Sequential aiming with one and two limbs: Effects of target size

It is well reported that movement times to the first target in a two-target sequence are slower than when a single target response is required. This one-target advantage has been shown to emerge when the two-target sequence is performed with the same limb and when the first and second segments within the sequence are performed with different limbs (i.e., when there is a switch between limbs at the first target). The present study examined the functional dependency between response segments in both single and two limb sequential aiming by varying the accuracy demands at the first and second target. Results revealed that, for both one and two limb conditions, the one-target advantage was present with large first targets but not with small first targets. Additionally, when the first target was large and the second target was small, spatial variability at the first target was significantly less (or constrained more) in both one and two limb conditions compared to conditions requiring only a single target response. These findings suggest that similar principles underlie the one-target advantage in both single and two limb sequential movements.     

Impact forces cannot explain the one-target advantage in rapid aimed hand movements

A pointing movement is executed faster when a subject is allowed to stop at the first target than when the subject has to proceed to a second target ("one-target advantage"). Our hypothesis was that this is because the impact at the target helps to stop the finger when the finger does not have to proceed to a second target. This hypothesis would predict that the horizontal force at contact with the first target should be larger when there is only one-target. Modelling smooth movements with larger forces at contact using a minimum-jerk model, shows that the peak velocity is slightly higher and it occurs later during the movement when there is only one target. Although the one-target advantage was present in our experiment, the horizontal force at contact in the one-target condition was not larger than in the two-target condition. The time of the maximum velocity did not differ, but the maximum velocity was higher in the one-target condition. Thus our hypothesis is rejected, favouring a non-mechanical explanation of the one-target advantage.     

Programming saccadic eye movements

This article addresses questions about the preparatory processes that immediately precede saccadic eye movements. Saccade latencies were measured in a task in which subjects were provided partial advance information about the spatial location of a target fixation. In one experiment, subjects were faster in initiating saccades when they knew either the direction or amplitude of the required movement in advance compared to a condition with equal uncertainty about the number of potential saccade targets but without knowledge of the parameters required to execute the movement. These results suggest that the direction and amplitude for an upcoming saccade were calculated separately, and not in a fixed serial order. In another experiment, subjects appear to have programmed the saccades more holistically--with computations of direction and amplitude parameters occurring simultaneously. The implications of these results for models of eye movement preparation are discussed.     

Transfer of adaptation between ocular saccades and arm movements

Previous studies found little or no transfer of adaptation from reactive saccades to arm pointing movements, which suggests that the two motor systems rely on distinct adaptive mechanisms. However, this conclusion is based on experiments about the adaptation of response amplitudes, which is known to follow somewhat different principles than the adaptation of response directions. In the present study, we therefore investigate whether adapting the direction of reactive saccades will transfer to arm movements. We also test transfer in the opposite direction, from the arm to the eyes. Participants executed aimed saccades or arm movements from a central starting point towards visual targets in the participants' frontal plane. Targets were presented in eight possible locations along a circle of 20 cm radius about the starting point; each remained for 200 ms in one position, and was then displaced along the circle by -15 degrees . Participants from group E adapted to these double-stepped targets while executing eye movements, and were then tested for transfer while executing arm movements. The reciprocal design was used in participants from group A. Adaptive change in group A was about 14 degrees , while in group E it was only about 7 degrees . Transfer of adaptation was substantial, and was more pronounced when using the arm (i.e., eye-to-arm transfer in group E) rather than the eyes (i.e., arm-to-eye transfer in group A). Strong aftereffects were yielded in both groups. This pattern of findings implies that the adaptive change observed in our study was mainly based on recalibration rather than on cognitive strategies (strong aftereffects), that eyes and arm had access to a common adaptive mechanism (substantial transfer), and that the arm had better access than the eyes (larger adaptation and transfer when using the arm). When considering this outcome along with the available literature, it appears that arm and eyes may rely sometimes on a common and sometimes on distinct adaptive mechanisms, depending on the adapted parameter and on the nature of the motor task.     

