The effect of a moving distractor on the initiation of smooth-pursuit eye movements

As a step toward understanding the mechanism by which targets are selected for smooth-pursuit eye movements, we examined the behavior of the pursuit system when monkeys were presented with two discrete moving visual targets. Two rhesus monkeys were trained to select a small moving target identified by its color in the presence of a moving distractor of another color. Smooth-pursuit eye movements were quantified in terms of the latency of the eye movement and the initial eye acceleration profile. We have previously shown that the latency of smooth pursuit, which is normally around 100 ms, can be extended to 150 ms or shortened to 85 ms depending on whether there is a distractor moving in the opposite or same direction, respectively, relative to the direction of the target. We have now measured this effect for a 360 deg range of distractor directions, and distractor speeds of 5-45 deg/s. We have also examined the effect of varying the spatial separation and temporal asynchrony between target and distractor. The results indicate that the effect of the distractor on the latency of pursuit depends on its direction of motion, and its spatial and temporal proximity to the target, but depends very little on the speed of the distractor. Furthermore, under the conditions of these experiments, the direction of the eye movement that is emitted in response to two competing moving stimuli is not a vectorial combination of the stimulus motions, but is solely determined by the direction of the target. The results are consistent with a competitive model for smooth-pursuit target selection and suggest that the competition takes place at a stage of the pursuit pathway that is between visual-motion processing and motor-response preparation.     

Temporal properties of visual motion signals for the initiation of smooth pursuit eye movements in monkeys

1. Our goal was to assess whether visual motion signals related to changes in image velocity contribute to pursuit eye movements. We recorded the smooth eye movements evoked by ramp target motion at constant speed. In two different kinds of stimuli, the onset of target motion provided either an abrupt, step change in target velocity or a smooth target acceleration that lasted 125 ms followed by prolonged target motion at constant velocity. We measured the eye acceleration in the first 100 ms of pursuit. Because of the 100-ms latency from the onset of visual stimuli to the onset of smooth eye movement, the eye acceleration in this 100-ms interval provides an estimate of the open-loop response of the visuomotor pathways that drive pursuit. 2. For steps of target velocity, eye acceleration in the first 100 ms of pursuit depended on the "motion onset delay," defined as the interval between the appearance of the target and the onset of motion. If the motion onset delay was > 100 ms, then the initial eye movement consisted of separable early and late phases of eye acceleration. The early phase dominated eye acceleration in the interval from 0 to 40 ms after pursuit onset and was relatively insensitive to image speed. The late phase dominated eye acceleration in the interval 40-100 ms after the onset of pursuit and had an amplitude that was proportional to image speed. If there was no delay between the appearance of the target and the onset of its motion, then the early component was not seen, and eye acceleration was related to target speed throughout the first 100 ms of pursuit. 3. For step changes of target velocity, the relationship between eye acceleration in the first 40 ms of pursuit and target velocity saturated at target speeds > 10 degrees /s. In contrast, the relationship was nearly linear when eye acceleration was measured in the interval 40-100 ms after the onset of pursuit. We suggest that the first 40 ms of pursuit are driven by a transient visual motion input that is related to the onset of target motion (motion onset transient component) and that the next 60 ms are driven by a sustained visual motion input (image velocity component). 4. When the target accelerated smoothly for 125 ms before moving at constant speed, the initiation of pursuit resembled that evoked by steps of target velocity. However, the latency of pursuit was consistently longer for smooth target accelerations than for steps of target velocity.(ABSTRACT TRUNCATED AT 400 WORDS)     

Postsaccadic enhancement of initiation of smooth pursuit eye movements in monkeys

