Stepping in circles: how locomotor signals of rotation adapt over time.

KEY POINTS: While it has been well described that prolonged rotational stepping will adapt the podokinetic sense of rotation, the mechanisms involved are not clearly understood. By studying podokinetic after-rotations following conditioning rotations not previously reported we have shown that slower rotational velocities are more readily adapted than faster velocities and adaptation occurs more quickly than previously thought. We propose a dynamic feedback model of vestibular and podokinetic adaptation that can fit rotation trajectories across multiple conditions and data sets. Two adaptation processes were identified that may reflect central and peripheral processes and the discussion unifies prior findings in the podokinetic literature under this new framework. The findings show the technique is feasible for people with locomotor turning problems. ABSTRACT: After a prolonged period stepping in circles, people walk with a curved trajectory when attempting to walk in a straight line without vision. Podokinetic adaptation shows promise in clinical populations to improve locomotor turning; however, the adaptive mechanisms involved are poorly understood. The first phase of this study asks: how does the podokinetic conditioning velocity affect the response velocity and how quickly can adaptation occur? The second phase of the study asks: can a mathematical feedback model account for the rotation trajectories across different conditioning parameters and different datasets? Twelve healthy participants stepped in place on the axis of a rotating surface ranging from 4 to 20 deg s-1 for durations of 1-10 min, while using visual cues to maintain a constant heading direction. Afterward on solid ground, participants were blindfolded and attempted to step without rotating. Participants unknowingly stepped in circles opposite to the direction of the prior platform rotation for all conditions. The angular velocity of this response peaked within 1 min and the ratio of the stimulus-to-response peak velocity fitted a decreasing power function. The response then decayed exponentially. The feedback model of podokinetic and vestibular adaptive processes had a good fit with the data and suggested that podokinetic adaptation is explained by a short (141 s) and a long (27 min) time constant. The podokinetic system adapts more quickly than previously thought and subjects adapt more readily to slower rotation than to faster rotation. These findings will have implications for clinical applications of the technique.

Physiological stress markers during breath-hold diving and SCUBA diving.

This study investigated the sources of physiological stress in diving by comparing SCUBA dives (stressors: hydrostatic pressure, cold, and hyperoxia), apneic dives (hydrostatic pressure, cold, physical activity, hypoxia), and dry static apnea (hypoxia only). We hypothesized that despite the hypoxia induces by a long static apnea, it would be less stressful than SCUBA dive or apneic dives since the latter combined high pressure, physical activity, and cold exposure. Blood samples were collected from 12SCUBA and 12 apnea divers before and after dives. On a different occasion, samples were collected from the apneic group before and after a maximal static dry apnea. We measured changes in levels of the stress hormones cortisol and copeptin in each situation. To identify localized effects of the stress, we measured levels of the cardiac injury markers troponin (cTnI) and brain natriuretic peptide (BNP), the muscular stress markers myoglobin and lactate), and the hypoxemia marker ischemia-modified albumin (IMA). Copeptin, cortisol, and IMA levels increased for the apneic dive and the static dry apnea, whereas they decreased for the SCUBA dive. Troponin, BNP, and myoglobin levels increased for the apneic dive, but were unchanged for the SCUBA dive and the static dry apnea. We conclude that hypoxia induced by apnea is the dominant trigger for the release of stress hormones and cardiac injury markers, whereas cold or and hyperbaric exposures play a minor role. These results indicate that subjects should be screened carefully for pre-existing cardiac diseases before undertaking significant apneic maneuvers.

Perceptions of Speed and Risk: Experimental Studies of Road Crossing by Older People.

