Not letting the left leg know what the right leg is doing: limb-specific locomotor adaptation to sensory-cue conflict

We investigated the phenomenon of limb-specific locomotor adaptation in order to adjudicate between sensory-cue-conflict theory and motor-adaptation theory. The results were consistent with cue-conflict theory in demonstrating that two different leg-specific hopping aftereffects are modulated by the presence of conflicting estimates of self-motion from visual and nonvisual sources. Experiment 1 shows that leg-specific increases in forward drift during attempts to hop in place on one leg while blindfolded vary according to the relationship between visual information and motor activity during an adaptation to outdoor forward hopping. Experiment 2 shows that leg-specific changes in performance on a blindfolded hopping-to-target task are similarly modulated by the presence of cue conflict during adaptation to hopping on a treadmill. Experiment 3 shows that leg-specific aftereffects from hopping additionally produce inadvertent turning during running in place while blindfolded. The results of these experiments suggest that these leg-specific locomotor aftereffects are produced by sensory-cue conflict rather than simple motor adaptation.     

Talking about walking: biomechanics and the language of locomotion

What drives humans around the world to converge in certain ways in their naming while diverging dramatically in others? We studied how naming patterns are constrained by investigating whether labeling of human locomotion reflects the biomechanical discontinuity between walking and running gaits. Similarity judgments of a student locomoting on a treadmill at different slopes and speeds revealed perception of this discontinuity. Naming judgments of the same clips by speakers of English, Japanese, Spanish, and Dutch showed lexical distinctions between walking and running consistent with the perceived discontinuity. Typicality judgments showed that major gait terms of the four languages share goodness-of-example gradients. These data demonstrate that naming reflects the biomechanical discontinuity between walking and running and that shared elements of naming can arise from correlations among stimulus properties that are dynamic and fleeting. The results support the proposal that converging naming patterns reflect structure in the world, not only acts of construction by observers.     

Target distance and adaptation in distance perception in the constancy of visual direction

Hay and Sawyer recently demonstrated that the constancy of visual direction (CVD) also operates for near targets. A luminous spot in the dark, 40 cm from the eyes, was perceived as stationary when S nodded his head. This implies that CVD takes target distance, as well as head rotation, into account as a stationary environment is perceived during head movements. Distance is a variable in CVD because, during a turning or nodding of the head, the eyes become displaced relative to the main target direction, the line between the target and the rotation axis of the head. This displacement of the eyes during head rotation causes an additional change in the target direction, i.e., a total angular change greater than the angle of the head rotation. The extent of this additional angular displacement is greater the nearer the target. We demonstrated that the natural combination of accommodation and convergence can supply the information needed by the nervous system to compensate for this additional target displacement. We also found that wearing glasses that alter the relation between these oculomotor adjustments and target distance produces an adaptation in CVD. An adaptation period of 1.5 h produced a large adaptation effect. This effect was not entirely accounted for by an adaptation in distance perception. Measurements of the alteration between oculomotor cues and registered distance with two kinds of tests for distance perception yielded effects significantly smaller than the effect measured with the CVD test. We concluded that the wearing of the glasses had also produced an adaptation within CVD.     

The effect of perceived locomotor constraints on distance estimation

Two experiments were conducted to examine the ways in which the visual estimation of distance to a target is affected by constraints perceived to be placed on the subsequent locomotion to the target without vision. We hypothesized that an appraisal of impending effort would play a role in ascertaining the distance to be walked. In Experiment 1, the amount of resistance to walking was variable and unpredictable. One group of subjects performed against relatively low resistance, whereas another group performed against substantially greater resistance. In the low-resistance condition, no significant differences in CE, VE, time to target, or number of steps to target were found between any of the eight combinations of predictable or unpredictable resistances during walking. In the high-resistance condition, however, significant differences were found for CE and number of strides to target when resistance varied unpredictably during walking. Experiment 2 was similar in design but required subjects to walk with combinations of normal or short steps after they had viewed the target knowing only the gait type that would be used to begin locomotion. No differences in CE, VE, or time to target were found between four different combinations of gait type and predictability, under subjectively controlled conditions. When the step constraints were externally imposed, however, differences were found for CE. None of the results from either experiment, in which the number of strides needed to reach the target or the predictability of gait did not change from normal, supported the hypothesis that motor output requirements are necessary in forming a mental representation of the target position that can be used to walk to the target with eyes closed. Whichever locomotor technique was used to walk the estimated distance in these cases, the representation was able to be used independently. When walking mechanics were altered by externally imposed constraints, however, the success at reaching the estimated target position was reduced. These latter results are consistent with those obtained using up, down, and level walking and support the premise that mental representations used in blind walking are linked to the locomotor mechanics afforded by environmental conditions.     

