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.@en

Vestibular signals, which encode head movement in space as well as orientation relative to gravity, contribute to the ongoing muscle activity required to stand. The strength of this vestibular contribution changes with the presence and quality of sensory cues of balance. Here we investigate whether the vestibular drive for standing balance also depends on different sensory cues of gravity by examining vestibular-evoked muscle responses when independently varying load and gravity conditions. Standing subjects were braced by a backboard structure that limited whole-body sway to the sagittal plane while load and vestibular cues of gravity were manipulated by: (a) loading the body downward at 1.5 and 2 times body weight (i.e., load cues), and/or (b) exposing subjects to brief periods (20 s) of micro- (<0.05 g) and hyper-gravity (∼1.8 g) during parabolic flights (i.e., vestibular cues). A stochastic electrical vestibular stimulus (0-25 Hz) delivered during these tasks evoked a vestibular-error signal and corrective muscles responses that were used to assess the vestibular drive to standing balance. With additional load, the magnitude of the vestibular-evoked muscle responses progressively increased, however, when these responses were normalized by the ongoing muscle activity, they decreased and plateaued at 1.5 times body weight. This demonstrates that the increased muscle activity necessary to stand with additional load is accompanied a proportionally smaller increase in vestibular input. This reduction in the relative vestibular contribution to balance was also observed when we varied the vestibular cues of gravity, but only during an absence (<0.05 g) and not an excess (∼1.8 g) of gravity when compared to conditions with normal 1 g gravity signals and equivalent load signals. Despite these changes, vestibular-evoked responses were observed in all conditions, indicating that vestibular cues of balance contribute to upright standing even in the near absence of a vestibular signal of gravity (i.e., micro-gravity). Overall, these experiments provide evidence that both load and vestibular cues of gravity influence the vestibular signal processing for the control of standing balance.@en

Background Effective sensorimotor integration is essential to modulate (adapt) neck stabilization strategies in response to varying tasks and disturbances. This study evaluates the hypothesis that relative to healthy controls cervical dystonia patients have an impaired ability to modulate afferent feedback for neck stabilization with changes in the frequency content of mechanical perturbations. Methods We applied anterior-posterior displacement perturbations (110 s) on the torso of seated subjects, while recording head-neck kinematics and muscular activity. We compared low bandwidth (0.2–1.2 Hz) and high bandwidth (0.2–8 Hz) perturbations where our previous research showed a profound modulation of stabilization strategies in healthy subjects. Cervical dystonia patients and age matched controls performed two tasks: (1) maintain head forward posture and (2) allow dystonia to dictate head posture. Findings Patients and controls demonstrated similar kinematic and muscular responses. Patient modulation was similar to that of healthy controls (P > 0.05); neck stiffness and afferent feedback decreased with high bandwidth perturbations. During the head forward task patients had an increased neck stiffness relative to controls (P @en

Botulinum toxin (BoNT) injections in the dystonic muscles is the preferred treatment for Cervical Dystonia (CD), but the proper identification of the dystonic muscles remains a challenge. Previous studies showed decreased 8–14 Hz autospectral power in the electromyography (EMG) of splenius muscles in CD patients. Cumulative distribution functions (CDF’s) of dystonic muscles showed increased CDF10 values, representing increased autospectral powers between 3 and 10 Hz, relative to power between 3 and 32 Hz. In this study, we evaluated both methods and investigated the effects of botulinum toxin. Intramuscular EMG recordings were obtained from the splenius, semispinalis, and sternocleidomastoid muscles during standardized isometric tasks in 4 BoNT-naïve CD patients, 12 BoNT-treated patients, and 8 healthy controls. BoNT-treated patients were measured 4–7 weeks after their last BoNT injections and again after 11–15 weeks. We found significantly decreased 8–14 Hz autospectral power in splenius muscles, but not in the semispinalis and sternocleidomastoid muscles of CD patients when compared to healthy controls. CDF10 analysis was superior in demonstrating subtle autospectral changes, and showed increased CDF10 values in all studied muscles of CD patients. These results did not change significantly after BoNT injections. Further studies are needed to investigate the origin of these autospectral changes in dystonia patients, and to assess their potential in muscle selection for BoNT treatment@en

