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To navigate complex environments, walking animals must detect and overcome unexpected perturbations. One technical challenge when investigating adaptive locomotion is measuring behavioral responses to precise perturbations during naturalistic walking; another is that manipulating neural activity in sensorimotor circuits often reduces spontaneous locomotion. To overcome these obstacles, we introduce miniature treadmill systems for coercing locomotion and tracking 3D kinematics of walking Drosophila. By systematically comparing walking in three experimental setups, we show that flies compelled to walk on the linear treadmill have similar stepping kinematics to freely walking flies, while kinematics of tethered walking flies are subtly different. Genetically silencing mechanosensory neurons altered step kinematics of flies walking on the linear treadmill across all speeds. We also discovered that flies can maintain a forward heading on a split-belt treadmill by specifically adapting the step distance of their middle legs. These findings suggest that proprioceptive feedback contributes to leg motor control irrespective of walking speed and that the fly's middle legs play a specialized role in stabilizing locomotion. [Display omitted] • A new actuated treadmill system captures 3D kinematics of flies compelled to walk • Flies walking on the treadmill have similar kinematics to freely walking flies • Proprioceptive feedback is important for leg motor control at all walking speeds • Flies on a split-belt treadmill use their middle legs to counteract perturbations Pratt et al. engineer tiny treadmills that enable 3D tracking of fruit fly walking kinematics. The authors systematically compare treadmills to free and tethered walking, evaluate the role of proprioceptive feedback across walking speeds, and reveal how flies adapt their stepping patterns to maintain course on a split-belt treadmill. [ABSTRACT FROM AUTHOR]

Locomotor adaptation to abrupt and gradual perturbations are likely driven by fundamentally different neural processes. The aim of this study was to quantify brain dynamics associated with gait adaptation to a gradually introduced gait perturbation, which typically results in smaller behavioral errors relative to an abrupt perturbation. Loss of balance during standing and walking elicits transient increases in midfrontal theta oscillations that have been shown to scale with perturbation intensity. We hypothesized there would be no significant change in anterior cingulate theta power (4–7 Hz) with respect to pre-adaptation when a gait perturbation is introduced gradually because the gradual perturbation acceleration and stepping kinematic errors are small relative to an abrupt perturbation. Using mobile electroencephalography (EEG), we measured gait-related spectral changes near the anterior cingulate, posterior cingulate, sensorimotor, and posterior parietal cortices as young, neurotypical adults (n = 30) adapted their gait to an incremental split-belt treadmill perturbation. Most cortical clusters we examined (>70%) did not exhibit changes in electrocortical activity between 2–50 Hz. However, we did observe gait-related theta synchronization near the left anterior cingulate cortex during strides with the largest errors, as measured by step length asymmetry. These results suggest gradual adaptation with small gait asymmetry and perturbation magnitude may not require significant cortical resources beyond normal treadmill walking. Nevertheless, the anterior cingulate may remain actively engaged in error monitoring, transmitting sensory prediction error information via theta oscillations.

Abstract Background Asymmetric walking gait impairs activities of daily living in neurological patient populations, increases their fall risk, and leads to comorbidities. Accessible, long-term rehabilitation methods are needed to help neurological patients restore symmetrical walking patterns. This study aimed to determine if a passive unilateral hip exosuit can modify an induced asymmetric walking gait pattern. We hypothesized that a passive hip exosuit would diminish initial- and post-split-belt treadmill walking after-effects in healthy young adults. Methods We divided 15 healthy young adults evenly between three experimental groups that each completed a baseline trial, an adaptation period with different interventions for each group, and a post-adaptation trial. To isolate the contribution of the exosuit we compared a group adapting to the exosuit and split-belt treadmill (Exo-Sb) to groups adapting to exosuit-only (Exo-only) and split-belt only (Sb-only) conditions. The independent variables step length, stance time, and swing time symmetry were analyzed across five timepoints (baseline, early- and late adaptation, and early- and late post-adaptation) using a 3 × 5 mixed ANOVA. Results We found significant interaction and time effects on step length, stance time and swing time symmetry. Sb-only produced increased step length asymmetry at early adaptation compared to baseline (p

Locomotor adaptation is crucial for daily gait adjustments to changing environmental demands and obstacle avoidance. Mobile brain imaging with high‐density electroencephalography (EEG) now permits quantification of electrocortical dynamics during human locomotion. To determine the brain areas involved in human locomotor adaptation, we recorded high‐density EEG from healthy, young adults during split‐belt treadmill walking. We incorporated a dual‐electrode EEG system and neck electromyography to decrease motion and muscle artefacts. Voluntary movement preparation and execution have been linked to alpha (8–13 Hz) and beta band (13–30 Hz) desynchronizations in the sensorimotor and posterior parietal cortices, whereas theta band (4–7 Hz) modulations in the anterior cingulate have been correlated with movement error monitoring. We hypothesized that relative to normal walking, split‐belt walking would elicit: (1) decreases in alpha and beta band power in sensorimotor and posterior parietal cortices, reflecting enhanced motor flexibility; and (2) increases in theta band power in anterior cingulate cortex, reflecting instability and balance errors that will diminish with practice. We found electrocortical activity in multiple regions that was associated with stages of gait adaptation. Data indicated that sensorimotor and posterior parietal cortices had decreased alpha and beta band spectral power during early adaptation to split‐belt treadmill walking that gradually returned to pre‐adaptation levels by the end of the adaptation period. Our findings emphasize that multiple brain areas are involved in adjusting gait under changing environmental demands during human walking. Future studies could use these findings on healthy, young participants to identify dysfunctional supraspinal mechanisms that may be impairing gait adaptation. Key points: Identifying the location and time course of electrical changes in the brain correlating with gait adaptation increases our understanding of brain function and provides targets for brain stimulation interventions.Using high‐density EEG in combination with 3D biomechanics, we found changes in neural oscillations localized near the sensorimotor, posterior parietal and cingulate cortices during split‐belt treadmill adaptation.These findings suggest that multiple cortical mechanisms may be associated with locomotor adaptation, and their temporal dynamics can be quantified using mobile EEG.Results from this study can serve as a reference model to examine brain dynamics in individuals with movement disorders that cause gait asymmetry and reduced gait adaptation. [ABSTRACT FROM AUTHOR]

