Sensory strategies in human postural control before and after unilateral vestibular neurotomy

Vestibular inputs tonically activate the anti-gravitative leg muscles during normal standing in humans, and visual information and proprioceptive inputs from the legs are very sensitive sensory loops for body sway control. This study investigated the postural control in a homogeneous population of 50 unilateral vestibular-deficient patients (Ménière's disease patients). It analyzed the postural deficits of the patients before and after surgical treatment (unilateral vestibular neurotomy) of their diseases and it focused on the visual contribution to the fine regulation of body sway. Static posturographic recordings on a stable force-plate were done with patients with eyes open (EO) and eyes closed (EC). Body sway and visual stabilization of posture were evaluated by computing sway area with and without vision and by calculating the percentage difference of sway between EC and EO conditions. Ménière's patients were examined when asymptomatic, 1 day before unilateral vestibular neurotomy, and during the time-course of recovery (1 week, 2 weeks, 1 month, 3 months, and 1 year). Data from the patients were compared with those recorded in 26 healthy, age- and sex-matched participants. Patients before neurotomy exhibited significantly greater sway area than controls with both EO (+52%) and EC (+93%). Healthy participants and Ménière's patients, however, displayed two different behaviors with EC. In both populations, 54% of the subjects significantly increased their body sway upon eye closure, whereas 46% exhibited no change or significantly swayed less without vision. This was statistically confirmed by the cluster analysis, which clearly split the controls and the patients into two well-identified subgroups, relying heavily on vision (visual strategy, V) or not (non-visual strategy, NV). The percentage difference of sway averaged +36.7%+/-10.9% and -6.2%+/-16.5% for the V and NV controls, respectively; +45.9%+/-16.8% and -4.2%+/-14.9% for the V and NV patients, respectively. These two distinct V and NV strategies seemed consistent over time in individual subjects. Body sway area was strongly increased in all patients with EO early after neurotomy (1 and 2 weeks) and regained preoperative values later on. In contrast, sway area as well as the percentage difference of sway were differently modified in the two subgroups of patients with EC during the early stage of recovery. The NV patients swayed more, whereas the V patients swayed less without vision. This surprising finding, indicating that patients switched strategies with respect to their preoperative behavior, was consistently observed in 45 out of the 50 Ménière's patients during the whole postoperative period, up to 1 year. We concluded that there is a differential weighting of visual inputs for the fine regulation of posture in both healthy participants and Ménière's patients before surgical treatment. This differential weighting was correlated neither with age or sex factors, nor with the clinical variables at our disposal in the patients. It can be accounted for by a different selection of sensory orientation references depending on the personal experience of the subjects, leading to a more or less heavy dependence on vision. The change of sensory strategy in the patients who had undergone neurotomy might reflect a reweighting of the visual and somatosensory cues controlling balance. Switching strategy by means of a new sensory selection of orientation references may be a fast adaptive response to the lesion-induced postural instability.     

Responsiveness and reliability of a pediatric strategy score for balance

Various motor patterns or 'strategies' can be used to maintain balance. The purpose of this study was to determine the responsiveness of a pediatric strategy score (PED-SS) compared to a standard strategy score (SS) as a measure of age-related changes in the force patterns used to maintain stance. Eighty-one healthy children between 3-6 years of age were tested during stance on a force platform while facing a visual surround. The platform, visual surround (or both) moved simultaneously with the child's body sway. Four sensory conditions that altered visual and somatosensory (support surface) inputs were presented. The PED-SS was found to be more responsive to age-related changes in balance behavior compared to the SS. The oldest children (aged 5 and 6 years) showed the greatest ability to utilize horizontal A/P shear force to maintain stance and this finding was reflected only in the PED-SS. The implications of evaluating force strategy as one component of balance in healthy children is discussed with respect to the early developmental assessment of vestibular and developmental coordination disorders.     

Aphysiologic performance on dynamic posturography

The remarkable ability of the body to maintain balance is the result of central nervous system integration of sophisticated inputs from the vestibular, visual, and somatosensory systems. Strategies by patients with balance dysfunction are aphysiologic when their performance is relatively better on more difficult conditions of sensory conflict than on easier ones. Twenty-two aphysiologic patterns on computerized dynamic posturography were compared with age-matched normal and vestibular patterns. The aphysiologic group performed significantly better than the patients in the vestibular dysfunction group on the most difficult subtests of computerized dynamic posturography, conditions 5 and 6, yet significantly poorer on the easier subtests, conditions 1 through 4. In addition, patients in the aphysiologic group tended to show greater intertrial variability compared with patients in both normal and vestibular system dysfunction groups. A stepwise linear discriminant analysis was used to determine a set of conditions that had significant value in discriminating between the three patient groups. Case studies are presented to further illustrate the clinical usefulness of computerized dynamic posturography testing in the evaluation of patients suspected of having a functional component to their on-feet balance problems.     

