The 'Postural Rhythm' of the Ground Reaction Force during Upright Stance and Its Conversion to Body Sway-The Effect of Vision, Support Surface and Adaptation to Repeated Trials.

The ground reaction force (GRF) recorded by a platform when a person stands upright lies at the interface between the neural networks controlling stance and the body sway deduced from centre of pressure (CoP) displacement. It can be decomposed into vertical (VGRF) and horizontal (HGRF) vectors. Few studies have addressed the modulation of the GRFs by the sensory conditions and their relationship with body sway. We reconsidered the features of the GRFs oscillations in healthy young subjects (n = 24) standing for 90 s, with the aim of characterising the possible effects of vision, support surface and adaptation to repeated trials, and the correspondence between HGRF and CoP time-series. We compared the frequency spectra of these variables with eyes open or closed on solid support surface (EOS, ECS) and on foam (EOF, ECF). All stance trials were repeated in a sequence of eight. Conditions were randomised across different days. The oscillations of the VGRF, HGRF and CoP differed between each other, as per the dominant frequency of their spectra (around 4 Hz, 0.8 Hz and <0.4 Hz, respectively) featuring a low-pass filter effect from VGRF to HGRF to CoP. GRF frequencies hardly changed as a function of the experimental conditions, including adaptation. CoP frequencies diminished to <0.2 Hz when vision was available on hard support surface. Amplitudes of both GRFs and CoP oscillations decreased in the order ECF > EOF > ECS ≈ EOS. Adaptation had no effect except in ECF condition. Specific rhythms of the GRFs do not transfer to the CoP frequency, whereas the magnitude of the forces acting on the ground ultimately determines body sway. The discrepancies in the time-series of the HGRF and CoP oscillations confirm that the body's oscillation mode cannot be dictated by the inverted pendulum model in any experimental conditions. The findings emphasise the robustness of the VGRF "postural rhythm" and its correspondence with the cortical theta rhythm, shed new insight on current principles of balance control and on understanding of upright stance in healthy and elderly people as well as on injury prevention and rehabilitation.

Vertical ground reaction force oscillation during standing on hard and compliant surfaces: The "postural rhythm".

When a person stands upright quietly, the position of the Centre of Mass (CoM), the vertical force acting on the ground and the geometrical configuration of body segments is accurately controlled around to the direction of gravity by multiple feedback mechanisms and by integrative brain centres that coordinate multi-joint movements. This is not always easy and the postural muscles continuously produce appropriate torques, recorded as ground reaction force by a force platform. We studied 23 young adults during a 90 s period, standing at ease on a hard (Solid) and on a compliant support (Foam) with eyes open (EO) and with eyes closed (EC), focusing on the vertical component of the ground reaction force (VGRF). Analysis of VGRF time series gave the amplitude of their rhythmic oscillations (the root mean square, RMS) and of their frequency spectrum. Sway Area and Path Length of the Centre of Pressure (CoP) were also calculated. VGRF RMS (as well as CoP sway measures) increased in the order EO Solid ≈ EC Solid < EO Foam < EC Foam. The VGRF frequency spectra featured prevailing frequencies around 4-5 Hz under all tested conditions, slightly higher on Solid than Foam support. Around that value, the VGRF frequencies varied in a larger range on hard than on compliant support. Sway Area and Path Length were inversely related to the prevailing VGRF frequency. Vision compared to no-vision decreased Sway Area and Path Length and VGRF RMS on Foam support. However, no significant effect of vision was found on VGRF mean frequency for either base of support condition. A description of the VGRF, at the interface between balance control mechanisms and sway of the CoP, can contribute information on how upright balance is maintained. Analysis of the frequency pattern of VGRF oscillations and its role in the maintenance of upright stance should complement the traditional measures of CoP excursions in the horizontal plane.

Incongruity of Geometric and Spectral Markers in the Assessment of Body Sway.

