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Studies investigating the effect of tread edge highlighters on descent speed differ, but collectively report the potential benefit of reduced fall risk. Here we examine the impact of adding high-contrast black vinyl striping to the front edge of each step's tread and its impact on descending gait speed (intervention), while controlling for illumination. Descending gait speed was estimated from 5,824 video observations using the stairway length and entry and exit times. A second stairway was unaltered (control) to compare to the intervention. Stair users were primarily 18-30 years old with a small percentage being middle-aged and older adults. Descending gait speed was significantly slower on the intervention stairway (Linear mixed effects model: standardised coefficient = -0.07, 95% CI = [-0.12, -0.02]

This study sought to examine stairway safety by identifying associations between fall-related events on stairways, distractions, gait speed, drifting, as well as handrail use and proximity.Video recordings captured 11,137 observations of stair users in two public stairways and recorded distractions (e.g., looking at a mobile device, talking on a mobile device, using earbuds or headphones, holding a mobile device, or talking with a peer), gait speed (m/s), drifting (change of direction), as well as handrail use and proximity to a handrail.In our sample, consisting of primarily young adults (observed 18-40 years old), we found that when a distraction was present, gait speed was reduced (p lt;.001), drifting increased (p lt;.001), and handrail use negatively impacted (p lt;.001) compared to stair users who were not distracted.These results indicate that distractions, such as mobile devices, used during stair negotiation can reduce handrail use and increase behaviors associated with fall-related events.Mobile device use during stairway negotiation increases the likelihood of distraction-induced events. Stair users should be encouraged to limit or avoid mobile device use in public stairway environments. Mobile manufacturers and mobile app developers could aim to develop strategies or mobile app alerts to reduce the impact of distractions (e.g., mobile device use) during stair negotiation to lessen the health and financial burden associated with fall-related events on stairways.

BackgroundVariability in the riser and depth of each step in a stairway can decrease step predictability, and increase users’ fall risk. Unfortunately, inconsistencies in inter-step riser height and depth are common, but their impact may be lessened by adding high-contrast tread edges and vertical, monochrome striping applied to the bottom and top steps’ faces. Such contrast enhancement may draw greater attention to the steps’ edges or enhance the precision of the edge’s estimated location.PurposeTo determine if greater inconsistency in inter-step riser height and depth are associated with more frequent slips, trips, and falls, and if these events are reduced in flights of stairs with contrast-enhanced step edges.MethodsStair users were videotaped on two public stairways. One stairway had black vinyl strips applied to the steps’ edges and black-and-white vertical stripes on the first and last steps’ faces. The intervention stairway was switched halfway through data collection. Stair users were coded for whether they experienced a fall-related event. A Monte Carlo simulation was used to determine the probability of observing a range of plausible distributions of fall-related events.ResultsFlights of stairs with riser height variability of 14 mm and tread depth variability of 38 mm were associated with 80% of the observed fall-related events. 13 of 16 (81%) fall-related events occurred on the control stairway compared to 3 of 16 (19%) on the intervention stairway. The distribution of fall-related events observed between conditions had a probability of occurring by random chance of less than 0.04.ConclusionThese data suggest a vision-based strategy (i.e., striping) may counteract fall risk associated with inter-step riser height and depth inconsistencies. While the mechanisms of its action remain unclear, the high-contrast striping appears to reduce the incidence of fall-related events in the presence of inter-step riser height and depth inconsistencies.

Falls on stairs occur frequently and pose a significant health and financial risk. Laboratory research has found that fall frequency can be reduced through contrast enhancement of the stairs by applying vinyl striping to the first and last steps’ faces, and all the steps’ top edges. Here we sought to determine if such step contrast enhancement can reduce the probability of fall-related events such as loss of balance, slips, trips, and falls in public university staircases. Cameras were used to capture stair users’ ascent and descent on contrast enhanced (‘striped’) and control stairways. Observed age group, observed gender, traverse direction (ascent, descent), fall-related events, and walking speed (m/s) were recorded. Fall-related events were reduced for the striped stairway compared to the control stairway (odds ratio = 2.87, average marginal effect = 0.002, p = .023) when controlling for observed gender, age category, and traverse direction. These results suggest contrast enhancement of first and last steps’ face and all steps’ edges may reduce fall- related events in public settings. Adding contrast enhancement to public stairways is a simple and cost-effective way to reduce the loss of balance, slips, and trips that could lead to catastrophic falls on stairs as well as the health and financial burden associated with such falls.PRECISContrast enhancement of stairway features may be a simple and cost-effective way to reduce falls on stairs. We show that monochrome striping on the first and last steps’ faces and black stripes on steps’ top edges reduce the probability of fall-related events in public stairways.

In young adults, performance on a test of response inhibition was recently found to be correlated with performance on a reactive balance test where automated stepping responses must occasionally be inhibited. The present study aimed to determine whether this relationship holds true in older adults, wherein response inhibition is typically deficient and the control of postural equilibrium presents a greater challenge. Ten participants (50+ years of age) completed a seated cognitive test (stop signal task) followed by a reactive balance test. Reactive balance was assessed using a modified lean-and-release system where participants were required to step to regain balance following perturbation, or suppress a step if an obstacle was present. The stop signal task is a standardized cognitive test that provides a measure of the speed of response inhibition called the Stop Signal Reaction Time (SSRT). Muscle responses in the legs were compared between conditions where a step was allowed or blocked to quantify response inhibition of the step. The SSRT was significantly related to leg muscle suppression during balance recovery in the stance leg. Thus, participants that were better at inhibiting their responses in the stop signal task were also better at inhibiting an unwanted leg response in favor of grasping a supportive handle. The relationship between a seated cognitive test using finger responses and leg muscle suppression when a step was blocked indicates a context-independent, generalized capacity for response inhibition. This suggests that a simple cognitive test such as the stop signal task could be used clinically to predict an individual’s capacity for adapting balance reactions and fall risk. The present results provide support for future studies, with larger samples, to verify this relationship between stop signal reaction time and leg response during balance recovery.

