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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]

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]

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]

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]

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]

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]

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]

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]

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]