Learning to stand with sensorimotor delays generalizes across directions and from hand to leg effectors.

Humans receive sensory information from the past, requiring the brain to overcome delays to perform daily motor skills such as standing upright. Because delays vary throughout the body and change over a lifetime, it would be advantageous to generalize learned control policies of balancing with delays across contexts. However, not all forms of learning generalize. Here, we use a robotic simulator to impose delays into human balance. When delays are imposed in one direction of standing, participants are initially unstable but relearn to balance by reducing the variability of their motor actions and transfer balance improvements to untrained directions. Upon returning to normal standing, aftereffects from learning are observed as small oscillations in control, yet they do not destabilize balance. Remarkably, when participants train to balance with delays using their hand, learning transfers to standing with the legs. Our findings establish that humans use experience to broadly update their neural control to balance with delays.

Vestibular control of deep and superficial lumbar muscles.

The active control of the lumbar musculature provides a stable platform critical for postures and goal-directed movements. Voluntary and perturbation-evoked motor commands can recruit individual lumbar muscles in a task-specific manner according to their presumed biomechanics. Here, we investigated the vestibular control of the deep and superficial lumbar musculature. Ten healthy participants were exposed to noisy electrical vestibular stimulation while balancing upright with their head facing forward, left, or right to characterize the differential modulation in the vestibular-evoked lumbar extensor responses in generating multidirectional whole body motion. We quantified the activation of the lumbar muscles on the right side using indwelling [deep multifidus, superficial multifidus, caudal longissimus (L4), and cranial longissimus (L1)] and high-density surface recordings. We characterized the vestibular-evoked responses using coherence and peak-to-peak cross-covariance amplitude between the vestibular and electromyographic signals. Participants exhibited responses in all lumbar muscles. The vestibular control of the lumbar musculature exhibited muscle-specific modulations: responses were larger in the longissimus (combined cranio-caudal) compared with the multifidus (combined deep-superficial) when participants faced forward (P < 0.001) and right (P = 0.011) but not when they faced left. The high-density surface recordings partly supported this observation: the location of the responses was more lateral when facing right compared with left (P < 0.001). The vestibular control of muscle subregions within the longissimus or the multifidus was similar. Our results demonstrate muscle-specific vestibular control of the lumbar muscles in response to perturbations of vestibular origin. The lack of differential activation of lumbar muscle subregions suggests the vestibular control of these subregions is co-regulated for standing balance.NEW & NOTEWORTHY We investigated the vestibular control of the deep and superficial lumbar extensor muscles using electrical vestibular stimuli. Vestibular stimuli elicited preferential activation of the longissimus muscle over the multifidus muscle. We did not observe clear regional activation of lumbar muscle subregions in response to the vestibular stimuli. Our findings show that the central nervous system can finely tune the vestibular control of individual lumbar muscles and suggest minimal regional variations in the activation of lumbar muscle subregions.

The Neural Basis for Biased Behavioral Responses Evoked by Galvanic Vestibular Stimulation in Primates.

Noninvasive electrical stimulation of the vestibular system in humans has become an increasingly popular tool with a broad range of research and clinical applications. However, common assumptions regarding the neural mechanisms that underlie the activation of central vestibular pathways through such stimulation, known as galvanic vestibular stimulation (GVS), have not been directly tested. Here, we show that GVS is encoded by VIIIth nerve vestibular afferents with nonlinear dynamics that differ markedly from those predicted by current models. GVS produced asymmetric activation of both semicircular canal and otolith afferents to the onset versus offset and cathode versus anode of applied current, that in turn produced asymmetric eye movement responses in three awake-behaving male monkeys. Additionally, using computational methods, we demonstrate that the experimentally observed nonlinear neural response dynamics lead to an unexpected directional bias in the net population response when the information from both vestibular nerves is centrally integrated. Together our findings reveal the neural basis by which GVS activates the vestibular system, establish that neural response dynamics differ markedly from current predictions, and advance our mechanistic understanding of how asymmetric activation of the peripheral vestibular system alters vestibular function. We suggest that such nonlinear encoding is a general feature of neural processing that will be common across different noninvasive electrical stimulation approaches.SIGNIFICANCE STATEMENT Here, we show that the application of noninvasive electrical currents to the vestibular system (GVS) induces more complex responses than commonly assumed. We recorded vestibular afferent activity in macaque monkeys exposed to GVS using a setup analogous to human studies. GVS evoked notable asymmetries in irregular afferent responses to cathodal versus anodal currents. We developed a nonlinear model explaining these GVS-evoked afferent responses. Our model predicts that GVS induces directional biases in centrally integrated head motion signals and establishes electrical stimuli that recreate physiologically plausible sensations of motion. Altogether, our findings provide new insights into how GVS activates the vestibular system, which will be vital to advancing new clinical and biomedical applications.