Background: Asymmetric walking gait impairs activities of daily living in neurological patient populations, increases their fall risk, and leads to comorbidities. Accessible, long-term rehabilitation methods are needed to help neurological patients restore symmetrical walking patterns. This study aimed to determine if a passive unilateral hip exosuit can modify an induced asymmetric walking gait pattern. We hypothesized that a passive hip exosuit would diminish initial- and post-split-belt treadmill walking after-effects in healthy young adults. Methods: We divided 15 healthy young adults evenly between three experimental groups that each completed a baseline trial, an adaptation period with different interventions for each group, and a post-adaptation trial. To isolate the contribution of the exosuit we compared a group adapting to the exosuit and split-belt treadmill (Exo-Sb) to groups adapting to exosuit-only (Exo-only) and split-belt only (Sb-only) conditions. The independent variables step length, stance time, and swing time symmetry were analyzed across five timepoints (baseline, early- and late adaptation, and early- and late post-adaptation) using a 3 × 5 mixed ANOVA. Results: We found significant interaction and time effects on step length, stance time and swing time symmetry. Sb-only produced increased step length asymmetry at early adaptation compared to baseline (p < 0.0001) and an after-effect with increased asymmetry at early post-adaptation compared to baseline (p < 0.0001). Exo-only increased step length asymmetry (in the opposite direction as Sb-only) at early adaptation compared to baseline (p = 0.0392) but did not influence the participants sufficiently to result in a post-effect. Exo-Sb produced similar changes in step length asymmetry in the same direction as Sb-only (p = 0.0014). However, in contrast to Sb-only there was no significant after-effect between early post-adaptation and baseline (p = 0.0885). Conclusion: The passive exosuit successfully diminished asymmetrical step length after-effects induced by the split-belt treadmill in Exo-Sb. These results support the passive exosuit's ability to alter walking gait patterns. [ABSTRACT FROM AUTHOR]

Adaptive human performance relies on the central nervous system to regulate the engagement of cognitive–motor resources as task demands vary. Despite numerous studies which employed a split-belt induced perturbation to examine biomechanical outcomes during locomotor adaptation, none concurrently examined the cerebral cortical dynamics to assess changes in mental workload. Additionally, while prior work suggests that optic flow provides critical information for walking regulation, a few studies have manipulated visual inputs during adaption to split-belt walking. This study aimed to examine the concurrent modulation of gait and Electroencephalography (EEG) cortical dynamics underlying mental workload during split-belt locomotor adaptation, with and without optic flow. Thirteen uninjured participants with minimal inherent walking asymmetries at baseline underwent adaptation, while temporal–spatial gait and EEG spectral metrics were recorded. The results revealed a reduction in step length and time asymmetry from early to late adaptation, accompanied by an elevated frontal and temporal theta power; the former being well corelated to biomechanical changes. While the absence of optic flow during adaptation did not affect temporal–spatial gait metrics, it led to an increase of theta and low-alpha power. Thus, as individuals adapt their locomotor patterns, the cognitive–motor resources underlying the encoding and consolidation processes of the procedural memory were recruited to acquire a new internal model of the perturbation. Also, when adaption occurs without optic flow, a further reduction of arousal is accompanied with an elevation of attentional engagement due to enhanced neurocognitive resources likely to maintain adaptive walking patterns. [ABSTRACT FROM AUTHOR]

Locomotion is a highly flexible process, requiring rapid changes to gait due to changes in the environment or goals. Here, we used a split-belt treadmill to examine how the central nervous system coordinates a novel gait pattern. Existing research has focused on summary measures, most often step lengths, when describing changes induced while walking on the split-belt treadmill and during subsequent aftereffects. Here, we asked how the nervous system adjusts individual joint motions and the coordination pattern of the legs when people walk with one leg moving at either 2×, 3×, or 4× the speed of the other leg. We found that relative to tied-belt walking, split-belt perturbations change the timing relationships between the legs while most joint angle peaks and range of motion change little. The kinematic changes over the course of adaptation (i.e., from the beginning to end of a single split-belt walking bout) were subtle, particularly when comparing individual joint motions. The magnitude of the belt speed differences impacted intralimb coordination but did not produce consistent differences in most other measures. Most significant changes in kinematics occurred in the fast leg. Overall, interlimb timing changes drove a large proportion of the differences observed between tied-belt and split-belt gaits. Thus, it appears that the central nervous system can produce novel gait patterns through changes in coordination between legs that lead to new configurations at significant time points. These patterns can use within-limb and within-joint patterns that closely resemble those of normal walking. NEW & NOTEWORTHY We studied how the nervous system coordinates limb movements during asymmetric gait. Using a split-belt treadmill, we found that most changes in motion occurred when comparing motions between limbs, rather than among joints within a limb. Individual joint patterns resembled speed-matched comparisons, but this meant that joint movements became asymmetric during split-belt walking. These findings demonstrate that the nervous system can use consistent joint motions that are reconfigured in time to achieve new gait patterns. [ABSTRACT FROM AUTHOR]

Abstract Introduction Split-belt treadmill training has been used to assist with gait rehabilitation following stroke. This method modifies a patient’s step length asymmetry by adjusting left and right tread speeds individually during training. However, current split-belt training approaches pay little attention to the individuality of patients by applying set tread speed ratios (e.g., 2:1 or 3:1). This generalization results in unpredictable step length adjustments between the legs. To customize the training, this study explores the capabilities of a live feedback system that modulates split-belt tread speeds based on real-time step length asymmetry. Materials and methods Fourteen healthy individuals participated in two 1.5-h gait training sessions scheduled 1 week apart. They were asked to walk on the Computer Assisted Rehabilitation Environment (CAREN) split-belt treadmill system with a boot on one foot to impose asymmetrical gait patterns. Each training session consisted of a 3-min baseline, 10-min baseline with boot, 10-min feedback with boot (6% asymmetry exaggeration in the first session and personalized in the second), 5-min post feedback with boot, and 3-min post feedback without boot. A proportional-integral (PI) controller was used to maintain a specified step-length asymmetry by changing the tread speed ratios during the 10-min feedback period. After the first session, a linear model between baseline asymmetry exaggeration and post-intervention asymmetry improvement was utilized to develop a relationship between target exaggeration and target post-intervention asymmetry. In the second session, this model predicted a necessary target asymmetry exaggeration to replace the original 6%. This prediction was intended to result in a highly symmetric post-intervention step length. Results and discussion Eleven out of 14 participants (78.6%) developed a successful relationship between asymmetry exaggeration and decreased asymmetry in the post-intervention period of the first session. Seven out of the 11 participants (63.6%) in this successful correlation group had second session post-intervention asymmetries of