Attentional localization prior to simple and directed manual responses

The relationship between attention and the programming of motor responses was investigated, using a paradigm in which the onsets of targets for movements were preceded by peripheral attentional cues. Simple (button release) and reaching manual responses were compared under conditions in which the subjects either made saccades toward the target location or refrained from making eye movements. The timing of the movement onset was used as the dependent measure for both simple and reaching manual responses. Eye movement latencies were also measured. A follow-up experiment measured the effect of the same peripheral cuing procedure on purely visual processes, using signal detection measures of visual sensitivity and response bias. The results of the first experiment showed that reaction time (RT) increased with the distance between the cued and the target locations. Stronger distance effects were observed when goal-directed responses were required, which suggests enhanced attentional localization of target positions under these conditions. The requirement to generate an eye movement response was found to delay simple manual RTs. However, mean reaching RTs were unaffected by the eye movement condition. Distance gradients on eye movement latencies were relatively shallow, as compared with those on goal-directed manual responses. The second experiment showed that the peripheral cue had only a very small effect on visual detection sensitivity in the absence of directed motor responses. It is concluded that cue-target distance effects with peripheral cues are modulated by the motor-programming requirements of the task. The effect of the peripheral cue on eye movement latencies was qualitatively different from that observed on manual RTs, indicating the existence of separate neural representations underlying both response types. At the same time, the interactions between response modalities are consistent with a supramodal representation of attentional space, within which different motor programs may interact.     

Attentional localization prior to simpleand directed manual responses

The relationship between attention and the programming of motor responses was investigated, using a paradigm in which the onsets of targets for movements were preceded by peripheral attentional cues. Simple (button release) and reaching manual responses were compared under conditions in which the subjects either made saccades toward the target location or refrained from making eye movements. The timing of the movement onset was used as the dependent measure for both simple and reaching manual responses. Eye movement latencies were also measured. A follow-up experiment measured the effect of the same peripheral cuing procedure on purely visual processes, using signal detection mea-sures of visual sensitivity and response bias. The results of the first experiment showed that reaction time (RT) increased with the distance between the cued and the target locations. Stronger distance ef-fects were observed when goal-directed responses were required, which suggests enhanced attentional localization of target positions under these conditions. The requirement to generate an eye movement response was found to delay simple manual RTs. However, mean reaching RTs were unaffected by the eye movement condition. Distance gradients on eye movement latencies were relatively shallow, as compared with those on goal-directed manual responses. The second experiment showed that the peripheral cue had only a very small effect on visual detection sensitivity in the absence of directed motor responses. It is concluded that cue-target distance effects with peripheral cues are modulated by the motor-programming requirements of the task. The effect of the peripheral cue on eye movement latencies was qualitatively different from that observed on manual RTs, indicating the existence of separate neural representations underlying both response types. At the same time, the interactions be-tween response modalities are consistent with a supramodal representation of attentional space, within which different motor programs may interact.     

Inhibition of return in single and dual tasks: examining saccadic, keypress, and pointing responses

Two experiments are reported in which inhibition of return (IOR)was examined wit h single-responsetasks (either manual responses alone or saccadic responses alone) and dual-response tasks (simultaneous manual and saccadic responses). The first experiment-using guided limb movements that require considerable spatial information-showed more IOR for saccades than for pointing responses. In addition, saccadic IOR was reduced with concurrent pointing movements, but manual IOR was not affected by concurrent saccades. Importantly, at the time of saccade initiation, the arm movements did not start yet, indicating that the influence on saccade IOR is due to arm-movement preparation. In the second experiment, using localization keypress responses that required only minimal spatial information, greater IOR was again found for saccadic than for manual responses, but no effect of concurrent movements was found. These findings add further support that there is a dissociation between oculomotor and skeletal-motor IOR. Moreover, the results show that the preparation manual responses tend to mediate saccadic behavior-but only when the manual responses require high levels of spatial accuracy-and that the superior colliculus is the likely neural substrate integrating IOR for eye and arm movements.     

Dissecting online control in Developmental Coordination Disorder: a kinematic analysis of double-step reaching

In a recent study, children with movement clumsiness (or Developmental Coordination Disorder-DCD) were shown to have difficulties making rapid online corrections when reaching, demonstrated by slower and less accurate movements to double-step targets (Hyde & Wilson, 2011). These results suggest that children with DCD have difficulty using predictive estimates of limb position when making rapid adjustments to movement, in-flight. However, chronometric data alone does not provide strong evidence for this hypothesis: it remains unclear whether early (and rapid) control parameters or post-correction stages of the movement trajectory are affected. Thus, the overarching aim of this study was to conduct a kinematic analysis of double-step reaching in order to isolate the different control parameters that might explain the slower and less accurate double-step reaching performance of children with DCD. Participants were a new sample of 13 children with DCD aged between 8-12 years and 13 age-matched controls. Children were required to reach and touch one of three possible targets presented at the coordinates -20°, 0° and 20° on a 17 in. LCD touch-screen. For most trials (80%) the target remained stationary for the duration of movement (non-jump trials), while for the remainder (20%), the target jumped randomly to one of two peripheral locations at movement onset (jump trials). Consistent with earlier work, children with DCD were slower to initiate reaching compared to controls and showed longer MT and more errors on jump trials. Kinematic data showed that while the two groups did not differ on time to peak velocity or acceleration, children with DCD were slower to correct reach trajectory on jump trials. No group differences were observed on late kinematic markers, e.g., post-correction time. The pattern of results support and extend earlier work showing deficits in ROC in DCD. From a computational perspective, delayed corrections to the reach trajectory suggests some difficulty integrating information about the target perturbation with a predictive (or forward) estimate of limb position relative to the initial target. These conclusions are discussed, along with directions for future research.     