Real-world characterization of vestibular contributions during locomotion.

Introduction: The vestibular system, which encodes our head movement in space, plays an important role in maintaining our balance as we navigate the environment. While in-laboratory research demonstrates that the vestibular system exerts a context-dependent influence on the control of balance during locomotion, differences in whole-body and head kinematics between indoor treadmill and real-world locomotion challenge the generalizability of these findings. Thus, the goal of this study was to characterize vestibular-evoked balance responses in the real world using a fully portable system. Methods: While experiencing stochastic electrical vestibular stimulation (0-20 Hz, amplitude peak ± 4.5 mA, root mean square 1.25 mA) and wearing inertial measurement units (IMUs) on the head, low back, and ankles, 10 participants walked outside at 52 steps/minute (∼0.4 m/s) and 78 steps/minute (∼0.8 m/s). We calculated time-dependent coherence (a measure of correlation in the frequency domain) between the applied stimulus and the mediolateral back, right ankle, and left ankle linear accelerations to infer the vestibular control of balance during locomotion. Results: In all participants, we observed vestibular-evoked balance responses. These responses exhibited phasic modulation across the stride cycle, peaking during the middle of the single-leg stance in the back and during the stance phase for the ankles. Coherence decreased with increasing locomotor cadence and speed, as observed in both bootstrapped coherence differences (p < 0.01) and peak coherence (low back: 0.23 ± 0.07 vs. 0.16 ± 0.14, p = 0.021; right ankle: 0.38 ± 0.12 vs. 0.25 ± 0.10, p < 0.001; left ankle: 0.33 ± 0.09 vs. 0.21 ± 0.09, p < 0.001). Discussion: These results replicate previous in-laboratory studies, thus providing further insight into the vestibular control of balance during naturalistic movements and validating the use of this portable system as a method to characterize real-world vestibular responses. This study will help support future work that seeks to better understand how the vestibular system contributes to balance in variable real-world environments.

A portable and low-cost solution for real-time manipulation of the vestibular sense.

BACKGROUND: The vestibular system encodes head motion in space which is naturally accompanied by other sensory cues. Electrical stimuli, applied across the mastoid processes, selectively activate primary vestibular afferents which has spurred clinical and biomedical applications of electrical vestibular stimulation (EVS). When properly matched to head motion, EVS may also manipulate the closed-loop relationship between actions and vestibular feedback to reveal the mechanisms of sensorimotor recalibration and learning. NEW METHOD: We designed a portable, low-cost real-time EVS system using an Arduino microcontroller programmed through Simulink that provides electrical currents based on head angular motion. We used well-characterized vestibular afferent physiological responses to head angular velocity and electrical current to compute head-motion equivalent of real-time modulatory EVS currents. We also examined if our system induced recalibration of the vestibular system during human balance control. RESULTS: Our system operated at 199.997 Hz ( ± 0.005 Hz) and delivered head-motion-equivalent electrical currents with ∼10 ms delay. The output driving the current stimulator matched the implemented linear model for physiological vestibular afferent dynamics with minimal background noise (<0.2% of ± 10 V range). Participants recalibrated to the modulated closed-loop vestibular feedback using visual cues during standing balance, replicating earlier findings. COMPARISON WITH EXISTING METHODS: EVS is typically used to impose external perturbations that are independent of one's own movement. We provided a solution using open-source hardware to implement a real-time, physiology based, and task-relevant vestibular modulations using EVS. CONCLUSIONS: Our portable, low-cost vestibular modulation system will make physiological closed-loop vestibular manipulations more accessible thus encouraging novel investigations and biomedical applications of EVS.