Different measurements of body oscillations in the time or frequency domain are being employed as markers of gait and balance abnormalities. This study investigates basic relationships within and between geometric and spectral measures in a population of young adult subjects. Twenty healthy subjects stood with parallel feet on a force platform with and without a foam pad. Adaptation effects to prolonged stance were assessed by comparing the first and last of a series of eight successive trials. Centre of Foot Pressure (CoP) excursions were recorded with Eyes Closed (EC) and Open (EO) for 90s. Geometric measures (Sway Area, Path Length), standard deviation (SD) of the excursions, and spectral measure (mean power Spectrum Level and Median Frequency), along the medio-lateral (ML) and antero-posterior (AP) direction were computed. Sway Area was more strongly associated than Path Length with CoP SD and, consequently, with mean Spectrum Level for both ML and AP, and both visual and surface conditions. The squared-SD directly specified the mean power Spectrum Level of CoP excursions (ML and AP) in all conditions. Median Frequency was hardly related to Spectrum Level. Adaptation had a confounding effect, whereby equal values of Sway Area, Path Length, and Spectrum Level corresponded to different Median Frequency values. Mean Spectrum Level and SDs of the time series of CoP ML and AP excursions convey the same meaning and bear an acceptable correspondence with Sway Area values. Shifts in Median Frequency values represent important indications of neuromuscular control of stance and of the effects of vision, support conditions, and adaptation. The Romberg Quotient EC/EO for a given variable is contingent on the compliance of the base of support and adaptation, and different between Sway Area and Path Length, but similar between Sway Area and Spectrum Level (AP and ML). These measures must be taken with caution in clinical studies, and considered together in order to get a reliable indication of overall body sway, of modifications by sensory and standing condition, and of changes with ageing, medical conditions and rehabilitation treatment. However, distinct measures shed light on the discrete mechanisms and complex processes underpinning the maintenance of stance.

Balance Adaptation While Standing on a Compliant Base Depends on the Current Sensory Condition in Healthy Young Adults.

Background: Several investigations have addressed the process of balance adaptation to external perturbations. The adaptation during unperturbed stance has received little attention. Further, whether the current sensory conditions affect the adaptation rate has not been established. We have addressed the role of vision and haptic feedback on adaptation while standing on foam. Methods: In 22 young subjects, the analysis of geometric (path length and sway area) and spectral variables (median frequency and mean level of both total spectrum and selected frequency windows) of the oscillation of the centre of feet pressure (CoP) identified the effects of vision, light-touch (LT) or both in the anteroposterior (AP) and mediolateral (ML) direction over 8 consecutive 90 s standing trials. Results: Adaptation was obvious without vision (eyes closed; EC) and tenuous with vision (eyes open; EO). With trial repetition, path length and median frequency diminished with EC (p < 0.001) while sway area and mean level of the spectrum increased (p < 0.001). The low- and high-frequency range of the spectrum increased and decreased in AP and ML directions, respectively. Touch compared to no-touch enhanced the rate of increase of the low-frequency power (p < 0.05). Spectral differences in distinct sensory conditions persisted after adaptation. Conclusion: Balance adaptation occurs during standing on foam. Adaptation leads to a progressive increase in the amplitude of the lowest frequencies of the spectrum and a concurrent decrease in the high-frequency range. Within this common behaviour, touch adds to its stabilising action a modest effect on the adaptation rate. Stabilisation is improved by favouring slow oscillations at the expense of sway minimisation. These findings are preliminary to investigations of balance problems in persons with sensory deficits, ageing, and peripheral or central nervous lesion.

Specific Posture-Stabilising Effects of Vision and Touch Are Revealed by Distinct Changes of Body Oscillation Frequencies.