Crossing a road safely is a complex task requiring good sensorimotor function and integration of information about traffic speed, distances and one's own speed. Poor judgement through age-related sensorimotor or cognitive impairment or a predisposition to take risks could lead to errors with serious consequences. On a simulated road, 85 participants (age ≥70 years) were asked to cross in front of an approaching car with a clearance as small as considered safe in two conditions; (1) with nothing else to attend to (free crossing) and (2) with an additional ball-gathering task while waiting to cross (task crossing). Participants were categorised according to their crossing outcome (failed to cross, 'hit', exact, safe, cautious). Participants also performed two sub-studies; (1) the perception of the time-to-arrival of moving objects and (2) the perception of own gait speed. Physical and cognitive function and everyday risk-taking behaviour were also assessed. In free crossing, clearances varied but no participants were "hit" by the car. In task crossing, participants allowed smaller clearances and 10% of participants would have been hit while 13% missed the opportunity to cross altogether. Across a wide range of physical and cognitive measures, including perceived and actual gait speed, a consistent pattern was observed in the task crossing condition. The exact group performed best, the 'hit', safe and cautious groups performed less well while those who missed the opportunity (fail) performed worst. The exact group reported taking the greatest risks in everyday life whereas the remaining groups reported being cautious. In conclusion, we found older people with poorer perceptual, physical and cognitive function made inappropriate and risky decisions in a divided attention road-crossing task despite self-reports of cautious behaviour in everyday life.

Activation of human inspiratory muscles in an upside-down posture.

During quiet breathing, activation of obligatory inspiratory muscles differs in timing and magnitude. To test the hypothesis that this coordinated activation can be modified, we determined the effect of the upside-down posture compared with standing and lying supine. Subjects (n=14) breathed through a pneumotachometer with calibrated inductance bands around the chest wall and abdomen. Surface electromyographic activity (EMG) was recorded from the scalene muscles. Crural diaphragmatic EMG and oesophageal and gastric pressures were measured in a subset of six subjects. Quiet breathing and standard lung function manoeuvres were performed. The upside-down posture reduced end-expiratory lung volume. During quiet breathing, for the same inspiratory airflow and tidal volume, ribcage contribution decreased, abdominal contribution increased and transdiaphragmatic pressure swing doubled in the upside-down posture compared to standing (p<0.05). Despite this, crural diaphragm EMG was unchanged, whereas scalene muscle EMG was reduced by ∼half (p<0.05). Thus, the mechanical effect of an upside-down posture differentially affects inspiratory muscle activation.

Cross-Modal Calibration of Vestibular Afference for Human Balance.

To determine how the vestibular sense controls balance, we used instantaneous head angular velocity to drive a galvanic vestibular stimulus so that afference would signal that head movement was faster or slower than actual. In effect, this changed vestibular afferent gain. This increased sway 4-fold when subjects (N = 8) stood without vision. However, after a 240 s conditioning period with stable balance achieved through reliable visual or somatosensory cues, sway returned to normal. An equivalent galvanic stimulus unrelated to sway (not driven by head motion) was equally destabilising but in this situation the conditioning period of stable balance did not reduce sway. Reflex muscle responses evoked by an independent, higher bandwidth vestibular stimulus were initially reduced in amplitude by the galvanic stimulus but returned to normal levels after the conditioning period, contrary to predictions that they would decrease after adaptation to increased sensory gain and increase after adaptation to decreased sensory gain. We conclude that an erroneous vestibular signal of head motion during standing has profound effects on balance control. If it is unrelated to current head motion, the CNS has no immediate mechanism of ignoring the vestibular signal to reduce its influence on destabilising balance. This result is inconsistent with sensory reweighting based on disturbances. The increase in sway with increased sensory gain is also inconsistent with a simple feedback model of vestibular reflex action. Thus, we propose that recalibration of a forward sensory model best explains the reinterpretation of an altered reafferent signal of head motion during stable balance.

Passive motion reduces vestibular balance and perceptual responses.