Walking patterns are adaptable in response to different environmental demands, which requires neural input from spinal and supraspinal structures. With an increase in age, there are changes in walking adaptation and in the neural control of locomotion, but the age-related changes in the neural control of locomotor adaptation is unclear. The purpose of this narrative review is to establish a framework where the age-related changes of neural control of human locomotor adaptation can be understood in terms of reactive feedback and predictive feedforward control driven by sensory feedback during locomotion. We parse out the effects of aging on (a) reactive adaptation to split-belt walking, (b) predictive adaptation to split-belt walking, (c) reactive visuomotor adaptation, and (d) predictive visuomotor adaptation, and hypothesize that specific neural circuits are influenced differentially with age, which influence locomotor adaptation. The differences observed in the age-related changes in walking adaptation across different locomotor adaptation paradigms will be discussed in light of the age-related changes in the neural mechanisms underlying locomotion. [ABSTRACT FROM AUTHOR]

Introduction: Split-belt treadmill training has been used to assist with gait rehabilitation following stroke. This method modifies a patient's step length asymmetry by adjusting left and right tread speeds individually during training. However, current split-belt training approaches pay little attention to the individuality of patients by applying set tread speed ratios (e.g., 2:1 or 3:1). This generalization results in unpredictable step length adjustments between the legs. To customize the training, this study explores the capabilities of a live feedback system that modulates split-belt tread speeds based on real-time step length asymmetry.Materials and Methods: Fourteen healthy individuals participated in two 1.5-h gait training sessions scheduled 1 week apart. They were asked to walk on the Computer Assisted Rehabilitation Environment (CAREN) split-belt treadmill system with a boot on one foot to impose asymmetrical gait patterns. Each training session consisted of a 3-min baseline, 10-min baseline with boot, 10-min feedback with boot (6% asymmetry exaggeration in the first session and personalized in the second), 5-min post feedback with boot, and 3-min post feedback without boot. A proportional-integral (PI) controller was used to maintain a specified step-length asymmetry by changing the tread speed ratios during the 10-min feedback period. After the first session, a linear model between baseline asymmetry exaggeration and post-intervention asymmetry improvement was utilized to develop a relationship between target exaggeration and target post-intervention asymmetry. In the second session, this model predicted a necessary target asymmetry exaggeration to replace the original 6%. This prediction was intended to result in a highly symmetric post-intervention step length.Results and Discussion: Eleven out of 14 participants (78.6%) developed a successful relationship between asymmetry exaggeration and decreased asymmetry in the post-intervention period of the first session. Seven out of the 11 participants (63.6%) in this successful correlation group had second session post-intervention asymmetries of < 3.5%.Conclusions: The use of a PI controller to modulate split-belt tread speeds demonstrated itself to be a viable method for individualizing split-belt treadmill training. [ABSTRACT FROM AUTHOR]

Gait dysfunction and fall risk have been well documented in people with Alzheimer's Disease (AD) and individuals with mild cognitive impairment (MCI). Normal locomotor adaptation may be an important prerequisite for normal and safe community walking function, especially in older adults with age-related neural, musculoskeletal, or cardiovascular changes and cognitive impairments. The split-belt walking task is a well-studied and robust method to evaluate locomotor adaptation (e.g., the ability to adjust stepping movements to changing environmental demands). Here, we capitalized on the split-belt adaptation task to test our hypothesis that a decreased capacity for locomotor adaptation may be an important contributing factor and indicator of increased fall risk and cognitive decline in older individuals with MCI and AD. The objectives of this study were to (1) compare locomotor adaptation capacity in MCI and AD compared to healthy older adults (HOA) during split-belt treadmill walking, and (2) evaluate associations between locomotor adaptation and cognitive impairments. Our results demonstrated a significant decrease in split-belt locomotor adaptation magnitude in older individuals with MCI and AD compared to HOA. In addition, we found significant correlations between the magnitude of early adaptation and de-adaptation vs. cognitive test scores, demonstrating that individuals with greater cognitive impairment also display a reduced capacity to adapt their walking in response to the split-belt perturbation. Our study takes an important step toward understanding mechanisms underlying locomotor dysfunction in older individuals with cognitive impairment. [ABSTRACT FROM AUTHOR]

Gait dysfunction and fall risk have been well documented in people with Alzheimer's Disease (AD) and individuals with mild cognitive impairment (MCI). Normal locomotor adaptation may be an important prerequisite for normal and safe community walking function, especially in older adults with age-related neural, musculoskeletal, or cardiovascular changes and cognitive impairments. The split-belt walking task is a well-studied and robust method to evaluate locomotor adaptation (e.g., the ability to adjust stepping movements to changing environmental demands). Here, we capitalized on the split-belt adaptation task to test our hypothesis that a decreased capacity for locomotor adaptation may be an important contributing factor and indicator of increased fall risk and cognitive decline in older individuals with MCI and AD. The objectives of this study were to (1) compare locomotor adaptation capacity in MCI and AD compared to healthy older adults (HOA) during split-belt treadmill walking, and (2) evaluate associations between locomotor adaptation and cognitive impairments. Our results demonstrated a significant decrease in split-belt locomotor adaptation magnitude in older individuals with MCI and AD compared to HOA. In addition, we found significant correlations between the magnitude of early adaptation and de-adaptation vs. cognitive test scores, demonstrating that individuals with greater cognitive impairment also display a reduced capacity to adapt their walking in response to the split-belt perturbation. Our study takes an important step toward understanding mechanisms underlying locomotor dysfunction in older individuals with cognitive impairment.

