Reinforcement-based processes actively regulate motor exploration along redundant solution manifolds.

From a baby's babbling to a songbird practising a new tune, exploration is critical to motor learning. A hallmark of exploration is the emergence of random walk behaviour along solution manifolds, where successive motor actions are not independent but rather become serially dependent. Such exploratory random walk behaviour is ubiquitous across species' neural firing, gait patterns and reaching behaviour. The past work has suggested that exploratory random walk behaviour arises from an accumulation of movement variability and a lack of error-based corrections. Here, we test a fundamentally different idea-that reinforcement-based processes regulate random walk behaviour to promote continual motor exploration to maximize success. Across three human reaching experiments, we manipulated the size of both the visually displayed target and an unseen reward zone, as well as the probability of reinforcement feedback. Our empirical and modelling results parsimoniously support the notion that exploratory random walk behaviour emerges by utilizing knowledge of movement variability to update intended reach aim towards recently reinforced motor actions. This mechanism leads to active and continuous exploration of the solution manifold, currently thought by prominent theories to arise passively. The ability to continually explore muscle, joint and task redundant solution manifolds is beneficial while acting in uncertain environments, during motor development or when recovering from a neurological disorder to discover and learn new motor actions.

Effect of Increasing Obstacle Distances Task on Postural Stability Variables During Gait Initiation in Older Nonfallers and Fallers.

OBJECTIVE: To compare the scaling of the postural stability variables between older nonfallers and fallers during gait initiation (GI) while stepping over increasing obstacle distances. DESIGN: Cross-sectional study. SETTING: University research laboratory. PARTICIPANTS: A sample of participants (N=24) divided into 2 groups: older nonfallers (n=12) and older fallers (n=12). Participants had no known neurologic, musculoskeletal, or cardiovascular conditions that could have affected their walking, and all were independent walkers. All the participants had an adequate cognitive function to participate as indicated by a score of more than 24 on the Mini-Mental State Examination. INTERVENTIONS: Not applicable. MAIN OUTCOME MEASURES: The primary dependent variables were peak anterior-posterior (AP) center of mass (CoM)-center of pressure (CoP) separation during anticipatory postural adjustments (APAs), AP CoM-CoP separation at the toe-off, and peak AP CoM-CoP separation during the swing. Secondary dependent variables were AP trunk angle during GI. Within- and between-repeated measures analysis of variance was used to compare means between groups across different task conditions for all the dependent variables. RESULTS: There was a main effect of group for peak AP CoM-CoP separation during APA (P=.018), an interaction effect between group and condition for AP CoM-CoP separation at toe-off (P=.009), and a main effect of condition for peak AP CoM-CoP separation during the swing (P<.001). We also found a main effect of group for peak AP trunk angle during the swing (P=.028). CONCLUSIONS: For GI while stepping over increasing obstacle distances, older fallers adopt a more conservative strategy of AP CoM-CoP separation than nonfallers prior to toe-off and demonstrate increased peak AP trunk lean during the swing. AP CoM-CoP separation prior to toe-off during the GI task may be a critical marker to identify fallers and warrants additional investigation.

Age of First Exposure to Soccer Heading and Sensory Reweighting for Upright Stance.

US Soccer eliminated soccer heading for youth players ages 10 years and younger and limited soccer heading for children ages 11-13 years. Limited empirical evidence associates soccer heading during early adolescence with medium-to-long-term behavioral deficits. The purpose of this study was to compare sensory reweighting for upright stance between college-aged soccer players who began soccer heading ages 10 years and younger (AFE ≤ 10) and those who began soccer heading after age 10 (AFE > 10). Thirty soccer players self-reported age of first exposure (AFE) to soccer heading. Sensory reweighting was compared between AFE ≤ 10 and AFE > 10. To evaluate sensory reweighting, we simultaneously perturbed upright stance with visual, vestibular, and proprioceptive stimulation. The visual stimulus was presented at two different amplitudes to measure the change in gain to vision, an intra-modal effect; and change in gain to galvanic vestibular stimulus (GVS) and vibration, both inter-modal effects. There were no differences in gain to vision (p=0.857, η2=0.001), GVS (p=0.971, η2=0.000), or vibration (p=0.974, η2=0.000) between groups. There were no differences in sensory reweighting for upright stance between AFE ≤ 10 and AFE > 10, suggesting that soccer heading during early adolescence is not associated with balance deficits in college-aged soccer players, notwithstanding potential deficits in other markers of neurological function.

