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Apparent motion cues distort object localisation in egocentric space.

The visual localisation of objects in space is thought to rely on retinal information defining the environmental context and non-retinal cues from proprioception and motor commands. Here, the influence of dynamic contextual cues on the perception of egocentric space in a reaching task was investigated. Compared to performances with realistic motion or static cues, target localisation was less accurate when apparent motion was used to provide contextual information about space between the hand and the target. This effect could not be explained by the 'presence' of motion, or a bias in depth perception. Since the distortion was connected with the reaching area it was concluded that cognitive factors can unconsciously influence the perception of egocentric space, in particular distance estimation. We propose a mechanism for this whereby signals from areas MT/MST (middle temporal/medial superior temporal) create a perceptual bias through cortico-cortical connections with posterior parietal cortex.

Adult↗

Motor planning: effect of directional uncertainty with discrete spatial cues.

We investigated the effect of spatial uncertainty on motor planning by using the cueing method in a reaching task (experiment 1). Discrete spatial cues indicated the different locations in which the target could be presented. The number of cues as well as their direction changed from trial to trial. We tested the adequacy of two models of motor planning to account for the data. The switching model assumes that only one motor response can be planned at a time, whereas the capacity-sharing model assumes that multiple motor responses can be planned in parallel. Both models predict the same relation between average reaction time (RT) and number of cues, but they differ in their prediction of the shape of the distribution of the reaction time. The results showed that RT increased with the number of cues independently from their spatial dispersion. This relation was well described by the function predicted by both models, whereas it was poorly described by the Hick-Hyman law. In addition, the distribution of RT conformed to the prediction of the capacity-sharing model and not to that of the switching model. We investigated the role that the requirement of a spatially directed motor response might have had on this pattern of results by testing subjects in a simple RT task (experiment 2) with the same cueing presentation as in experiment 1. The results contrasted with those in experiment 1 and showed that RT was dependent on the spatial dispersion of the cues and not on their number. The results of the two experiments suggest that the mode of processing of potential targets is dependent on the spatial constraints of the task. The processing resources can be either divided relative to the spatial distribution of possible targets or across multiple independent discrete representations of these targets.

Adult↗

Cancellation of visuoparietal lesion-induced spatial neglect.

In humans lesions of right visuoparietal cortex induce a neglect of the contralesional visual field that is characterized in its mild form by inattentiveness to objects and events and, in its more severe form, by a condition that has many features that are indistinguishable from blindness. Here we show that spatial neglect can be induced in cats by lesions of posterior and inferior visuoparietal cortex, and that the lesion-induced neglect can be cancelled by cooling deactivation of the same region in the opposite hemisphere.

Animals↗

Spatial tuning and dynamics of vestibular semicircular canal afferents in rhesus monkeys.

Rotational head motion in vertebrates is detected by the three semicircular canals of the vestibular system whose innervating primary afferent fibers encode movement information in specific head planes. In order to further investigate the nature of vestibular central processing of rotational motion in rhesus monkeys, it was first necessary to quantify afferent information coding in this species. Extracellular recordings were performed to determine the spatial and dynamic properties of semicircular canal afferents to rotational motion in awake rhesus monkeys. We found that the afferents innervating specific semicircular canals had maximum sensitivity vectors that were mutually orthogonal. Similar to other species, afferent response dynamics varied, with regular firing afferents having increased long time constants ( t(1)), decreased cupula velocity time constants ( t(v)), and decreased fractional order dynamic operator values ( s(k)) as compared to irregular firing afferents.

Action Potentials↗

The relative effects of external spatial and motoric factors on the bimanual coordination of discrete movements.

The ability to coordinate the two hands effectively is a fundamental requirement for many everyday tasks. To investigate how bimanual coordination is achieved we asked subjects to perform discrete bimanual key-press responses under conditions in which the motoric (i.e., muscles employed) and external spatial (i.e., direction of movement in external space) relationships between the actions of the left and right index fingers were systematically varied. Subjects made simultaneous right and left index finger key-presses in response to an auditory tone. The right finger always made downward flexion movements whilst the left finger either flexed in a downward/upward direction, or extended in a downward/upward direction. Unimanual control trials of each movement type for both hands were also performed. Reaction times for each hand (RTs) and the inter-response interval (IRI) were recorded. Right hand RTs were significantly affected only when the left finger performed motorically different actions, but were unaffected by the external spatial direction in which the left hand's actions were made. The IRI results followed a similar pattern with the worst coordination (highest IRI) occurring when the left finger performed different motor actions to the right finger regardless of the direction of the left hand movement. In contrast to other recent results from experiments examining oscillatory tasks (e.g., Mechsner et al. 2001), our results suggest that in discrete tasks there is a dominance of the motor relationship between the hands over the external spatial relationship.