Effects of motor intention on the perception of somatosensory events: A behavioural and functional magnetic resonance imaging study

The intention to execute a movement can modulate our perception of sensory events, and this modulation is observed ahead of both ocular and upper limb movements. However, theoretical accounts of these effects, and also the empirical data, are often contradictory. Accounts of “active touch”, and the premotor theory of attention, have emphasized how movement intention leads to enhanced perceptual processing at the target of a movement, or on the to-be-moved effector. By contrast, recent theories of motor control emphasize how internal “forward” model (FM) estimates may be used to cancel or attenuate sensory signals that arise as a result of self-generated movements. We used behavioural and functional brain imaging (functional magnetic resonance imaging, fMRI) to investigate how perception of a somatosensory stimulus differed according to whether it was delivered to a hand that was about to execute a reaching movement or the alternative, nonmoving, hand. The results of our study demonstrate that a somatosensory stimulus delivered to a hand that is being prepared for movement is perceived to have occurred later than when that same stimulus is delivered to a nonmoving hand. This result indicates that it takes longer for a tactile stimulus to be detected when it is delivered to a moving limb and may correspond to a change in perceptual threshold. Our behavioural results are paralleled by the results of our fMRI study that demonstrated that there were significantly reduced blood-oxygen-level-dependent (BOLD) responses within the parietal operculum and insula following somatosensory stimulation of the hand being prepared for movement, compared to when an identical stimulus was delivered to a nonmoving hand. These findings are consistent with the prediction of FM accounts of motor control that postulate that central sensory suppression of somatosensation accompanies self-generated limb movements, and with previous reports indicating that effects of sensory suppression are observed in higher order somatosensory regions.     

Speed and accuracy of saccadic eye movements: characteristics of impulse variability in the oculomotor system

Dynamic characteristics observed in the trajectories of saccadic eye movements reveal systematic variability of the force pulses used to move the eyes. This variability causes saccades to exhibit a linear speed-accuracy trade-off: As the average distance and duration of saccades toward specified target points increase, the standard deviations of saccadic-movement endpoints increase linearly with the saccades' average velocity. The linear trade-off, and other observed stochastic properties of saccades, may be attributed to noise in neuromotor processes and may be described in terms of an impulse-variability model originally designed for characterizing limb movements. According to the model, both eye and limb movements are controlled through stochastic force and time parameters that govern movement kinematics. Such an account may promote a unified conceptual framework for understanding a wide range of motor behavior.     

When two actions are easier than one: How inhibitory control demands affect response processing

Numerous studies showed that the simultaneous execution of multiple actions is associated with performance costs. Here, we demonstrate that when highly automatic responses are involved, performance in single-response conditions can actually be worse than in dual-response conditions. Participants responded to peripheral visual stimuli with an eye movement (saccade), a manual key press, or both. To manipulate saccade automaticity, a central fixation cross either remained present throughout the trial (overlap condition, lower automaticity) or disappeared 200 ms before visual target onset (gap condition, greater automaticity). Crucially, single-response conditions yielded more performance errors than dual-response conditions (i.e., dual-response benefit), especially in gap trials. This was due to difficulties associated with inhibiting saccades when only manual responses were required, suggesting that response inhibition (remaining fixated) can be even more resource-demanding than overt response execution (saccade to peripheral target).     