Sensory-Challenge Balance Exercises Improve Multisensory Reweighting in Fall-Prone Older Adults.

BACKGROUND AND PURPOSE: Multisensory reweighting (MSR) deficits in older adults contribute to fall risk. Sensory-challenge balance exercises may have value for addressing the MSR deficits in fall-prone older adults. The purpose of this study was to examine the effect of sensory-challenge balance exercises on MSR and clinical balance measures in fall-prone older adults. METHODS: We used a quasi-experimental, repeated-measures, within-subjects design. Older adults with a history of falls underwent an 8-week baseline (control) period. This was followed by an 8-week intervention period that included 16 sensory-challenge balance exercise sessions performed with computerized balance training equipment. Measurements, taken twice before and once after intervention, included laboratory measures of MSR (center of mass gain and phase, position, and velocity variability) and clinical tests (Activities-specific Balance Confidence Scale, Berg Balance Scale, Sensory Organization Test, Limits of Stability test, and lower extremity strength and range of motion). RESULTS: Twenty adults 70 years of age and older with a history of falls completed all 16 sessions. Significant improvements were observed in laboratory-based MSR measures of touch gain (P = 0.006) and phase (P = 0.05), Berg Balance Scale (P = 0.002), Sensory Organization Test (P = 0.002), Limits of Stability Test (P = 0.001), and lower extremity strength scores (P = 0.005). Mean values of vision gain increased more than those for touch gain, but did not reach significance. DISCUSSION AND CONCLUSIONS: A balance exercise program specifically targeting multisensory integration mechanisms improved MSR, balance, and lower extremity strength in this mechanistic study. These valuable findings provide the scientific rationale for sensory-challenge balance exercise to improve perception of body position and motion in space and potential reduction in fall risk.

A central processing sensory deficit with Parkinson's disease.

Parkinson's disease (PD) is a progressive degenerative disease manifested by tremor, rigidity, bradykinesia, and postural instability. Deficits in proprioceptive integration are prevalent in individuals with PD, even at early stages of the disease. These deficits have been demonstrated primarily during investigations of reaching. Here, we investigated how PD affects sensory fusion of multiple modalities during upright standing. We simultaneously perturbed upright stance with visual, vestibular, and proprioceptive stimulation, to understand how these modalities are reweighted so that overall feedback remains suited to stabilizing upright stance in individuals with PD. Eight individuals with PD stood in a visual cave with a moving visual scene at 0.2 Hz while an 80-Hz vibratory stimulus was applied bilaterally to their Achilles tendons (stimulus turns on-off at 0.28 Hz) and a ±1 mA bilateral monopolar galvanic stimulus was applied at 0.36 Hz. The visual stimulus was presented at different amplitudes (0.2°, 0.8° rotation about ankle axis) to measure: the change in gain (weighting) to vision, an intramodal effect; and a simultaneous change in gain to vibration and galvanic stimulation, both intermodal effects. Trunk/leg gain relative to vision decreased when visual amplitude was increased, reflecting an intramodal visual effect. In contrast, when vibration was turned on/off, leg gain relative to vision was equivalent in individuals with PD, indicating no reweighting of visual information when proprioception was disrupted through vibration (i.e., no intermodal effect). Trunk and leg angle gain relative to GVS also showed no reweighting in individuals with PD. These results are in contrast to previous results with healthy adults, who showed clear intermodal effects in the same paradigm, suggesting that individuals with PD not only have a proprioceptive deficit during standing, but also have a cross-modal sensory fusion deficit that is crucial for upright stance control.