We addressed postural instability during stance with eyes closed (EC) on a compliant surface in healthy young people. Spectral analysis of the centre of foot pressure oscillations was used to identify the effects of haptic information (light-touch, EC-LT), or vision (eyes open, EO), or both (EO-LT). Spectral median frequency was strongly reduced by EO and EO-LT, while spectral amplitude was reduced by all "stabilising" sensory conditions. Reduction in spectrum level by EO mainly appeared in the high-frequency range. Reduction by LT was much larger than that induced by the vision in the low-frequency range, less so in the high-frequency range. Touch and vision together produced a fall in spectral amplitude across all windows, more so in anteroposterior (AP) direction. Lowermost frequencies contributed poorly to geometric measures (sway path and area) for all sensory conditions. The same subjects participated in control experiments on a solid base of support. Median frequency and amplitude of the spectrum and geometric measures were largely smaller when standing on solid than on foam base but poorly affected by the sensory conditions. Frequency analysis but not geometric measures allowed to disclose unique tuning of the postural control mode by haptic and visual information. During standing on foam, the vision did not reduce low-frequency oscillations, while touch diminished the entire spectrum, except for the medium-high frequencies, as if sway reduction by touch would rely on rapid balance corrections. The combination of frequency analysis with sensory conditions is a promising approach to explore altered postural mechanisms and prospective interventions in subjects with central or peripheral nervous system disorders.

Basic Spatiotemporal Gait Variables of Young and Older Healthy Volunteers Walking Along a Novel Figure-of-8 Path.

Background: Locomotion along curved trajectories requires fine coordination among body segments. Elderly people may adopt a cautious attitude when steering. A simple, expeditious, patient-friendly walking protocol can be a tool to help clinicians. We evaluated the feasibility of a procedure based upon a newly designed Figure-of-eight (nFo8) path and an easy measurement operation. Methods: Sixty healthy volunteers, aged from 20 to 86 years, walked three times at self-selected speed along a 20 m linear (LIN) and the 20 m nFo8 path. Number of steps, mean speed and walk ratio (step length/cadence) were collected. Data were analysed for the entire cohort and for the groups aged 20-45, 46-65, and >65 years. Results: There was no difference in mean LIN walking speed between the two younger groups but the oldest was slower. During nFo8, all groups were slower (about 16%) than during LIN. Cadence was not different across groups but lower during nFo8 in each group. Step length was about 8% shorter in the two younger groups and 14% shorter in the oldest during nFo8 compared to LIN. Walk ratio was the smallest in the oldest group for both LIN and nFo8. Conclusions: A complex nFo8 walking path, with fast and easy measurement of a simple set of variables, detects significant differences with moderate and large effects in gait variables in people >65 years. This challenging trajectory is more revealing than LIN. Further studies are needed to develop a quick clinical tool for assessment of gait conditions or outcome of rehabilitative treatments.

Adaptation of balancing behaviour during continuous perturbations of stance. Supra-postural visual tasks and platform translation frequency modulate adaptation rate.

When humans are administered continuous and predictable perturbations of stance, an adaptation period precedes the steady state of balancing behaviour. Little information is available on the modulation of adaptation by vision and perturbation frequency. Moreover, performance of supra-postural tasks may modulate adaptation in as yet unidentified ways. Our purpose was to identify differences in adaptation associated to distinct visual tasks and perturbation frequencies. Twenty non-disabled adult volunteers stood on a platform translating 10 cm in antero-posterior (AP) direction at low (LF, 0.18 Hz) and high frequency (HF, 0.56 Hz) with eyes open (EO) and closed (EC). Additional conditions were reading a text fixed to platform (EO-TP) and reading a text stationary on ground (EO-TG). Peak-to-peak (PP) displacement amplitude and AP position of head and pelvis markers were computed for each of 27 continuous perturbation cycles. The time constant and extent of head and pelvis adaptation and the cross-correlation coefficients between head and pelvis were compared across visual conditions and frequencies. Head and pelvis mean positions in space varied little across conditions and perturbation cycles but the mean head PP displacements changed over time. On average, at LF, the PP displacement of the head and pelvis increased progressively. Adaptation was rapid or ineffective with EO, but slower with EO-TG, EO-TP, EC. At HF, the head PP displacement amplitude decreased progressively with fast adaptation rates, while the pelvis adaptation was not apparent. The results show that visual tasks can modulate the adaptation rate, highlight the effect of the perturbation frequency on adaptation and provide evidence of priority assigned to pelvis stabilization over visual tasks at HF. The effects of perturbation frequency and optic flow and their interaction with other sensory inputs and cognitive tasks on the adaptation strategies should be investigated in impaired individuals and considered in the design of rehabilitation protocols.