Counteradaptation after exposure to displaced visual direction

Counteradaptation, previously demonstrated in connection with adaptation in distance perception, was obtained after exposure to displaced visual direction. When S adapted to a laterally displacing wedge prism by walking during the exposure period, there was not only a change in the perceived visual direction, but also a change m the proprioceptively perceived walking direction. When S adapts to lateral displacement of the visual direction by looking at his stationary or his moving arm, visual adaptation is obtained in the latter, but not in the former, case (Held & Hein, 1958). We obtained a change in the proprioceptively perceived position of the arm when it was stationary during the exposure period, a condition which had not yielded visual adaptation, and a much smaller, not significant, change in the felt position in the case of the actively moved arm. In the present experiments, changes in proprioceptively perceived direction or position amounted to counteradaptation.     

Differences in the dissipation of the effect of adaptation to two kinds of field displacement during head movements

When Ss were simultaneously adapted to horizontal and to vertical target displacements of equal rate during head turning about a vertical axis, the adaptation effects measured by one-trial tests immediately after the adaptation period were about equal. But retests after a time lapse of 10 and 20 min, during which S sat immobile and with eyes closed, showed a greatly different rate of dissipation of the two adaptation effects. After a lapse of 20 min, the effect of adaptation to horizontal target displacements had been reduced to 37%, whereas the effect of adaptation to vertical displacements at this final test still stood at 80% of the initial measurement. The decline over 20 min in the latter case was so smail that it could readily be ascribed to an effect of the two tests that preceded the final test. These two tests represented an effective exposure to natural viewing conditions and hence caused an unlearning of the adaptation, an effect whose existence we had demonstrated in previous work with the one-trial test.     

The Effect of Perceived Locomotor Constraints on Distance Estimation

Two experiments were conducted to examine the ways in which the visual estimation of distance to a target is affected by constraints perceived to be placed on the subsequent locomotion to the target without vision. We hypothesized that an appraisal of impending effort would play a role in ascertaining the distance to be walked. In Experiment 1, the amount of resistance to walking was variable and unpredictable. One group of subjects performed against relatively low resistance, whereas another group performed against substantially greater resistance. In the low-resistance condition, no significant differences in CE, VE, time to target, or number of steps to target were found between any of the eight combinations of predictable or unpredictable resistances during walking. In the high-resistance condition, however, significant differences were found for CE and number of strides to target when resistance varied unpredictably during walking. Experiment 2 was similar in design but required subjects to walk with combinations of normal or short steps after they had viewed the target knowing only the gait type that would be used to begin locomotion. No differences in CE, VE, or time to target were found between four different combinations of gait type and predictability, under subjectively controlled conditions. When the step constraints were externally imposed, however, differences were found for CE. None of the results from either experiment, in which the number of strides needed to reach the target or the predictability of gait did not change from normal, supported the hypothesis that motor output requirements are necessary in forming a mental representation of the target position that can be used to walk to the target with eyes closed. Whichever locomotor technique was used to walk the estimated distance in these cases, the representation was able to be used independently. When walking mechanics were altered by externally imposed constraints, however, the success at reaching the estimated target position was reduced. These latter results are consistent with those obtained using up, down, and level walking and support the premise that mental representations used in blind walking are linked to the locomotor mechanics afforded by environmental conditions.     

Multifractality of Unperturbed and Asymmetric Locomotion

The purpose of this study was to explore the extent of multifractality in unperturbed and constrained locomotion, and to determine if multifractality predicted gait adaptability. Young, healthy participants (n?=?15) walked at preferred and slow speeds, as well as asymmetrically (one leg at half speed) on a split-belt treadmill. Stride time multifractality was assessed via local detrended fluctuation analysis, which evaluates the evolution of fluctuations both spatially and temporally. Unperturbed walking exhibited monofractal behavior. Asymmetric walking displayed greater multifractality in the faster moving limb, indicating more intermittent periods of extreme high or low variance. Multifractality was not associated with adaptation to asymmetric walking. These findings further suggest that unperturbed locomotion is monofractal and establish that perturbed walking yields multifractal behavior.     

Walking and Running on the Circular Treadmill: Transition Speed and Podokinetic Aftereffects

The author compared 10 participants' self-selected walk-to-run transition speeds on a standard treadmill with those on a circular treadmill. The speed of the outer limb at walk-to-run transition on the circular treadmill and on the standard treadmill were very similar. Adaptive aftereffects from running and walking on the circular treadmill were also similar. When asked to step in place without vision, all participants inadvertently turned in circles following walking or running on the treadmill. The results of the present study suggest that the mechanisms controlling walk-to-run transitions are similar for the standard and circular treadmills and demonstrate the robust generalizability of locomotor aftereffects from running to walking. Adaptive control of speed, form, and direction may therefore share similar mechanisms for walking and running.     