Key points Considerable debate exists regarding whether electrical vestibular stimuli encoded by vestibular afferents induce a net signal of linear acceleration, rotation or a combination of the two. This debate exists because an isolated signal of head rotation encoded by the vestibular afferents can cause perceptions of both linear and angular motion. We recorded participants' perceptions in different orientations relative to gravity and predicted their responses by modelling the effect of electrical vestibular stimuli on vestibular afferents and a current model of central vestibular processing. We show that, even if electrical vestibular stimuli are encoded as a net signal of head rotation, participants perceive both linear acceleration and rotation motions, provided the electrical stimulation-induced rotational vector has a component orthogonal to gravity. The emergence of a perception of linear acceleration from a single rotational input signal clarifies the origins of the neural mechanisms underlying electrical vestibular stimulation. Abstract Electrical vestibular stimulation (EVS) is an increasingly popular biomedical tool for generating sensations of virtual motion in humans, for which the mechanism of action is a topic of considerable debate. Contention surrounds whether the evoked vestibular afferent activity encodes a signal of net rotation and/or linear acceleration. Central processing of vestibular self-motion signals occurs through an internal representation of gravity that can lead to inferred linear accelerations in absence of a true inertial acceleration. Applying this model to virtual signals of rotation evoked by EVS, we predict that EVS will induce behaviours attributed to both angular and linear motion, depending on the head orientation relative to gravity. To demonstrate this, 18 subjects indicated their perceived motion during sinusoidal EVS when in one of four head/body positions orienting the gravitational vector parallel or orthogonal to the EVS rotation vector. During stimulation, participants selected one simulated movement from seven that corresponded best to what they perceived. Participants' responses in each orientation were predicted by a model combining the influence of EVS on vestibular afferents with known mechanisms of vestibular processing. When the EVS rotation vector had a component orthogonal to gravity, human perceptual responses were consistent with a non-zero central estimate of interaural or superior-inferior linear acceleration. The emergence of a perception of linear acceleration from a single rotational input signal clarifies the origins of the neural mechanisms underlying EVS, which has important implications for its use in human biomedical or sensory augmentation applications.

The ability to move and maintain posture is critically dependent on motion and orientation information provided by the vestibular system. When this system delivers noisy or erred information it can, in some cases, be attenuated through habituation. Here we investigate whether multiple mechanisms of attenuation act to decrease vestibular gain due to noise added using supra-threshold random-waveform galvanic vestibular stimulation (GVS). Forty-five participants completed one of three conditions. Each condition consisted of two 4-min standing periods with stimulation surrounding a 1-h period of either walking with stimulation, walking without stimulation, or sitting quietly. An instrumented treadmill recorded horizontal forces at the feet during standing and walking. We quantified response attenuation to GVS by comparing vestibular stimulus-horizontal force gain between conditions. First stimulus exposure caused an 18% decrease in gain during the first 40 s of standing. Attenuation recommenced only when subjects walked with stimulation, resulting in a 38% decrease in gain over 60 min that did not transfer to standing following walking. The disparity in attenuation dynamics and absent carry over between standing and walking suggests that two mechanisms of attenuation, one associated with first exposure to the stimulus and another that is task specific, may act to decrease vestibulomotor gain.

The vestibular system encodes motion and orientation of the head in space and is essential for negotiating in and interacting with the world. Recently, random waveform electric vestibular stimulation has become an increasingly common means of probing the vestibular system. However, many of the methods used to analyze the behavioral response to this type of stimulation assume a linear relationship between frequencies in the stimulus and its associated response. Here we examine this stimulus-response frequency linearity to determine the validity of this assumption. Forty-five university-aged subjects stood on a force-plate for 4 min while receiving vestibular stimulation. To determine the linearity of the stimulus-response relationship we calculated the cross-frequency power coupling between a 0 and 25 Hz bandwidth limited white noise stimulus and induced postural responses, as measured using the horizontal forces acting at the feet. Ultimately, we found that, on average, the postural response to a random stimulus is linear across stimulation frequencies. This result supports the use of analysis methods that depend on the assumption of stimulus-response frequency linearity, such as coherence and gain, which are commonly used to analyze the body’s response to random waveform electric stimuli.

Recent studies have shown balance recovery can be enhanced via task-specific training, referred to as perturbation-based balance training (PBT). These interventions rely on principles of motor learning where repeated exposure to task-relevant postural perturbations results in more effective compensatory balance responses. Evidence indicates that compensatory responses trained using PBT can be retained for many months and can lead to a reduction in falls in community-dwelling older adults. A notable shortcoming with PBT is that it does not transfer well to similar but contextually different scenarios (e.g., falling sideways versus a forward trip). Given that it is not feasible to train all conditions in which someone could fall, this limited transfer presents a conundrum; namely, how do we best use PBT to appropriately equip people to deal with the enormous variety of fall-inducing scenarios encountered in daily life? In this perspective article, we draw from fields of research that explore how general learning can be promoted. From this, we propose a series of methods, gleaned from parallel streams of research, to inform and hopefully optimize this emerging field where people receive training to specifically improve their balance reactions.