With the hypothesis that vestibular sensitivity is regulated to deal with a range of environmental motion conditions, we explored the effects of passive whole-body motion on vestibular perceptual and balance responses. In 10 subjects, vestibular responses were measured before and after a period of imposed passive motion. Vestibulospinal balance reflexes during standing evoked by galvanic vestibular stimulation (GVS) were measured as shear reaction forces. Perceptual tests measured thresholds for detecting angular motion, perceptions of suprathreshold rotation and perceptions of GVS-evoked illusory rotation. The imposed conditioning motion was 10 min of stochastic yaw rotation (0.5-2.5 Hz ≤ 300 deg s(-2) ) with subjects seated. This conditioning markedly reduced reflexive and perceptual responses. The medium latency galvanic reflex (300-350 ms) was halved in amplitude (48%; P = 0.011) but the short latency response was unaffected. Thresholds for detecting imposed rotation more than doubled (248%; P < 0.001) and remained elevated after 30 min. Over-estimation of whole-body rotation (30-180 deg every 5 s) before conditioning was significantly reduced (41.1 to 21.5%; P = 0.033). Conditioning reduced illusory vestibular sensations of rotation evoked by GVS (mean 113 deg for 10 s at 1 mA) by 44% (P < 0.01) and the effect persisted for at least 1 h (24% reduction; P < 0.05). We conclude that a system of vestibular sensory autoregulation exists and that this probably involves central and peripheral mechanisms, possibly through vestibular efferent regulation. We propose that failure of these regulatory mechanisms at different levels could lead to disorders of movement perception and balance control during standing.

Ability versus hazard: risk-taking and falls in older people.

BACKGROUND: Among older people, undue risk taking could lead to falls, irrespective of physical ability. We investigated the interaction between risk-taking behavior and physical ability and its contribution to falls. METHODS: Participants (N = 294, age ≥ 70) were asked to walk as quickly as possible to a visible destination by choosing one of six paths. Each contained a raised plank that had to be walked along without falling. The shortest path had the narrowest and tallest plank and the longest had the widest and lowest. Behavioral risk was defined as the probability of falling off the chosen plank. This was estimated from a ground path walking task because, for safety, participants were stopped before crossing the plank. Self-reported everyday risk-taking behavior, fear of falling, physical functioning, and 1-year prospective fall rates were measured. RESULTS: Older participants and those with poor physical ability chose easier planks to cross. Participants with good physical ability consistently took a slight behavioral risk, whereas those with poor physical ability took either very-high behavioral risks or chose the overly safe path with no risk. Unexpectedly, participants reporting cautious behavior on the everyday risk-taking behavior scale took greater behavioral risks. Independent of physical performance, behavioral risk was significantly associated with falls during the subsequent year. CONCLUSIONS: Assessing behavioral choice in relation to physical ability can identify risk-taking but neither the difficulty of a chosen action nor self-reports of risk-taking behavior are sufficient. Risk-taking behavior is an independent risk factor for falls and management of undue risk-taking might complement existing fall prevention strategies.

Blood pressure and the contractility of a human leg muscle.

These studies investigate the relationships between perfusion pressure, force output and pressor responses for the contracting human tibialis anterior muscle. Eight healthy adults were studied. Changing the height of tibialis anterior relative to the heart was used to control local perfusion pressure. Electrically stimulated tetanic force output was highly sensitive to physiological variations in perfusion pressure showing a proportionate change in force output of 6.5% per 10 mmHg. This perfusion-dependent change in contractility begins within seconds and is reversible with a 53 s time constant, demonstrating a steady-state equilibrium between contractility and perfusion pressure. These stimulated contractions did not produce significant cardiovascular responses, indicating that the muscle pressor response does not play a major role in cardiovascular regulation at these workloads. Voluntary contractions at forces that would require constant motor drive if perfusion pressure had remained constant generated a central pressor response when perfusion pressure was lowered. This is consistent with a larger cortical drive being required to compensate for the lost contractility with lower perfusion pressure. The relationship between contractility and perfusion for this large postural muscle was not different from that of a small hand muscle (adductor pollicis) and it responded similarly to passive peripheral and active central changes in arterial pressure, but extended over a wider operating range of pressures. If we consider that, in a goal-oriented motor task, muscle contractility determines central motor output and the central pressor response, these results indicate that muscle would fatigue twice as fast without a pressor response. From its extent, timing and reversibility we propose a testable hypothesis that this change in contractility arises through contraction- and perfusion-dependent changes in interstitial K(+) concentration.

Force-controlled balance perturbations associated with falls in older people: a prospective cohort study.