Ionic currents and electromotility in inner ear hair cells from humans

The upright posture and rich vocalizations of primates place demands on their senses of balance and hearing that differ from those of other animals. There is a wealth of behavioral, psychophysical, and CNS measures characterizing these senses in primates, but no prior recordings from their inner ear sensory receptor cells. We harvested human hair cells from patients undergoing surgical removal of life-threatening brain stem tumors and measured their ionic currents and electromotile responses. The hair cells were either isolated or left in situ in their sensory epithelium and investigated using the tight-seal, whole cell technique. We recorded from both type I and type II vestibular hair cells under voltage clamp and found four voltage-dependent currents, each of which has been reported in hair cells of other animals. Cochlear outer hair cells demonstrated electromotility in response to voltage steps like that seen in rodent animal models. Our results reveal many qualitative similarities to hair cells obtained from other animals and justify continued investigations to explore quantitative differences that may be associated with normal or pathological human sensation.     

Age-associated differences in sensori-motor function and balance in community dwelling women

Tests of visual, vestibular, sensori-motor and balance function were administered to 550 women, aged between 20 and 99 years at a Balance and Gait Laboratory. All of the sensory, motor and balance system measures showed significant age-associated differences. Multiple regression analyses revealed that the measures of lower limb sensation were the consistent sensori-motor factors contributing to balance under normal conditions (standing on a firm surface with eyes open or closed). Under more challenging conditions (standing on foam with eyes open) vision, strength and reaction time played significant roles, whilst when standing on foam with eyes closed, vestibular function also made a significant contribution. Analysis of percentage increases in sway under conditions where visual and peripheral sensation systems were removed or diminished, compared with sway under optimal conditions, indicated that up until age 65 there was an increased reliance on vision for balance control. Beyond this age, the contribution made by vision declined, so that in the oldest age-groups reduced vision was less able to supplement peripheral input, resulting in increased sway areas. Peripheral sensation however was the most important sensory system in the maintenance of static postural stability at all ages.     

Influence of visual control, conduction, and central integration on static and dynamic balance in healthy older adults

Aging is associated with decreased balance abilities, resulting in an increased risk of fall. In order to appreciate the visual, somatosensory, and central signals involved in balance control, sophisticated methods of posturography assessment have been developed, using static and dynamic tests, eventually associated with electromyographic measurements. We applied such methods to a population of healthy older adults in order to appreciate the respective importance of each of these sensorial inputs in aging individuals. Posture control parameters were recorded on a force-measuring platform in 41 healthy young (age 28.5 +/- 5.9 years) and 50 older (age 69.8 +/- 5.9 years) adults, using a static test and two dynamic tests performed by all individuals first with eyes open, then with eyes closed. The distance covered by the center of foot pressure, sway area, and anteroposterior oscillations were significantly higher, with eyes open or closed, in older people than in young subjects. Significant differences were noted in dynamic tests with longer latency responses in the group of old people. Dynamic recordings in a sinusoidal test had a more regular pattern when performed eyes open in both groups and evidenced significantly greater instability in old people. These data suggest that vision remains important in maintaining postural control while conduction and central integration become less efficient with age.     