Step-ramp target motion evokes a characteristic sequence of presaccadic smooth eye movement in the direction of the target ramp, catch-up targets to bring eye position close to the position of the moving target, and postsaccadic eye velocities that nearly match target velocity. I have analyzed this sequence of eye movements in monkeys to reveal a strong postsaccadic enhancement of pursuit eye velocity and to document the conditions that lead to that enhancement. Smooth eye velocity was measured in the last 10 ms before and the first 10 ms after the first saccade evoked by step-ramp target motion. Plots of eye velocity as a function of time after the onset of the target ramp revealed that eye velocity at a given time was much higher if measured after versus before the saccade. Postsaccadic enhancement of pursuit was recorded consistently when the target stepped 3 degrees eccentric on the horizontal axis and moved upward, downward, or away from the position of fixation. To determine whether postsaccadic enhancement of pursuit was invoked by smear of the visual scene during a saccade, I recorded the effect of simulated saccades on the presaccadic eye velocity for step-ramp target motion. The 3 degrees simulated saccade, which consisted of motion of a textured background at 150 degrees/s for 20 ms, failed to cause any enhancement of presaccadic eye velocity. By using a strategically selected set of oblique target steps with horizontal ramp target motion, I found clear enhancement for saccades in all directions, even those that were orthogonal to target motion. When the size of the target step was varied by up to 15 degrees along the horizontal meridian, postsaccadic eye velocity did not depend strongly either on the initial target position or on whether the target moved toward or away from the position of fixation. In contrast, earlier studies and data in this paper show that presaccadic eye velocity is much stronger when the target is close to the center of the visual field and when the target moves toward versus away from the position of fixation. I suggest that postsaccadic enhancement of pursuit reflects activation, by saccades, of a switch that regulates the strength of transmission through the visual-motor pathways for pursuit. Targets can cause strong visual motion signals but still evoke low presaccadic eye velocities if they are ineffective at activating the pursuit system.     

Initiation of disjunctive smooth pursuit in monkeys: evidence that Hering's law of equal innervation is not obeyed by the smooth pursuit system

Monkeys generated disjunctive smooth pursuit eye movements when they tracked visual targets that moved toward or away from them. Eye acceleration was computed during the initial 100 msec of pursuit (the open-loop interval) for various target trajectories. The initial acceleration of either eye was a function of the target's motion with respect to that eye, regardless of whether or not the pursuit was conjugate or disjunctive, or performed with one eye occluded. Eye movements produced by fusional vergence could be separated temporally from eye movements produced by smooth pursuit using step-ramp paradigms. The separation of the two responses demonstrates that the fusional vergence system operates in parallel with the smooth pursuit system, presumably to minimize disparity, but not to generate disjunctive components of smooth pursuit eye movements.     

The oculomanual coordination control center takes into account the mechanical properties of the arm

When the eyes and arm are involved in a tracking task, the characteristics of each system differ from those observed when they act alone: smooth pursuit (SP) latency decreases from 130 ms in external target tracking tasks to 0 ms in self-moved target tracking tasks. Two models have been proposed to explain this coordination. The common command model suggests that the same command be addressed to the two sensorimotor systems, which are otherwise organized in parallel, while the coordination control model proposes that coordination is due to a mutual exchange of information between the motor systems. In both cases, the interaction should take into account the dynamic differences between the two systems. However, the nature of the adaptation depends on the model. During self-moved target tracking a perturbation was applied to the arm through the use of an electromagnetic brake. A randomized perturbation of the arm increased the arm motor reaction time without affecting SP. In contrast, a constant perturbation produced an adaptation of the coordination control characterized by a decrease in arm latency and an increase in SP latency relative to motor command. This brought the arm-to-SP latency back to 0 ms. These results support the coordination control model.     

Delayed retinal feedback of eye movements: A dynamic basis of perceptual disabilities.

Investigated temporal factors in vision in relation to the delay of the retinal feedback of ocular movements in eye tracking. A hybrid real-time computer system and dynamic programing methods were used to calibrate photoelectric eye-movement transducers in viewing visual targets, to yoke these targets to eye motion, to introduce feedback delays in eye-movement-retinal interaction, and to measure error in eye tracking. Results indicate that feedback delay affected the accuracy of both the compensatory and the pursuit tracking in a significant way, with a somewhat greater effect being found for pursuit movements. Since delay reduced smooth pursuit motions to saccadic reactions that varied in size with the delay interval, it is suggested that ocular dynamics and guidance in space perception are governed by time-specific neuron mechanisms of the central visual system. Findings negate classical theory of ocular dynamics and perception of direction by proving that directional guidance of the eyes is determined by directional specificity and temporal specificity of the feedback processes of pursuit and saccadic movements of the eyes and is not caused primarily by learned temporal association between visual and tactual sensory processes. It is concluded that major disabilities and distortions in vision, which are not reducible to traditionally defined optometric and ophthalmologic factors, may be produced by built-in developmental perturbations of ocular feedback timing. Findings emphasize dynamic optometric measurements in understanding common and elusive distortions of visual perception. (19 ref.) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Fixation cells in monkey superior colliculus. I. Characteristics of cell discharge