Adolescents with idiopathic scoliosis show decreased intermuscular coherence in lumbar paraspinal muscles: A new pathophysiological perspective.

OBJECTIVE: Although adolescent idiopathic scoliosis is thought to be an orthopedic disorder, sensorimotor deficits resulting in asymmetric neural drive to the axial musculature have been proposed as contributing factors. Asymmetry in the vestibular control of spinal motoneurons can cause spine deformation reminiscent of idiopathic scoliosis in animal models. METHODS: To examine the neural control of axial muscles, we compared common oscillatory drive to bilateral lumbar muscles between 19 participants with adolescent idiopathic scoliosis and 19 healthy adolescents. We measured right and left paraspinal muscle activity during steady isometric back extensions at 15% or 30% of their maximum voluntary contraction. RESULTS: The variance in exerted force and symmetry in bilateral muscle activation were similar between groups. We estimated the strength of common oscillations between muscle motoneuron pools using intermuscular coherence. Compared to controls, participants with adolescent idiopathic scoliosis exhibited smaller intermuscular coherence between paraspinal muscles in the alpha and beta bands. To identify the cause of the observed decreased in intermuscular coherence, we quantified variability of electromyography power ratio and relative activation timing between the paraspinal muscle. Intermuscular phase between muscle oscillations across the alpha band demonstrated larger variability in adolescent with idiopathic scoliosis. The variability of the ratio of lumbar muscles power was similar between groups in the alpha and beta bands. CONCLUSION: Our results suggest that altered bilateral control of axial muscles characterized by increased variability in the timing of alpha oscillations may be linked to spine deformation in adolescents. SIGNIFICANCE: Our findings provide a new perspective on neural factors associated with a common spine deformation, adolescent idiopathic scoliosis.

Unperceived motor actions of the balance system interfere with the causal attribution of self-motion.

The instability of human bipedalism demands that the brain accurately senses balancing self-motion and determines whether movements originate from self-generated actions or external disturbances. Here, we challenge the longstanding notion that this process relies on a single representation of the body and world to accurately perceive postural orientation and organize motor responses to control balance self-motion. Instead, we find that the conscious sense of balance can be distorted by the corrective control of upright standing. Using psychophysics, we quantified thresholds to imposed perturbations and balance responses evoking cues of self-motion that are (in)distinguishable from corrective balance actions. When standing immobile, participants clearly perceived imposed perturbations. Conversely, when freely balancing, participants often misattributed their own corrective responses as imposed motion because their balance system had detected, integrated, and responded to the perturbation in the absence of conscious perception. Importantly, this only occurred for perturbations encoded ambiguously with balance-correcting responses and that remained below the natural variability of ongoing balancing oscillations. These findings reveal that our balance system operates on its own sensorimotor principles that can interfere with causal attribution of our actions, and that our conscious sense of balance depends critically on the source and statistics of induced and self-generated motion cues.

Learning to stand with unexpected sensorimotor delays.

Human standing balance relies on self-motion estimates that are used by the nervous system to detect unexpected movements and enable corrective responses and adaptations in control. These estimates must accommodate for inherent delays in sensory and motor pathways. Here, we used a robotic system to simulate human standing about the ankles in the anteroposterior direction and impose sensorimotor delays into the control of balance. Imposed delays destabilized standing, but through training, participants adapted and re-learned to balance with the delays. Before training, imposed delays attenuated vestibular contributions to balance and triggered perceptions of unexpected standing motion, suggesting increased uncertainty in the internal self-motion estimates. After training, vestibular contributions partially returned to baseline levels and larger delays were needed to evoke perceptions of unexpected standing motion. Through learning, the nervous system accommodates balance sensorimotor delays by causally linking whole-body sensory feedback (initially interpreted as imposed motion) to self-generated balance motor commands.