Adapting reflexes controlling the human posture

Doubt about the role of stretch reflexes in movement and posture control has remained in part because the questions of reflex "usefulness" and the postural "set" have not been adequately considered in the design of experimental paradigms. The intent of this study was to discover the stabilizing role of stretch reflexes acting upon the ankle musculature while human subjects performed stance tasks requiring several different postural "sets". Task specific differences of reflex function were investigated by experiments in which the role of stretch reflexes to stabilize sway doing stance could be altered to be useful, of no use, or inappropriate. Because the system has available a number of alternate inputs to posture (e.g., vestibular and visual), stretch reflex responses were in themselves not necessary to prevent a loss of balance. Nevertheless, 5 out of 12 subjects in this study used long-latency (120 msec) stretch reflexes to help reduce postural sway. Following an unexpected change in the usefulness of stretch reflexes, the 5 subjects progressively altered reflex gain during the succeeding 3-5 trials. Adaptive changes in gain were always in the sense to reduce sway, and therefore could be attenuating or facilitating the reflex response. Comparing subjects using the reflex with those not during so, stretch reflex control resulted in less swaying when the task conditions were unchanging. However, the 5 subjects using reflex controls oftentimes swayed more during the first 3-5 trials after a change, when inappropriate responses were elicited. Four patients with clinically diagnosed cerebellar deficits were studied briefly. Among the stance tasks, their performance was similar to normal in some and significantly poorer in others. Their most significant deficit appeared to be the inability to adapt long-latency reflex gain following changes in the stance task. The study concludes with a discussion of the role of stretch reflexes within a hierarchy of controls ranging from muscle stiffness up to centrally initiated responses.     

Effects on platelet aggregation and fibrinolytic activity during upright posture and exercise in healthy men

The circadian variation of acute myocardial infarction suggests that daily activities such as assuming the upright posture and performing different daily activities may trigger the onset of coronary thrombosis. Such triggering may result from unfavorable alterations in the balance between the prothrombotic and antithrombotic properties of the blood. The present study compares the effects of 2 common daily activities, assuming the upright posture and exercise, on platelet aggregation and fibrinolytic activity. In healthy male subjects, assuming the upright posture in the morning significantly increased platelet aggregation and produced only a moderate increase in fibrinolytic activity within 10 minutes. These changes were still present after 90 minutes in the upright posture. Supine posture for 45 minutes resulted in levels of fibrinolytic activity and platelet aggregation comparable to that observed before initially assuming the upright posture in the morning. Return to the supine posture for 45 minutes resulted in levels of fibrinolytic activity and platelet aggregation comparable to that observed before the initial assumption of upright posture. The changes recurred when upright posture was taken later in the day. Exercise did not increase platelet aggregation to levels beyond that produced by the upright posture, but was associated with a marked increase in fibrinolytic activity. Thus, exercise and upright posture produce distinctive alterations in the thrombogenic potential of the blood that may influence the timing of clinical vascular events.     

Gravitational inputs modulate visuospatial neglect

Right brain-damaged patients with left visuospatial neglect were required to bisect a line placed in front of them in two different body positions (upright and supine) and two different light conditions (light and dark). The neglect patients, unlike right brain-damaged patients without neglect, strongly reduced their rightward directional error in the supine compared with the upright position. No systematic changes were produced by the light-dark manipulation. The present result cannot be explained with an attentional interpretation of hemispatial neglect. We suggest that the present data provide further evidence that hemineglect is the consequence of a mismatch between different afferent information integrated into an egocentric space representation. According to this model, the presence of a lateralized brain lesion produces asymmetries in some intermediate spatial representations (eye-head, head-trunk, body-environment) but not in the retinotopic one. Any experimental manipulation that reduces the asymmetry of the intermediate representation such as the reduction of gravitational inputs may improve the dynamic integration of the egocentric coordinates.     

Adaptive Utilization of Optical Variables During Postural and Suprapostural Dual-Task Performance: Comment on Stoffregen, Smart, Bardy, and Pagulayan (1999).

T. A. Stoffregen, L. J. Smart, B. G. Bardy, and R. J. Pagulayan (1999) combined a postural task (upright stance) with a suprapostural task (visual fixation) to show that sway variability was not driven by optic flow in a task-independent manner (autonomous control) but governed by the demands of the supra-postural task (facilitatory control). The present study used a novel combination of Stoffregen et al.'s task conditions but obtained clear evidence of autonomous control and no indication of facilitatory control. The theoretical adequacy of the stabilization-by-looking versus stabilization-of-looking contrast was examined, as was emerging evidence that posture control and common cognitive tasks place concurrent demands on the same capacity-limited resources. An adaptive resource-sharing view of postural-suprapostural multitasking was proposed as an alternative to both the autonomous- and facilitatory-control views. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Auditory and visual interactions in postural stabilization

The interaction and subsequent interpretation of sensory feedback from different modalities are important determinants in the regulation of balance. The importance of sound in this respect is not, as yet, fully understood. The aim of the present study was to determine the interaction of specific auditory frequencies and vision on postural sway behaviour. The frequencies employed represent the geometrical mean of 23 of the 25 critical bandwidths of sound, each presented at two loudness levels (70 and 90 phones). Postural sway was recorded using a biomechanical measuring platform. As expected vision had a highly significant stabilizing effect on most sway parameters. The frequency of the sound, however, appeared to influence the regulation of anteroposterior sway, while increasing loudness tended to increase mediolateral sway. At some frequencies the sound appeared to compensate for the lack of visual feedback. The interaction of sound and vision, particularly in combinations that lead to increased sway behaviour, may have implications in the occurrence, and possible prevention, of industrial accidents.     