We tested the hypothesis that the brain uses a variance-based weighting of multisensory cues to estimate head rotation to perceive which way is up. The hypothesis predicts that the known bias in perceived vertical, which occurs when the visual environment is rotated in a vertical-plane, will be reduced by the addition of visual noise. Ten healthy participants sat head-fixed in front of a vertical screen presenting an annulus filled with coloured dots, which could rotate clockwise or counter-clockwise at six angular velocities (1, 2, 4, 6, 8, 16°/s) and with six levels of noise (0, 25, 50, 60, 75, 80%). Participants were required to keep a central bar vertical by rotating a hand-held dial. Continuous adjustments of the bar were required to counteract low-amplitude low-frequency noise that was added to the bar's angular position. During visual rotation, the bias in verticality perception increased over time to reach an asymptotic value. Increases in visual rotation velocity significantly increased this bias, while the addition of visual noise significantly reduced it, but did not affect perception of visual rotation velocity. The biasing phenomena were reproduced by a model that uses a multisensory variance-weighted estimate of head rotation velocity combined with a gravito-inertial acceleration signal (GIA) from the vestibular otoliths. The time-dependent asymptotic behaviour depends on internal feedback loops that act to pull the brain's estimate of gravity direction towards the GIA signal. The model's prediction of our experimental data furthers our understanding of the neural processes underlying human verticality perception.

Many diseases, disabilities, and mental health conditions associated with aging can be delayed or prevented through regular exercise. Several barriers to exercise, many of which are exacerbated in rural communities, prevent mid-life and older adults from accessing its benefits. However, recently, a racquet sport named pickleball has become popular among older adults, and it appears to overcome some of these barriers. We conducted a feasibility study to evaluate the impact of a six-week pickleball intervention on measures of muscle function, cognitive function, perceived pain, and cardio-metabolic risk, as well as several psychosocial factors contributing to adherence in sedentary rural participants. Participants improved their vertical jump, cognitive performance, and reported a decrease in self-reported pain, suggesting improved physical and cognitive health across the sample. Participants also reported high levels of satisfaction and demonstrated good adherence over the duration of the study. Perhaps of greatest value was the overwhelmingly positive response from participants to the intervention and follow-up interviews reporting a desire to continue pickleball play beyond the study period. Overall, pickleball appears to be a promising intervention to, (1) elicit functional- and cognitive-related improvements, and (2) motivate mid-life and older adults to adhere to exercise sufficiently long to benefit their health.

During walking, the vestibular influence on locomotor activity is phase-dependent and modulated in both limbs with changes in velocity. It is unclear, however, whether this bilateral modulation is due to a coordinated mechanism between both limbs or instead through limb-specific processes that remain masked by the symmetric nature of locomotion. Here, human subjects walked on a split-belt treadmill with one belt moving at 0.4 m s−1 and the other moving at 0.8 m s−1 while exposed to an electrical vestibular stimulus. Muscle activity was recorded bilaterally around the ankles of each limb and used to compare vestibulo-muscular coupling between velocity-matched and unmatched tied-belt walking. In general, response magnitudes decreased by ∼20–50% and occurred ∼13–20% earlier in the stride cycle at the higher belt velocity. This velocity-dependent modulation of vestibular-evoked muscle activity was retained during split-belt walking and was similar within each limb to velocity-matched tied-belt walking. These results demonstrate that the vestibular influence on ankle muscles during locomotion can be adapted independently to each limb. Furthermore, modulation of vestibular-evoked muscle responses occurred rapidly (∼13–34 strides) after onset of split-belt walking. This rapid adaptation contrasted the prolonged adaptation in step length symmetry (∼128 strides) as well as EMG magnitude and timing (∼40–100 and ∼20–80 strides, respectively). These results suggest that vestibular influence on ankle muscle control is adjusted rapidly in sensorimotor control loops as opposed to longer-term error correction mechanisms commonly associated split-belt adaptation. Rapid limb-specific sensorimotor feedback adaptation may be advantageous for asymmetric overground locomotion, such as navigating irregular terrain or turning.

The soleus (Sol) and medial gastrocnemius (mGas) muscles have different patterns of activity during standing balance and may have distinct functional roles. Using surface electromyography we previously observed larger responses to galvanic vestibular stimulation (GVS) in the mGas compared with the Sol muscle. However, it is unclear whether this difference is an artifact that reflects limitations associated with surface electromyography recordings or whether a compensatory balance response to a vestibular error signal activates the mGas to a greater extent than the Sol. In the present study, we compared the effect of GVS on the discharge behavior of 9 Sol and 21 mGas motor units from freely standing subjects. In both Sol and mGas motor units, vestibular stimulation induced biphasic responses in measures of discharge timing [11 ± 5.0 (mGas) and 5.6 ± 3.8 (Sol) counts relative to the sham (mean ± SD)], and frequency [0.86 ± 0.6 Hz (mGas), 0.34 ± 0.2 Hz (Sol) change relative to the sham]. Peak-to-trough response amplitudes were significantly larger in the mGas (62% in the probability-based measure and 160% in the frequency-based measure) compared with the Sol (multiple P < 0.05). Our results provide direct evidence that vestibular signals have a larger influence on the discharge activity of motor units in the mGas compared with the Sol. More tentatively, these results indicate the mGas plays a greater role in vestibular-driven balance corrections during standing balance.

To interact successfully with an uncertain environment, organisms must be able to respond to both unanticipated and anticipated events. For unanticipated events, organisms have evolved stereotyped motor behaviors mapped to the statistical regularities of the environment, which can be trigged by specific sensory stimuli. These "reflexive" responses are more or less hardwired to prevent falls and represent, maybe, the best available solution to maintaining posture given limited available time and information. With the gift of foresight, however, motor behaviors can be tuned or prepared in advance, improving the ability of the organism to compensate for, and interact with, the changing environment. Indeed, foresight's improvement of our interactive capacity occurs through several means, such as better action selection, processing, and conduction delay compensation and by providing a prediction with which to compare our actual behaviors to, thereby facilitating error identification and learning. Here we review the various roles foresight (prediction) plays in maintaining our postural equilibrium. We start by describing some of the more recent findings related to the prediction of instability. Specifically, we cover recent advancements in the understanding of anticipatory postural behaviors that are used broadly to stabilize volitional movement and compensate for impending postural disturbances. We also describe anticipatory changes in the state, or set, of the nervous system that may facilitate anticipatory behaviors. From changes in central set, we briefly discuss prediction of postural instability online before moving into a discussion of how predictive mechanisms, such as internal models, permit us to tune, perhaps our highest level predictive behaviors, namely the priming associated with motor affordances. Lastly, we explore methods best suited to expose the contribution of prediction to postural equilibrium control across a variety of contexts.