When standing, neurons in the brain send signals to skeletal muscles so we can adjust our movements to stay upright based on the requirements from the surrounding environment. The long nerves needed to connect our brain, muscles and sensors lead to considerable time delays (up to 160 milliseconds) between sensing the environment and the generation of balance-correcting motor signals. Such delays must be accounted for by the brain so it can adjust how it regulates balance and compensates for unexpected movements. Aging and neurological disorders can lead to lengthened neural delays, which may result in poorer balance. Computer modeling suggests that we cannot maintain upright balance if delays are longer than 300-340 milliseconds. Directly assessing the destabilizing effects of increased delays in human volunteers can reveal how capable the brain is at adapting to this neurological change. Using a custom-designed robotic balance simulator, Rasman et al. tested whether healthy volunteers could learn to balance with delays longer than the predicted 300-340 millisecond limit. In a series of experiments, 46 healthy participants stood on the balance simulator which recreates the physical sensations and neural signals for balancing upright based on a computer-driven virtual reality. This unique device enabled Rasman et al. to artificially impose delays by increasing the time between the generation of motor signals and resulting whole-body motion. The experiments showed that lengthening the delay between motor signals and whole-body motion destabilized upright standing, decreased sensory contributions to balance and led to perceptions of unexpected movements. Over five days of training on the robotic balance simulator, participants regained their ability to balance, which was accompanied by recovered sensory contributions and perceptions of expected standing, despite the imposed delays. When a subset of participants was tested three months later, they were still able to compensate for the increased delay. The experiments show that the human brain can learn to overcome delays up to 560 milliseconds in the control of balance. This discovery may have important implications for people who develop balance problems because of older age or neurologic diseases like multiple sclerosis. It is possible that robot-assisted training therapies, like the one in this study, could help people overcome their balance impairments.

Potential Mechanisms of Acute Standing Balance Deficits After Concussions and Subconcussive Head Impacts: A Review.

Standing balance deficits are prevalent after concussions and have also been reported after subconcussive head impacts. However, the mechanisms underlying such deficits are not fully understood. The objective of this review is to consolidate evidence linking head impact biomechanics to standing balance deficits. Mechanical energy transferred to the head during impacts may deform neural and sensory components involved in the control of standing balance. From our review of acute balance-related changes, concussions frequently resulted in increased magnitude but reduced complexity of postural sway, while subconcussive studies showed inconsistent outcomes. Although vestibular and visual symptoms are common, potential injury to these sensors and their neural pathways are often neglected in biomechanics analyses. While current evidence implies a link between tissue deformations in deep brain regions including the brainstem and common post-concussion balance-related deficits, this link has not been adequately investigated. Key limitations in current studies include inadequate balance sampling duration, varying test time points, and lack of head impact biomechanics measurements. Future investigations should also employ targeted quantitative methods to probe the sensorimotor and neural components underlying balance control. A deeper understanding of the specific injury mechanisms will inform diagnosis and management of balance deficits after concussions and subconcussive head impact exposure.

Soleus responses to Achilles tendon stimuli are suppressed by heel and enhanced by metatarsal cutaneous stimuli during standing.

KEY POINTS: We examined the influence of cutaneous feedback from the heel and metatarsal regions of the foot sole on the soleus stretch reflex pathway during standing. We found that heel electrical stimuli suppressed and metatarsal stimuli enhanced the soleus vibration response. Follow-up experiments indicated that the interaction between foot sole cutaneous feedback and the soleus vibration response was likely not mediated by presynaptic inhibition and was contingent upon a modulation at the ⍺-motoneuron pool level. The spatially organized interaction between cutaneous feedback from the foot sole and the soleus vibration response provides information about how somatosensory information is combined to appropriately respond to perturbations during standing. ABSTRACT: Cutaneous feedback from the foot sole provides balance-relevant information and has the potential to interact with spinal reflex pathways. In this study, we examined how cutaneous feedback from the foot sole (heel and metatarsals) influenced the soleus response to proprioceptive stimuli during standing. We delivered noisy vibration (10-115 Hz) to the right Achilles tendon while we intermittently applied electrical pulse trains (five 1-ms pulses at 200 Hz, every 0.8-1.0 s) to the skin under either the heel or the metatarsals of the ipsilateral foot sole. We analysed time-dependent (referenced to cutaneous stimuli) coherence and cross-correlations between the vibration acceleration and rectified soleus EMG. Vibration-EMG coherence was observed across a bandwidth of ∼10-80 Hz, and coherence was suppressed by heel but enhanced by metatarsal cutaneous stimuli. Cross-correlations showed soleus EMG was correlated with the vibration (∼40 ms lag) and cross-correlations were also suppressed by heel (from 104-155 ms) but enhanced by metatarsal (from 76-128 ms) stimuli. To examine the neural mechanisms mediating this reflex interaction, we conducted two further experiments to probe potential contributions from (1) presynaptic inhibition, and (2) modulations at the ⍺- and γ-motoneuron pools. Results suggest the cutaneous interactions with the stretch reflex pathway required a modulation at the ⍺-motoneuron pool and were likely not mediated by presynaptic inhibition. These findings demonstrate that foot sole cutaneous information functionally tunes the stretch reflex pathway during the control of upright posture and balance.