Background: Gait impairments are common in healthy older adults (HOA) and people with Parkinson's disease (PwPD), especially when adaptations to the environment are required. Traditional rehabilitation programs do not typically address these adaptive gait demands in contrast to repeated gait perturbation training (RGPT). RGPT is a novel reactive form of gait training with potential for both short and long-term consolidation in HOA and PwPD. The aim of this systematic review with meta-analysis is to determine whether RGPT is more effective than non-RGPT gait training in improving gait and balance in HOA and PwPD in the short and longer term. Methods: This review was conducted according to the PRISMA-guidelines and pre-registered in the PROSPERO database (CRD42020183273). Included studies tested the effects of any form of repeated perturbations during gait in HOA and PwPD on gait speed, step or stride length. Studies using balance scales or sway measures as outcomes were included in a secondary analysis. Effects of randomized controlled trials (RCT) on RGPT were pooled using a meta-analysis of final measures. Results: Of the 4421 studies, eight studies were deemed eligible for review, of which six could be included in the meta-analysis, totaling 209 participants (159 PwPD and 50 HOA). The studies were all of moderate quality. The meta-analysis revealed no significant effects of RGPT over non-RGPT training on gait performance (SMD = 0.16; 95% CI = −0.18, 0.49; Z = 0.92; P = 0.36). Yet, in some individual studies, favorable effects on gait speed, step length and stride length were observed immediately after the intervention as well as after a retention period. Gait variability and asymmetry, signifying more direct outcomes of gait adaptation, also indicated favorable RGPT effects in some individual studies. Conclusion: Despite some promising results, the pooled effects of RGPT on gait and balance were not significantly greater as compared to non-RGPT gait training in PwPD and HOA. However, these findings could have been driven by low statistical power. Therefore, the present review points to the imperative to conduct sufficiently powered RCT's to verify the true effects of RGPT on gait and balance in HOA and PwPD. Systematic Review Registration: https://www.crd.york.ac.uk/prospero/display%5frecord.php? Identifier: CRD42020183273. [ABSTRACT FROM AUTHOR]

Background: Gait impairments are common in healthy older adults (HOA) and people with Parkinson's disease (PwPD), especially when adaptations to the environment are required. Traditional rehabilitation programs do not typically address these adaptive gait demands in contrast to repeated gait perturbation training (RGPT). RGPT is a novel reactive form of gait training with potential for both short and long-term consolidation in HOA and PwPD. The aim of this systematic review with meta-analysis is to determine whether RGPT is more effective than non-RGPT gait training in improving gait and balance in HOA and PwPD in the short and longer term.Methods: This review was conducted according to the PRISMA-guidelines and pre-registered in the PROSPERO database (CRD42020183273). Included studies tested the effects of any form of repeated perturbations during gait in HOA and PwPD on gait speed, step or stride length. Studies using balance scales or sway measures as outcomes were included in a secondary analysis. Effects of randomized controlled trials (RCT) on RGPT were pooled using a meta-analysis of final measures.Results: Of the 4421 studies, eight studies were deemed eligible for review, of which six could be included in the meta-analysis, totaling 209 participants (159 PwPD and 50 HOA). The studies were all of moderate quality. The meta-analysis revealed no significant effects of RGPT over non-RGPT training on gait performance (SMD = 0.16; 95% CI = −0.18, 0.49; Z = 0.92; P = 0.36). Yet, in some individual studies, favorable effects on gait speed, step length and stride length were observed immediately after the intervention as well as after a retention period. Gait variability and asymmetry, signifying more direct outcomes of gait adaptation, also indicated favorable RGPT effects in some individual studies.Conclusion: Despite some promising results, the pooled effects of RGPT on gait and balance were not significantly greater as compared to non-RGPT gait training in PwPD and HOA. However, these findings could have been driven by low statistical power. Therefore, the present review points to the imperative to conduct sufficiently powered RCT's to verify the true effects of RGPT on gait and balance in HOA and PwPD.Systematic Review Registration:https://www.crd.york.ac.uk/prospero/display_record.php? Identifier: CRD42020183273.

Humans continuously adapt locomotor patterns. Whether metabolic cost reduction is the primary objective or a by-product of the observed biomechanical changes during adaptation is not known. The main goal of our study is to determine if perception of task duration affects the adaptation of locomotor patterns to reduce energetic cost during split-belt walking. We tested the hypothesis that individuals who believe they will sustain a locomotor adaptation task for a prolonged time will reduce metabolic cost by adapting toward a walking pattern associated with lower mechanical work. N=14 participants walked on a split-belt treadmill for 10 minutes with knowledge of task duration (group K), while N=15 participants performed the task under the assumption that they would walk for 30 minutes (group U). Both groups walked for 10 minutes with the belts moving at 1.5 and 0.5 m/s, followed by 6 minutes of walking with both belts at 1.0 m/s. We observed a significant main effect of Time (p<0.001, observed power 1.0) and the interaction of Time×Group (p=0.004, observed power 0.84) on metabolic cost. Participants in the U group had a metabolic cost that was 12% lower during adaptation compared to the K group, which did not reduce metabolic cost during adaptation. The metabolic cost reduction observed in group U was not associated with biomechanical changes during adaptation. Our results indicate that metabolic cost reduction has a primary role in tasks that need to be sustained for a prolonged time, and this reduction is not only related to biomechanical factors.

Locomotor adaptation to abrupt and gradual perturbations are likely driven by fundamentally different neural processes. The aim of this study was to quantify brain dynamics associated with gait adaptation to a gradually introduced gait perturbation, which typically results in smaller behavioral errors relative to an abrupt perturbation. Loss of balance during standing and walking elicits transient increases in midfrontal theta oscillations that have been shown to scale with perturbation intensity. We hypothesized there would be no significant change in anterior cingulate theta power (4-7 Hz) with respect to pre-adaptation when a gait perturbation is introduced gradually because the gradual perturbation acceleration and stepping kinematic errors are small relative to an abrupt perturbation. Using mobile electroencephalography (EEG), we measured gait-related spectral changes near the anterior cingulate, posterior cingulate, sensorimotor, and posterior parietal cortices as young, neurotypical adults ( n  = 30) adapted their gait to an incremental split-belt treadmill perturbation. Most cortical clusters we examined (>70%) did not exhibit changes in electrocortical activity between 2-50 Hz. However, we did observe gait-related theta synchronization near the left anterior cingulate cortex during strides with the largest errors, as measured by step length asymmetry. These results suggest gradual adaptation with small gait asymmetry and perturbation magnitude may not require significant cortical resources beyond normal treadmill walking. Nevertheless, the anterior cingulate may remain actively engaged in error monitoring, transmitting sensory prediction error information via theta oscillations., Competing Interests: The authors declare there are no competing interests., (©2024 Jacobsen et al.)