Adult↗

Amplitude and direction errors in kinesthetic pointing.

We investigated the accuracy with which, in the absence of vision, one can reach again a 2D target location that had been previously identified by a guided movement. A robotic arm guided the participant's hand to a target (locating motion) and away from it (homing motion). Then, the participant pointed freely toward the remembered target position. Two experiments manipulated separately the kinematics of the locating and homing motions. Some robot motions followed a straight path with the bell-shaped velocity profile that is typical of natural movements. Other motions followed curved paths, or had strong acceleration and deceleration peaks. Current motor theories of perception suggest that pointing should be more accurate when the homing and locating motion mimics natural movements. This expectation was not borne out by the results, because amplitude and direction errors were almost independent of the kinematics of the locating and homing phases. In both experiments, participants tended to overshoot the target positions along the lateral directions. In addition, pointing movements towards oblique targets were attracted by the closest diagonal (oblique effect). This error pattern was robust not only with respect to the manner in which participants located the target position (perceptual equivalence), but also with respect to the manner in which they executed the pointing movements (motor equivalence). Because of the similarity of the results with those of previous studies on visual pointing, it is argued that the observed error pattern is basically determined by the idiosyncratic properties of the mechanisms whereby space is represented internally.

Adult↗

Strategies and determinants for selection of alternate foot placement during human locomotion: influence of spatial and temporal constraints.

During locomotion in a cluttered terrain, certain terrain surfaces such as an icy one are not appropriate for foot placement; an alternate choice is required. In a previous study we showed that the selection of foot placement is not random but systematic; the dominant choices made are not uniquely defined by the available or predicted sensory inputs. We argued that selection is guided by specific rules and involves minimal displacement of the foot from its normal landing spot. The experimental protocol involved implicit spatial constraint by requiring individuals to step on the force plate that could trigger a lighted area to be avoided, thereby requiring individuals to respond within one step-cycle. Alternate foot placement was visually identified, but not measured. The purpose of this study was to directly measure foot placement, validate and/or refine the rules used to guide selection, and identify whether the alternate foot placement choices are influenced by spatial and temporal constraints on response selection. The area to be avoided was visible from the start and therefore individuals could plan and implement appropriate avoidance strategies without any temporal constraint. Spatial constraint introduced in this experiment included requirement both to step on a specific location and to avoid stepping on a specific location on the next step. The results provide support for the rules previously identified in guiding foot placement to an alternate location. Minimal displacement of the foot from its normal landing spot was validated as an important factor for selecting alternate foot placement. When several choices satisfied this factor, additional factors guide alternate foot placement. Modifications in the plane of progression are preferred while stepping wide is avoided. When no temporal constraints are imposed on the response selection, enhancing forward progression of the body becomes the dominant determinant followed by stability and lastly by energy costs associated with the modifications. A decision algorithm for selecting foot placement is proposed based on these findings. It is clear that while visual input plays a critical role in guiding foot placement, it is not entirely based on reactive control. This has implications for implementing visually guided adaptive locomotion in legged robots.

Adolescent↗

Maintaining spatial body alignment on a rotating platform by means of active counter-circling: role of vestibular and podokinesthetic afferents.

We investigated the behaviour of vision-deprived human subjects who try to maintain their horizontal alignment in space on a rotating platform by stepping about their own axis in counter-direction ('podomotor counter-rotation'), and we ask which of two alternative hypotheses best explains this behaviour. (1) The feedback hypothesis assumes that the podomotor counter-rotation is controlled by negative feedback of vestibular signals from the horizontal canals. (2) The reconstruction hypothesis holds that the vestibular cue first is combined with leg proprioceptive afferents signalling the individual's rotation on the platform ('podokinesthetic cue') in a way that reconstructs the platform's motion in space for internal representation; a negative (direction-inverted) copy of this representation then would drive the counter-rotation. Subjects were exposed to three different velocity profiles of platform rotation: VC, constant velocity rotation with sudden onset and offset; VS, sinusoidal rotation; VN, pseudorandom noise sequences. The subjects' response (i.show $132#e., their active self-rotation on the platform) to the onset and offset of VC rotations was reminiscent of a first-order lead system. Specifically, after rotation onset subjects immediately began to step on the platform in opposite direction; initially, the velocity of this response matched that of platform rotation, leading to a fairly good stabilisation of subjects' alignment in space. However, this response declined exponentially; consequently, subjects began to increasingly rotate in space along with the platform, ultimately stepping in place on the platform. After rotation offset, subjects immediately began to step around on the now stationary platform so as to continue their previous rotation in space; this response again declined exponentially until subjects became gradually stable again with respect to space. Within subjects, the time constant (tau) of these responses was similar for onset and offset. Across subjects it exhibited a conspicuous variability, ranging from 7 s to virtually infinity. The responses to VS and VN rotations were closely correlated to what could be predicted for each individual from his tau during VC on the assumption of a first-order lead system. We conclude that the mechanism stabilising body orientation basically is linear (no prediction with sinusoidal rotation, no extrapolation of constant velocity rotation). A comparison of the experimental results with simulations of the feedback hypothesis and of the reconstruction hypothesis suggests that the reconstruction hypothesis is a more likely description of the underlying processing of the vestibular and podokinesthetic cues.