The neural control of transition between posture and movement encompasses the regulation of reflex-stabilizing mechanisms to enable motion. Optimal feedback theory suggests that such transitions require the disengagement of one motor control policy before the implementation of another. To test this possibility, we investigated the continuity of the vestibular control of balance during transitions between quiet standing and locomotion and between two standing postures. Healthy subjects initiated and terminated locomotion or shifted the distribution of their weight between their feet, while exposed to electrical vestibular stimuli (EVS). The relationship between EVS and ground reaction forces was quantified using time-frequency analyses. Discontinuities corresponding to null coherence periods were observed preceding the onset of movement initiation and during the step preceding locomotion termination. These results show humans interrupt the vestibular balance stabilizing mechanisms to transition between motor states, suggesting a discrete change between motor control policies, as predicted by optimal feedback theory., eLife digest Crossing Abbey Road is something of a paradox in neuroscientific terms. As you stand waiting to cross, tiny movements of your body – such as those due to breathing – cause you to sway by small amounts. To prevent you from falling over, your brain makes active corrections to your posture. These posture-correcting mechanisms oppose movements such as sway and keep you standing upright. But what happens when you want to cross the road? To get you moving, your brain has two options. It could temporarily suppress the posture-correcting mechanisms. Or it could reconfigure them so that they work in a different way. The posture-correcting mechanisms rely upon sensory input from various sources. These include the vestibular system of the inner ear. The vestibular system tells the brain about the position and movement of the head in space and relative to gravity. Monitoring vestibular system activity as a person starts to move should thus reveal what is happening to the posture-correcting mechanisms. Tisserand et al. asked healthy volunteers to transition between standing still and walking, or to shift their weight from one foot to the other. At the same time, small non-painful electric currents were applied to the bones behind the volunteers' ears. These currents induced small changes in vestibular system activity. Sensors in the floor measured the forces the volunteers generated while standing or walking, thereby revealing how they adjusted their balance. The results showed that the brain suppresses its posture-correcting mechanisms before people start or stop moving. These findings have implications for robotics. They could make it easier to program robots to show smooth transitions into and out of movement. The findings are also relevant to movement disorders such as Parkinson's disease. One common symptom of this disorder is freezing of gait, in which patients suddenly feel as though their feet are glued to the ground. Understanding how the brain controls movement transitions may reveal how such symptoms arise.

Gravity is a defining force that governs the evolution of mechanical forms, shapes and anchors our perception of the environment, and imposes fundamental constraints on our interactions with the world. Within the animal kingdom, humans are relatively unique in having evolved a vertical, bipedal posture. Although a vertical posture confers numerous benefits, it also renders us less stable than quadrupeds, increasing susceptibility to falls. The ability to accurately and precisely estimate our orientation relative to gravity is therefore of utmost importance. Here we review sensory information and computational processes underlying gravity estimation and verticality perception. Central to gravity estimation and verticality perception is multisensory cue combination, which serves to improve the precision of perception and resolve ambiguities in sensory representations by combining information from across the visual, vestibular, and somatosensory systems. We additionally review experimental paradigms for evaluating verticality perception, and discuss how particular disorders affect the perception of upright. Together, the work reviewed here highlights the critical role of multisensory cue combination in gravity estimation, verticality perception, and creating stable gravity-centered representations of our environment.

Rectification of surface electromyographic (EMG) recordings prior to their correlation with other signals is a widely used form of preprocessing. Recently this practice has come into question, elevating the subject of EMG rectification to a topic of much debate. Proponents for rectifying suggest it accentuates the EMG spike timing information, whereas opponents indicate it is unnecessary and its nonlinear distortion of data is potentially destructive. Here we examine the necessity of rectification on the extraction of muscle responses, but for the first time using a known oscillatory input to the muscle in the form of electrical vestibular stimulation. Participants were exposed to sinusoidal vestibular stimuli while surface and intramuscular EMG were recorded from the left medial gastrocnemius. We compared the unrectified and rectified surface EMG to single motor units to determine which method best identified stimulus-EMG coherence and phase at the single-motor unit level. Surface EMG modulation at the stimulus frequency was obvious in the unrectified surface EMG. However, this modulation was not identified by the fast Fourier transform, and therefore stimulus coherence with the unrectified EMG signal failed to capture this covariance. Both the rectified surface EMG and single motor units displayed significant coherence over the entire stimulus bandwidth (1–20 Hz). Furthermore, the stimulus-phase relationship for the rectified EMG and motor units shared a moderate correlation ( r = 0.56). These data indicate that rectification of surface EMG is a necessary step to extract EMG envelope modulation due to motor unit entrainment to a known stimulus.

Key points The vestibular system is an important sensory contributor to the control of standing balance. Fear, anxiety and arousal are thought to influence the excitability of the vestibular system, but it is not clear if these changes lead to altered vestibular-evoked balance reflexes. Low and high standing surface heights were used to manipulate fear and anxiety in this study, while stochastic vestibular stimulation was used to evoke balance reflexes. High surface heights lead to greater coupling between vestibular inputs and balance reflexes, as well as larger responses. These results support the idea that the manner in which vestibular information is processed is altered when people are exposed to a threat to their balance, and this altered processing may explain why normal balance behaviour is different in threatening scenarios.