1. We studied the role of the superior colliculus (SC) in the control of visual fixation by recording from cells in the rostral pole of the SC in awake monkeys that were trained to perform fixation and saccade tasks. 2. We identified a subset of neurons in three monkeys that we refer to as fixation cells. These cells increased their tonic discharge rate when the monkey actively fixated a visible target spot to obtain a reward. This sustained activity persisted when the visual stimulation of the target spot was momentarily removed but the monkey was required to continue fixation. 3. The fixation cells were in the rostral pole of the SC. As the electrode descended through the SC, we encountered visual cells with foveal and parafoveal receptive fields most superficially, saccade-related burst cells with parafoveal movement fields below these visual cells, and fixation cells below the burst cells. From this sequence in depth, the fixation cells appeared to be centered in the deeper reaches of the intermediate layers, and this was confirmed by small marking lesions identified histologically. 4. During saccades, the tonically active fixation cells showed a pause in their rate of discharge. The duration of this pause was correlated to the duration of the saccade. Many cells did not decrease their discharge rate for small-amplitude contraversive saccades. 5. The saccade-related pause in fixation cell discharge always began before the onset of the saccade. The mean time from pause onset to saccade onset for contraversive saccades and ipsiversive saccades was 36.2 and 33.0 ms, respectively. Most fixation cells were reactivated before the end of contraversive saccades. The mean time from saccade terminatioN to pause end was -2.6 ms for contraversive saccades and 9.9 ms for ipsiversive saccades. The end of the saccade-related pause in fixation cell discharge was more tightly correlated to saccade termination, than pause onset was to saccade onset. 6. After the saccade-related pause in discharge, many fixation cells showed an increased discharge rate exceeding that before the pause. This increased postsaccadic discharge rate persisted for several hundred milliseconds. 7. The discharge rate of fixation cells was not consistently altered when the monkey actively fixated targets requiring different orbital positions. 8. Fixation cells discharged during smooth pursuit eye movements as they did during fixation. They maintained a steady tonic discharge during pursuit at different speeds and in different directions, provided the monkey looked at the moving target.(ABSTRACT TRUNCATED AT 400 WORDS)     

Neuronal responses in visual areas MT and MST during smooth pursuit target selection

We recorded the activity of single neurons in the middle temporal (MT) and middle superior temporal (MST) visual areas in two macaque monkeys while the animals performed a smooth pursuit target selection task. The monkeys were presented with two moving stimuli of different colors and were trained to initiate smooth pursuit to the stimulus that matched the color of a previously given cue. We designed these experiments so that we could separate the component of the neuronal response that was driven by the visual stimulus from an extraretinal component that predicted the color or direction of the selected target. We found that for all cells in MT and MST the response was primarily determined by the visual stimulus. However, 14% (8 of 58) of MT neurons and 26% (22 of 84) of MST neurons had a small predictive component that was significant at the P < or = 0.05 level. In some cells, the predictive component was clearly related to the color of the intended target, but more often it was correlated with the direction of the target. We have previously documented a systematic shift in the latency of smooth pursuit that depends on the relative direction of motion of the two stimuli. We found that neither the latency nor the amplitude of neuronal responses in MT or MST was correlated with behavioral latency. These results are consistent with a model for target selection in which a weak selection bias for the intended target is amplified by a competitive network that suppresses motion signals related to the nonintended stimulus. It is possible that the predictive component of neuronal responses in MT and MST contributes to the selection bias. However, the strength of the selection bias in MT and MST is not sufficient to account for the high degree of selectivity shown by pursuit behavior.     

Effects of aging on planning and implementing arm movements.