Walking and running on the circular treadmill: transition speed and podokinetic aftereffects

The author compared 10 participants' self-selected walk-to-run transition speeds on a standard treadmill with those on a circular treadmill. The speed of the outer limb at walk-to-run transition on the circular treadmill and on the standard treadmill were very similar. Adaptive aftereffects from running and walking on the circular treadmill were also similar. When asked to step in place without vision, all participants inadvertently turned in circles following walking or running on the treadmill. The results of the present study suggest that the mechanisms controlling walk-to-run transitions are similar for the standard and circular treadmills and demonstrate the robust generalizability of locomotor aftereffects from running to walking. Adaptive control of speed, form, and direction may therefore share similar mechanisms for walking and running.     

Progressive locomotor recalibration during blind walking

Blind walking has become a common measure of perceived target location. This article addresses the possibility that blind walking might vary systematically within an experimental session as participants accrue exposure to nonvisual locomotion. Such variations could complicate the interpretation of blind walking as a measure of perceived location. We measured walked distance, velocity, and pace length in indoor and outdoor environments (1.5-16.0 m target distances). Walked distance increased over 37 trials by approximately 9.33% of the target distance; velocity (and to a lesser extent, pace length) also increased, primarily in the first few trials. In addition, participants exhibited more unintentional forward drift in a blindfolded marching-in-place task after exposure to nonvisual walking. The results suggest that participants not only gain confidence as blind-walking exposure increases, but also adapt to nonvisual walking in a way that biases responses toward progressively longer walked distances.     

Effects of visual focus and gait speed on walking balance in the frontal plane

We investigated how head position and gait speed influenced frontal plane balance responses to external perturbations during gait. Thirteen healthy participants walked on a treadmill at three different gait speeds. Visual conditions included either focus downward on lower extremities and walking surface only or focus forward on a stationary scene with horizontal and vertical lines. The treadmill was positioned on a platform that was stationary (non-perturbed) or moving in a pattern that appeared random to the subjects (perturbed). In non-perturbed walking, medial–lateral upper body motion was very similar between visual conditions. However, in perturbed walking, there was significantly less body motion when focus was on the stationary visual scene, suggesting visual feedback of stationary vertical and horizontal cues are particularly important when balance is challenged. Sensitivity of body motion to perturbations was significantly decreased by increasing gait speed, suggesting that faster walking was less sensitive to frontal plane perturbations. Finally, our use of external perturbations supported the idea that certain differences in balance control mechanisms can only be detected in more challenging situations, which is an important consideration for approaches to investigating sensory contribution to balance during gait.     

Perception of whether an object can be carried through an aperture depends on anticipated speed

We investigated whether anticipated speed of locomotion through an aperture influences perception of whether an object can be carried through that aperture. Participants reported whether they would be able to carry objects through an aperture (a) if they were to attempt to walk through the aperture and (b) if they were to attempt to run through the aperture. Furthermore, they did so when the object was held but not seen and when the object was seen but not held. In general, perception was influenced by object width and by anticipated speed but not by perceptual modality. Perceptual boundaries occurred at smaller object widths when participants anticipated running through the aperture than when they anticipated walking through the aperture. The results build on work showing that perception of affordances is influenced by kinetic potential as well as geometric properties and that perception may be supported by the detection of modality-neutral stimulation patterns.     

Perception of visual speed while moving

During self-motion, the world normally appears stationary. In part, this may be due to reductions in visual motion signals during self-motion. In 8 experiments, the authors used magnitude estimation to characterize changes in visual speed perception as a result of biomechanical self-motion alone (treadmill walking), physical translation alone (passive transport), and both biomechanical self-motion and physical translation together (walking). Their results show that each factor alone produces subtractive reductions in visual speed but that subtraction is greatest with both factors together, approximating the sum of the 2 separately. The similarity of results for biomechanical and passive self-motion support H. B. Barlow's (1990) inhibition theory of sensory correlation as a mechanism for implementing H. Wallach's (1987) compensation for self-motion.     

Effects of velocity and limb loading on the coordination between limb movements during walking

The authors investigated the effects of velocity (increasing from 0.5 to 5.0 km/hr in steps of 0.5 km/hr) and limb loading on the coordination between arm and leg movements during treadmill walking in 7 participants. Both the consistency of the individual limb movements and the stability of their coordination increased with increasing velocity; the frequency coordination between arm and leg movements was 2:1 at the lower velocities and 1:1 at the higher velocities. The mass manipulation affected the individual limb movements but not their coordination, indicating that a stable walking pattern was preserved. The results differed qualitatively from those obtained in studies on bimanual interlimb coordination, implying that the dynamical principles identified therein are not readily applicable to locomotion.     