Influence of sensory manipulation on postural control in Parkinson's disease

Postural control was assessed on a tilting platform system in 20 patients with idiopathic Parkinson's disease and 20 age-matched controls. The amount of information provided by vision and lower limb proprioception was varied during the experiment to investigate the influence of changes in sensory cues on postural control. The patient group with clinical evidence of impaired postural control (Hoehn and Yahr III) had significantly higher sway scores over all sensory conditions than either the Hoehn and Yahr II group or controls. The pattern of sway scores indicated that no obvious deficit in the quality, or processing, of sensory information was responsible for the postural instability observed in this group. The patients in both Hoehn and Yahr groups were also able to respond appropriately to potentially destabilising sensory conflict situations and significantly improved their sway scores when provided with visual feedback of body sway. The results indicate that in Parkinson's disease, the main site of dysfunction in postural control is likely to be at a central motor level.     

Clinimetrics of postural instability in Parkinson's disease

Judgement of the ability to recover balance after a sudden shoulder pull is used as a clinical measure of postural instability in Parkinson's disease. To further evaluate its merits, we compared this 'retropulsion test' with dynamic posturography in 23 Parkinson patients. Dynamic posturography involved 20 serial 'toe-up' support surface rotations, which induced backward body sway. We found a moderate correlation (Spearman's p = 0.54; P < 0.05) between the retropulsion test and body sway after platform rotations during the 'off' phase, but no correlation during the 'on' phase (Spearman's p = 0.43; P = 0.11). These results cast doubt on the use of the retropulsion test as a measure of postural instability in Parkinson's disease.     

Functional cortico-muscular coupling during upright standing in athletes and nonathletes: A coherence electroencephalographic-electromyographic study.

We tested the hypothesis that functional cortico-muscular coupling of brain rhythms is implied in the control of lower limb muscles for upright standing. Electroencephalographic (EEG; Be-plus Eb-Neuro) and electromyographic (EMG) data were recorded in 18 fencing and 19 karate elite athletes, 14 karate amateurs, and 9 non-athletes, during quiet upright standing with open and closed eyes conditions. Cortico-muscular coupling was evaluated by computing EEG-EMG spectral coherence and directed transfer function (DTF). Body sway area did not differ among the groups. In non-athletes, the EEG-EMG coherence (gastrocnemius lateralis) at centro-parietal and parasylvian alpha rhythms (about 8-12 Hz) was higher during the open than closed eyes condition. This was not true in the elite athletes. At the same alpha rhythms, the sport amateurs presented values halfway between the non-athletes and elite athletes. Finally, the DTF was higher for cortico-muscular than muscular-cortical direction. These results suggest that visual information affects cortico-muscular coherence at 8-12 Hz in non-athletes and amateur athletes but not in elite athletes. In elite athletes, this might be due to a long training for the control of equilibrium based on proprioceptive and tactile inputs. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Postural stability in diabetic polyneuropathy

OBJECTIVE: To examine whether sensory changes in lower limbs associated with diabetic sensory polyneuropathy compromise postural stability in different visual sensory conditions. RESEARCH DESIGN AND METHODS: The presence and severity of sensory neuropathy was evaluated with a clinical scale and measures of nerve conduction velocity in the lower limbs. Balance control was evaluated by testing subjects' postural stability (with a force platform) with vision, without vision, and during a recovery period after being without vision. RESULTS: Neuropathic patients showed larger ranges of sway, a faster sway speed, and a greater dispersion of sway than control subjects in all conditions. They also exhibited similar or less stable postural performance with vision than that of control subjects without vision. There was a strong relationship between the severity of the neuropathy and the postural stability. CONCLUSIONS: This experiment highlights that even with vision, the postural stability of neuropathic patients is impaired and may put them at higher risk of falling when performing more challenging daily tasks.     