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.@en

Neck muscle activity evoked by vestibular stimuli is a clinical measure for evaluating the function of the vestibular apparatus. Cervical vestibular-evoked myogenic potentials (cVEMP) are most commonly measured in the sternocleidomastoid muscle (and more recently the splenius capitis muscle) in response to air-conducted sound, bone-conducted vibration or electrical vestibular stimuli. It is currently unknown, however, whether and how other neck muscles respond to vestibular stimuli. Here we measured activity bilaterally in the sternocleidomastoid, splenius capitis, sternohyoid, semispinalis capitis, multifidus, rectus capitis posterior, and obliquus capitis inferior using indwelling electrodes in two subjects exposed to binaural bipolar electrical vestibular stimuli. All recorded neck muscles responded to the electrical vestibular stimuli (0-100 Hz) provided they were active. Furthermore, the evoked responses were inverted on either side of the neck, consistent with a coordinated contribution of all left-right muscle pairs acting as antagonists in response to the electrically-evoked vestibular error of head motion. Overall, our results suggest that, as previously observed in cat neck muscles, broad connections exist between the human vestibular system and neck motoneurons and highlight the need for future investigations to establish their neural connections.@en

The human head-neck system requires continuous stabilization in the presence of gravity and trunk motion. We investigated contributions of the vestibulocollic reflex (VCR), the cervicocollic reflex (CCR), and neck muscle co-contraction to head-in-space and head-on-trunk stabilization, and investigated modulation of the stabilization strategy with the frequency content of trunk perturbations and the presence of visual feedback. We developed a multisegment cervical spine model where reflex gains (VCR and CCR) and neck muscle co-contraction were estimated by fitting the model to the response of young healthy subjects, seated and exposed to anterior-posterior trunk motion, with frequency content from 0.3 up to 1, 2, 4 and 8 Hz, with and without visual feedback. The VCR contributed to head-in-space stabilization with a strong reduction of head rotation (<8 Hz) and a moderate reduction of head translation (>1 Hz). The CCR contributed to head-on-trunk stabilization with a reduction of head rotation and head translation relative to the trunk (<2 Hz). The CCR also proved essential to stabilize the individual intervertebral joints and prevent neck buckling. Co-contraction was estimated to be of minor relevance. Control strategies employed during low bandwidth perturbations most effectively reduced head rotation and head relative displacement up to 3 Hz while control strategies employed during high bandwidth perturbations reduced head global translation between 1 and 4 Hz. This indicates a shift from minimizing head-on-trunk rotation and translation during low bandwidth perturbations to minimizing head-in-space translation during high bandwidth perturbations. Presence of visual feedback had limited effects suggesting increased usage of vestibular feedback. @en

Objective To identify effects of a deviant motor drive in the autospectral power of dystonic muscles during voluntary contraction in cervical dystonia patients. Methods Submaximal (20%) isometric head-neck tasks were performed with the head fixed, measuring surface EMG of the sternocleidomastoid, splenius capitis and semispinalis capitis in CD patients and controls. Autospectral power of muscle activity, and head forces was analyzed using cumulative distribution functions (CDF). A downward shift between the theta/low alpha-band (3-10 Hz) and the high alpha/beta-band (10-30 Hz) was detected using the CDF10, defined as the cumulative power from 3-10 Hz relative to power from 3-30 Hz. Results CDF10 was increased in dystonic muscles compared to controls and patient muscles unaffected by dystonia, due to a 3-10 Hz power increase and a 10-30 Hz decrease. CDF10 also increased in patient head forces. Conclusions Submaximal isometric contractions with the head fixed provided a well-defined test condition minimizing effects of reflexive feedback and tremor. We associate shifts in autospectral power with prokinetic sensorimotor control. Significance Analysis of autospectral power in isometric tasks with the head fixed is a promising approach in research and diagnostics of cervical dystonia.@en

Background: While standing upright, the brain must accurately accommodate for delays between sensory feedback and self-generated motor commands. Natural aging may limit adaptation to sensorimotor delays due to age-related decline in sensory acuity, neuromuscular capacity and cognitive function. This study examined balance learning in young and older adults as they stood with robot-induced sensorimotor delays. Methods: A cohort of community dwelling young (mean = 23.6 years, N = 20) and older adults (mean = 70.1 years, N = 20) participated in this balance learning study. Participants stood on a robotic balance simulator which was used to artificially impose a 250 ms delay into their control of standing. Young and older adults practiced to balance with the imposed delay either with or without visual feedback (i.e., eyes open or closed), resulting in four training groups. We assessed their balance behavior and performance (i.e., variability in postural sway and ability to maintain upright posture) before, during and after training. We further evaluated whether training benefits gained in one visual condition transferred to the untrained condition. Results: All participants, regardless of age or visual training condition, improved their balance performance through training to stand with the imposed delay. Compared to young adults, however, older adults had larger postural oscillations at all stages of the experiments, exhibited less relative learning to balance with the delay and had slower rates of balance improvement. Visual feedback was not required to learn to stand with the imposed delay, but it had a modest effect on the amount of time participants could remain upright. For all groups, balance improvements gained from training in one visual condition transferred to the untrained visual condition. Conclusion: Our study reveals that while advanced age partially impairs balance learning, the older nervous system maintains the ability to recalibrate motor control to stand with initially destabilizing sensorimotor delays under differing visual feedback conditions.@en