Adult↗

Accuracy of spatial localization depending on head posture in a perturbed gravitoinertial force field.

Spatial orientation is crucial when subjects have to accurately reach memorized visual targets. In previous studies modified gravitoinertial force fields were used to affect the accuracy of pointing movements in complete darkness without visual feedback of the moving limb. Target mislocalization was put forward as one hypothesis to explain this decrease in accuracy of pointing movements. The aim of this study was to test this hypothesis by determining the accuracy of spatial localization of memorized visual targets in a perturbed gravitoinertial force field. As head orientation is involved in localization tasks and carrying relevant sensory systems (visual, vestibular and neck muscle proprioceptive), we also tested the effect of head posture on the accuracy of localization. Subjects (n=10) were seated off-axis on a rotating platform (120 degrees s(-1)) in complete darkness with the head fixed (head-fixed session) or free to move (head-free session). They were required to report verbally the egocentric spatial localization of visual memorized targets. They gave the perceived target location in direction (i.e. left or right) and in amplitude (in centimeters) relative to the direction they thought to be straight ahead. Results showed that the accuracy of visual localization decreased when subjects were exposed to inertial forces. Moreover, subjects localized the memorized visual targets more to the right than their actual position, that was in the direction of the inertial forces. With further analysis, it appeared that this shift of localization was concomitant with a shift of the visual straight ahead (VSA) in the opposite direction. Thus, the modified gravitoinertial force field led to a modification in the orientation of the egocentric reference frame. Furthermore, this shift of localization increased when the head was free to move while the head was tilted in roll toward the center of rotation of the platform and turned in yaw in the same direction. It is concluded that the orientation of the egocentric reference frame was influenced by the gravitoinertial vector.

Adolescent↗

Velocity and curvature in human locomotion along complex curved paths: a comparison with hand movements.

There is extensive experimental evidence linking instantaneous velocity to curvature in drawing and hand-writing movements. The empirical relationship between these characteristics of motion and path is well described by a power law in which the velocity varies in proportion to the one-third power of the radius of curvature. It was recently shown that a similar relationship can be observed during locomotion along curved elliptical paths raising the possibility that these very different motor activities might, at some level, share the same planning strategies. It has, however, been noted that the ellipse is a special case with respect to the one-third power law and therefore these previous results might not provide strong evidence that the one-third power law is a general feature of locomotion around curved paths. For this reason the experimental study of locomotion and its comparison with hand writing is extended here to non-elliptical paths. Subjects walked along predefined curved paths consisting of two complex shapes drawn on the ground: the cloverleaf and the limacon. It was found that the data always supported a close relationship between instantaneous velocity and curvature. For these more complex paths, however, the relationship is shape-dependent--although velocity and curvature can still be linked by a power law, the exponent depends on the geometrical form of the path. The results demonstrate the existence of a close relationship between instantaneous velocity and curvature in locomotion that is more general than the one-third power law. The origins of this relationship and its possible explanation in the mechanical balance of forces and in central planning are discussed.

Adult↗

Manual tracking in three dimensions.

