due to a sensorimotor process, using motor output signals, and resulting in an optimized trajectory supporting a successful grasp. Analysis of the grasp component of prehension showed that perturbing object location influenced the movement of the fingers suggesting a kinematic coupling of the two components. However, forthcoming work shows that, when object size changes, and location remains constant, there is a clear temporal dissociation of the two components of prehension.(ABSTRACT TRUNCATED AT 400 WORDS)

Acquisition of co-ordination between posture and movement in a bimanual task.

The acquisition of co-ordination between posture and movement was investigated in human subjects performing a load lifting task. Sitting subjects held their left (postural) forearm in a horizontal position while supporting a 1 kg load via an electromagnet. Perturbation of the postural forearm position consisted of the load release triggered either by the experimenter (control) or by the subject voluntarily moving the other arm. In the latter case, the movement involved the elbow joint (load lifting (A), isometric force change at the wrist level (B), elbow rotation (C) and pressing a button with the wrist (D] or the fingers (grip isometric force change). We recorded the maximal amplitude and maximal velocity of the rotation of the postural forearm, the EMG of the forearm flexors on both sides and the force exerted either by the load on the postural arm or by the isometric contraction of the moving arm. The maximal forearm angular velocity after unloading was known to be related to the level of muscle contraction before unloading. 1. In the control situation, repetition of the imposed unloading test resulted in a progressive reduction in the maximal forearm rotation without any decrease in the maximal velocity. The amplitude and duration of the unloading reflex were found to increase in parallel. These results suggest that an adaptive mechanism took place which increased the gain of the unloading reflex loop and reduced the mechanical effect of the perturbation. This mechanism was found to come into play not only in the control situation but also in other paradigms where the perturbation was expected by the subjects. 2. A decrease in both maximal amplitude and velocity of forearm rotation together with a weak "anticipatory" deactivation of the forearm postural flexors was observed when the unloading was caused by an elbow movement (situations A, B, C) which indicates that a feedforward postural control took place. An interlimb coordination was built up and stabilized after 40-60 trials. Pressing a button with the wrist (weak force and displacement) was a less effective means of inducing an anticipatory control of the flexors of the postural forearm, which indicates that the intensity of the central control plays a role in the building up of the coordination. 3. A distal grip action exerting either weak (100 g) or a high (1 kg) force was able to reduce the maximal amplitude of the forearm rotation, but not the maximal velocity, which indicates that an improved reflex action takes place, but not a feedforward anticipatory postural control.(ABSTRACT TRUNCATED AT 400 WORDS)

Integrated control of hand transport and orientation during prehension movements.

At a descriptive level, prehension movements can be partitioned into three components ensuring, respectively, the transport of the arm to the vicinity of the target, the orientation of the hand according to object tilt, and the grasp itself. Several authors have suggested that this analytic description may be an operational principle for the organization of the motor system. This hypothesis, called "visuomotor channels hypothesis," is in particular supported by experiments showing a parallelism between the reach and grasp components of prehension movements. The purpose of the present study was to determine whether or not the generalization of the visuomotor channels hypothesis, from its initial form, restricted to the grasp and transport components, to its actual form, including the reach orientation and grasp components, may be well founded. Six subjects were required to reach and grasp cylindrical objects presented at a given location, with different orientations. During the movements, object orientation was either kept constant (unperturbed trials) or modified at movement onset (perturbed trials). Results showed that both wrist path (sequence of positions that the hand follows in space), and wrist trajectory (time sequence of the successive positions of the hand) were strongly affected by object orientation and by the occurrence of perturbations. These observations suggested strongly that arm transport and hand orientation were neither planned nor controlled independently. The significant linear regressions observed, with respect to the time, between arm displacement (integral of the magnitude of the velocity vector) and forearm rotation also supported this view. Interestingly, hand orientation was not implemented at only the distal level, demonstrating that all the redundant degrees of freedom available were used by the motor system to achieve the task. The final configuration reached by the arm was very stable for a given final orientation of the object to grasp. In particular, when object tilt was suddenly modified at movement onset, the correction brought the upper limb into the same posture as that obtained when the object was initially presented along the final orientation reached after perturbation. Taken together, the results described in the present study suggest that arm transport and hand orientation do not constitute independent visuomotor channels. They also further suggest that prehension movements are programmed, from an initial configuration, to reach smoothly a final posture that corresponds to a given "location and orientation" as a whole.

Postural and synergic control for three-dimensional movements of reaching and grasping.

1. A fundamental question about motor control is related to the nature of the representations used by the nervous system to program the movement. Theoretically, arm displacement can be encoded either in task (extrinsic) or in joint (intrinsic) space. 2. The present study investigated the organization of complex movements consisting of reaching and grasping a cylindrical object presented along different orientations in space. In some trials, object orientation was suddenly modified at movement onset. 3. At a static level, the final limb angles were highly predictable despite the wide range of possible postures allowed by articular redundancy. Moreover, when object orientation was unexpectedly modified at movement onset, the final angular configuration of the limb was identical to that obtained when the object was initially presented along the orientation reached after the perturbation. 4. At a dynamical level, a generalized synergy was observed, and tight correlations were noted between all joint angles implicated in the movement with the exception of elbow flexion. For this joint angle, which did not vary monotonically, strong partial correlations were however observed before and after movement reversal. 5. These results suggest that natural movements are mostly carried out in joint space by postural transitions.