Rhythmic limb movements during locomotion are controlled by central pattern generator (CPG) circuits located in the spinal cord. It is considered that these circuits are composed of individual rhythm generators (RGs) for each limb interacting with each other through multiple commissural and long propriospinal circuits. The organization and operation of each RG are not fully understood, and different competing theories exist about interactions between its flexor and extensor components, as well as about left–right commissural interactions between the RGs. The central idea of circuit organization proposed in this study is that with an increase of excitatory input to each RG (or an increase in locomotor speed) the rhythmogenic mechanism of the RGs changes from “flexor-driven” rhythmicity to a “classical half-center” mechanism. We test this hypothesis using our experimental data on changes in duration of stance and swing phases in the intact and spinal cats walking on the ground or tied-belt treadmills (symmetric conditions) or split-belt treadmills with different left and right belt speeds (asymmetric conditions). We compare these experimental data with the results of mathematical modeling, in which simulated CPG circuits operate in similar symmetric and asymmetric conditions with matching or differing control drives to the left and right RGs. The obtained results support the proposed concept of state-dependent changes in RG operation and specific commissural interactions between the RGs. The performed simulations and mathematical analysis of model operation under different conditions provide new insights into CPG network organization and limb coordination during locomotion.

Rhythmic limb movements during locomotion are controlled by central pattern generator (CPG) circuits located in the spinal cord. It is considered that these circuits are composed of individual rhythm generators (RGs) for each limb interacting with each other through multiple commissural and long propriospinal circuits. The organization and operation of each RG are not fully understood, and different competing theories exist about interactions between its flexor and extensor components, as well as about left–right commissural interactions between the RGs. The central idea of circuit organization proposed in this study is that with an increase of excitatory input to each RG (or an increase in locomotor speed) the rhythmogenic mechanism of the RGs changes from "flexor-driven" rhythmicity to a "classical half-center" mechanism. We test this hypothesis using our experimental data on changes in duration of stance and swing phases in the intact and spinal cats walking on the ground or tied-belt treadmills (symmetric conditions) or split-belt treadmills with different left and right belt speeds (asymmetric conditions). We compare these experimental data with the results of mathematical modeling, in which simulated CPG circuits operate in similar symmetric and asymmetric conditions with matching or differing control drives to the left and right RGs. The obtained results support the proposed concept of state-dependent changes in RG operation and specific commissural interactions between the RGs. The performed simulations and mathematical analysis of model operation under different conditions provide new insights into CPG network organization and limb coordination during locomotion. [ABSTRACT FROM AUTHOR]

Split-belt treadmills that move the legs at different speeds are thought to update internal representations of the environment, such that this novel condition generates a new locomotor pattern with distinct spatio-temporal features compared to those of regular walking. It is unclear the degree to which such recalibration of movements in the spatial and temporal domains is interdependent. In this study, we explicitly altered subjects' limb motion in either space or time during split-belt walking to determine its impact on the adaptation of the other domain. Interestingly, we observed that motor adaptation in the spatial domain was susceptible to altering the temporal domain, whereas motor adaptation in the temporal domain was resilient to modifying the spatial domain. This non-reciprocal relation suggests a hierarchical organization such that the control of timing in locomotion has an effect on the control of limb position. This is of translational interest because clinical populations often have a greater deficit in one domain compared to the other. Our results suggest that explicit changes to temporal deficits cannot occur without modifying the spatial control of the limb.

Treadmills used for gait training in clinical rehabilitation and experimental settings are commonly fitted with handrails to assist or support persons in locomotor tasks. However, the effects of balance support through handrail holding on locomotor learning are unknown. Locomotor learning can be studied on split-belt treadmills, where participants walk on two parallel belts with asymmetric left and right belt speeds, to which they adapt their stepping pattern within a few minutes. The aim of this study was to determine how handrail holding affects the walking pattern during split-belt adaptation and after-effects in able-bodied persons. Fifty healthy young participants in five experimental groups were instructed to hold handrails, swing arms freely throughout the experiment or hold handrails during adaptation and swing arms freely during after-effects. Step length asymmetry and double support asymmetry were measured to assess the spatiotemporal walking pattern. The results showed that holding handrails during split-belt adaptation reduces magnitude of initial perturbation of step length asymmetry and reduces after-effects in step length asymmetry upon return to symmetric belt speeds. The findings of this study imply that balance support during gait training reduces locomotor learning, which should be considered in daily clinical gait practice and future research on locomotor learning. [ABSTRACT FROM AUTHOR]

Split-belt treadmills that move the legs at different speeds are thought to update internal representations of the environment, such that this novel condition generates a new locomotor pattern with distinct spatio-temporal features compared to those of regular walking. It is unclear the degree to which such recalibration of movements in the spatial and temporal domains is interdependent. In this study, we explicitly altered subjects' limb motion in either space or time during split-belt walking to determine its impact on the adaptation of the other domain. Interestingly, we observed that motor adaptation in the spatial domain was susceptible to altering the temporal domain, whereas motor adaptation in the temporal domain was resilient to modifying the spatial domain. This non-reciprocal relation suggests a hierarchical organization such that the control of timing in locomotion has an effect on the control of limb position. This is of translational interest because clinical populations often have a greater deficit in one domain compared to the other. Our results suggest that explicit changes to temporal deficits cannot occur without modifying the spatial control of the limb. [ABSTRACT FROM AUTHOR]

Abstract Clinical locomotor research seeks to facilitate adaptation or retention of new walking patterns by providing feedback. Within a split-belt treadmill paradigm, sagittal plane feedback improves adaptation but does not affect retention. Representation of error in this manner is cognitively demanding. However, it is unknown in this paradigm how frontal plane feedback, which may utilize a unique learning process, impacts locomotor adaptation. Frontal plane movement feedback has been shown to impact retention of novel running mechanics but has yet to be evaluated in gait conditions widely applicable within neurorehabilitation, such as walking. The purpose of this study was to investigate the effects of frontal plane mirror feedback on gait adaptation and retention during split-belt treadmill walking. Forty healthy young adults were divided into two groups: one group received mirror feedback during the first split-belt exposure and the other received no mirror feedback. Individuals in the mirror feedback group were asked to look at their legs in the mirror, but no further instructions were given. Individuals with mirror feedback displayed more symmetric stance time during the first strides of adaptation and maintained this pattern into the second split-belt exposure when no feedback was provided. Individuals with mirror feedback also demonstrated more symmetric double support time upon returning to normal walking. Lastly, the mirror feedback also allowed individuals to walk with smaller gait variability during the final steps of both split-belt exposures. Overall, mirror feedback allowed individuals to reduce their stance time asymmetry and led to a more consistent adapted pattern, suggesting this type of feedback may have utility in gait training that targets symmetry and consistency in movement. [ABSTRACT FROM AUTHOR]