In a standing position, the vertical projection of the center of mass passes in front of the ankle, which requires active plantar-flexor torque from the triceps surae to maintain balance. We recorded motor unit (MU) activity in the medial (MG) and lateral (LG) gastrocnemius muscle and the soleus (SOL) in standing balance and voluntary isometric contractions to understand the effect of functional requirements and descending drive from different neural sources on motoneuron behavior. Single MU activity was recorded in seven subjects with wire electrodes in the triceps surae. Two 3-min standing balance trials and several ramp-and-hold contractions were performed. Lateral gastrocnemius MU activity was rarely observed in standing. The lowest thresholds for LG MUs in ramp contractions were 20–35 times higher than SOL and MG MUs ( P < 0.001). Compared with MUs from the SOL, MG MUs were intermittently active ( P < 0.001), had higher recruitment thresholds ( P = 0.022), and greater firing rate variability ( P < 0.001); this difference in firing rate variability was present in standing balance and isometric contractions. In SOL and MG MUs, both recruitment of new MUs ( R2 = 0.59–0.79, P < 0.01) and MU firing rates ( R2 = 0.05–0.40, P < 0.05) were associated with anterior-posterior and medio-lateral torque in standing. Our results suggest that the two heads of the gastrocnemius may operate in different ankle ranges with the larger MG being of primary importance when standing, likely due to its fascicle orientation. These differences in MU discharge behavior were independent of the type of descending neural drive, which points to a muscle-specific optimization of triceps surae motoneurons.

Our perception of verticality relies on combining sensory information from multiple sources. Neuronal recordings in animals implicate the cerebellum in the process, yet disease of the human cerebellum was not found to affect this perception. Here we show that a perceptual disturbance of verticality is indeed present in people with a genetically determined and pure form of cerebellar degeneration (spinocerebellar ataxia type 6; SCA 6), but is only revealed under dynamic visual conditions. Participants were required to continuously orient a visually displayed bar to vertical while the bar angle was perturbed by a low-frequency random signal and a random dot pattern rotated in their visual periphery. The random dot pattern was rotated at one of two velocities (4°/s and 16°/s), traveling with either coherent or noisy motion. Perceived vertical was biased by visual rotation in healthy participants, particularly in a more elderly group, but SCA 6 participants were biased more than both groups. The bias was reduced by visual noise, but more so for SCA 6 participants than young controls. Distortion of verticality by visual rotation stems from the stimulus creating an illusion of self-rotation. We modeled this process using a maximum-likelihood sensory cue-combination model operating on noisy visual- and vestibular-rotation signals. The observed effects of visual rotation and visual noise could be compellingly explained by cerebellar degeneration, and to a lesser extent aging, causing an increase in central vestibular noise. This is consistent with the human cerebellum operating on dynamic vestibular signals to inform the process that estimates which way is up.

Fear of falling (FoF) is an autonomic, cognitive and behavioural response to a real or imminent threat of a fall (Hadjistavropoulos et al. 2011). Given its multidimensional nature, the assessment of fear usually relies on self-report and converging evidence from a number of independent but related measures, such as anxiety (Davis et al. 2010), confidence (self-efficacy) and arousal (Hadjistavropoulos et al. 2011). Recent studies, in which young healthy adults have been exposed to a postural threat (e.g. standing on an elevated platform where the consequences of a fall are severe), have shown that fear and related factors directly affect control of quiet standing and dynamic balance responses (Brown & Frank, 1997; Carpenter et al. 2001, 2004; Sibley et al. 2010); however, the mechanisms underlying these fear-related postural changes are yet to be understood fully. Of recent debate is whether FoF-related changes in vestibular function, as evidenced by modulation (or lack thereof) of vestibular-evoked balance responses, might contribute to threat-related adjustments in balance behaviours (Osler et al. 2013; Horslen et al. 2014). Our position is that FoF does influence vestibular-evoked balance responses.

Vestibular pathways form short-latency disynaptic connections with neck motoneurons, whereas they form longer-latency disynaptic and polysynaptic connections with lower limb motoneurons. We quantified frequency responses of vestibular reflexes in neck, back, and lower limb muscles to explain between-muscle differences. Two hypotheses were evaluated: 1) that muscle-specific motor-unit properties influence the bandwidth of vestibular reflexes; and 2) that frequency responses of vestibular reflexes differ between neck, back, and lower limb muscles because of neural filtering. Subjects were exposed to electrical vestibular stimuli over bandwidths of 0–25 and 0–75 Hz while recording activity in sternocleidomastoid, splenius capitis, erector spinae, soleus, and medial gastrocnemius muscles. Coherence between stimulus and muscle activity revealed markedly larger vestibular reflex bandwidths in neck muscles (0–70 Hz) than back (0–15 Hz) or lower limb muscles (0–20 Hz). In addition, vestibular reflexes in back and lower limb muscles undergo low-pass filtering compared with neck-muscle responses, which span a broader dynamic range. These results suggest that the wider bandwidth of head-neck biomechanics requires a vestibular influence on neck-muscle activation across a larger dynamic range than lower limb muscles. A computational model of vestibular afferents and a motoneuron pool indicates that motor-unit properties are not primary contributors to the bandwidth filtering of vestibular reflexes in different muscles. Instead, our experimental findings suggest that pathway-dependent neural filtering, not captured in our model, contributes to these muscle-specific responses. Furthermore, gain-phase discontinuities in the neck-muscle vestibular reflexes provide evidence of destructive interaction between different reflex components, likely via indirect vestibular-motor pathways.