Balance recovery from an unpredictable postural perturbation can be a challenging task for many older people and poor recovery could contribute to their risk of falls. This study examined associations between responses to unpredictable perturbations and fall risk in older people. 242 older adults (80.0 ± 4.4 years) underwent assessments of stepping responses to multi-directional force-controlled waist-pull perturbations. Participants returned monthly falls calendars for the subsequent 12 months. Future falls were associated with lower force thresholds for stepping in the posterior and lateral but not anterior directions. Those with lower posterior force thresholds for stepping were 68% more likely to fall at home than those with higher force thresholds for stepping. These results suggest that amount of force that can be withstood following an unpredictable balance perturbation predicts future falls in community-dwelling older adults. Perturbations in the posterior direction best discriminated between future fallers and non-fallers.

The galvanic whole-body sway response in health and disease.

OBJECTIVE: To explore the galvanic-evoked vestibulospinal reflex in health and disease. METHODS: Vestibular-evoked whole-body movement was measured in 60 controls and eight patients with bilateral vestibular impairment, using 1mA × 2s Galvanic Vestibular Stimulation (GVS). The displacement (s) and velocity (v) of 7 markers placed on the head, neck, shoulders, mid-thorax and hips and ground reaction forces (F) were recorded. RESULTS: Monaural GVS evoked anterolateral whole-body movement away from the cathode. For controls, peak response magnitudes were 22.3 ± 12.4mm, 25.9 ± 14.3 mms(-1) and 3.2 ± 1.3N for s, v and F. Left right asymmetry measured using the Jongkee's formula was 12.59 ± 8.64%, 11.94 ± 11.09% and 11.68 ± 8.99% for s, v and F. All three measures were symmetrical for left and right mastoid stimulation and showed good to excellent test retest reliability. Patients with bilateral vestibular failure had absent or attenuated responses. For this patient group, the ("R/B") ratios between the "response period" and baseline body-sway for s, v and F were 2.1 ± 0.8, 1.7 ± 1.1 and 1.3 ± 0.6, which were significantly lower than 5.4 ± 4.2, 4.1 ± 2.5 and 2.3 ± 0.8 for controls, with s and v providing better separation between controls and patients. There were no significant correlations between the amplitudes of galvanic body sway responses and vestibular-evoked myogenic potentials (VEMPs). CONCLUSION: GVS enables non-invasive assessment of the vestibulospinal reflex. SIGNIFICANCE: This method offers a clinically applicable, test of vestibular contributions to standing balance.

Perception of the postural vertical and falls in older people.

BACKGROUND: Research on the relationship between vestibular function and falls in older people is sparse. The perception of the postural vertical (PPV) provides an indicator measure of vestibular (otolith) function in the absence of visual input and diminished somatosensory feedback. OBJECTIVE: This study examined whether impaired PPV is associated with falls in this group. METHODS: One hundred and ninety-five people aged 70 plus years stood blindfolded on a motorised platform that could be tilted in the roll plane and attempted to adjust it so that their bodies were aligned to the vertical. Somatosensory feedback was minimised as the base and vertical support surfaces on the tilting platform were covered in thick soft foam rubber. PPV error from true vertical and PPV variability (°) were calculated. Participants also underwent an assessment of distal tactile sensitivity and the physiological profile assessment (PPA); fallers were defined as those who had one or more falls during a prospective 12-month follow-up period. RESULTS: Eighty-eight participants (45%) reported falling in the follow-up year. Increased PPV error and variability were correlated with increased lateral sway in a condition of absent visual input and reduced foot somatosensory feedback (eyes closed/foam; r range = 0.16-0.20, p < 0.05) and with composite PPA fall risk scores (r range = 0.22-0.26, p < 0.05). PPV variability was a significant and independent predictor of falls after adjusting for the composite PPA scores, age and gender [adjusted RR = 1.42 (1.01-1.98)]. CONCLUSIONS: Older people with increased PPV variability are at increased risk of falls. These findings indicate that assessment of PPV may augment fall risk assessments in older people.

Sensorimotor and neuropsychological correlates of force perturbations that induce stepping in older adults.