Task- and phase-dependent reflex modulation during locomotion is well established, but we do not know the signals driving this modulation. To determine whether signals related to left-right symmetry of the locomotor pattern modulate cutaneous reflexes, we stimulated the superficial peroneal nerve in five intact female cats and in four spinal-transected cats (spinal cats, two males and two females) during split-belt locomotion at different left-right speeds. We compared cutaneous reflexes evoked in three ipsilateral and two contralateral hindlimb muscles during split-belt locomotion with those evoked during tied-belt (equal left-right speeds) locomotion at matched speeds of the slow and fast limbs. Our results showed similar phase-dependent modulation of cutaneous reflexes during tied-belt and split-belt locomotion in intact and spinal cats. During tied-belt locomotion in intact cats, an increase in speed significantly increased reflex modulation from minimum to maximum values, whereas in spinal cats, we observed a significant decrease. However, in all muscles of intact and spinal cats, split-belt locomotion significantly reduced reflex modulation compared with tied-belt locomotion independently of which limb was stepping on the slow or fast belt. Additionally, reflex modulation correlated more with spatial left-right symmetry, as opposed to a temporal one, in intact and spinal cats. Our results indicate that signals related to left-right symmetry reduce cutaneous reflex modulation independendy of speed via a spinal mechanism. We propose that asymmetric sensory feedback from the left and right legs alters the state of the spinal network, thereby reducing cutaneous reflexes to prevent inputs from destabilizing a potentially unstable pattern. [ABSTRACT FROM AUTHOR]

Gait adaptation is a task that requires fine-tuned coordination of all degrees of freedom in the lower limbs by the central nervous system. However, when individuals change their gait it is unknown how this coordination is organized, and how it can be influenced by contextual interference during practice. Such knowledge could provide information about measurement of gait adaptation during rehabilitation. Able-bodied individuals completed an acute bout of asymmetric split-belt treadmill walking, where one limb was driven at a constant velocity and the other according to one of three designed practice paradigms: serial practice, where the variable limb belt velocity increased over time; random blocked practice, where every 20 strides the variable limb belt velocity changed randomly; random practice, where every stride the variable limb belt velocity changed randomly. On the second day, subjects completed one of two different transfer tests; one with a belt asymmetry close to that experienced on the acquisition day (transfer 1; 1.5:1), and one with a greater asymmetry (transfer 2; 2:1) . To reduce this inherently high-dimensional dataset, principal component analyses were used for kinematic data collected throughout the acquisition and transfer phases; resulting in extraction of the first two principal components (PCs). For acquisition, PC1 and PC2 were related to sagittal and frontal plane control. For transfer 1, PC1 and PC2 were related to frontal plane control of the base of support and whole-body center of mass. For transfer 2, PC1 did not have any variables with high enough coefficients deemed to be relevant, and PC2 was related to sagittal plane control. Observations of principal component scores indicate that variance structuring differs among practice groups during acquisition and transfer 1, but not transfer 2. These results demonstrate the main kinematic coordinative structures that exist during gait adaptation, and that control of sagittal plane and frontal plane motion are perhaps a trade-off during acquisition of a novel asymmetric gait pattern. [ABSTRACT FROM AUTHOR]

Very little is known about the effects of specific practice on motor learning of predictive balance control during novel bipedal gait. This information could provide an insight into how the direction and magnitude of predictive errors during acquisition of a novel gait task influence transfer of balance control, as well as yield a practice protocol for the restoration of balance for those with locomotor impairments. This study examined the effect of a variable practice paradigm on transfer of a novel asymmetric gait pattern in able-bodied individuals. Using a split-belt treadmill, one limb was driven at a constant velocity (constant limb) and the other underwent specific changes in velocity (variable limb) during practice according to one of three prescribed practice paradigms: serial, where the variable limb velocity increased linearly; random blocked, where variable limb underwent random belt velocity changes every 20 strides; and random practice, where the variable limb underwent random step-to-step changes in velocity. Random practice showed the highest balance control variability during acquisition compared to serial and random blocked practice which demonstrated the best transfer of balance control on one transfer test. Both random and random blocked practices showed significantly less balance control variability during a second transfer test compared to serial practice. These results indicate that random blocked practice may be best for generalizability of balance control while learning a novel gait, perhaps, indicating that individuals who underwent this practice paradigm were able to find the most optimal balance control solution during practice. [ABSTRACT FROM AUTHOR]

Locomotor adaptation is commonly studied using split-belt treadmill walking, in which each foot is placed on a belt moving at a different speed. As subjects adapt to split-belt walking, they reduce metabolic power, but the biomechanical mechanism behind this improved efficiency is unknown. Analyzing mechanical work performed by the legs and joints during split-belt adaptation could reveal this mechanism. Because ankle work in the step-to-step transition is more efficient than hip work, we hypothesized that control subjects would reduce hip work on the fast belt and increase ankle work during the step-to-step transition as they adapted. We further hypothesized that subjects with unilateral, trans-tibial amputation would instead increase propulsive work from their intact leg on the slow belt. Control subjects reduced hip work and shifted more ankle work to the step-to-step transition, supporting our hypothesis. Contrary to our second hypothesis, intact leg work, ankle work and hip work in amputees were unchanged during adaptation. Furthermore, all subjects increased collisional energy loss on the fast belt, but did not increase propulsive work. This was possible because subjects moved further backward during fast leg single support in late adaptation than in early adaptation, compensating by reducing backward movement in slow leg single support. In summary, subjects used two strategies to improve mechanical efficiency in split-belt walking adaptation: a CoM displacement strategy that allows for less forward propulsion on the fast belt; and, an ankle timing strategy that allows efficient ankle work in the step-to-step transition to increase while reducing inefficient hip work. [ABSTRACT FROM AUTHOR]