Function dictates the phase dependence of vision during human locomotion.

In human and animal locomotion, sensory input is thought to be processed in a phase-dependent manner. Here we use full-field transient visual scene motion toward or away from subjects walking on a treadmill. Perturbations were presented at three phases of walking to test 1) whether phase dependence is observed for visual input and 2) whether the nature of phase dependence differs across body segments. Results demonstrated that trunk responses to approaching perturbations were only weakly phase dependent and instead depended primarily on the delay from the perturbation. Recording of kinematic and muscle responses from both right and left lower limb allowed the analysis of six distinct phases of perturbation effects. In contrast to the trunk, leg responses were strongly phase dependent. Leg responses during the same gait cycle as the perturbation exhibited gating, occurring only when perturbations were applied in midstance. In contrast, during the postperturbation gait cycle, leg responses occurred at similar response phases of the gait cycle over a range of perturbation phases. These distinct responses reflect modulation of trunk orientation for upright equilibrium and modulation of leg segments for both hazard accommodation/avoidance and positional maintenance on the treadmill. Overall, these results support the idea that the phase dependence of responses to visual scene motion is determined by different functional tasks during walking.

Imaging the lower limb network in Parkinson's disease.

BACKGROUND: Despite the significant impact of lower limb symptoms on everyday life activities in Parkinson's disease (PD), knowledge of the neural correlates of lower limb deficits is limited. OBJECTIVE: We ran an fMRI study to investigate the neural correlates of lower limb movements in individuals with and without PD. METHODS: Participants included 24 PD and 21 older adults who were scanned while performing a precisely controlled isometric force generation task by dorsiflexing their ankle. A novel MRI-compatible ankle dorsiflexion device that limits head motion during motor tasks was used. The PD were tested on their more affected side, whereas the side in controls was randomized. Importantly, PD were tested in the off-state, following overnight withdrawal from antiparkinsonian medication. RESULTS: The foot task revealed extensive functional brain changes in PD compared to controls, with reduced fMRI signal during ankle dorsiflexion within the contralateral putamen and M1 foot area, and ipsilateral cerebellum. The activity of M1 foot area was negatively correlated with the severity of foot symptoms based on the Movement Disorder Society-Sponsored Revision of the Unified Parkinson's Disease Rating Scale (MDS-UPDRS-III). CONCLUSION: Overall, current findings provide new evidence of brain changes underlying motor symptoms in PD. Our results suggest that pathophysiology of lower limb symptoms in PD appears to involve both the cortico-basal ganglia and cortico-cerebellar motor circuits.

Rate control deficits during pinch grip and ankle dorsiflexion in early-stage Parkinson's disease.

BACKGROUND: Much of our understanding of the deficits in force control in Parkinson's disease (PD) relies on findings in the upper extremity. Currently, there is a paucity of data pertaining to the effect of PD on lower limb force control. OBJECTIVE: The purpose of this study was to concurrently evaluate upper- and lower-limb force control in early-stage PD and a group of age- and gender-matched healthy controls. METHODS: Twenty individuals with PD and twenty-one healthy older adults participated in this study. Participants performed two visually guided, submaximal (15% of maximum voluntary contractions) isometric force tasks: a pinch grip task and an ankle dorsiflexion task. PD were tested on their more affected side and after overnight withdrawal from antiparkinsonian medication. The tested side in controls was randomized. Differences in force control capacity were assessed by manipulating speed-based and variability-based task parameters. RESULTS: Compared with controls, PD demonstrated slower rates of force development and force relaxation during the foot task, and a slower rate of relaxation during the hand task. Force variability was similar across groups but greater in the foot than in the hand in both PD and controls. Lower limb rate control deficits were greater in PD with more severe symptoms based on the Hoehn and Yahr stage. CONCLUSIONS: Together, these results provide quantitative evidence of an impaired capacity in PD to produce submaximal and rapid force across multiple effectors. Moreover, results suggest that force control deficits in the lower limb may become more severe with disease progression.