An intensity matched comparison of laser- and contact heat evoked potentials.

Previous studies comparing laser (LEPs) and contact heat evoked potentials (CHEPs) consistently reported higher amplitudes following laser compared to contact heat stimulation. However, none of the studies matched the perceived pain intensity, questioning if the observed difference in amplitude is due to biophysical differences between the two methods or a mismatch in stimulation intensity. The aims of the current study were twofold: (1) to directly compare the brain potentials induced by intensity matched laser and contact heat stimulation and (2) investigate how capsaicin-induced secondary hyperalgesia modulates LEPs and CHEPs. Twenty-one healthy subjects were recruited and measured at four experimental sessions: (1) CHEPs + sham, (2) LEPs + sham, (3) CHEPs + capsaicin, and (4) LEPs + capsaicin. Baseline (sham) LEPs latency was significantly shorter and amplitude significantly larger compared to CHEPs, even when matched for perceived pain. Neither CHEPs nor LEPs was sensitive enough to detect secondary hyperalgesia. These differences provide evidence that a faster heating rate results in an earlier and more synchronized LEPs than CHEPs. To our knowledge, this was the first study to match perceived intensity of contact heat and laser stimulations, revealing distinct advantages associated with the acquisition of LEPs.

A task-relevant experimental pain model to target motor adaptation.

KEY POINTS: Motor adaptation is thought to be a strategy to avoid pain. Current experimental pain models do not allow for consistent modulation of pain perception depending on movement. We showed that low-frequency sinusoidal stimuli delivered at painful intensity result in minimal habituation of pain perception (over 60 s) and minimal stimulation artefacts on electromyographic signals. When the amplitude of the low-frequency sinusoidal stimuli was modulated based on the vertical force participants applied to the ground with their right leg while standing upright, we demonstrated a strong association between perceived pain and motor adaptation. By enabling task-relevant modulation of perceived pain intensity and the recording electromyographic signals during electrical painful stimulation, our novel pain model will permit direct experimental testing of the relationship between pain and motor adaptation. ABSTRACT: Contemporary pain adaptation theories predict that motor adaptation occurs to limit pain. Current experimental pain models, however, do not allow for pain intensity modulation according to one's posture or movements. We developed a task-relevant experimental pain model using low-frequency sinusoidal electrical stimuli applied over the infrapatellar fat pad. In fourteen participants, we compared perceived pain habituation and stimulation-induced artefacts in vastus medialis electromyographic recordings elicited by sinusoidal (4, 10, 20 and 50 Hz) and square electrical waveforms delivered at constant peak stimulation amplitude. Next, we simulated a clinical condition where perceived knee pain intensity is proportional to the load applied on the leg by controlling sinusoidal current amplitude (4 Hz) according to the vertical force the participants applied with their right leg to the ground while standing upright. Pain ratings habituated over a 60 s period for 50 Hz sinusoidal and square waveforms but not for low-frequency sinusoidal stimuli (P < 0.001). EMG filters removed most stimulation artefacts for low-frequency sinusoidal stimuli (4 Hz). While balancing upright, participants' pain ratings were correlated with the force applied by the right leg (R2  = 0.65), demonstrating task-relevant changes in perceived pain intensity. Low-frequency sinusoidal stimuli can induce knee pain of constant intensity for 60 s with minimal EMG artefacts while enabling task-relevant pain modulation when controlling current amplitude. By enabling task-dependent modulation of perceived pain intensity, our novel experimental model replicates key temporal aspects of clinical musculoskeletal pain while allowing quantification of neuromuscular activation during painful electrical stimulation. This approach will enable researchers to test the predicted relationship between movement strategies and pain.