Little is known about the manual tracking of targets that move in three dimensions. In the present study, human subjects followed, with the tip of a hand-held pen, a virtual target moving four times (period 5 s) around a novel, unseen path. Two basic types of target paths were used: a peanut-shaped Cassini ellipse and a quasi-spherical shape where four connected semicircles lay in orthogonal planes. The quasi-spherical shape was presented in three different sizes, and the Cassini shape was varied in spatial orientation and by folding it along one of the three bend axes. During the first cycle of Cassini shapes, the hand lagged behind the target by about 150 ms on average, which decreased to 100 ms during the last three cycles. Tracking performance gradually improved during the first 3 s of the first cycle and then stabilized. Tracking was especially good during the smooth, planar sections of the shapes, and time lag was significantly shorter when the tracking of a low-frequency component was compared to performance at a higher frequency (-88 ms at 0.2 Hz vs. -101 ms at 0.6 Hz). Even after the appropriate adjustment of the virtual target path to a virtual shape tracing condition, tracking in depth was poor compared to tracking in the frontal plane, resulting in a flattening of the hand path. In contrast to previous studies where target trajectories were linear or sinusoidal, these complex trajectories may have involved estimation of the overall shape, as well as prediction of target velocity.

Feedback↗

Impedance is modulated to meet accuracy demands during goal-directed arm movements.

The neuromuscular system is inherently noisy and joint impedance may serve to filter this noise. In the present experiment, we investigated whether individuals modulate joint impedance to meet spatial accuracy demands. Twelve subjects were instructed to make rapid, time constrained, elbow extensions to three differently sized targets. Some trials (20 out of 140 for each target, randomly assigned) were perturbed mechanically at 75% of movement amplitude. Inertia, damping and stiffness were estimated from the torque and angle deviation signal using a forward simulation and optimization routine. Increases in endpoint accuracy were not always reflected in a decrease in trajectory variability. Only in the final quarter of the trajectory the variability decreased as target width decreased. Stiffness estimates increased significantly with accuracy constraints. Damping estimates only increased for perturbations that were initially directed against the movement direction. We concluded that joint impedance modulation is one of the strategies used by the neuromuscular system to generate accurate movements, at least during the final part of the movement.

Adult↗

Multisensory information for postural control: sway-referencing gain shapes center of pressure variability and temporal dynamics.

The authors investigated the multisensory control of posture by altering sensory information across the visual and somatosensory systems. The support surface and visual surround were sway-referenced to anterior/posterior center of mass sway and the gain between postural sway and degree of sway referencing was manipulated (gain settings were 0.2, 1.0, and 1.8). These alterations in the sensory environment lead to observed changes in the temporal structure of the center of pressure (COP) trajectories. COP path length increased across gain settings while COP coefficient of variation decreased. The COP became increasingly more deterministic across more challenging sensory organization test (SOT) conditions and with increasing gain, and more nonstationary across more challenging SOT conditions and when the support surface was sway-referenced using a 1.8 gain setting. These findings indicate that changes in the responsiveness of the support surface and/or visual surround within each of the sway-referenced SOT conditions had functional consequences for the control of posture as evidenced by the variations in postural sway dynamics.

Adolescent↗

Relative stability improves with experience in a dynamic standing task.

This study tested the hypothesis that subjects improve their relative stability as they learn a dynamic pulling task. Healthy adult subjects practiced making brief horizontal pulls (<300 ms) on a handle to a range of target forces ranging from 20 to 80% of their estimated maximum for 5 days. They were instructed to always keep their feet flat and begin and end their motion in an upright posture. In order to do this, subjects had to develop the appropriate body momentum prior to the pull and then recover their balance following the pull. We analyzed relative stability during balance recovery, using two measures: spatial safety margin (minimum distance of the center of pressure, COP, to the edges of the feet) and temporal safety margin (minimum extrapolated time for the COP to reach the edges of the feet). We hypothesized that: (1) spatial and temporal safety margins would be uncorrelated; (2) safety-margin means would increase with practice; and (3) safety-margin standard deviations would decrease with practice. Two experiments were conducted: one where subjects practiced three force targets and positioned their initial COP in a small window, and one where subjects practiced two force targets with no initial COP constraint. Results showed that spatial and temporal safety margins were correlated but shared less than 6% variance, indicating that they reflected different aspects of control. Safety-margin averages increased with practice and standard deviations decreased with practice, indicating that the stability of balance control in the execution of this task became more robust. We suggest that the nervous system could use safety margins in both feedback and feedforward control of balance.

Adult↗

Spatial orientation in humans: perception of angular whole-body displacements in two-dimensional trajectories.