Inappropriate stepping in response to unexpected balance perturbations is more prevalent in older people and in those at risk of falling. This study examined responses to force-controlled waist pulls in young and older people, and sought to identify physiological and cognitive correlates of the force threshold for stepping. 242 older (79.7±4.2 years) and 15 young (29.5±5.3 years) adults underwent waist pull perturbations and assessments of physiological and neuropsychological functioning, general health and falls efficacy. Perturbation force that induced stepping, stepping strategy and number of steps were measured. The older group withstood less forceful perturbations with a feet-in-place strategy, compared to young. Likewise, older adults with high falls risk withstood less force than those with low risk. After controlling for body weight and gender, sway and lower limb strength were independent predictors of anterior stepping thresholds, reaction time was an independent predictor of posterior thresholds, and executive functioning and lower limb strength were independent predictors of the lateral thresholds. These results suggest that balance, strength and agility training, in addition to cognitive exercises may enhance the ability to withstand unexpected balance perturbations and reduce the risk of falls in older people.

Reach distance but not judgment error is associated with falls in older people.

BACKGROUND: Reaching is a vital action requiring precise motor coordination and attempting to reach for objects that are too far away can destabilize balance and result in falls and injury. This could be particularly important for many elderly people with age-related loss of sensorimotor function and a reduced ability to recover balance. Here, we investigate the interaction between reaching ability, errors in judging reach, and the incidence of falling (retrospectively and prospectively) in a large cohort of older people. METHODS: Participants (n = 415, 70-90 years) had to estimate the furthest distance they could reach to retrieve a broomstick hanging in front of them. In an iterative dialog with the experimenter, the stick was moved until it was at the furthest distance they estimated to be reached successfully. At this point, participants were asked to attempt to retrieve the stick. Actual maximal reach was then measured. The difference between attempted reach and actual maximal reach provided a measure of judgment error. One-year retrospective fall rates were obtained at initial assessment and prospective falls were monitored by monthly calendar. RESULTS: Participants with poor maximal reach attempted shorter reaches than those who had good reaching ability. Those with the best reaching ability most accurately judged their maximal reach, whereas poor performers were dichotomous and either underestimated or overestimated their reach with few judging exactly. Fall rates were significantly associated with reach distance but not with reach judgment error. CONCLUSIONS: Maximal reach but not error in perceived reach is associated with falls in older people.

The fusimotor and reafferent origin of the sense of force and weight.

Signals associated with the command the brain sends to muscles are thought to create the sensation of heaviness when we lift an object. Thus, as a muscle is weakened by fatigue or partial paralysis (neuromuscular blockade), the increase in the motor command needed to lift a weight is thought to explain the increasing subjective heaviness of the lifted object.With different fatiguing contractions we approximately halved the force output of the thumb flexor muscles, which were then used to lift an object. For two deafferented subjects the perceived heaviness of the lifted object approximately doubled, in keeping with the central-signal theory. However, for normal subjects this resulted in objects feeling the same or lighter, inconsistent with the central-signal theory but consistent with the expected effects of the conditioning contractions on the sensitivity of peripheral receptors. In separate experiments we subjected the forearm muscles to complete paralysis with a non-depolarising neuromuscular blocking agent and then allowed them to recover to approximately half-force output. This also resulted in objects feeling lighter when lifted by the semi-paralysed thumb, even though the motor command to the motoneurons must have been greater. This is readily explained by reduced lift-related reafference caused by the prolonged paralysis of muscle spindle intrafusal fibres.We conclude that peripheral signals, including a major contribution from muscle spindles, normally give rise to the sense of exerted force. In concept, however, reafference from peripheral receptors may also be considered a centrally generated signal that traverses efferent and then afferent pathways to feed perceptual centres rather than one confined entirely to the central nervous system. These results therefore challenge the distinction between central- and peripheral-based perception, and the concept that muscle spindles provide only information about limb position and movement.

The human semicircular canal model of galvanic vestibular stimulation.