Effect of coordination training on proprioception of the functionally unstable ankle

Exercises to improve joint proprioception and coordination of the functionally unstable ankle are advocated throughout the literature, yet there is little evidence that these exercise have any effect on proprioception and balance. The purpose of this study was to determine the effects of a 6-week coordination and balance training program on proprioception of subjects with functional ankle instability. Forty-five subjects (age = 22.53 +/- 3.95 years, height = 172.04 +/- 10.0 cm, weight = 71.72 +/- 15.7 kg) were randomly placed into a control (Group 1), sham (Group 2), or experimental (Group 3) group. The experimental group trained 3 days per week, 10 minutes each day, performing various balance and proprioception exercises. Postural sway and active and passive joint position sense were assessed. Analysis of variance for postural sway modified equilibrium score for anterior and posterior sway, as well as medial and lateral sway revealed significant four-way interactions. Tukey post hoc analyses revealed that Group 3 performed significantly better (p < .05) than Group 1 and Group 2 on the posttests. There were no significant differences for joint position sense or postural sway index. Results suggest that balance and coordination training can improve some measures of postural sway. It is still unclear if joint position sense can be improved in the functionally unstable ankle.     

Multisensory training of standing balance in older adults: I. Postural stability and one-leg stance balance

BACKGROUND: The effects of standing balance training on the ability to maintain stability in both static two-leg and one-leg stance were tested in healthy older adults. METHODS: Subjects (age range 65-90 years) were randomly assigned to a training (n = 12) or control group (n = 12). Training subjects received a 10-hour balance training program which selectively manipulated sensory inputs from the visual, vestibular, and somatosensory systems. RESULTS: Training subjects showed significantly improved stability (root-mean-square values of anteroposterior platform torque) after training in five of the eight training conditions (when somatosensory inputs were changed or when two or more sensory systems were simultaneously manipulated) (p < .006). When tested 4 weeks after completion of training, subjects (a) fell less frequently when the ankle/foot somatosensory inputs were minimized and (b) stood longer on one leg than the control group (p < .001). CONCLUSIONS: Balance training designed to improve intersensory interaction could effectively improve balance performance in healthy older adults.     

Stiffness control of balance in quiet standing

Our goal was to provide some insights into how the CNS controls and maintains an upright standing posture, which is an integral part of activities of daily living. Although researchers have used simple performance measures of maintenance of this posture quite effectively in clinical decision making, the mechanisms and control principles involved have not been clear. We propose a relatively simple control scheme for regulation of upright posture that provides almost instantaneous corrective response and reduces the operating demands on the CNS. The analytic model is derived and experimentally validated. A stiffness model was developed for quiet standing. The model assumes that muscles act as springs to cause the center-of-pressure (COP) to move in phase with the center-of-mass (COM) as the body sways about some desired position. In the sagittal plane this stiffness control exists at the ankle plantarflexors, in the frontal plane by the hip abductors/adductors. On the basis of observations that the COP-COM error signal continuously oscillates, it is evident that the inverted pendulum model is severely underdamped, approaching the undamped condition. The spectrum of this error signal is seen to match that of a tuned mass, spring, damper system, and a curve fit of this "tuned circuit" yields omega n the undamped natural frequency of the system. The effective stiffness of the system, Ke, is then estimated from Ke = I omega n2, and the damping B is estimated from B = BW X I, where BW is the bandwidth of the tuned response (in rad/s), and I is the moment of inertia of the body about the ankle joint. Ten adult subjects were assessed while standing quietly at three stance widths: 50% hip-to-hip distance, 100 and 150%. Subjects stood for 2 min in each position with eyes open; the 100% stance width was repeated with eyes closed. In all trials and in both planes, the COP oscillated virtually in phase (within 6 ms) with COM, which was predicted by a simple 0th order spring model. Sway amplitude decreased as stance width increased, and Ke increased with stance width. A stiffness model would predict sway to vary as Ke-0.5. The experimental results were close to this prediction: sway was proportional to Ke(-0.55). Reactive control of balance was not evident for several reasons. The visual system does not appear to contribute because no significant difference between eyes open and eyes closed results was found at 100% stance width. Vestibular (otolith) and joint proprioceptive reactive control were discounted because the necessary head accelerations, joint displacements, and velocities were well below reported thresholds. Besides, any reactive control would predict that COP would considerably lag (150-250 ms) behind the COM. Because the average COP was only 4 ms delayed behind the COM, reactive control was not evident; this small delay was accounted for by the damping in the tuned mechanical system.     