Vision Does Not Necessarily Stabilize the Head in Space During Continuous Postural Perturbations.

Vision favors head stabilization in space during perturbations of standing balance. This is particularly obvious under conditions of continuous predictable perturbations as during sinusoidal antero-posterior (A-P) translations of the supporting platform. We tested here the hypothesis that under this condition the head can instead undergo large A-P oscillations, when a precision visual task is concurrently performed. We compared the head oscillations across four conditions while standing on a continuously translating platform. Eyes open (EO, no visual task), EO while reading a text fixed to the moving platform (EO-TP), EO while reading a text fixed to earth-ground (EO-TG), eyes-closed (EC). The platform translated at 0.2 and 0.6 Hz. Participants were young adult subjects, who received no particular instruction except reading the text aloud when required. Markers fixed on head, platform and text-sheet were captured by an optoelectronic device. We found that head oscillations were larger with EC than under all EO conditions. The oscillations were the least with EO and EO-TG, and intermediate with EO-TP. This was true under both low and high translation frequency, in spite of broadly smaller head oscillations at high frequency, common to all visual conditions. The distance between the head and the text was quite constant with EO-TP but fluctuated with EO-TG. The basic whole-body coordination features were moderately similar under all conditions, as assessed by the head-platform correlation coefficients and time lags. It appears that vision does not produce head stabilization in space when a concurrent visual task requiring focusing on a reading-text moving with the platform is performed. Contrary to traditional views centered on the stabilizing effect of vision under both static and dynamic conditions, the results show that head stabilization, normally ensuring a reference for inertial guidance for body balance, can be revoked by the CNS to allow performance of a non-postural task. This novel paradigm can shift long-standing views on the effect of vision on equilibrium control and be considered a potential exercise treatment for enhancing the multisensory integration process in people with balance problems.

Podokinetic After-Rotation Is Transiently Enhanced or Reversed by Unilateral Axial Muscle Proprioceptive Stimulation.

Unilateral axial muscle vibration, eliciting a proprioceptive volley, is known to incite steering behavior. Whole-body rotation while stepping in place also occurs as an after-effect of stepping on a circular treadmill (podokinetic after-rotation, PKAR). Here, we tested the hypothesis that PKAR is modulated by axial muscle vibration. If both phenomena operate through a common pathway, enhancement or cancellation of body rotation would occur depending on the stimulated side when vibration is administered concurrently with PKAR. Seventeen subjects participated in the study. In one session, subjects stepped in place eyes open on the center of a platform that rotated counterclockwise 60°/s for 10 min. When the platform stopped, subjects continued stepping in place blindfolded. In other session, a vibratory stimulus (100 Hz, 2 min) was administered to right or left paravertebral muscles at lumbar level at two intervals during the PKAR. We computed angular body velocity and foot step angles from markers fixed to shoulders and feet. During PKAR, all subjects rotated clockwise. Decreased angular velocity was induced by right vibration. Conversely, when vibration was administered to the left, clockwise rotation velocity increased. The combined effect on body rotation depended on the time at which vibration was administered during PKAR. Under all conditions, foot step angle was coherent with shoulder angular velocity. PKAR results from continuous asymmetric input from the muscles producing leg rotation, while axial muscle vibration elicits a proprioceptive asymmetric input. Both conditioning procedures appear to produce their effects through a common mechanism. We suggest that both stimulations would affect our straight ahead by combining their effects in an algebraic mode.