Neck Muscle and Head/Neck Kinematic Responses While Bracing Against the Steering Wheel During Front and Rear Impacts.

Drivers often react to an impending collision by bracing against the steering wheel. The goal of the present study was to quantify the effect of bracing on neck muscle activity and head/torso kinematics during low-speed front and rear impacts. Eleven seated subjects (3F, 8 M) experienced multiple sled impacts (Δv = 0.77 m/s; apeak = 19.9 m/s2, Δt = 65.5 ms) with their hands on the steering wheel in two conditions: relaxed and braced against the steering wheel. Electromyographic activity in eight neck muscles (sternohyoid, sternocleidomastoid, splenius capitis, semispinalis capitis, semispinalis cervicis, multifidus, levator scapulae, and trapezius) was recorded unilaterally with indwelling electrodes and normalized by maximum voluntary contraction (MVC) levels. Head and torso kinematics (linear acceleration, angular velocity, angular rotation, and retraction) were measured with sensors and motion tracking. Muscle and kinematic variables were compared between the relaxed and braced conditions using linear mixed models. We found that pre-impact bracing generated only small increases in the pre-impact muscle activity (< 5% MVC) when compared to the relaxed condition. Pre-impact bracing did not increase peak neck muscle responses during the impacts; instead it reduced peak trapezius and multifidus muscle activity by about half during front impacts. Bracing led to widespread changes in the peak amplitude and timing of the torso and head kinematics that were not consistent with a simple stiffening of the head/neck/torso system. Instead pre-impact bracing served to couple the torso more rigidly to the seat while not necessarily coupling the head more rigidly to the torso.

Influence of age on the frequency characteristics of the soleus muscle response to Achilles tendon vibration during standing.

KEY POINTS: Proprioceptive sensory information from the ankle joint is critical for the control of upright posture and balance. We examined the influence of age (n = 54 healthy adults, 20-82 years old) on lower limb muscle responses to proprioceptive perturbations evoked by Achilles tendon vibration during standing. The frequency bandwidth of the muscle response became narrower, and the gain (the muscle response relative to the stimulus) and scaling (increases in response amplitude with increases in stimulus amplitude) decreased with age. Mechanics of the muscle-tendon unit (mechanical admittance) did not differ with age during standing, and thus probably did not mediate the age-related changes observed in soleus muscle responses to vibration. These findings add to our understanding of how altered proprioceptive responses may contribute to impaired mobility and falls with ageing. ABSTRACT: Proprioceptive information from the ankle joint plays an important role in the control of upright posture and balance. Ageing influences many components of the sensorimotor system, which leads to poor mobility and falls. However, little is known about the influence of age on the characteristics of short latency muscle responses to proprioceptive stimuli during standing across frequencies that are encoded by muscle spindles. We examined the frequency characteristics of the soleus muscle response to noisy (10-115 Hz) Achilles tendon vibration during standing in 54 healthy adults across a broad age range (20-82 years). The results showed the frequency bandwidth of the soleus response (vibration-electromyography coherence) became progressively narrower with ageing. Coherence was significantly lower in middle-aged relative to young adults between ∼7-11 and 28-62 Hz, lower in older relative to middle-aged adults between ∼30-50 Hz and lower in older relative to young adults between ∼7-64 Hz. Muscle response gain was similar between age groups at low frequencies, although gain was lower in older relative to young adults between ∼28-54 Hz. Across the age range, the response amplitude (peak-to-peak cross-covariance) and the scaling of the response with stimulus amplitude were both negatively correlated with age. Muscle-tendon mechanics (admittance) did not differ with age, suggesting this did not mediate differences in soleus responses. Our findings suggest there is a progressive change in the soleus response to proprioceptive stimuli with ageing during standing, which could contribute to poorer mobility and falls.

Lower-limb muscle responses evoked with noisy vibrotactile foot sole stimulation.