Background: The pathophysiology behind gait impairments seen in Parkinson's disease (PD), in particular freezing of gait (FOG), is not fully understood. Here we study the interplay between several gait features related to FOG during different split-belt treadmill (SBTM) conditions.Methods: We investigated the spatiotemporal properties, the phenomenon of sequence effect and the inter-limb symmetry and temporal coordination of gait during different split-belt conditions in 20 patients with advanced Parkinson's disease and different severities of freezing. Subjects were tested in four belt configurations: tied, split while reducing the velocity of leg with the shorter (worst side reduction, WSR) and longer (best side reduction, BSR) step length, and tied again to measure the after-effect.Results: We found that in spite of an improvement of spatial symmetry, the BSR led to a worsening of coordination (i.e. the left-right anti-phased stepping) and an increased sequence effect (i.e. progressive shortening of the step length). By contrast, in spite of a worsened spatial symmetry, WSR improved inter-limbs coordination and reduced the sequence effect. After prolonged split-belt walking gait was differently modulated according to the reduction of the best or worst leg velocity: BSR led to positive after effects in symmetry, bilateral coordination and sequence effect.Conclusions: These findings support the hypothesis that the irregularity of inter-limb coordination and defective amplitude generation leading to sequence effect might be coupled and result from the same maladaptive motor behavior. Furthermore, our results show that SBTM can be an effective tool to improve parkinsonian gait. [ABSTRACT FROM AUTHOR]

The hip and ankle strategies that affect learning of a novel gait have not been fully determined, and could be of importance in design of clinical gait interventions. The authors' purpose was to determine the effects of asymmetric split-belt treadmill walking on ankle and hip work during propulsion. Participants were randomized into either a gradual training group or a sudden training group and later returned for a retention test. The gradual training group performed significantly more work at the hip joint of the slow limb during acquisition, and decreased the hip joint work performed during retention. These findings reveal the hip joint on the slow limb during initial swing as a possible site of adaptation to a novel locomotor pattern. [ABSTRACT FROM PUBLISHER]

Stepping along curvilinear paths produces speed differences between the inner and outer limb(s). This can be reproduced experimentally by independently controlling left and right speeds with split-belt locomotion. Here we provide additional details on the pattern of the four limbs during quadrupedal split-belt locomotion in intact cats. Six cats performed tied-belt locomotion (same speed bilaterally) and split-belt locomotion where one side (constant side) stepped at constant treadmill speed while the other side (varying side) stepped at several speeds. Cycle, stance, and swing durations changed in parallel in homolateral limbs with shorter and longer stance and swing durations on the fast side, respectively, compared with the slow side. Phase variations were quantified in all four limbs by measuring the slopes of the regressions between stance and cycle durations (rSTA) and between swing and cycle durations (rSW). For a given limb, rSTA and rSW were not significantly different from one another on the constant side whereas on the varying side rSTA increased relative to tied-belt locomotion while rSW became more negative. Phase variations were similar for homolateral limbs. Increasing left-right speed differences produced a large increase in homolateral double support on the slow side, while triple-support periods decreased. Increasing left-right speed differences altered homologous coupling, homolateral coupling on the fast side, and coupling between the fast hindlimb and slow forelimb. Results indicate that homolateral limbs share similar control strategies, only certain features of the interlimb pattern adjust, and spinal locomotor networks of the left and right sides are organized symmetrically. [ABSTRACT FROM AUTHOR]

Faster relearning of an external perturbation, savings, offers a behavioral linkage between motor learning and memory. To explain savings effects in reaching adaptation experiments, recent models suggested the existence of multiple learning components, each shows different learning and forgetting properties that may change following initial learning. Nevertheless, the existence of these components in rhythmic movements with other effectors, such as during locomotor adaptation, has not yet been studied. Here, we study savings in locomotor adaptation in two experiments; in the first, subjects adapted to speed perturbations during walking on a split-belt treadmill, briefly adapted to a counter-perturbation and then readapted. In a second experiment, subjects readapted after a prolonged period of washout of initial adaptation. In both experiments we find clear evidence for increased learning rates (savings) during readaptation. We show that the basic error-based multiple timescales linear state space model is not sufficient to explain savings during locomotor adaptation. Instead, we show that locomotor adaptation leads to changes in learning parameters, so that learning rates are faster during readaptation. Interestingly, we find an intersubject correlation between the slow learning component in initial adaptation and the fast learning component in the readaptation phase, suggesting an underlying mechanism for savings. Together, these findings suggest that savings in locomotion and in reaching may share common computational and neuronal mechanisms; both are driven by the slow learning component and are likely to depend on cortical plasticity. [ABSTRACT FROM AUTHOR]

[Display omitted] • The utility of SSA to decompose and assess motor adaptation time series is tested. • Trend reconstruction performance of SSA was assessed and compared with filter and fitting methods. • SSA reconstructed the simulated global trends more accurately than the filter and fitting methods. • SSA was able to reconstruct global trends from idiosyncratic split-belt adaptation time series. Motor adaptation is a form of learning to re-establish desired movements in novel situations. To probe motor adaptation, one can replicate such conditions experimentally by imposing a sustained perturbation during movement. Exposure to such perturbations initially causes an abrupt change in relevant performance variables, followed by a gradual return to baseline behaviour. The resulting time series exhibit persistent properties related to structural changes in underlying motor control and transitory properties related to trial-to-trial variations. The global trend, signifying the structural change, is often assessed by averaging the time series in predefined bins or nonlinear model fitting. However, these methods to study motor adaptation require a priori decisions to produce accurate fits. Here, we test a data-driven approach called Singular Spectrum Analysis (SSA) to assess the global trend. In SSA, we first decompose the adaptation time series into components that represent either a global trend or additional variations, and secondly , select the component(s) corresponding to the global trend using spectral analysis. In this paper, we will use simulated data to compare the reconstruction performance of SSA with often applied filter and fitting methods in motor adaptation studies and apply SSA to real data obtained during split-belt adaptation. In the simulations, we show that SSA reconstructed the fast-initial component and entire global trends more accurately than the filtering and fitting methods. In addition, we show that SSA also successfully reconstructed global trends from real data. Therefore, the SSA might be useful in motor learning studies to decompose and assess adaptation time series. [ABSTRACT FROM AUTHOR]

Let the machine do the work: learning to reduce the energetic cost of walking on a split-belt treadmill In recent decades, there have been numerous attempts to exploit this phenomenon in the form of wearable devices like powered exoskeletons, which can reduce the energy cost of walking (e.g. Malcolm I et al i . Sánchez I et al i .'s study demonstrates that humans can quickly learn to take advantage of external mechanical work, and that the learning process is driven by an attempt to reduce energy cost. [Extracted from the article]