In Exps 1 and 2, aiming movements were performed with and without visual feedback in young and elderly adults. The initial (acceleration and deceleration phases) and secondary movement components were analyzed. Although deceleration phase accuracy decreased without visual feedback in both age groups, accuracy diminished as movement amplitude increased only in the elderly. This suggested that the elderly were more dependent on visual feedback to modify motor programs for longer duration movements. Velocity also increased less with increasing amplitude and target size in the elderly, which was related to impaired preprogramming (acceleration phase) and implementation (deceleration phase) of higher forces. This conclusion was confirmed directly in Exp 2 because only the deceleration phase was affected by the removal of a visual feedback of arm position when availability of visual information could not be predicted before movement. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

The combination of interaural information across frequencies: the effects of number and spacing of components, onset asynchrony, and harmonicity

Threshold interaural delays were measured for a single interaurally delayed low-frequency target component presented against a background of two, four, six, or eight diotic "distractor" components. In the first experiment, a 753-Hz target and the flanking distractor components were gated on and off simultaneously. In subsequent experiments, the distractors were gated on 25-200 ms prior to the target. In addition, the target and distractor components were given various harmonic configurations. In general, threshold interaural delays were higher in all conditions in which distractors were present relative to thresholds obtained for the target component in isolation. Subjects reported that the pitch of the target component was more salient when an onset asynchrony between the target and distractors was present, but the components were perceived as occupying a single intracranial position in spite of the various interaural delays across the frequency domain. These results suggest that binaural processing of stimuli consisting of a small number of low-frequency temporally overlapping components occurs in a spectrally synthetic manner in which interaural information is combined across the spectrum, even in situations in which the segregation of pitch information occurs.     

Centripetal force draws the eyes, not memory of the target, toward the center.

Many observers believe that a target will continue on a curved trajectory after exiting a spiral tube. Similarly, when observers were asked to localize the final position of a target moving on a circular orbit, displacement of the judged position in the direction of forward motion ("representational momentum") and toward the center of the orbit was observed (cf. T. L. Hubbard, 1996). The present study shows that memory displacement of targets on a circular orbit is affected by eye movements. Forward displacement was larger with ocular pursuit of the target, whereas inward displacement was larger with motionless eyes. The results challenge an account attributing forward and inward displacement to mental analogues of momentum and centripetal force, respectively. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Effects of feedback delay on eye-hand synchronism in steering behavior.

Used real-time computer methods of controlling feedback factors in eye tracking to compare accuracy in tracking environmentally-generated and hand-generated visual targets in steering behavior of 5 undergraduates. Feedback delays of .1 and .2 sec. between hand and target movement produced a time lag of eye motion with respect to the hand-produced target action. Results confirm the assumption that steering and stimulus tracking represent different modes of response and are subject to different conditions of delay and displacement of action feedback of body movements. The main effect of feedback delays on eye tracking in steering was to restrict the normal capability of the eye to predict the course of self-generated stimulus movements by reducing the interval of time between hand action and eye response beyond the magnitude of the actual delay interval. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Spatio-temporal firing patterns in the frontal cortex of behaving monkeys

Recording the activity of several neurons in parallel in the frontal cortex of behaving monkeys reveals that firing times of neurons can maintain +/- 1 ms accuracy even after delays of over 400 ms. The accurate firing structures were associated with behavior. Neural networks that can sustain such accuracy can learn 'learn' to bind with each other and thus may serve as building blocks for cognitive processes.     

Asymmetrical hemispheric control of visual-spatial attention in adults with attention deficit hyperactivity disorder.

As neuropsychological mechanisms for attention have been hypothesized to be located in the right hemisphere of the brain, several investigators have begun to conceptualize attention deficit hyperactivity disorder (ADHD)-related attentional deficits as involving right-hemispheric abnormalities. The authors evaluated and compared adult patients diagnosed with ADHD with a non-ADHD group of patients using a chronometric visual-spatial attention task that is sensitive to hemispheric differences in efficiency of information processing. Reaction times across different cuing conditions, cue-target delays, and visual fields were assessed. When participants' attention was misdirected with cues in the right visual field and attention had to be switched to a target on the left visual field, there was a longer delay among ADHD adults than non-ADHD adults, specifically when the interval between the cue and target was 800 ms as compared with 100 ms. This specific pattern of dysfunction was interpreted as a difficulty with maintaining attention possibly associated with anterior attention mechanisms in the right hemisphere. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Directional organization of eye movement and visual signals in the floccular lobe of the monkey cerebellum