AIM: Cutaneous feedback from the foot sole contributes to the control of standing balance in two ways: it provides perceptual awareness of tactile perturbations at the interface with the ground (e.g., shifts in the pressure distribution, slips, etc.) and it reflexively activates lower-motor neurons to trigger stabilizing postural responses. Here we focus on the latter, cutaneous (or cutaneomotor) reflex coupling in the lower limb. These reflexes have been studied most-frequently with electrical pulse trains that bypass natural cutaneous mechanotransduction, stimulating cutaneous afferents in a largely non-physiological manner. Harnessing the mechanical filtering properties of cutaneous afferents, we take a novel mechanical approach by applying supra-threshold continuous noisy vibrotactile stimulation (NVS) to the medial forefoot. METHODS: Using NVS, we characterized the time and frequency domain properties of cutaneomotor reflexes in the Tibialis Anterior. We additionally measured stimulus-triggered average muscle responses to repeated discrete sinusoidal pulses for comparison. To investigate cutaneomotor reflex gain scaling, stimuli were delivered at 3- or 10-times perceptual threshold (PT), while participants held 12.5% or 25% of maximum voluntary contraction (MVC). RESULTS: Peak responses in the time domain were observed at lags reflecting transmission delay through a polysynaptic reflex pathway (~90-100 ms). Increasing the stimulus amplitude enhanced cutaneomotor coupling, likely by increasing afferent firing rates. Although greater background muscle contraction increased the overall amplitude of the evoked responses, it did not increase the proportion of the muscle response attributable to cutaneous input. CONCLUSION: Taken together, our findings support the use of NVS as a novel tool for probing the physiological properties of cutaneomotor reflex pathways.

Frequency characteristics of heteronymous responses evoked by Achilles tendon vibration during quiet stance.

Primary (Ia) sensory afferents that innervate muscle spindles provide strong synaptic input to homonymous motoneurons and are thought to play a role in balance control. In addition, Ia afferents have broad heteronymous connections; i.e., projections to motoneurons that innervate other muscles that act at the same joint as well as at different joints. The purpose of the current study was to investigate heteronymous Ia afferent connections from the triceps surae muscles to lower limb and back muscles during quiet standing in humans. We applied supra-threshold noisy vibration (10-115 Hz) to the right Achilles tendon of 12 participants maintaining quiet stance and recorded EMG activity bilaterally from homonymous (Soleus) and heteronymous muscles (Semitendinosus, Vastus Lateralis, Erector Spinae). We estimated coherence, phase, and gain between the tendon probe acceleration and rectified EMG from each muscle. We found significant coherence between the probe acceleration and EMG in ipsilateral Soleus (5-100 Hz), Semitendinosus (10-75 Hz), Vastus Lateralis (5-70 Hz), and bilateral Erector Spinae muscles (10-70 Hz). These results provide evidence that triceps surae muscle spindle afferents can influence the activity of muscles proximal to the ankle joint across a broad frequency band during quiet standing.

A comparison of anti-whiplash seats during low/moderate speed rear-end collisions.

Objectives: The Insurance Institute for Highway Safety (IIHS) rates automotive seats as good, acceptable, marginal, and poor on their abilities to prevent whiplash injuries during rear-end collisions. The goal of this study was to compare the performance of some good- and poor-rated seats at speed changes below 16 km/h where some whiplash injuries occur.Methods: A BioRID II anthropometric test device (ATD) underwent rear-end collisions from 2 to 14 km/h while seated on one of two Volvo Whiplash Prevention seats (WHIPS), a Saab Active Head Restraint seat (SAHR), or a General Motors High Retention seat (GMHR). The WHIPS and SAHR seats were rated good whereas the GMHR seat was rated poor by the IIHS. The ATD's kinematics, kinetics and three neck injury criteria were evaluated across the range of collision severities.Results: Most of the head and torso kinematics, kinetics and injury criteria exhibited graded responses with increasing collision severities. Only head extension angle remained relatively similar across all speed changes. Differences between the good- and poor-rated seats were most apparent in the upper neck loads and moments, and head retraction for speed changes greater than 6 km/h.Conclusions: The relatively similar occupant responses across all seats could explain the marginal reductions in whiplash injury risk between good- and poor-rated seats in field studies. Further research into the design of anti-whiplash devices is required to better understand the link between occupant response and injury, and to better mitigate the risk of whiplash injuries during rear-end collisions.

