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.     

Changes in the amplitude and direction of goal-directed hand movements in the lack of visual information

The goal of the present investigation was to determine the precision of goal-directed hand movements in the lack of visual information. The movement amplitude and direction was examined under different experimental conditions. Subjects were ten female and ten male university students. The motor test was drawing 10 cm long straight line and 24 cm long zigzag line in four different experimental conditions. 1) The drawing with open eyes was followed immediately with drawing with closed eyes. 2) The drawing was executed from memory in the lack of visual information. 3) Drawing with restricted amplitude or direction. 4) Drawing with verbal feedback. The errors of the target distance and the lateral deviations from the target were different under the different experimental conditions. The largest errors and underestimation of the target distance occurred in drawing horizontal straight line with closed eyes. No statistically significant gender differences were found. It is concluded that the practice, adjustment of single movement parameter to the target, and the verbal feedback assist better the accuracy of unseen goal-directed hand movement than the recent visual memory.     

Disruption of state estimation in the human lateral cerebellum

The cerebellum has been proposed to be a crucial component in the state estimation process that combines information from motor efferent and sensory afferent signals to produce a representation of the current state of the motor system. Such a state estimate of the moving human arm would be expected to be used when the arm is rapidly and skillfully reaching to a target. We now report the effects of transcranial magnetic stimulation (TMS) over the ipsilateral cerebellum as healthy humans were made to interrupt a slow voluntary movement to rapidly reach towards a visually defined target. Errors in the initial direction and in the final finger position of this reach-to-target movement were significantly higher for cerebellar stimulation than they were in control conditions. The average directional errors in the cerebellar TMS condition were consistent with the reaching movements being planned and initiated from an estimated hand position that was 138 ms out of date. We suggest that these results demonstrate that the cerebellum is responsible for estimating the hand position over this time interval and that TMS disrupts this state estimate.     

Motor control of voluntary arm movements

The motor control of pointing and reaching-to-grasp movements was investigated using two different approaches (kinematic and modelling) in order to establish whether the type of control varies according to modifications of arm kinematics. Kinematic analysis of arm movements was performed on subjects' hand trajectories directed to large and small stimuli located at two different distances. The subjects were required either to grasp and to point to each stimulus. The kinematics of the subsequent movement, during which subject's hand came back to the starting position, were also studied. For both movements, kinematic analysis was performed on hand linear trajectories as well as on joint angular trajectories of shoulder and elbow. The second approach consisted in the parametric identification of the black box (ARMAX) model of the controller driving the arm movement. Such controller is hypothesized to work for the correct execution of the motor act. The order of the controller ARMAX model was analyzed with respect to the different experimental conditions (distal task, stimulus size and distance). Results from kinematic analysis showed that target distance and size influenced kinematic parameters both of angular and linear displacements. Nevertheless, the structure of the motor program was found to remain constant with distane and distal task, while it varied with precision requirements due to stimulus size. The estimated model order of the controller confirmed the invariance of the control law with regard to movement amplitude, whereas it was sensitive to target size.     

Concatenation of Observed Grasp Phases with Observer’s Distal Movements: A Behavioural and TMS Study

The present study aimed at determining how actions executed by two conspecifics can be coordinated with each other, or more specifically, how the observation of different phases of a reaching-grasping action is temporary related to the execution of a movement of the observer. Participants observed postures of initial finger opening, maximal finger aperture, and final finger closing of grasp after observation of an initial hand posture. Then, they opened or closed their right thumb and index finger (experiments 1, 2 and 3). Response times decreased, whereas acceleration and velocity of actual finger movements increased when observing the two late phases of grasp. In addition, the results ruled out the possibility that this effect was due to salience of the visual stimulus when the hand was close to the target and confirmed an effect of even hand postures in addition to hand apparent motion due to the succession of initial hand posture and grasp phase. In experiments 4 and 5, the observation of grasp phases modulated even foot movements and pronunciation of syllables. Finally, in experiment 6, transcranial magnetic stimulation applied to primary motor cortex 300 ms post-stimulus induced an increase in hand motor evoked potentials of opponens pollicis muscle when observing the two late phases of grasp. These data suggest that the observation of grasp phases induced simulation which was stronger during observation of finger closing. This produced shorter response times, greater acceleration and velocity of the successive movement. In general, our data suggest best concatenation between two movements (one observed and the other executed) when the observed (and simulated) movement was to be accomplished. The mechanism joining the observation of a conspecific’s action with our own movement may be precursor of social functions. It may be at the basis for interactions between conspecifics, and related to communication between individuals.     