Identification of neural feedback for upright stance in humans: stabilization rather than sway minimization.

A fundamental issue in motor control is how to determine the task goals for a given behavior. Here, we address this question by separately identifying the musculoskeletal and feedback components of the human postural control loop. Eighteen subjects were perturbed by two mechanical perturbations (gentle pulling from behind at waist and shoulder levels) and one sensory perturbation (movement of a virtual visual scene). Body kinematics was described by the leg and trunk segment angles in the sagittal plane. Muscle activations were described by ankle and hip EMG signals, with each EMG signal computed as a weighted sum of rectified EMG signals from multiple muscles at the given joint. The mechanical perturbations were used to identify feedback, defined as the mapping from the two segment angles to the two EMG signals. The sensory perturbation was used to estimate parameters in a mechanistic model of the plant, defined as the mapping from the two EMG signals to the two segment angles. Using the plant model and optimal control theory, we compared identified feedback to optimal feedback for a range of cost functions. Identified feedback was similar to feedback that stabilizes upright stance with near-minimum muscle activation, but was not consistent with feedback that substantially increases muscle activation to reduce movements of the body's center of mass or center of pressure. The results suggest that the common assumption of reducing sway may not apply to musculoskeletal systems that are inherently unstable.

Dynamics of inter-modality re-weighting during human postural control.

Flexible and stable postural control requires adaptation to changing environmental conditions, a process which requires re-weighting of multisensory stimuli. Recent studies, as well as predictions from a computational model, have indicated a reciprocal re-weighting relationship between modalities when a sensory stimulus changes amplitude. As one modality is down-weighted, another is up-weighted to compensate (and vice versa). The purpose of this study was to investigate the dynamics of intra- and inter-modality re-weighting process by examining postural responses to manipulation of proprioception and visual modalities simultaneously. Twenty-two young adults were placed in a visual cave and stood on a variable-pitch platform for thirteen trials of 250 s apiece. The platform was rotated at constant frequency of 0.4 Hz and amplitudes of 0.3 (low) or 1.5 (high) degrees. Platform amplitude was manipulated in two conditions: low-to-high or high-to-low. The visual stimulus was displayed at constant frequency of 0.35 Hz and amplitude of 0.08 degrees. The results showed both fast and slow changes in center of mass (CoM) response to the switch in platform amplitude. On both timescales, CoM response changed in a reciprocal manner relative to platform amplitude. When the platform amplitude increased (low-to-high condition), CoM response decreased relative to the platform and increased relative to the visual stimulus, indicating both intra-modality and inter-modality sensory re-weighting. In the high-to-low condition, however, there was no change in CoM response relative to visual stimulus, indicating that re-weighting may also be dependent on the absolute level of gain. Sway variability at frequencies other than the stimulus frequency also showed a reciprocal relationship with CoM gain relative to platform. Overall, these results indicate that dynamics of multisensory re-weighting is clearly more complicated than the schemes proposed by current adaptive models of human postural control.

Persistent Visual and Vestibular Impairments for Postural Control Following Concussion: A Cross-Sectional Study in University Students.