The floccular lobe of the monkey is critical for the generation of visually-guided smooth eye movements. The present experiments reveal physiological correlates of the directional organization in the primate floccular lobe by examining the selectivity for direction of eye motion and visual stimulation in the firing of individual Purkinje cells (PCs) and mossy fibers. During tracking of sinusoidal target motion along different axes in the frontoparallel plane, PCs fell into two classes based on the axis that caused the largest modulation of simple-spike firing rate. For "horizontal" PCs, the response was maximal during horizontal eye movements, with increases in firing rate during pursuit toward the side of recording (ipsiversive). For "vertical" PCs, the response was maximal during eye movement along an axis just off pure vertical, with increases in firing rate during pursuit directed downward and slightly contraversive. During pursuit of target motion at constant velocity, PCs again fell into horizontal and vertical classes that matched the results from sinusoidal tracking. In addition, the directional tuning of the sustained "eye velocity" and transient "visual" components of the neural responses obtained during constant velocity tracking were very similar. PCs displayed very broad tuning approximating a cosine tuning curve; the mean half-maximum bandwidth of their tuning curves was 170-180 degrees. Other cerebellar elements, related purely to eye movement and presumed to be mossy fibers, exhibited tuning approximately 40 degrees narrower than PCs and had best directions that clustered around the four cardinal directions. Our data indicate that the motion signals encoded by PCs in the monkey floccular lobe are segregated into channels that are consistent with a coordinate system defined by the vestibular apparatus and eye muscles. The differences between the tuning properties exhibited by PCs compared with mossy fibers indicate that a spatial transformation occurs within the floccular lobe.     

Central modifications of reflex parameters may underlie the fastest arm movements

Descending and reflex pathways usually converge on common interneurons and motoneurons. This implies that active movements may result from changes in reflex parameters produced by control signals conveyed by descending systems. Specifically, according to the lambda-model, a fast change in limb position is produced by a rapid change in the threshold of the stretch reflex. Consequently, external perturbations may be ineffective in eliciting additional reflex modifications of electromyographic (EMG) patterns unless the perturbations are relatively strong. In this way, the model accounts for the relatively weak effects of perturbations on the initial agonist EMG burst (Ag1) usually observed in fast movements. On the other hand, the same model permits robust reflex modifications of the timing and shape of the Ag1 in response to strong perturbations even in the fastest movements. To test the model, we verified the suggestion that the onset time of the Ag1, even in the fastest movements, depends on proprioceptive feedback in a manner consistent with a stretch reflex. In control trials, subjects (n = 6) made fast unopposed elbow flexion movements of approximately 60 degrees (peak velocity 500-700 degrees/s) in response to an auditory signal. In random test trials, a brief (50 ms) torque of 8-15 Nm either assisting or opposing the movement was applied 50 ms after this signal. Subjects had no visual feedback and were instructed not to correct arm deflections in case of perturbations. In all subjects, the onset time of the Ag1 depended on the direction of perturbation: it was 25-60 ms less in opposing compared with assisting load conditions. Assisting torques caused, at a short latency of 37 ms, an additional antagonist EMG burst preceding the Ag1. The direction-dependent effects of the perturbation persisted when cutaneous feedback was suppressed. It was concluded that the direction-dependent changes in the onset time and duration of the Ag1 as well as the antagonist activation preceding the Ag1 resulted from stretch reflex activity elicited by the perturbations rather than from a change in the control strategy or cutaneous reflexes. The results support the hypothesis on the hierarchical scheme of sensorimotor integration in which EMG patterns and movement emerge from the modification of the thresholds and other parameters of proprioceptive reflexes by control systems.     

On-line control of rapid aiming movements: Unexpected target perturbations and movement kinematics.

Two experiments with a total of 11 university students (aged 23–47 yrs) assessed the use of on-line visual information during rapid goal-directed aiming movements. In both experiments, participants were required to complete a discrete aiming movement to a single target on a graphics tablet. In Exp 1, during 76% of the experimental trials, the target width remained constant throughout the movement. The remaining 24% of the trials were evenly divided between 2 target perturbations in which the width of the target unexpectedly increased or decreased in size upon movement initiation. During Exp 2, the spatial location of the single target was perturbed to a new location closer to, or farther away from, the original target position. The proportion of perturbation trials remained constant across experiments. Results indicate that peak velocity was determined prior to movement initiation in order to meet the speed–accuracy demands of the original target width or movement amplitude. In contrast, during deceleration, participants modified their movement trajectories to account for a perturbation in target width or movement amplitude. These data suggest that on-line monitoring of visual information can be used to modify the latter half of a movement trajectory. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Human head-free gaze saccades to targets flashed before gaze-pursuit are spatially accurate