Interleukin-15 is an autocrine/paracrine viability factor for cutaneous T-cell lymphoma cells

Prenatal exposure to alcohol is known to affect gross motor functioning. Animal studies have shown that balance is particularly affected, and there is some evidence that similar deficits exist in alcohol-exposed children. In the current study, postural balance, or the ability to maintain equilibrium, was assessed in a group of alcohol-exposed children (ALC group; n = 11) and controls (NC group; n = 11) individually matched for age and sex. Balance was measured across six conditions designed to systematically manipulate or eliminate visual or somatosensory information. Equilibrium and strategy scores for each condition and a derived composite balance score were analyzed. Although the ALC group had a lower mean composite balance score, their performance was similar to that of the NC group on all conditions where somatosensory input was reliable. However, when somatosensory input was manipulated, and when both somatosensory and visual input were inaccurate, the ALC group performed more poorly than controls. Interestingly, there were no differences between the ALC group and NC group in the type of control strategy used to maintain balance. These results suggest that alcohol-exposed children are overly reliant on somatosensory input. When this input is atypical, alcohol-exposed children display significantly greater anterior-posterior body sway and are unable to compensate using available visual or vestibular information. These deficits may be related to cerebellar anomalies previously reported in fetal alcohol syndrome children.     

Interactions between vestibular and proprioceptive inputs triggering and modulating human balance-correcting responses differ across muscles

Interactions between proprioceptive and vestibular inputs contributing to the generation of balance corrections may vary across muscles depending on the availability of sensory information at centres initiating and modulating muscle synergies, and the efficacy with which the muscle action can prevent a fall. Information which is not available from one sensory system may be obtained by switching to another. Alternatively, interactions between sensory systems and the muscle to which this interaction is targeted may be fixed during neural development and not switchable. To investigate these different concepts, balance corrections with three different sets of proprioceptive trigger signals were examined under eyes-open and eyes-closed conditions in the muscles of normal subjects and compared with those of subjects with bilateral peripheral vestibular loss. The different sets of early proprioceptive inputs were obtained by employing three combinations of support surface rotation and translation, for which ankle inputs were nulled, normal or enhanced, the knees were either locked or in flexion, and the trunk was either in flexion or extension. Three types of proprioceptive and vestibulospinal interactions were identified in muscles responses. These interactions were typified by the responses of triceps surae, quadriceps, and paraspinal muscles. The amplitudes of stretch responses at 50 ms after the onset of ankle flexion in triceps surae muscles were related to the velocity of ankle stretch. The amplitude of balance-correcting responses at 100 ms corresponded more with stretch of the biarticular gastrocnemius when the knee was re-extended at 60 ms. Absent stretch reflexes at 50 ms in triceps surae with nulled ankle inputs caused a minor, 12-ms delay in the onset of balance-correcting responses in triceps surae muscles. Vestibular loss caused no change in the amplitude of balance-correcting responses, but a negligible decrease in onset latency in triceps surae even with nulled ankle inputs. Stretch responses in quadriceps at 80 ms increased with the velocity of knee flexion but were overall lower in amplitude in vestibular loss subjects. Balance-correcting responses in quadriceps had amplitudes which were related to the directions of initial trunk movements, were still present when knee inputs were negligible and were also altered after vestibular loss. Stretch and unloading responses in paraspinals at 80 ms were consistent with the direction of initial trunk flexion and extension. Subsequent balance-correcting responses in paraspinals were delayed 20 ms in onset and altered in amplitude by vestibular loss. The changes in the amplitudes of ankle (tibialis anterior), knee (quadriceps) and trunk (paraspinal) muscle responses with vestibular loss affected the amplitudes and timing of trunk angular velocities, requiring increased stabilizing tibialis anterior, paraspinal and trapezius responses post 240 ms as these subjects attempted to remain upright. The results suggest that trunk inputs provide an ideal candidate for triggering balance corrections as these would still be present when vestibular, ankle and knee inputs are absent. The disparity between the amplitudes of stretch reflex and automatic balance-correcting responses in triceps surae and the insignificant alteration in the timing of balance-correcting responses in these muscles with nulled ankle inputs indicates that ankle inputs do not trigger balance corrections. Furthermore, modulation of balance corrections normally performed by vestibular inputs in some but not all muscles is not achieved by switching to another sensory system on vestibular loss. We postulate that a confluence of trunk and upper-leg proprioceptive input establishes the basic timing of automatic, triggered balance corrections which is then preferentially weighted by vestibular modulation in muscles that prevent falling. (ABSTRACT TRUNCATED)