Standing balance relies on the integration of multiple sensory inputs to generate the motor commands required to stand. Mechanical and sensory perturbations elicit compensatory postural responses that are interpreted as a window into the sensorimotor processing involved in balance control. Popular methods involve imposed external perturbations that disrupt the control of quiet stance. Although these approaches provide critical information on how the balance system responds to external disturbances, the control mechanisms involved in correcting for these errors may differ from those responsible for the regulation of quiet standing. Alternative approaches use manipulations of the balance control loop to alter the relationship between sensory and motor cues. Coupled with imposed perturbations, these manipulations of the balance control loop provide unique opportunities to reveal how sensory and motor signals are integrated to control the upright body. In this review, we first explore imposed perturbation approaches that have been used to investigate the neural control of standing balance. We emphasize imposed perturbations that only elicit balance responses when the disturbing stimuli are relevant to the balance task. Next, we highlight manipulations of the balance control loop that, when carefully implemented, replicate and/or alter the sensorimotor dynamics of quiet standing. We further describe how manipulations of the balance control loop can be used in combination with imposed perturbations to characterize mechanistic principles underlying the control of standing balance. We propose that recent developments in the use of robotics and sensory manipulations will continue to enable new possibilities for simulating and/or altering the sensorimotor control of standing beyond compensatory responses to imposed external perturbations.@en

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.@en

During standing balance, vestibular signals encode head movement and are transformed into coordinates that are relevant to maintaining upright posture of the whole body. This transformation must account for head-on-body orientation as well as the muscle actions generating the postural response. Here, we investigate whether this transformation is dependent upon a muscle’s ability to stabilize the body along the direction of a vestibular disturbance. Subjects were braced on top of a robotic balance system that simulated the mechanics of standing while being exposed to an electrical vestibular stimulus that evoked a craniocentric vestibular error of head roll. The balance system was limited to move in a single plane while the vestibular error direction was manipulated by having subjects rotate their head in yaw. Vestibular-evoked muscle responses were greatest when the vestibular error was aligned with the balance direction and decreased to zero as the two directions became orthogonal. This demonstrates that muscles respond only to the component of the error that is aligned with the balance direction and thus relevant to the balance task, not to the cumulative afferent activity, as expected for vestibulospinal reflex loops. When we reversed the relationship between balancing motor commands and associated vestibular sensory feedback, the direction of vestibular-evoked ankle compensatory responses was also reversed. This implies that the nervous system quickly reassociates new relationships between vestibular sensory signals and motor commands related to maintaining balance. These results indicate that vestibular-evoked muscle activity is a highly flexible balance response organized to compensate for vestibular disturbances.@en

During standing balance, vestibular signals encode head movement and are transformed into coordinates that are relevant to maintaining upright posture of the whole body. This transformation must account for head-on-body orientation as well as the muscle actions generating the postural response. Here, we investigate whether this transformation is dependent upon a muscle’s ability to stabilize the body along the direction of a vestibular disturbance. Subjects were braced on top of a robotic balance system that simulated the mechanics of standing while being exposed to an electrical vestibular stimulus that evoked a craniocentric vestibular error of head roll. The balance system was limited to move in a single plane while the vestibular error direction was manipulated by having subjects rotate their head in yaw. Vestibular-evoked muscle responses were greatest when the vestibular error was aligned with the balance direction and decreased to zero as the two directions became orthogonal. This demonstrates that muscles respond only to the component of the error that is aligned with the balance direction and thus relevant to the balance task, not to the cumulative afferent activity, as expected for vestibulospinal reflex loops. When we reversed the relationship between balancing motor commands and associated vestibular sensory feedback, the direction of vestibular-evoked ankle compensatory responses was also reversed. This implies that the nervous system quickly reassociates new relationships between vestibular sensory signals and motor commands related to maintaining balance. These results indicate that vestibular-evoked muscle activity is a highly flexible balance response organized to compensate for vestibular disturbances.@en