OBJECTIVE: To examine how concussion may impair sensory processing for control of upright stance. METHODS: Participants were recruited from a single university into 3 groups: 13 participants (8 women, 21 ± 3 years) between 2 weeks and 6 months post-injury who initiated a return-to-play progression (under physician management) by the time of testing (recent concussion group), 12 participants (7 women, 21 ± 1 years) with a history of concussion (concussion history group, > 1 year post-injury), and 26 participants (8 women, 22 ± 3 years) with no concussion history (control group). We assessed sensory reweighting by simultaneously perturbing participants' visual, vestibular, and proprioceptive systems and computed center of mass gain relative to each modality. The visual stimulus was a sinusoidal translation of the visual scene at 0.2 Hz, the vestibular stimulus was ± 1 mA binaural monopolar galvanic vestibular stimulation (GVS) at 0.36 Hz, the proprioceptive stimulus was Achilles' tendon vibration at 0.28 Hz. RESULTS: The recent concussion (95% confidence interval 0.078-0.115, p = 0.001) and the concussion history (95% confidence interval 0.056-0.094, p = 0.038) groups had higher gains to the vestibular stimulus than the control group (95% confidence interval 0.040-0.066). The recent concussion (95% confidence interval 0.795-1.159, p = 0.002) and the concussion history (95% confidence interval 0.633-1.012, p = 0.018) groups had higher gains to the visual stimulus than the control group (95% confidence interval 0.494-0.752). There were no group differences in gains to the proprioceptive stimulus or in sensory reweighting. CONCLUSION: Following concussion, participants responded more strongly to visual and vestibular stimuli during upright stance, suggesting they may have abnormal dependence on visual and vestibular feedback. These findings may indicate an area for targeted rehabilitation interventions.

Posterior fall-recovery training applied to individuals with chronic stroke: A single-group intervention study.

BACKGROUND: To assess the effects of the initial stepping limb on posterior fall recovery in individuals with chronic stroke, as well as to determine the benefits of fall-recovery training on these outcomes. METHODS: This was a single-group intervention study of 13 individuals with chronic stroke. Participants performed up to six training sessions, each including progressively challenging, treadmill-induced perturbations from a standing position. Progressions focused on initial steps with the paretic or non-paretic limb. The highest perturbation level achieved, the proportion of successful recoveries, step and trunk kinematics, as well as stance-limb muscle activation about the ankle were compared between the initial stepping limbs in the first session. Limb-specific outcomes were also compared between the first and last training sessions. FINDINGS: In the first session, initial steps with the non-paretic limb were associated with a higher proportion of success and larger perturbations than steps with the paretic limb (p = 0.02, Cohen's d = 0.8). Paretic-limb steps were wider relative to the center of mass (CoM; p = 0.01, d = 1.3), likely due to an initial standing position with the CoM closer to the non-paretic limb (p = 0.01, d = 1.4). In the last training session, participants recovered from a higher proportion of perturbations and advanced to larger perturbations (p < 0.05, d > 0.6). There were no notable changes in kinematic or electromyography variables with training (p > 0.07, d < 0.5). INTERPRETATION: The skill of posterior stepping in response to a perturbation can be improved with practice in those with chronic stroke, we were not able to identify consistent underlying kinematic mechanisms behind this adaptation.

Effects of purposeful soccer heading on circulating small extracellular vesicle concentration and cargo.

BACKGROUND: Considering the potential cumulative effects of repetitive head impact (HI) exposure, we need sensitive biomarkers to track short- and long-term effects. Circulating small extracellular vesicles (sEVs) (<200 nm) traffic biological molecules throughout the body and may have diagnostic value as biomarkers for disease. The purpose of this study was to identify the microRNA (miRNA) profile in circulating sEVs derived from human plasma following repetitive HI exposure. METHODS: Healthy adult (aged 18-35 years) soccer players were randomly assigned to one of 3 groups: the HI group performed 10 standing headers, the leg impact group performed 10 soccer ball trapping maneuvers over 10 min, and the control group did not participate in any soccer drills. Plasma was collected before testing and 24 h afterward, and sEVs were isolated and characterized via nanoparticle tracking analysis. Next-generation sequencing was utilized to identify candidate miRNAs isolated from sEVs, and candidate microRNAs were analyzed via quantitative polymerase chain reaction. In silico target prediction was performed using TargetScan (Version 7.0; targetscan.org) and miRWalk (http://mirwalk.umm.uni-heidelberg.de/) programs, and target validation was performed using luciferase reporter vectors with a miR-7844-5p mimic in human embryonic kidney (HEK) 293T/17 cells. RESULTS: Plasma sEV concentration and size were not affected across time and group following repetitive HI exposure. After 24 h, the HI read count from next-generation sequencing showed a 4-fold or greater increase in miR-92b-5p, miR-423-5p, and miR-24-3p and a 3-fold or greater decrease in miR-7844-5p, miR-144-5p, miR-221-5p, and miR-22-3p. Analysis of quantitative polymerase chain reaction revealed that leg impact did not alter the candidate miRNA levels. To our knowledge, miR-7844-5p is a previously unknown miRNA. We identified 8 miR-7844-5p mRNA targets: protein phosphatase 1 regulatory inhibitor subunit 1B (PPP1R1B), LIM and senescent cell antigen-like domains 1 (LIMS1), autophagy-related 12 (ATG12), microtubule-associated protein 1 light chain 3 beta (MAP1LC3B), integrin subunit alpha-1 (ITGA1), mitogen-activated protein kinase 1 (MAPK1), glycogen synthase kinase 3ß (GSK3ß), and mitogen-activated protein kinase 8 (MAPK8). CONCLUSION: Collectively, these data indicate repetitive HI exposure alters plasma sEV miRNA content, but not sEV size or number. Furthermore, for the first time we demonstrate that previously unknown miR-7844-5p targets mRNAs known to be involved in mitochondrial apoptosis, autophagy regulation, mood disorders, and neurodegenerative disease.