Measuring 3D Hand and Finger Kinematics—A Comparison between Inertial Sensing and an Opto-Electronic Marker System

Objective analysis of hand and finger kinematics is important to increase understanding of hand function and to quantify motor symptoms for clinical diagnosis. The aim of this paper is to compare a new 3D measurement system containing multiple miniature inertial sensors (PowerGlove) with an opto-electronic marker system during specific finger tasks in three healthy subjects. Various finger movements tasks were performed: flexion, fast flexion, tapping, hand open/closing, ab/adduction and circular pointing. 3D joint angles of the index finger joints and position of the thumb and index were compared between systems. Median root mean square differences of the main joint angles of interest ranged between 3.3 and 8.4deg. Largest differences were found in fast and circular pointing tasks, mainly in range of motion. Smallest differences for all 3D joint angles were observed in the flexion tasks. For fast finger tapping, the thumb/index amplitude showed a median difference of 15.8mm. Differences could be explained by skin movement artifacts caused by relative marker movements of the marker system, particularly during fast tasks; large movement accelerations and angular velocities which exceeded the range of the inertial sensors; and by differences in segment calibrations between systems. The PowerGlove is a system that can be of value to measure 3D hand and finger kinematics and positions in an ambulatory setting. The reported differences need to be taken into account when applying the system in studies understanding the hand function and quantifying hand motor symptoms in clinical practice.     

Structural basis of compound recognition by adenosine deaminase

Structural snapshots corresponding to various states enable elucidation of the molecular recognition mechanism of enzymes. Adenosine deaminase has two distinct conformations, an open form and a closed form, although it has so far been unclear what factors influence adaptation of the alternative conformations. Herein, we have determined the first nonligated structure as an initial state, which was the open form, and have thereby rationally deduced the molecular recognition mechanism. Inspection of the active site in the nonligated and ligated states indicated that occupancy at one of the water-binding positions in the nonligated state was highly significant in determining alternate conformations. When this position is empty, subsequent movement of Phe65 toward the space induces the closed form. On the other hand, while occupied, the overall conformation remains in the open form. This structural understanding should greatly assist structure-oriented drug design and enable control of the enzymatic activity.     

Motor Activity Improves Temporal Expectancy

Certain brain areas involved in interval timing are also important in motor activity. This raises the possibility that motor activity might influence interval timing. To test this hypothesis, we assessed interval timing in healthy adults following different types of training. The pre- and post-training tasks consisted of a button press in response to the presentation of a rhythmic visual stimulus. Alterations in temporal expectancy were evaluated by measuring response times. Training consisted of responding to the visual presentation of regularly appearing stimuli by either: (1) pointing with a whole-body movement, (2) pointing only with the arm, (3) imagining pointing with a whole-body movement, (4) simply watching the stimulus presentation, (5) pointing with a whole-body movement in response to a target that appeared at irregular intervals (6) reading a newspaper. Participants performing a motor activity in response to the regular target showed significant improvements in judgment times compared to individuals with no associated motor activity. Individuals who only imagined pointing with a whole-body movement also showed significant improvements. No improvements were observed in the group that trained with a motor response to an irregular stimulus, hence eliminating the explanation that the improved temporal expectations of the other motor training groups was purely due to an improved motor capacity to press the response button. All groups performed a secondary task equally well, hence indicating that our results could not simply be attributed to differences in attention between the groups. Our results show that motor activity, even when it does not play a causal or corrective role, can lead to improved interval timing judgments.     

Distinct motor plans form and retrieve distinct motor memories for physically identical movements

We can adapt movements to a novel dynamic environment (e.g., tool use, microgravity, and perturbation) by acquiring an internal model of the dynamics. Although multiple environments can be learned simultaneously if each environment is experienced with different limb movement kinematics, it is controversial as to whether multiple internal models for a particular movement can be learned and flexibly retrieved according to behavioral contexts. Here, we address this issue by using a novel visuomotor task. While participants reached to each of two targets located at a clockwise or counter-clockwise position, a gradually increasing visual rotation was applied in the clockwise or counter-clockwise direction, respectively, to the on-screen cursor representing the unseen hand position. This procedure implicitly led participants to perform physically identical pointing movements irrespective of their intentions (i.e., movement plans) to move their hand toward two distinct visual targets. Surprisingly, if each identical movement was executed according to a distinct movement plan, participants could readily adapt these movements to two opposing force fields simultaneously. The results demonstrate that multiple motor memories can be learned and flexibly retrieved, even for physically identical movements, according to distinct motor plans in a visual space.     