Daily activities, such as walking, may require dynamic modulation of vestibular input onto motoneurons. This dynamic modulation is difficult to identify in humans due to limitations in the delivery and analysis of current vestibular probes, such as galvanic vestibular stimulation. Stochastic vestibular stimulation, however, provides an alternative method to extract human vestibular reflexes. Here, we used time-dependent coherence and time-dependent cross-correlation, coupled with stochastic vestibular stimulation, to investigate the phase dependency of human vestibular reflexes during locomotion. We found that phase-dependent activity from the medial gastrocnemius muscles is correlated with the vestibular signals over the 2- to 20-Hz bandwidth during the stance phase of locomotion. Vestibular-gastrocnemius coherence and time-dependent cross-correlations reached maximums at 21 ± 4 and 23 ± 8% of the step cycle following heel contact and before the period of maximal electromyographic activity (38 ± 5%). These results demonstrate 1) the effectiveness of these techniques in extracting the phase-dependent modulation of vestibulomuscular coupling during a cyclic task; 2) that vestibulomuscular coupling is phasically modulated during locomotion; and 3) that the period of strongest vestibulomuscular coupling does not correspond to the period of maximal electromyographic activity in the gastrocnemius. Therefore, we have shown that stochastic vestibular stimulation, coupled with time-frequency decomposition, provides an effective tool to assess the contribution of vestibular ex-afference to the muscular control during locomotion.

Passing current through mastoid electrodes (conventionally termed galvanic vestibular stimulation; GVS) evokes a balance response containing a short- and a medium-latency response. The origins of these two responses are debated. Here we test the hypotheses that they originate from net signals evoked by stimulation of otolith and semi-circular canal afferents, respectively. Based on anatomy and function, we predicted the directions of the stimulus-evoked net head rotation vector from the canals and the linear acceleration net vector from the otoliths. We tested these predictions in healthy adults by obtaining responses with the head in strategic postures to alter the relevance of the signals to the balance system. Cross-covariance between a stochastic waveform of stimulating current and motor output was used to assess the balance responses. Consistent with the canal hypothesis, with the head pitched down the medium-latency EMG response was abolished while the short-latency EMG response was maintained. The results, however, did not support the otolith hypothesis. The direction of the linear acceleration signal from the otoliths was predicted to change substantially when using monaural stimuli compared to binaural stimuli. In contrast, short-latency response direction measured from ground-reaction forces was not altered. It was always directed along the inter-aural axis irrespective of whether the stimulus was applied binaurally or monaurally, whether the head was turned in yaw through 90 deg, whether the head was pitched down through 90 deg, or combinations of these manipulations. We conclude that a net canal signal evoked by GVS contributes to the medium-latency response whilst a net otolith signal does not make a significant contribution to either the short- or medium-latency responses.

Galvanic vestibular stimulation (GVS) results in characteristic muscle and whole-body responses in humans maintaining standing balance. However, the relationship between these two vestibular-evoked responses remains elusive. This study seeks to determine whether mechanical filtering from conversion of lower-limb muscle activity to body sway, during standing balance, can be used to attenuate sway while maintaining biphasic lower-limb muscle responses using frequency-limited stochastic vestibular stimulation (SVS). We hypothesized that SVS deprived of frequencies 2 Hz to standing humans do not cause a destabilizing whole-body sway response but are associated with the typical biphasic lower-limb muscle responses.

Presenting a startling stimulus in a simple reaction time (RT) task, can involuntarily trigger the pre-programmed response. However, this effect is not seen when the response is programmed following the imperative stimulus (IS) providing evidence that a startle can only trigger pre-programmed responses. In a "Go/No-go" (GNG) RT task the response may be programmed in advance of the IS because there exists only a single predetermined response. The purpose of the current investigation was to examine if startle could elicit a response in a GNG task. Participants completed a wrist extension task in response to a visual stimulus. A startling acoustic stimulus (124dB) was presented in both Go and No-go trials with Go probability manipulated between groups. The inclusion of a startle did not significantly speed RT and led to more response errors. This result is similar to that observed in a startled choice RT task, indicating that in a GNG task participants waited until the IS complete motor programming.

Stochastic vestibular stimulation (SVS) can be used to study the postural responses to unpredictable vestibular perturbations. The present study seeks to determine if stochastic vestibular stimulation elicits lower limb muscular responses and to estimate the frequency characteristics of these vestibulo-motor responses in humans. Fourteen healthy subjects were exposed to unpredictable galvanic currents applied on their mastoid processes while quietly standing (±3 mA, 0–50 Hz). The current amplitude and stimulation configuration as well as the subject's head position relative to their feet were manipulated in order to determine that: (1) the muscle responses evoked by stochastic currents are dependent on the amplitude of the current, (2) the muscle responses evoked by stochastic currents are specific to the percutaneous stimulation of vestibular afferents and (3) the lower limb muscle responses exhibit polarity changes with different head positions as previously described for square-wave galvanic vestibular stimulation (GVS) pulses. Our results revealed significant coherence (between 0 and 20 Hz) and cumulant density functions (peak responses at 65 and 103 ms) between SVS and the lower limbs' postural muscle activity. The polarity of the cumulant density functions corresponded to that of the reflexes elicited by square-wave GVS pulses. The SVS–muscle activity coherence and time cumulant functions were modulated by current amplitude, electrode position and head orientation with respect to the subject's feet. These findings strongly support the vestibular origin of the lower limb muscles evoked by SVS. In addition, specific frequency bandwidths in the stochastic vestibular signal contributed to the early (12–20 Hz) and late components (2–10 Hz) of the SVS-evoked muscular responses. These frequency-dependent SVS-evoked muscle responses support the view that the biphasic muscle response is conveyed by two distinct physiological processes.