Previous studies have shown that accurate saccades can be generated, in the dark, that compensate for movements of the visual axis that result from movements of either the eyes alone or the head alone that intervene between target presentation and saccade onset. We have carried out experiments with human subjects to test whether gaze saccades (gaze = eye-in-space = eye-in-head + head-in-space) can be generated that compensate for smooth pursuit movements of gaze that intervene between target onset and gaze-saccade onset. In both head-unrestrained (head-free) and -restrained (head-fixed) conditions, subjects were asked to make gaze shifts, in the dark, to the remembered location of a briefly flashed target. On most trials, during the memory period, the subjects carried out intervening head-free gaze pursuit or head-fixed ocular pursuit along the horizontal meridian. On the remaining (control) trials, subjects did not carry out intervening pursuit movements during the memory period; this was the classical memory-guided saccade task. We found that the subjects accurately compensated for intervening movements of the visual axis in both the head-free and head-fixed conditions. We conclude that the human gaze-motor system is able to monitor on-line changes in gaze position and add them to initial retinal error, to program spatially accurate gaze saccades.     

Independent control of reflex and volitional EMG modulation during sinusoidal pursuit tracking in humans

It is well known that during volitional sinusoidal tracking the long-latency reflex modulates in parallel with the volitional EMG activity. In this study, a series of experiments are reported demonstrating several conditions in which an uncoupling of reflex from volitional activity occurs. The paradigm consists of a visually guided task in which the subject tracked a sinusoid with the wrist. The movement was perturbed by constant torque or controlled velocity perturbations at 45 degrees intervals of the tracking phase. Volitional and reflex-evoked EMG and wrist displacement as functions of the tracking phase were recorded. The relationship of both short-latency (30-60 ms) and longer-latency (60-100 ms) reflex components to the volitional EMG was evaluated. In reflex tracking, the peak reflex amplitude occurs at phases of tracking which correspond to a maximum of wrist joint angular velocity in the direction of homonymous muscle shortening and a minimum of wrist compliance. Uncoupling of the reflex and volitional EMG was observed in three situations. First, during passive movement of the wrist through the sinusoidal tracking cycle perturbation-evoked long-latency stretch reflex peak is modulated as for normal, volitional tracking. However, with passive joint movement the volitional EMG modulation is undetectable. Second, a subset of subjects demonstrate a normally modulated and positioned long-latency reflex with a single peak. However, these subjects have distinct bimodal peaks of volitional EMG. Third, the imposition of an anti-elastic load (positive position feedback) shifts the volitional EMG envelope by as much as 180 degrees along the tracking phase when compared with conventional elastic loading. Yet the long-latency reflex peak remains at its usual phase in the tracking cycle, corresponding to the maximal velocity in the direction of muscle shortening. Furthermore, comparison of the results from elastic and anti-elastic loads reveals a dissociation of short- and long-latency reflex activity, with the short-latency reflex shifting with the volitional EMG envelope. Comparable results were also obtained for controlled velocity perturbations used to control for changes in joint compliance. The uncoupling of the reflex and volitional EMG activity in the present series of experiments points to a flexible relationship between reflex and volitional control systems, altered by peripheral input and external load.     

Movement prediction and movement production.

The prediction of future positions of moving objects occurs in cases of actively produced and passively observed movement. Additionally, the moving object may or may not be tracked with the eyes. The authors studied the difference between active and passive movement prediction by asking observers to estimate displacements of an occluded moving target, where the movement was driven by the observer's manual action or was passively observed. In the absence of eye tracking, they found that in the active condition, estimates are more anticipatory than in the passive conditions. Decreasing the congruence between motor action and visual feedback diminished but did not eliminate the anticipatory effect of action. When the target was tracked with the eyes, the effect of manual action disappeared. Results indicate distinct contributions of hand and eye movement signals to the prediction of trajectories of moving objects. (PsycINFO Database Record (c) 2010 APA, all rights reserved)