Benefit of Bi-Ocular Visual Stimulation for Postural Control in Children with Strabismus

Vision is important for postural control as is shown by the Romberg quotient (RQ): with eyes closed, postural instability increases relative to eyes open (RQ = 2). Yet while fixating at far distance, postural stability is similar with eyes open and eyes closed (RQ = 1). Postural stability can be better with both eyes viewing than one eye, but such effect is not consistent among healthy subjects. The first goal of the study is to test the RQ as a function of distance for children with convergent versus divergent strabismus. The second goal is to test whether vision from two eyes relative to vision from one eye provides better postural stability. Thirteen children with divergent strabismus and eleven with convergent strabismus participated in this study. Posturtography was done with the Techno concept device. Experiment 1, four conditions: fixation at 40 cm and at 200 cm both with eyes open and eyes covered (evaluation of RQ). Experiment 2, six conditions: fixation at 40 cm and at 200 cm, with both eyes viewing or under monocular vision (dominant and non-dominant eye). For convergent strabismus, the groups mean value of RQ was 1.3 at near and 0.94 at far distance; for divergent, it was 1.06 at near and 1.68 at far. For all children, the surface of body sway was significantly smaller under both eyes viewing than monocular viewing (either eye). Increased RQ value at near for convergent and at far for divergent strabismus is attributed to the influence of the default strabismus angle and to better use of ocular motor signals. Vision with the two eyes improves postural control for both viewing distances and for both types of strabismus. Such benefit can be due to complementary mechanisms: larger visual field, better quality of fixation and vergence angle due to the use of visual inputs from both eyes.     

Manuo-ocular coordination in target tracking. II. Comparing the model with human behavior

Several studies have shown that humans track a moving visual target with their eyes better if the movement of this target is directly controlled by the observer's hand. The improvement in performance has been attributed to coordination control between the arm motor system and the smooth pursuit (SP) system. In such a task, the SP system shows characteristics that differ from those observed during eye-alone tracking: latency (between the target-arm and the eye motion onsets) is shorter, maximum SP velocity is higher and the maximum target motion frequency at which the SP can function effectively is also higher. The aim of this article is to qualitatively evaluate the behavior of a dynamical model simulating the oculomotor system and the arm motor system when both are involved in tracking visual targets. The evaluation is essentially based on a comparison of the behavior of the model with the behavior of human subjects tracking visual targets under different conditions. The model has been introduced and quantitatively evaluated in a companion paper. The model is based on an exchange of internal information between the two sensorimotor systems, mediated by sensory signals (vision, arm muscle proprioception) and motor signals (arm motor command copy). The exchange is achieved by a specialized structure of the central nervous system, previously identified as a part of the cerebellum. Computer simulation of the model yielded results that fit the behavior of human subjects observed during previously reported experiments, both qualitatively and quantitatively. The parallelism between physiology and human behavior on the one hand, and structure and simulation of the model on the other hand, is discussed. Received: 6 March 1997 / Accepted in revised form: 15 July 1997     

Object Grasping using the Minimum Variance Model

Reaching-to-grasp has generally been classified as the coordination of two separate visuomotor processes: transporting the hand to the target object and performing the grip. An alternative view has recently been formed that grasping can be explained as pointing movements performed by the digits of the hand to target positions on the object. We have previously implemented the minimum variance model of human movement as an optimal control scheme suitable for control of a robot arm reaching to a target. Here, we extend that scheme to perform grasping movements with a hand and arm model. Since the minimum variance model requires that signal-dependent noise be present on the motor commands to the actuators of the movement, our approach is to plan the reach and the grasp separately, in line with the classical view, but using the same computational model for pointing, in line with the alternative view. We show that our model successfully captures some of the key characteristics of human grasping movements, including the observations that maximum grip size increases with object size (with a slope of approximately 0.8) and that this maximum grip occurs at 60–80% of the movement time. We then use our model to analyse contributions to the digit end-point variance from the two components of the grasp (the transport and the grip). We also briefly discuss further areas of investigation that are prompted by our model.     

Fast corrections of movements with a computer mouse

When we reach out for an object with our hand, we transform visual information about the object's position into muscle contractions that will bring our digits to that position. If we reach out with a tool the transformation is different, because the muscle contractions must bring the critical part of the tool to the object, rather than the digits. The difference between the motion of the hand and that of the tool can be quite large, as when moving a computer mouse across a table to bring a cursor to a position on a screen. We examined the responses to unpredictable visual perturbations during such movements. People responded about as quickly to changes in the position of the target when pointing with the mouse as when doing so with their hand. They also responded about as quickly when the cursor was displaced as when the target was displaced. We show that this is not because the visually perceived separation between target and cursor is transformed into a desired displacement of the hand. Our conclusion is that our actions are controlled by the judged positions of the end-effector and the target, even when the former is quite detached from the muscles and joints that are involved in the action.     