Vestibular perception of whole-body passive rotation in the horizontal plane was studied by applying two-dimensional (2D) motion to eight blindfolded healthy volunteers: pure rotations in place, corner-like trajectories and arcs of a circular trajectory were randomly applied by means of a remotely controlled robot. Angles embedded in the 2D trajectories were 45 degrees, 90 degrees, 135 degrees and 180 degrees. Stimulation of semicircular canals was the same for all trajectories but was accompanied by concurrent otolith stimulation during circular motion. Subjects participated in two successive experimental sessions. In the first session they were instructed to use a pointer to reproduce the total angular displacement after the motion (REPRODUCTION); in the second session they had to keep pointing towards a remote (15 m) memorised target during the motion (TRACKING). In REPRODUCTION subjects tended to overestimate their rotation angle by 28 +/- 11% (mean +/- SD). There was no systematic effect of the trajectory. Overestimation also occurred when subjects were required to rotate in darkness by 180 degrees (by controlling a joystick). In TRACKING there was virtually no overestimation (6 +/- 17%) and the movement of the pointer matched the dynamics of angular motion. We conclude that (a) the brain can separate and memorise the angular component of complex 2D motion; however, a large inter-individual variability in estimating its amplitude exists; (b) in the range of linear accelerations used in the study, no appreciable effect of otolith-canal perceptual interaction was shown; (c) angular displacements can be dynamically transformed into matched pointing movements; (d) overestimation seems to be typical of delayed judgements of angular displacement and of self-controlled rotations in place. This could be due to the characteristics of the physiological calibration of the vestibular input.

Humans↗

Sinusoidal forearm tracking with delayed visual feedback. I. Dependence of the tracking error on the relative delay.

During forearm tracking of a sinusoidally moving target matching of proprioceptive and visual feedback was altered by introducing different visual delays. In five normal subjects seven target frequencies were tested ranging from 0.3 to 1.5 Hz. For each target frequency nine different delays were used ranging from 0 to 120% of the cycle duration with target frequency and delay being varied randomly. Tracking error revealed a cyclic behaviour with an increase up to delays of about 50% of the target cycle duration and an improvement for delays larger than 50%. Modulation of response frequency was less pronounced compared with tracking error variation but also was dependent on relative phase. The response frequency matched the target frequency at delays of 0 and 100% of cycle duration and was slightly lower than the target frequency with relative delays of about 50%. The introduction of a visual delay during sinusoidal forearm tracking leads to a spatial in addition to the temporal mismatch between proprioception and vision. The temporal and the spatial incompatibility influence the tracking performance differentially.

Adult↗

Coordination of hand aperture with the spatial path of hand transport.

We have investigated the coordination of hand aperture with the spatial path of hand transport in prehensile movement by comparing straight prehensile movements with curved movements, in which subjects had to pass over a "via point" marked on the work surface before picking up an object in the target location. Spatial plots of hand aperture against hand transport showed that the preshaping of the hand to prepare an appropriate grasp was delayed in the curved movements relative to the straight movements, with most of the preshaping of the hand occurring after passing the via point, even when the via point occurred late in the course of the movement. The postponement of hand preshaping was apparently not due to subjects' segmenting the movement into two completely separate portions preceding and following the via point, since some degree of hand opening often occurred before the via point. We suggest that the delay in hand opening in curved movements involves a scheduling process, which uses information about hand transport to set an appropriate hand aperture.

Adult↗

Motor learning in the "podokinetic" system and its role in spatial orientation during locomotion.

The present study characterizes a previously reported adaptive phenomenon in a somatosensory-motor system involved in directional control of locomotor trajectory through foot contact with the floor. We call this the "podokinetic" (PK) system. Podokinetic adaptation was induced in six subjects by stepping in-place over the axis of a horizontally rotating disc over a range of disc angular velocities (11.25-90 degrees/s) and durations (7.5-60 min). After adaptation, subjects were blindfolded and attempted to step in-place on the floor without turning. Instead they all rotated relative to space. The rate of the "podokinetic afterrotation" (PKAR) was linearly related to stimulus amplitude up to 45 degrees/s, and the ratio of initial PKAR velocity to that of the adaptive stimulus was approximately 1:3. PKAR exhibited exponential decay, which was composed of "short-" and "long-term" components with "discharging" time constants on the order of 6-12 min and 1-2 h, respectively. The effect of stimulus duration on PKAR revealed a "charging" time constant that approximated that of the short-term component. A significant suppression of PKAR occurred during the 1 st min of the postadaptive response, suggesting functional interaction between the PK and vestibular systems during the period of vestibular stimulation. During PKAR subjects perceived no self-rotation, indicating that perception as well as locomotor control of spatial orientation were remodeled by adaptation of the PK system.

Adaptation, Physiological↗