Using Variance to Explore the Diagnostic Utility of Baseline Concussion Testing.

The Graded Symptom Checklist (GSC), Standardized Assessment of Concussion (SAC), Balance Error Scoring System (BESS), and King-Devick Test (KDT) are considered important components of concussion assessment. Whether baseline testing improves the diagnostic utility of these tests remains unclear. We performed an observational cohort study to investigate the within-subject and between-subjects variability of these tests over repeated assessments during two football seasons to examine whether baseline testing reduces variability in test performance. Thirty-five male collegiate football players completed weekly clinical concussion assessments over two seasons. Within-subject (week-to-week) and between-subjects (player-to-player) variability for each test were compared using a bootstrap analysis. Within-subject and between-subjects proportions of overall variance for each test score were calculated. Mixed-model analyses were used to quantify practice effects resulting from repeated testing. For the GSC and BESS, within-subject and between-subjects variability did not significantly differ. For the KDT, the proportion of within-subject variance (20.2%) was significantly less than the between-subjects variance (79.8%). For SAC, however, the proportion of within-subject variance (66.8%) was significantly greater than the between-subjects variance (33.8%). A small, but significant, practice effect was observed for the BESS and KDT tests. When athletes are evaluated during a football season for concussion using the GSC, SAC, and BESS, comparing their scores to baseline performance is likely no more beneficial than comparing them to normative population data for identifying neurological changes associated with concussion. For the KDT, comparison to baseline testing is likely beneficial because of significantly higher between-subjects variability.

Neural Mechanisms Underlying High-Frequency Vestibulocollic Reflexes In Humans And Monkeys.

The vestibulocollic reflex is a compensatory response that stabilizes the head in space. During everyday activities, this stabilizing response is evoked by head movements that typically span frequencies from 0 to 30 Hz. Transient head impacts, however, can elicit head movements with frequency content up to 300-400 Hz, raising the question whether vestibular pathways contribute to head stabilization at such high frequencies. Here, we first established that electrical vestibular stimulation modulates human neck motor unit (MU) activity at sinusoidal frequencies up to 300 Hz, but that sensitivity increases with frequency up to a low-pass cutoff of ∼70-80 Hz. To examine the neural substrates underlying the low-pass dynamics of vestibulocollic reflexes, we then recorded vestibular afferent responses to the same electrical stimuli in monkeys. Vestibular afferents also responded to electrical stimuli up to 300 Hz, but in contrast to MUs their sensitivity increased with frequency up to the afferent resting firing rate (∼100-150 Hz) and at higher frequencies afferents tended to phase-lock to the vestibular stimulus. This latter nonlinearity, however, was not transmitted to neck motoneurons, which instead showed minimal phase-locking that decreased at frequencies >75 Hz. Similar to human data, we validated that monkey muscle activity also exhibited low-pass filtered vestibulocollic reflex dynamics. Together, our results show that neck MUs are activated by high-frequency signals encoded by primary vestibular afferents, but undergo low-pass filtering at intermediate stages in the vestibulocollic reflex. These high-frequency contributions to vestibular-evoked neck muscle responses could stabilize the head during unexpected head transients.SIGNIFICANCE STATEMENT Vestibular-evoked neck muscle responses rely on accurate encoding and transmission of head movement information to stabilize the head in space. Unexpected transient events, such as head impacts, are likely to push the limits of these neural pathways since their high-frequency features (0-300 Hz) extend beyond the frequency bandwidth of head movements experienced during everyday activities (0-30 Hz). Here, we demonstrate that vestibular primary afferents encode high-frequency stimuli through frequency-dependent increases in sensitivity and phase-locking. When transmitted to neck motoneurons, these signals undergo low-pass filtering that limits neck motoneuron phase-locking in response to stimuli >75 Hz. This study provides insight into the neural dynamics producing vestibulocollic reflexes, which may respond to high-frequency transient events to stabilize the head.