The Role of the Caudal Superior Parietal Lobule in Updating Hand Location in Peripheral Vision: Further Evidence from Optic Ataxia

Patients with optic ataxia (OA), who are missing the caudal portion of their superior parietal lobule (SPL), have difficulty performing visually-guided reaches towards extra-foveal targets. Such gaze and hand decoupling also occurs in commonly performed non-standard visuomotor transformations such as the use of a computer mouse. In this study, we test two unilateral OA patients in conditions of 1) a change in the physical location of the visual stimulus relative to the plane of the limb movement, 2) a cue that signals a required limb movement 180° opposite to the cued visual target location, or 3) both of these situations combined. In these non-standard visuomotor transformations, the OA deficit is not observed as the well-documented field-dependent misreach. Instead, OA patients make additional eye movements to update hand and goal location during motor execution in order to complete these slow movements. Overall, the OA patients struggled when having to guide centrifugal movements in peripheral vision, even when they were instructed from visual stimuli that could be foveated. We propose that an intact caudal SPL is crucial for any visuomotor control that involves updating ongoing hand location in space without foveating it, i.e. from peripheral vision, proprioceptive or predictive information.     

Effects of head position on errors in 3-D pointing to remembered locations

The accuracy of pointing movements performed under different head positions to remembered target locations in 3-D space was studied in healthy persons. The subjects fixated a visual target, then closed their eyes and after 1.0 sec performed the targeted movement with their right arm. The target (a point light source) was presented in random order by a programmable robot arm at one of five space locations. The accuracy of pointing movements was examined in a spherical coordinate system centered in respect with the shoulder of the responding arm. The pointing movements were most accurate under natural eye-head coordination. With the head fixed in the straight-ahead position, both the 3-D absolute error and its standard deviation increased significantly. At the same time, individual components of spatial error (directional and radial) did not change significantly. With the head turned to the rightmost or leftmost position, the pointing accuracy was disturbed within larger limits than under head-fixed condition. The main contributors to the 3-D absolute error were the changes in the azimuth error. The latter depended on the direction of the head-turn: the rightmost turn either increased leftward or decreased rightward shift, and conversely, the left turn increased rightward shift or decreased leftward shift of the target-directed movements.It is suggested that the increased inaccuracy of pointing under head-fixed condition reflected the impairment of the eye-head coordination underlying gaze orientation, and increased inaccuracy under the head-turned condition may be explained by changes in the internal representation of the head and target position in space.Neirofiziologiya/Neurophysiology, Vol. 26, No. 2, pp. 122–131, March–April, 1994.     

Modification of the strategy of intentional movement after lesions of the substantia nigra in the cat

Bilateral lesions of the substantia nigra were carried out in cats previously overtrained at performing a visually guided forepaw movement towards a moving target. Both their reaction time (RT) and movement time (MT) were impaired postoperatively. On the other hand, their pointing precision was unimpaired after lesion and even improved relative to the preoperative level. This improved accuracy persisted even when the duration of the movement had returned to normal. It is suggested that lesioned animals develop and keep a new strategy using visual feed-back throughout the movement, while normal cats, even on such a complex pointing task, mainly use visual information to trigger their movement.     

Structure of the Plasmodium falciparum triosephosphate isomerase-phosphoglycolate complex in two crystal forms: characterization of catalytic loop open and closed conformations in the ligand-bound state

Triosephosphate isomerase (TIM) has been the subject of many structural and mechanistic studies. At position 96, there is a highly conserved Ser residue, which is proximal to the catalytic site. Thus far, no specific role has been ascribed to this residue. Plasmodium falciparum TIM (PfTIM), a fully catalytically active enzyme, is unique in possessing a Phe residue at position 96. The structure of PfTIM complexed to phosphoglycolate (PG), a transition state analogue, has been determined in an effort to probe the effects of the mutation at residue 96 on the nature of inhibitor-enzyme interactions and the orientation of the critical catalytic loop (loop 6, residues 166-176) in TIM. Crystal structures of PfTIM complexed to phosphoglycolate in orthorhombic (P2(1)2(1)2(1)) and monoclinic (C2) forms were determined and refined at resolutions of 2.8 and 1.9 A, respectively. The P2(1)2(1)2(1) form contains two dimers in the asymmetric unit. In the C2 form, the molecular and crystal 2-fold axes are coincident, leading to a monomer in the asymmetric unit. The catalytic loop adopts the open state in the P2(1)2(1)2(1) form and the closed conformation in the C2 crystal. The open conformation of the loop in the P2(1)2(1)2(1) form appears to be a consequence of the Ser to Phe mutation at residue 96. The steric clash between Phe96 and Ile172 probably impedes loop closure in PfTIM-ligand complexes. The PfTIM-PG complex is the first example of a TIM-ligand complex being observed in both loop open and closed forms. In the C2 form (loop closed), Phe96 and Leu167 adopt alternative conformations that are different from the ones observed in the open form, permitting loop closure. These structures provide strong support for the view that loop closure is not essential for ligand binding and that dynamic loop movement may occur in both free and ligand-bound forms of the enzyme.     