Our vertical bipedal posture is one of the defining traits that separates us from our phylogenetic peers. Not surprisingly, the mechanisms underlying how we maintain this orientation have come under intense scientific scrutiny. This article is protected by copyright. All rights reserved

Electrical vestibular stimulation is often used to assess vestibulo-motor and postural responses in both clinical and research settings. Stochastic vestibular stimulation (SVS) is a recently established technique with many advantages over its square-wave counterpart; however, the evoked muscle responses remain relatively small. Although the vestibular-evoked responses can be enhanced by increasing the stimulus amplitude, subjects often perceive these higher intensity electrical stimuli as noxious or painful. Here, we developed multisine vestibular stimulation (MVS) signals that include precise frequency contributions to increase signal-to-noise ratios (SNR) of stimulus-evoked muscle and motor responses. Subjects were exposed to three different MVS stimuli to establish that: 1) MVS signals evoke equivalent vestibulo-motor responses compared to SVS while improving subject comfort and reducing experimentation time, 2) stimulus-evoked vestibulo-motor responses are reliably estimated as a linear system and 3) specific components of the cumulant density time domain vestibulo-motor responses can be targeted by controlling the frequency content of the input stimulus. Our results revealed that in comparison to SVS, MVS signals increased the SNR 3–6 times, reduced the minimum experimentation time by 85% and improved subjective measures of comfort by 20–80%. Vestibulo-motor responses measured using both EMG and force were not substantially affected by nonlinear distortions. In addition, by limiting the contribution of high frequencies within the MVS input stimulus, the magnitude of the medium latency time domain motor output response was increased by 58%. These results demonstrate that MVS stimuli can be designed to target and enhance vestibulo-motor output responses while simultaneously improving subject comfort, which should prove beneficial for both research and clinical applications.

Patients with vestibular dysfunction have visual, perceptual, and postural deficits. While there is considerable evidence that a semicircular canal prosthesis that senses angular head velocity and stimulates canal ampullary nerves can improve vision by augmenting the vestibulo-ocular reflex, no information is available regarding the potential utility of a canal prosthesis to improve perceptual deficits. In this study we investigated the possibility that electrical stimulation of canal afferents could be used to modify percepts of head orientation. Two rhesus monkeys were trained to align a light bar parallel to gravity, and were tested in the presence and absence of electrical stimulation provided by an electrode implanted in the right posterior canal. While the monkeys aligned the light bar close to the true earth-vertical without stimulation, when the right posterior canal was stimulated their responses deviated towards their left ear, consistent with a misperception of head tilt towards the right. The deviation of the light bar from the earth-vertical exceeded the torsional deviation of the eyes, indicating that the perceptual changes were not simply visual in origin. Eye movements recorded during electrical stimulation in the dark were consistent with isolated activation of right posterior canal afferents, with no evidence of otolith stimulation. These results demonstrate that electrical stimulation of canal afferents affects the perception of head orientation, and therefore suggest that motion-modulated stimulation of canal afferents by a vestibular prosthesis could potentially improve vestibular percepts in patients lacking normal vestibular function.

Vestibular information is one of the many sensory signals used to stabilize the body during locomotion. When locomotor velocity increases, the influence of these signals appears to wane. It is unclear whether vestibular signals are globally attenuated with velocity or are influenced by factors such as whether a muscle is contributing to balance control. Here we investigate how vestibular sensory signals influence muscles of the leg during locomotion and what causes their attenuation with increasing locomotor velocity. We hypothesized that 1) vestibular signals influence the activity of all muscles engaged in the maintenance of medio-lateral stability during locomotion and 2) increases in both cadence and velocity would be associated with attenuation of these signals. We used a stochastic vestibular stimulus and recorded electromyographic signals from muscles of the ankle, knee, and hip. Participants walked using two cadences (52 and 78 steps/min) and two walking velocities (0.4 and 0.8 m/s). We observed phase-dependent modulation of vestibular influence over ongoing muscle activity in all recorded muscles. Within a stride, reversals of the muscle responses were observed in the biceps femoris, tibialis anterior, and rectus femoris. Vestibular-muscle coupling decreases with increases in both cadence and walking velocity. These results show that the observed vestibular suppression is muscle- and phase dependent. We suggest that the phase- and muscle-specific influence of vestibular signals on locomotor activity is organized according to each muscle's functional role in body stabilization during locomotion.

Electrical vestibular stimulation produces biphasic responses in muscles maintaining balance. The two components of these muscle responses (termed the short latency and medium latency components) are believed to be independent and elicited by vestibular stimuli of different frequencies. We tested these hypotheses by determining (a) if frequency-specific stimulation protocols could evoke independently the short and medium latency responses and (b) whether these two components are triggered by distinct brain regions with a fixed time delay, interacting around 10 Hz. First, subjects were provided 10–25 Hz, 0–10 Hz, and 0–25 Hz vestibular stimuli to selectively modulate the short latency, medium latency, or both components of the response; and second, they were provided twenty sinusoidal stimuli from 1 to 20 Hz with a 0–20 Hz control trial, designed to determine whether an interaction between the short and medium latency responses occurs at a specific stimulation frequency. Both the 0–10 Hz and 10–25 Hz vestibular stimuli elicited multiphasic waveforms, suggesting the short and medium latency components were not modulated independently by the frequency-specific stimuli. Sinusoidal vestibular stimuli evoked responses at the stimulated frequency but no evidence of a reflex component interaction was observed. Instead, summation of the responses evoked by each of the sinusoidal stimuli resembled the biphasic response to broad bandwidth stimuli. Due to the lack of interaction and linear contribution of all stimulus frequencies to both the short and medium latency responses, the present results support the use of broad bandwidth electrical vestibular signal for physiological or clinical testing.