Human upright posture of both the head and body is facilitated by the CNS’s ability to integrate multiple sensory feedback signals, as well as its discernibility of the motor commands that maintain this stabilization. The vestibular organ in particular detects motion of the head-in-space, which is transformed according to on-going head and body orientation into appropriate motor responses. However, when motor commands do not contribute to the control of standing posture, and are incongruent with their expected sensory consequences, vestibulomuscular responses in the lower limb undergo unconscious suppression. In this study, we investigated whether vestibular response suppression occurs in neck muscles under conditions where the muscles are active but not engaged in a task to balance the head. In addition, we examined the effects of head orientation to identify spatial transformation of vestibular reflex responses. Eight subje cts were exposed to stochastic vestibular stimulation (0-75 Hz) in a seated condition while their head was either free or fixed, and rotated at either 0 or 60°. In head-free conditions, subjects were asked to rotate their head 60° to the left in order to activate agonist neck muscle pairs (sternocleidomastoid - SCM and splenius capitis - SPL). In headfixed conditions, subjects performed isometric neck muscle contractions in yaw at orientations of 0° and 60°, as well as flexion, extension and co-contraction at an orientation of 0°. Intramuscular EMG was collected bilaterally in SCM and SPL muscles. Muscle responses correlated to the in put stimuli were significant (P < 0.05) for all conditions provided the muscle was used in contraction. Neither muscle underwent the expected vestibulomuscular suppression when not engaged in the balance task (i.e. head-fixed). Nevertheless, the magnitude of the SPL responses decreased by 22% when the head was fixed whereas SCM responses were unaffected. The effect of head fixation only in SPL suggests differences in neural pathways across muscles, possibly via alternative pathways known to exist in the SPL from the well-established monosynaptic vestibulospinal inputs in SCM and SPL. For both muscles, the effect of orientation and force direction had no effect on muscles responses. Since the stimulation is fixed relative to the head, the same muscles are activated to respond to the input stimulus at both orientations and all force directions. These results indicate that the vestibular pathways connecting neck muscles are less susceptible to suppression than lower limb muscles, most likely because the monosynaptic inputs innervating them are subject to less central control.@en

Human upright posture of both the head and body is facilitated by the CNS’s ability to integrate multiple sensory feedback signals, as well as its discernibility of the motor commands that maintain this stabilization. The vestibular organ in particular detects motion of the head-in-space, which is transformed according to on-going head and body orientation into appropriate motor responses. However, when motor commands do not contribute to the control of standing posture, and are incongruent with their expected sensory consequences, vestibulomuscular responses in the lower limb undergo unconscious suppression. In this study, we investigated whether vestibular response suppression occurs in neck muscles under conditions where the muscles are active but not engaged in a task to balance the head. In addition, we examined the effects of head orientation to identify spatial transformation of vestibular reflex responses. Eight subje cts were exposed to stochastic vestibular stimulation (0-75 Hz) in a seated condition while their head was either free or fixed, and rotated at either 0 or 60°. In head-free conditions, subjects were asked to rotate their head 60° to the left in order to activate agonist neck muscle pairs (sternocleidomastoid - SCM and splenius capitis - SPL). In headfixed conditions, subjects performed isometric neck muscle contractions in yaw at orientations of 0° and 60°, as well as flexion, extension and co-contraction at an orientation of 0°. Intramuscular EMG was collected bilaterally in SCM and SPL muscles. Muscle responses correlated to the in put stimuli were significant (P < 0.05) for all conditions provided the muscle was used in contraction. Neither muscle underwent the expected vestibulomuscular suppression when not engaged in the balance task (i.e. head-fixed). Nevertheless, the magnitude of the SPL responses decreased by 22% when the head was fixed whereas SCM responses were unaffected. The effect of head fixation only in SPL suggests differences in neural pathways across muscles, possibly via alternative pathways known to exist in the SPL from the well-established monosynaptic vestibulospinal inputs in SCM and SPL. For both muscles, the effect of orientation and force direction had no effect on muscles responses. Since the stimulation is fixed relative to the head, the same muscles are activated to respond to the input stimulus at both orientations and all force directions. These results indicate that the vestibular pathways connecting neck muscles are less susceptible to suppression than lower limb muscles, most likely because the monosynaptic inputs innervating them are subject to less central control.@en

Galvanic vestibular stimulation (GVS) uses the external application of electrical current to selectively target the vestibular system in humans. Despite its recent popularity for the assessment/treatment of clinical conditions, exactly how this non-invasive tool activates the vestibular system remains an open question. Here we directly investigate single vestibular afferent responses to GVS applied to the mastoid processes of awake-behaving monkeys. Transmastoid GVS produces robust and parallel activation of both canal and otolith afferents. Notably, afferent activation increases with intrinsic neuronal variability resulting in constant GVS-evoked neuronal detection thresholds across all afferents. Additionally, afferent tuning differs for GVS versus natural self-motion stimulation. Using a stochastic model of repetitive activity in afferents, we largely explain the main features of GVS-evoked vestibular afferent dynamics. Taken together, our results reveal the neural substrate underlying transmastoid GVS-evoked perceptual, ocular and postural responses—information that is essential to advance GVS applicability for biomedical uses in humans.@en