The many roles of vision during walking.

Vision can improve bipedal upright stability during standing and locomotion. However, during locomotion, vision supports additional behaviors such as gait cycle modulation, navigation, and obstacle avoidance. Here, we investigate how the multiple roles of vision are reflected in the dynamics of trunk control as the neural control problem changes from a fixed to a moving base of support. Subjects were presented with either low- or high-amplitude broadband visual stimuli during standing posture or while walking on a treadmill at 1 km/h and 5 km/h. Frequency response functions between visual scene motion (input) and trunk kinematics (output) revealed little or no change in the gain of trunk orientation in the standing posture and walking conditions. However, a dramatic increase in gain was observed in trunk (hip and shoulder) horizontal displacement from posture to locomotion. Such increases in gain may be interpreted as an increased coupling to visual scene motion. However, we believe that the increased gain reflects a decrease in stability due to a change of the control problem from standing to locomotion. Indeed, keeping the body upright with the use of vision during walking is complicated by the additional locomotor processes at work. Unlike during standing, vision plays many roles during locomotion, providing information for upright stability as well as body position relative to the external environment.

Interactions Between Different Age-Related Factors Affecting Balance Control in Walking.

Maintaining balance during walking is a continuous sensorimotor control problem. Throughout the movement, the central nervous system has to collect sensory data about the current state of the body in space, use this information to detect possible threats to balance and adapt the movement pattern to ensure stability. Failure of this sensorimotor loop can lead to dire consequences in the form of falls, injury and death. Such failures tend to become more prevalent as people get older. While research has established a number of factors associated with higher risk of falls, we know relatively little about age-related changes of the underlying sensorimotor control loop and how such changes are related to empirically established risk factors. This paper approaches the problem of age-related fall risk from a neural control perspective. We begin by summarizing recent empirical findings about the neural control laws mapping sensory input to motor output for balance control during walking. These findings were established in young, neurotypical study populations and establish a baseline of sensorimotor control of balance. We then review correlates for deteriorating balance control in older adults, of muscle weakness, slow walking, cognitive decline, and increased visual dependency. While empirical associations between these factors and fall risk have been established reasonably well, we know relatively little about the underlying causal relationships. Establishing such causal relationships is hard, because the different factors all co-vary with age and are difficult to isolate empirically. One option to analyze the role of an individual factor for balance control is to use computational models of walking comprising all levels of the sensorimotor control loop. We introduce one such model that generates walking movement patterns from a short list of spinal reflex modules with limited supraspinal modulation for balance. We show how this model can be used to simulate empirical studies, and how comparison between the model and empirical results can indicate gaps in our current understanding of balance control. We also show how different aspects of aging can be added to this model to study their effect on balance control in isolation.

Foot and Ankle Somatosensory Deficits Affect Balance and Motor Function in Children With Cerebral Palsy.