The purpose of the study was to determine whether the central nervous system (CNS) requires the sensory feedback generated by balance perturbations in order to trigger postural responses (PRs). In Experiment 1, twenty-one participants experienced toes-up support-surface tilts in two blocks. Control blocks involved unexpected balance perturbations whereas an auditory tone cued the onset of balance perturbations in Conditioning blocks. A single Cue-Only trial followed each block (Cue-Only Control and Cue-Only Conditioning trials) in the absence of balance perturbations. Cue-Only Conditioning trials were used to determine whether postural perturbations were required in order to trigger PRs. Counter-balancing the order of Control and Conditioning blocks allowed Cue-Only Control trials to examine both the audio–spinal/acoustic startle effects of the auditory cue and the carryover effects of the initial conditioning procedure. In Experiment 2, six participants first experienced five consecutive Tone-Only trials that were followed by twenty-five conditioning trials. After conditioning, five Tone-Only trials were again presented consecutively to first elicit and then extinguish the conditioned PRs. Surface electromyography (EMG) recorded muscle activity in soleus (SOL), tibialis anterior (TA) and rectus femoris (RF). EMG onset latencies and amplitudes were calculated together with the onset latency, peak and time-to-peak of shank angular accelerations. Results indicated that an auditory cue could be conditioned to initiate PRs in multiple muscles without balance-relevant sensory triggers generated by balance perturbations. Postural synergies involving excitation of TA and RF and inhibition of SOL were observed following the Cue-Only Conditioning trials that resulted in shank angular accelerations in the direction required to counter the expected toes-up tilt. Postural synergies were triggered in response to the auditory cue even 15 min post-conditioning. Furthermore, conditioned PRs were quickly extinguished as participants became unresponsive by the third trial in extinction. In conclusion, our results reveal that the CNS does not require sensory feedback from postural perturbations in order to trigger PRs.

Stochastic vestibular stimulation (SVS) can be used to study the postural responses to unpredictable vestibular perturbations. The present study seeks to determine if stochastic vestibular stimulation elicits lower limb muscular responses and to estimate the frequency characteristics of these vestibulo-motor responses in humans. Fourteen healthy subjects were exposed to unpredictable galvanic currents applied on their mastoid processes while quietly standing (+/-3 mA, 0-50 Hz). The current amplitude and stimulation configuration as well as the subject's head position relative to their feet were manipulated in order to determine that: (1) the muscle responses evoked by stochastic currents are dependent on the amplitude of the current, (2) the muscle responses evoked by stochastic currents are specific to the percutaneous stimulation of vestibular afferents and (3) the lower limb muscle responses exhibit polarity changes with different head positions as previously described for square-wave galvanic vestibular stimulation (GVS) pulses. Our results revealed significant coherence (between 0 and 20 Hz) and cumulant density functions (peak responses at 65 and 103 ms) between SVS and the lower limbs' postural muscle activity. The polarity of the cumulant density functions corresponded to that of the reflexes elicited by square-wave GVS pulses. The SVS-muscle activity coherence and time cumulant functions were modulated by current amplitude, electrode position and head orientation with respect to the subject's feet. These findings strongly support the vestibular origin of the lower limb muscles evoked by SVS. In addition, specific frequency bandwidths in the stochastic vestibular signal contributed to the early (12-20 Hz) and late components (2-10 Hz) of the SVS-evoked muscular responses. These frequency-dependent SVS-evoked muscle responses support the view that the biphasic muscle response is conveyed by two distinct physiological processes.

The topic of debate is whether fear influences vestibular-evoked balance responses. Opposing arguments from Reynolds et al. (2015) rely on the reported lack of changes in the early ( 800 ms) responses. The evidence drawn from the lack of early response change is unconvincing due to an increased likelihood of type II error caused by a small sample size and poor resolution resulting from the following factors: (i) use of centre of mass position and velocity estimates to detect vestibular-evoked responses in leg muscles, especially as tandem-stance requires multijoint control; (ii) the limited number of stimulations; and (iii) small early responses due to low (1 mA) stimulus intensity (Ali et al. 2003). Likewise, interpretation of late responses is confounded by the potential influence of fear on reactions to non-vestibular sensory feedback related to the vestibular perturbation (Day & Guerraz, 2007) and conscious strategies (Reynolds, 2010) within this time period. We disagree with Reynolds et al. (2015) that fear-related increases in gain and coupling of ground reaction forces (GRFs) to stochastic vestibular stimulation (SVS) are irrelevant to balance. Early responses are clearly distinguishable using GRFs (Marsden et al. 2002), with less mechanical filtering of the neuromuscular response affecting GRFs than kinematics (Forbes et al. 2015). In our study, SVS was engineered to highlight changes in early balance responses to vestibular stimulation that are not confounded by later perturbation-induced sensorimotor responses (Horslen et al. 2014). The significant SVS gain changes observed between 2 and 16 Hz coincides with natural head frequencies recorded during gait and whole-body perturbations (for review, see Forbes et al. 2015). After establishing threat-related effects on early vestibular-evoked balance responses, we confirmed these effects in GRFs and EMG from posturally engaged muscles using an SVS stimulus that included the 0–2 Hz bandwidth and with vestibular-evoked myogenic potentials (see Horslen et al. 2015). The notion that we have confused statistical and functional significance is false; we view evidence of a true effect, beyond chance or bias, as a necessary prerequisite to considering functional significance. Furthermore, functional significance should not be discounted because of small absolute changes, which would be expected given the small virtual rotational velocity induced by the stimulus (Peters et al. 2015) and are consistent with magnitudes reported elsewhere (Reynolds, 2010; Mian & Day, 2014). In contrast, we believe that the relative change in gain of early vestibular-evoked responses provides more meaningful insight into functional significance. Thus, fear-related increases in gain (81–231%) are non-trivial and functionally significant, particularly when translated to real-life head accelerations.

Recent experiments pairing a startling stimulus with a simple reaction time (RT) task have shown that when participants are startled, a prepared movement was initiated earlier in comparison to voluntary initiation. It has been argued that the startle acts to trigger the response involuntarily. However, an alternative explanation is that the decrease in RT may be due to stimulus intensity effects, not involuntary triggering. Thus the aim of the current investigation was to determine if RT simply declined in a linear fashion with increasing stimulus intensity, or if there was a point at which RT dramatically decreased. In the present experiment participants completed 50 active wrist extension trials to a target in response to an auditory stimulus of varying stimulus intensity (83–123 dB). The presented data show that RTs associated with a startle response are separate from stimulus intensity facilitated responses. Furthermore, this startle facilitation is more highly associated with sternocleidomastoid electromyographic (EMG) activity, rather than the EMG from the widely used startle response indicator muscle orbicularis oculi.