A vector summation model of the action of galvanic stimuli on the semicircular canals has been shown to explain empirical balance and perceptual responses to binaural-bipolar stimuli. However, published data suggest binaural-monopolar stimuli evoke responses that are in the reverse direction of the model prediction. Here, we confirm this by measuring balance responses to binaural-monopolar stimulation as movements of the upper trunk. One explanation for the discrepancy is that the galvanic stimulus might evoke an oppositely directed balance response from the otolith organs that sums with and overrides the semicircular canal response. We tested this hypothesis by measuring sway responses across the full range of head pitch. The results showed some modulation of sway with pitch such that the maximal response occurred with the head in the primary position. However, the effect fell a long way short of that required to reverse the canal sway response. This indicates that the model is incomplete. Here, we examine alterations to the model that could explain both the bipolar and monopolar-evoked behavioural responses. An explanation was sought by remodelling the canal response with more recent data on the orientation of the individual canals. This improved matters but did not reverse the model prediction. However, the model response could be reversed by either rotating the entire labyrinth in the skull or by altering the gains of the individual canals. The most parsimonious solution was to use the more recent canal orientation data coupled with a small increase in posterior canal gain.

Adaptation of vestibular signals for self-motion perception.

A fundamental concern of the brain is to establish the spatial relationship between self and the world to allow purposeful action. Response adaptation to unvarying sensory stimuli is a common feature of neural processing, both peripherally and centrally. For the semicircular canals, peripheral adaptation of the canal-cupula system to constant angular-velocity stimuli dominates the picture and masks central adaptation. Here we ask whether galvanic vestibular stimulation circumvents peripheral adaptation and, if so, does it reveal central adaptive processes. Transmastoidal bipolar galvanic stimulation and platform rotation (20 deg s−1) were applied separately and held constant for 2 min while perceived rotation was measured by verbal report. During real rotation, the perception of turn decayed from the onset of constant velocity with a mean time constant of 15.8 s. During galvanic-evoked virtual rotation, the perception of rotation initially rose but then declined towards zero over a period of ∼100 s. For both stimuli, oppositely directed perceptions of similar amplitude were reported when stimulation ceased indicating signal adaptation at some level. From these data the time constants of three independent processes were estimated: (i) the peripheral canal-cupula adaptation with time constant 7.3 s, (ii) the central 'velocity-storage' process that extends the afferent signal with time constant 7.7 s, and (iii) a long-term adaptation with time constant 75.9 s. The first two agree with previous data based on constant-velocity stimuli. The third component decayed with the profile of a real constant angular acceleration stimulus, showing that the galvanic stimulus signal bypasses the peripheral transformation so that the brainstem sees the galvanic signal as angular acceleration. An adaptive process involving both peripheral and central processes is indicated. Signals evoked by most natural movements will decay peripherally before adaptation can exert an appreciable effect, making a specific vestibular behavioural role unlikely. This adaptation appears to be a general property of the internal coding of self-motion that receives information from multiple sensory sources and filters out the unvarying components regardless of their origin. In this instance of a pure vestibular sensation, it defines the afferent signal that represents the stationary or zero-rotation state.

The sense of self-motion, orientation and balance explored by vestibular stimulation.

The sense of orientation during locomotion is derived from our spatial relationship with the external environment, sensed predominantly by sight and sound, and from internal signals of motion, generated by the vestibular sense and the pattern of efferent and afferent signals to the muscles and joints. The sensory channels operate in different reference frames and have different time-dependent adaptive properties and yet the inputs are combined by the central nervous system to create an internal representation of self-motion. In normal circumstances vestibular, visual and proprioceptive cues provide congruent information on locomotor trajectory; however, in cases of sensory discord there must be a recalibration of sensory signals to provide a unitary representation. We develop a means of studying these fusion processes by perturbing each channel in isolation about a consistent behavioural axis. This review focuses on creating the vestibular perturbation of the orientation sense by transmastoidal galvanic stimulation, a technique generally used to evoke balance reflexes. Vector summation across the population of semicircular canal afferents creates a net signal that is interpreted by the brain as a vector of angular acceleration in a craniocentric reference frame. The signal feeds perceptual processes of orientation after transformation that resolves the 3-D signal onto the terrestrial or behavioural plane. Changing head posture changes the interpretation of the galvanic vestibular signal for balance and orientation responses. With appropriate head alignments during locomotion, the galvanic stimulus can be used to either steer trajectory over the terrestrial plane or perturb balance.

Lack of otolith involvement in balance responses evoked by mastoid electrical stimulation.

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.