Sensory dysfunction is prevalent in cerebral palsy (CP). Evidence suggests that sensory deficits can contribute to manual ability impairments in children with CP, yet it is still unclear how they contribute to balance and motor performance. Therefore, the objective of this study was to investigate the relationship between lower extremity (LE) somatosensation and functional performance in children with CP. Ten participants with spastic diplegia (Gross Motor Function Classification Scale: I-III) and who were able to stand independently completed the study. Threshold of light touch pressure, two-point discriminatory ability of the plantar side of the foot, duration of cutaneous vibration sensation, and error in the joint position sense of the ankle were assessed to quantify somatosensory function. The balance was tested by the Balance Evaluation System Test (BESTest) and postural sway measures during a standing task. Motor performance was evaluated by using a battery of clinical assessments: (1) Gross Motor Function Measure (GMFM-66-IS) to test gross motor ability; (2) spatiotemporal gait characteristics (velocity, step length) to evaluate walking ability; (3) Timed Up and Go (TUG) and 6 Min Walk (6MWT) tests to assess functional mobility; and (4) an isokinetic dynamometer was used to test the Maximum Volitional Isometric Contraction (MVIC) of the plantar flexor muscles. The results showed that the light touch pressure measure was strongly associated only with the 6MWT. Vibration and two-point discrimination were strongly related to balance performance. Further, the vibration sensation of the first metatarsal head demonstrated a significantly strong relationship with motor performance as measured by GMFM-66-IS, spatiotemporal gait parameters, TUG, and ankle plantar flexors strength test. The joint position sense of the ankle was only related to one subdomain of the BESTest (Postural Responses). This study provides preliminary evidence that LE sensory deficits can possibly contribute to the pronounced balance and motor impairments in CP. The findings emphasize the importance of developing a thorough LE sensory test battery that can guide traditional treatment protocols toward a more holistic therapeutic approach by combining both motor and sensory rehabilitative strategies to improve motor function in CP.

Sensory Reweighting for Upright Stance in Soccer Players: A Comparison of High and Low Exposure to Soccer Heading.

The purpose of this study was to compare sensory reweighting for upright stance between soccer players who report higher soccer heading exposure to those who report lower soccer heading exposure. Thirty participants completed a self-reported questionnaire to estimate the number of soccer headers experienced over the previous year and were divided into "low exposure" and "high exposure" groups based on their responses. Sensory reweighting for upright stance was assessed by simultaneously perturbing visual, vestibular, and proprioceptive systems. The visual stimulus was a sinusoidal translation of the visual scene at 0.2 Hz, the vestibular stimulus was ±1mA binaural monopolar galvanic vestibular stimulation (GVS) at 0.36 Hz, and the proprioceptive stimulus was Achilles tendon vibration at 0.28 Hz. The visual stimulus was presented at two amplitudes (0.2 m, 0.8 m). Center of mass (COM) gain/phase to each modality, total power, 95% area and velocity were compared between low exposure (N = 15, six males, 21.5 ± 1.9 years, 27.7 ± 31.6 headers) and high exposure groups (N = 15, 10 males, 22.1 ± 3.5years, 734.9 ± 877.7 headers). Without vibration, COM 95% area (F = 5.861, p = 0.022*, partial η2 = 0.173), velocity (F = 14.198, p = 0.001, partial η2 = 0.336), and total power (F = 13.491, p = 0.001, partial η2 = 0.325) for the "high exposure" group were higher than for the "low exposure" group, and postural sway lagged the vestibular stimulus in the "high exposure" group rather than leading it as in the "low exposure" group (F = 4.765, p = 0.038, partial η2 = 0.145). There were no differences in sensory reweighting and no differences in COM gain/phase between groups. These findings lend empirical evidence to a detrimental effect of soccer heading exposure on balance control during upright stance.

Optimal motor control may mask sensory dynamics.