Gait adaptation is crucial for adults at risk for mobility disability, and executive function and physical function may be important for adaptation performance. Gait adaptation can be measured using a treadmill with two belts, known as a split-belt treadmill. Increasing evidence supports that gait adaptability, executive function, and physical function are interrelated in older adults. The purpose of this study was to determine if: a) executive function and measures of relative effort of the ankle and knee relate to split-belt treadmill adaptation; b) older adults classified as fast adapters display differences in relative effort, executive function, and propulsive impulse (push-off) compared to slow adapters; and c) spatial and temporal control differ between individuals with faster rate of adaptation compared to those with slower rates of adaptation. Greater effort of the knee on the slow belt was related to faster early adaptation (r = 0.650, p = 0.005) indicating its importance for adapting quickly to the perturbation. We did not observe a relationship between cognitive tests and adaptation performance. We did not detect any statistical differences in cognitive tests performance, push-off, spatial or temporal control between fast adapters compared to slow adapters. Our results suggest that in older adults at risk for mobility disability, higher effort at the knee is important for early split-belt adaptation. • Greater effort generated at the knee relates to faster adaptation. • Physical but not cognitive function influences adaptation in older adults. • Higher-order cognitive functions do not relate to split-belt adaptation. [ABSTRACT FROM AUTHOR]

Human movements are flexible as they continuously adapt to changes in the environment. The recalibration of corrective responses to sustained perturbations (e.g., constant force) altering one's movement contributes to this flexibility. We asked whether the recalibration of corrective actions involve cerebral structures using stroke as a disease model. We characterized changes in muscle activity in stroke survivors and control subjects before, during, and after walking on a split-belt treadmill moving the legs at different speeds. The recalibration of corrective muscle activity was comparable between stroke survivors and control subjects, which was unexpected given the known deficits in feedback responses poststroke. Also, the intact recalibration in stroke survivors contrasted their limited ability to adjust their muscle activity during steady-state split-belt walking. Our results suggest that the recalibration and execution of motor commands are partially dissociable: cerebral lesions interfere with the execution, but not the recalibration, of motor commands on novel movement demands., (Copyright © 2020 de Kam et al.)

Background: Persons with anterior cruciate ligament reconstruction (ACLR) show deficits in gait and neuromuscular control following rehabilitation. This altered behavior extends to locomotor adaptation and learning, however the contributing factors to this observed behavior have yet to be investigated., Objective: The purpose of this study was to assess differences in locomotor adaptation and learning between ACLR and controls, and identify underlying contributors to motor adaptation in these individuals., Methods: Twenty ACLR individuals and 20 healthy controls (CON) agreed to participate in this study. Participants performed four cognitive and dexterity tasks (local version of Trail Making Test, reaction time test, electronic pursuit rotor test, and the Purdue pegboard). Three-dimensional kinematics were also collected while participants walked on a split-belt treadmill., Results: ACLR individuals completed the local versions of Trails A and Trails B significantly faster than CON. During split-belt walking, ACLR individuals demonstrated smaller step length asymmetry during EARLY and LATE adaptation, smaller double support asymmetry during MID adaptation, and larger stance time asymmetry during DE-ADAPT compared with CON., Conclusions: ACLR individuals performed better during tasks that required visual attention and task switching and were less perturbed during split-belt walking compared to controls. Persons with ACLR may use different strategies than controls, cognitive or otherwise, to adapt locomotor patterns.

Background: Despite the strong implications for rehabilitation design, the capability of individuals with anterior cruciate ligament reconstruction (ACLR) to adapt and store novel gait patterns have not been well studied., Purpose: To investigate how reconstructive surgery may affect the ability to adapt and store novel gait patterns in persons with ACLR while walking on a split-belt treadmill., Study Design: Controlled laboratory study., Methods: Gait adaptation was compared between 20 participants with ACLR and 20 healthy controls during split-belt treadmill walking. Gait adaptation was assessed in slow- and fast-adapting parameters by (1) the magnitude of symmetry during late adaptation and (2) the amount of the asymmetry during de-adaptation., Results: Healthy individuals adapted a new walking pattern and stored the new walking pattern equally in both the dominant and nondominant limbs. Conversely, individuals with ACLR displayed impairments in both slow-adapting and fast-adapting derived gait adaptation and significant differences in behavior between the reconstructed and uninjured limb., Conclusion: While surgical reconstruction and physical therapy are aimed at improving mechanical stability to the knee, the study data suggest that fundamental features of motor control remain altered. After ACLR, participants display an altered ability to learn and store functional gait patterns.

Increased practice difficulty during asymmetrical split-belt treadmill rehabilitation has been shown to improve gait outcomes during retention and transfer tests. However, research in this area has been limited by manual treadmill operation. In the case of variable practice, which requires stride-by-stride changes to treadmill belt velocities, the treadmill control must be automated. This paper presents a method for automation of asymmetrical split-belt treadmill walking, and evaluates how well this method performs with regards to timing of gait events. One participant walked asymmetrically for 100 strides, where the non-dominant limb was driven at their self-selected walking speed, while the other limb was driven randomly on a stride-by-stride basis. In the control loop, the key factors to insure that the treadmill belt had accelerated to its new velocity safely during the swing phase were the sampling rate of the A/D converter, processing time within the controller software, and acceleration of the treadmill belt. The combination of these three factors resulted in a total control loop time during each swing phase that satisfied these requirements with a factor of safety that was greater than 4. Further, a polynomial fit indicated that belt acceleration was the largest contributor to changes in this total time. This approach appears to be safe and reliable for stride-by-stride adjustment of treadmill belt speed, making it suitable for future asymmetrical split-belt walking studies. Further, it can be incorporated into virtual reality rehabilitation paradigms that utilize split-belt treadmill walking., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

Although significant effort is concentrated toward gait retraining during stroke rehabilitation; 33% of community-dwelling individuals following stroke continue to demonstrate gait asymmetries following participation in conventional rehabilitation. Recent studies utilizing the split-belt treadmill indicate that subjects after stroke retain the ability to learn a novel locomotor pattern. Through the use of error augmentation, this locomotor pattern can provide a temporary improvement in symmetry, which can be exploited through repetitive task specific locomotor training. This article reviews findings from this experimental paradigm in chronic stroke survivors and discusses the future questions to be addressed in order to provide optimal rehabilitation interventions., (Copyright © 2015 Elsevier Inc. All rights reserved.)