Influence of adaptation to horizontal body position on pointing errors and visual estimation of a target position in humans

The central program of a targeted movement includes a component intended for to compensate for the weight of the arm; this is why the accuracy of pointing to a memorized position of the visual target in darkness depends on orientation of the moving limb in relation to the vertical axis. Transition from the vertical to the horizontal body position is accompanied by a shift of the final hand position along the body axis towards the head. We studied how pointing errors and visual localization of the target are modified due to adaptation to the horizontal body position; targeted movements to a real target were repeatedly performed during the adaptation period. Three types of experiments were performed: a basic experiment, and two different experiments with adaptation realized under somewhat dissimilar conditions. In the course of the first adaptation experiment, subjects received no visual information on the hand’s position in space, and targeted movements of the arm to a luminous target could be corrected using proprioceptive information only. With such a paradigm, the accuracy of pointing to memorized visual targets showed no adaptation-related changes. In the second adaptation experiment, subjects were allowed to continuously view a marker (a light-emitting diode taped to the fingertip). After such adaptation practice, the accuracy of pointing movements to memorized targets increased: both constant and variational errors, as well as both components of constant error (i.e.,X andY errors) significantly dropped. Testing the accuracy of visual localization of the targets by visual/verbal adjustment, performed after this adaptation experiment, showed that the pattern of errors did not change compared with that in the basic experiment. Therefore, we can conclude that sensorimotor adaptation to the horizontal position develops much more successfully when the subject obtains visual information about the working point position; such adaptation is not related to modifications in the system of visual localization of the target.     

Motor commands for fast point-to-point arm movements are customized for small changes in inertial load

For repeated point-to-point arm movements it is often assumed that motor commands are customized in a trial-to-trial manner, based on previous endpoint error. To test this assumption, we perturbed movement execution without affecting the endpoint error by using a modest manipulation of inertia. Participants made point-to-point elbow flexion and extension movements in the horizontal plane, under the instruction to move as fast as possible from one target area to another. In selected trials the moment of inertia of the lower arm was increased or decreased by 25%. First, we found that an unexpected increase or decrease of inertia did not affect the open loop controlled part of the movement path (and thus endpoint error was not affected). Second, we found that when the increased or decreased inertia was presented repeatedly, after 5-11 trials motor commands were customized: the first 100ms of agonistic muscle activity in the smoothed and rectified electromyographic signal of agonistic muscles was higher for the high inertia compared to the low inertia. We conclude that endpoint error is not the only parameter that is used to evaluate if motor commands lead to movements as planned.     

Oculocentric frames of reference for limb movement

We have reviewed evidence that suggests that the target for limb motion is encoded in a retinocentric frame of reference. Errors in pointing that are elicited by an illusion that distorts the perceived motion of a target are strongly correlated with errors in gaze position. The modulations in the direction and speed of ocular smooth pursuit and of the hand show remarkable similarities, even though the inertia of the arm is much larger than that of the eye. We have suggested that ocular motion is constrained so that gaze provides an appropriate target signal for the hand. Finally, ocular and manual tracking deficits in patients with cerebellar ataxia are very similar. These deficits are also consistent with the idea that a gaze signal provides the target for hand motion; in some cases limb ataxia would be a consequence of optic ataxia rather than reflecting a deficit in the control of limb motion per se. These results, as well as neurophysiological data summarized here, have led us to revise a hypothesis we have previously put forth to account for the initial stages of sensorimotor transformations underlying targeted limb motions. In the original hypothesis, target location and initial arm posture were ultimately encoded in a common frame of reference tied to somatosensation, i.e. a body-centered frame of reference, and a desired change in posture was derived from the difference between the two. In our new scheme, a movement vector is derived from the difference between variables encoded in a retinocentric frame of reference. Accordingly, gaze, with its exquisite ability to stabilize a target image even under dynamic conditions, would be used as a reference signal. Consequently, this scheme would facilitate the processing of information under conditions in which the body and the target are moving relative to each other.