Coupling of Eye,Finger, Elbow,and Shoulder Movements During Manual Aiming

Temporal and spatial coupling of point of gaze (PG) and movements of the finger, elbow, and shoulder during a speeded aiming task were examined. Ten participants completed 40-cm aiming movements with the right arm, in a situation that allowed free movement of the eyes, head, arm, and trunk. On the majority of trials, a large initial saccade undershot the target slightly, and 1 or more smaller corrective saccades brought the eyes to the target position. The finger, elbow, and shoulder exhibited a similar pattern of undershooting their final positions, followed by small corrective movements. Eye movements usually preceded limb movements, and the eyes always arrived at the target well in advance of the finger. There was a clear temporal coupling between primary saccade completion and peak acceleration of the finger, elbow, and shoulder. The initiation of limb-segment movement usually occurred in a proximal-to-distal pattern. Increased variability in elbow and shoulder position as the movement progressed may have served to reduce variability in finger position. The spatial-temporal coupling of PG with the 3 limb segments was optimal for the pick up of visual information about the position of the finger and the target late in the movement.     

Coupling of eye, finger, elbow, and shoulder movements during manual aiming

Temporal and spatial coupling of point of gaze (PG) and movements of the finger, elbow, and shoulder during a speeded aiming task were examined. Ten participants completed 40-cm aiming movements with the right arm, in a situation that allowed free movement of the eyes, head, arm, and trunk. On the majority of trials, a large initial saccade undershot the target slightly, and 1 or more smaller corrective saccades brought the eyes to the target position. The finger, elbow, and shoulder exhibited a similar pattern of undershooting their final positions, followed by small corrective movements. Eye movements usually preceded limb movements, and the eyes always arrived at the target well in advance of the finger. There was a clear temporal coupling between primary saccade completion and peak acceleration of the finger, elbow, and shoulder. The initiation of limb-segment movement usually occurred in a proximal-to-distal pattern. Increased variability in elbow and shoulder position as the movement progressed may have served to reduce variability in finger position. The spatial-temporal coupling of PG with the 3 limb segments was optimal for the pick up of visual information about the position of the finger and the target late in the movement.     

Spatial Error Detection and Assimilation Effects in Rapid Single and Bimanual Aiming Movements

In 2 experiments, spatial error detection capability and movement accuracy were investigated in both single and bimanual rapid aiming movements. In both experiments, right-handed college-age participants (N = 40 [Experiment 1]; N = 24 [Experiment 2]) used light, aluminum levers to make quick single and dual reversal movements in the sagittal plane in a time to reversal of 210 ms to either the same or different target locations involving identical (Experiment 1) or mirror-image (Experiment 2) movements. In Experiment 1, the shorter-distance limb overshot the target by 15-23&percent; when paired with a limb traveling at least 20 degrees farther, but no spatial assimilations were shown when movements differed by 20 degrees or less. In Experiment 2, the shorter-distance limb overshot 22-29&percent; when paired with a limb traveling 20 degrees farther, but spatial assimilations were not mitigated when both limbs moved to the same target position. Participants underestimated movement amplitude in all dual conditions but particularly when spatial assimilations were noted. Correlations between actual and estimated errors decreased from single to dual trials in both experiments. The findings suggest that spatial assimilations are caused by bimanual differences in movement amplitude, regardless of movement direction, and that individuals have greater difficulty identifying errors in simultaneous actions, especially when spatial assimilations are present, than identifying errors in single-limb actions.     

Coordinated control of eye and hand movements in dynamic reaching

In the present study, we integrated two recent, at first sight contradictory findings regarding the question whether saccadic eye movements can be generated to a newly presented target during an ongoing hand movement. Saccades were measured during so-called adaptive and sustained pointing conditions. In the adapted pointing condition, subjects had to direct both their gaze and arm movements to a displaced target location. The results showed that the eyes could fixate the new target during pointing. In addition, a temporal coupling of these corrective saccades was found with changes in arm movement trajectories when reaching to the new target. In the sustained pointing condition, however, the same subjects had to point to the initial target, while trying to deviate their gaze to a new target that appeared during pointing. It was found that the eyes could not fixate the new target before the hand reached the initial target location. Together, the results indicate that ocular gaze is always forced to follow the target intended by a manual arm movement. A neural mechanism is proposed that couples ocular gaze to the target of an arm movement. Specifically, the mechanism includes a reach neuron layer besides the well-known saccadic layer in the primate superior colliculus. Such a tight, sub-cortical coupling of ocular gaze to the target of a reaching movement can explain the contrasting behavior of the eyes in dependency of whether the eye and hand share the same target position or attempt to move to different locations.     