Ground reaction force adaptations during cross-slope walking and running

Though transversely inclined (cross-sloped) surfaces are prevalent, our understanding of the biomechanical adaptations required for cross-slope locomotion is limited. The purpose of this study was to examine ground reaction forces (GRF) in cross-sloped and level walking and running. Nine young adult males walked and ran barefoot along an inclinable walkway in both level (0°) and cross-slope (10°) configurations. The magnitude and time of occurrence of selected features of the GRF were extracted from the force plate data. GRF data were collected in level walking and running (LW and LR), inclined walking and running up-slope (IWU and IRU), and down-slope (IWD and IRD), respectively. The GRF data were then analyzed using repeated measures MANOVA. In the anteroposterior direction, the timing of the peak force values differed across conditions during walking (p=.041), while the magnitude of forces were modified across conditions for running (p=.047). Most significant differences were observed in the mediolateral direction, where generally force values were up to 390% and 530% (p<.001) larger during the cross-slope conditions compared to level for walking and running, respectively. The maximum force peak during running occurred earlier at IRU compared to the other conditions (p≤.031). For the normal axis a significant difference was observed in the first maximum force peak during walking (p=.049). The findings of this study showed that compared to level surfaces, functional adaptations are required to maintain forward progression and dynamic stability in stance during cross-slope walking and running.     

Gradual increase of perturbation load induces a longer retention of locomotor adaptation in children with cerebral palsy

The goal of this study is to determine whether the size and the variability of error have an impact on the retention of locomotor adaptation in children with cerebral palsy (CP). Eleven children with CP, aged 7–16 years old, were recruited to participate in this study. Three types of force perturbations (i.e., abrupt, gradual and noisy loads) were applied to the right leg above the ankle starting from late stance to mid-swing in three test sessions while the subject walked on a treadmill. Spatial-temporal gait parameters were recorded using a custom designed 3D position sensor during treadmill walking. We observed that children with CP adapted to the resistance force perturbation and showed an aftereffect consisting of increased step length after load release. Further, we observed a longer retention of the aftereffect for the condition with a gradual load than that with an abrupt load. Results from this study suggested that the size of error might have an impact on the retention of motor adaptation in children with CP with a longer retention of motor adaptation for the condition with a small size of error than that with a large error. In addition, enhanced variability of error seems facilitate motor learning during treadmill training. Results from this study may be used for the development of force perturbation based training paradigms for improving walking function in children with CP.     

Visual perception and the guidance of locomotion without vision to previously seen targets

Two experiments were performed to assess the accuracy and precision with which adults perceive absolute egocentric distances to visible targets and coordinate their actions with them when walking without vision. In experiment 1 subjects stood in a large open field and attempted to judge the midpoint of self-to-target distances of between 4 and 24 m. In experiment 2 both highly practiced and unpracticed subjects stood in the same open field, viewed the same targets, and attempted to walk to them without vision or other environmental feedback under three conditions designed to assess the effects on accuracy of time-based memory decay and of walking at an unusually rapid pace. In experiment 1 the visual judgments were quite accurate and showed no systematic constant error. The small variable errors were linearly related to target distance. In experiment 2 the briskly paced walks were accurate, showing no systematic constant error, and the small, variable errors were a linear function of target distance and averaged about 8% of the target distance. Unlike Thomson's (1983) findings, there was not an abrupt increase in variable error at around 9 m, and no significant time-based effects were observed. The results demonstrate the accuracy of people's visual perception of absolute egocentric distances out to 24 m under open field conditions. The accuracy of people's walking without vision to previously seen targets shows that efferent and proprioceptive information about locomotion is closely calibrated to visually perceived distance. Sensitivity to the correlation of optical flow with efferent/proprioceptive information while walking with vision may provide the basis for this calibration when walking without vision.     

Inconsistent locomotion inhibits vection

We measured the strength of illusory self-motion perception (vection) with and without locomotion on a treadmill. The results revealed that vection was inhibited by inconsistent locomotion, but facilitated by consistent locomotion.     

Changes in motion perception following oculomotor smooth pursuit adaptation

The hypothesis that oculomotor smooth pursuit (SP) adaptation is accompanied by alterations in velocity perception was tested by assessing coherence thresholds, using random-dot kinematograms before and after the adaptation paradigm. The results showed that the sensitivity to coherent motion at 10 deg/sec (the initial target velocity during adaptation) was reduced after the SP adaptation, ending up at a level that was between those normally observed for velocities of 10 and 20 deg/sec. This is consistent with an overestimation of the velocity of the coherent motion and suggests that SP adaptation alters not only the oculomotor output, but also the perception of target velocity.