Properties of neural controllers for closed-loop sensorimotor behavior can be inferred with system identification. Under the standard paradigm, the closed-loop system is perturbed (input), measurements are taken (output), and the relationship between input and output reveals features of the system under study. Here we show that under common assumptions made about such systems (e.g. the system implements optimal control with a penalty on mechanical, but not sensory, states) important aspects of the neural controller (its zeros mask the modes of the sensors) remain hidden from standard system identification techniques. Only by perturbing or measuring the closed-loop system "between" the sensor and the control can these features be exposed with closed-loop system identification methods; while uncommon, there exist noninvasive techniques such as galvanic vestibular stimulation that perturb between sensor and controller in this way.

Phase-Dependency of Medial-Lateral Balance Responses to Sensory Perturbations During Walking.

The human body is mechanically unstable during walking. Maintaining upright stability requires constant regulation of muscle force by the central nervous system to push against the ground and move the body mass in the desired way. Activation of muscles in the lower body in response to sensory or mechanical perturbations during walking is usually highly phase-dependent, because the effect any specific muscle force has on the body movement depends upon the body configuration. Yet the resulting movement patterns of the upper body after the same perturbations are largely phase-independent. This is puzzling, because any change of upper-body movement must be generated by parts of the lower body pushing against the ground. How do phase-dependent muscle activation patterns along the lower body generate phase-independent movement patterns of the upper body? We hypothesize that when a sensory system detects a deviation of the body in space from a desired state that indicates the onset of a fall, the nervous system generates a functional response by pushing against the ground in any way possible with the current body configuration. This predicts that the changes in the ground reaction force patterns following a balance perturbation should be phase-independent. Here we test this hypothesis by disturbing upright balance in the frontal plane using Galvanic vestibular stimulation at three different points in the gait cycle. We measure the resulting changes in whole-body center of mass movement and the location of the center of pressure of the ground reaction force. We find that the magnitude of the initial center of pressure shift in the direction of the perceived fall is larger for perturbations late in the gait cycle, while there is no statistically significant difference in onset time. These results contradict our hypothesis by showing that even the initial CoP shift in response to a balance perturbation depends upon the phase of the gait cycle. Contrary to expectation, we also find that the whole-body balance response is not phase-independent. Both the onset time and the magnitude of the whole-body center of mass shift depend on the phase of the perturbation. We conclude that the central nervous system recruits any available mechanism to generate a functional balance response by pushing against the ground as fast as possible in response to a perturbation, but that the different mechanisms available at different phases in the gait cycle are not equally strong, leading to phase-dependent differences in the overall response.

Anterior fall-recovery training applied to individuals with chronic stroke.

BACKGROUND: To study the effects of the initial stepping limb on anterior fall-recovery performance and kinematics, as well as to determine the benefits of fall-recovery training on those outcomes in individuals with chronic stroke. METHODS: Single-group intervention of 15 individuals with chronic stroke who performed up to six sessions of fall-recovery training. Each session consisted of two progressions of treadmill-induced perturbations to induce anterior falls from a standing position. Progressions focused on initial steps with the paretic or non-paretic limb. Fall-recovery performance (the highest disturbance level achieved and the proportion of successful recoveries), as well as step and trunk kinematics were compared between the initial stepping limbs on the first session. Limb-specific outcomes were also compared between the first and last training sessions. FINDINGS: There were no between-limb differences in fall-recovery performance in the first session. With training, participants successfully recovered from a higher proportion of falls (p's = 0.01, Cohen's d's > 0.7) and progressed to larger perturbation magnitudes (p's < 0.06, d's > 0.5). Initial steps with the paretic limb were wider and shorter relative to the center of mass (p's < 0.06, d's > 0.5). With training, initial paretic-limb steps became longer relative to the CoM (p = 0.03, d = 0.7). Trunk forward rotation was reduced when first stepping with the non-paretic limb (p = 0.03, d = 0.6). INTERPRETATION: The initial stepping limb affects relevant step kinematics during anterior fall recovery. Fall-recovery training improved performance and select kinematic outcomes in individuals with chronic stroke.