Visual regulation of manual aiming: A comparison of methods

Visual regulation of upper limb movements occurs throughout the trajectory and is not confined to discrete control in the target area. Early control is based on the dynamic relationship between the limb, the target, and the environment. Despite robust outcome differences between protocols involving visual manipulations, it remains difficult to identify the kinematic events that characterize these differences. In this study, participants performed manual aiming movements with and without vision. We compared several traditional approaches to movement analysis with two new methods of quantifying online limb regulation. As expected, participants undershot the target and their movement endpoints were more variable when vision was not available. Although traditional measures such as reaction time, time after peak velocity, and the presence of discontinuities in acceleration were sensitive to the visual manipulation, measures quantifying the trial-to-trial spatial variability throughout the trajectory were the most effective in isolating the time course of online regulation.     

Oculomotor interference during manual response preparation: evidence from the response-cueing paradigm

Preparation provided by visual location cues is known to speed up behavior. However, the role of concurrent saccades in response to visual cues remains unclear. In this study, participants performed a spatial precueing task by pressing one of four response keys with one of four fingers (two of each hand) while eye movements were monitored. Prior to the stimulus, we presented a neutral cue (baseline), a hand cue (corresponding to left vs. right positions), or a finger cue (corresponding to inner vs. outer positions). Participants either remained fixated on a central fixation point or moved their eyes freely. The results demonstrated that saccades during the cueing interval altered the pattern of cueing effects. Finger cueing trials in which saccades were spatially incompatible (vs. compatible) with the subsequently required manual response exhibited slower manual RTs. We propose that interference between saccades and manual responses affects manual motor preparation.     

The dual role of vision in sequential aiming movements

Previous research has demonstrated that movement times to the first target in sequential aiming movements are influenced by the properties of subsequent segments. Based on this finding, it has been proposed that individual segments are not controlled independently. The purpose of the current study was to investigate the role of visual feedback in the interaction between movement segments. In contrast to past research in which participants were instructed to minimize movement time, participants were set a criterion movement time and the resulting errors and limb trajectory kinematics were examined under vision and no vision conditions. Similar to single target movements, the results indicated that vision was used within each movement segment to correct errors in the limb trajectory. In mediating the transition between segments, visual feedback from the first movement segment was used to adjust the parameters of the second segment. Hence, increases in variability that occurred from the first to the second target in the no vision condition were curtailed when visual feedback was available. These results are discussed along the lines of the movement constraint and movement integration hypotheses.     

Speed-accuracy characteristics of saccadic eye movements

Speed-accuracy trade-off characteristic of horizontal saccadic eye movements were examined in this study. Unlike limb movements, saccadic eye movements are preprogrammed, unidimensional, and do not involve target impact. Hence, they provide an optimal test of the impulse variability account of the speed-accuracy trade-off in rapid movements. Subjects were required to alternately look at two target lights as fast and as accurately as possible for a period of 10 s. Target lights subtended angles of 5, 10, 15, and 20 degrees. By restricting target distances to less than 20 degrees of arc, the speed-accuracy relation was examined for single horizontal saccadic movements of the eye. movement of the dominant eye was tracked with an infra-red eye monitoring device. Fifty saccadic movements of the eye were recorded for each target distance and used to compute the average amplitude, duration, and velocity of eye movements, as well as, movement endpoint variability. An increase in both average velocity and movement endpoint variability with increasing movement amplitude was found. This, together with the unique features of the eye movement system, support the impulse variability account of the speed-accuracy trade-off in rapid movements.     

Bimanual coupling effects during arm immobilization and passive movements

When humans simultaneously perform different movements with both hands, each limb movement interferes with the contralateral limb movement (bimanual coupling). Previous studies on both healthy volunteers and patients with central or peripheral nervous lesions suggested that such motor constraints are tightly linked to intentional motor programs, rather than to movement execution. Here, we aim to investigate this phenomenon, by using a circles-lines task in which, when subjects simultaneously draw lines with the right hand and circles with the left hand, both the trajectories tend to become ovals (bimanual coupling effect). In a first group, we immobilized the subjects’ left arm with a cast and asked them to try to perform the bimanual task. In a second group, we passively moved the subjects’ left arm and asked them to perform voluntary movements with their right arm only. If the bimanual coupling arises from motor intention and planning rather than spatial movements, we would expect different results in the two groups. In the Blocked group, where motor intentionality was required but movements in space were prevented by immobilization of the arm, a significant coupling effect (i.e., a significant increase of the ovalization index for the right hand lines) was found. On the contrary, in the Passive group, where movements in space were present but motor intentionality was not required, no significant coupling effect was observed. Our results confirmed, in healthy subjects, the central role of the intentional and predictive operations, already evidenced in pathological conditions, for the occurrence of bimanual coupling.