Balance in Blind Subjects: Cane and Fingertip Touch Induce Similar Extent and Promptness of Stance Stabilization.

Subjects with low vision often use a cane when standing and walking autonomously in everyday life. One aim of this study was to assess differences in the body stabilizing effect produced by the contact of the cane with the ground or by the fingertip touch of a firm surface. Another aim was to estimate the promptness of balance stabilization (or destabilization) on adding (or withdrawing) the haptic input from cane or fingertip. Twelve blind subjects and two subjects with severe visual impairment participated in two experimental protocols while maintaining the tandem Romberg posture on a force platform. In one protocol, subjects lowered the cane to a second platform on the ground and lifted it in sequence at their own pace. In the other protocol, they touched an instrumented pad with the index finger and withdrew the finger from the pad in sequence. In both protocols, subjects were asked to exert a force not granting mechanical stabilization. Under steady-state condition, the finger touch or the contact of the cane with the ground significantly reduced (to ∼78% and ∼86%, respectively) the amplitude of medio-lateral oscillation of the centre of foot pressure (CoP). Oscillation then increased when haptic information was removed. The delay to the change in body oscillation after the haptic shift was longer for addition than withdrawal of the haptic information (∼1.4 s and ∼0.7 s, respectively; p < 0.001), but was not different between the two haptic conditions (finger and cane). Similar stabilizing effects of input from cane on the ground and from fingertip touch, and similar latencies to integrate haptic cue from both sources, suggest that the process of integration of the input for balance control is initiated by the haptic stimulus at the interface cane-hand. Use of a tool is as helpful as the fingertip input, and does not produce different stabilization. Further, the latencies to haptic cue integration (from fingertip or cane) are similar to those previously found in a group of sighted subjects, suggesting that integration delays for automatic balance stabilization are not modified by visual impairment. Haptic input from a tool is easily exploited by the neural circuits subserving automatic balance stabilization in blind people, and its use should be enforced by sensory-enhancing devices and appropriate training.

Body Sway Increases After Functional Inactivation of the Cerebellar Vermis by cTBS.

Balance stability correlates with cerebellar vermis volume. Furthermore, the cerebellum is involved in precise timing of motor processes by fine-tuning the sensorimotor integration. We tested the hypothesis that any cerebellar action in stance control and in timing of visuomotor integration for balance is impaired by continuous theta-burst stimulation (cTBS) of the vermis. Ten subjects stood quietly and underwent six sequences of 10-min acquisition of center of foot pressure (CoP) data after cTBS, sham stimulation, and no stimulation. Visual shifts from eyes closed (EC) to eyes open (EO) and vice versa were presented via electronic goggles. Mean anteroposterior and mediolateral CoP position and oscillation, and the time delay at which body sway changed after visual shift were calculated. CoP position under both EC and EO condition was not modified after cTBS. Sway path length was greater with EC than EO and increased in both visual conditions after cTBS. CoP oscillation was also larger with EC and increased under both visual conditions after cTBS. The delay at which body oscillation changed after visual shift was longer after EC to EO than EO to EC, but unaffected by cTBS. The time constant of decrease or increase of oscillation was longer in EC to EO shifts, but unaffected by cTBS. Functional inactivation of the cerebellar vermis is associated with increased sway. Despite this, cTBS does not detectably modify onset and time course of the sensorimotor integration process of adaptation to visual shifts. Cerebellar vermis normally controls oscillation, but not timing of adaptation to abrupt changes in stabilizing information.

Haptic Cues for Balance: Use of a Cane Provides Immediate Body Stabilization.

Haptic cues are important for balance. Knowledge of the temporal features of their effect may be crucial for the design of neural prostheses. Touching a stable surface with a fingertip reduces body sway in standing subjects eyes closed (EC), and removal of haptic cue reinstates a large sway pattern. Changes in sway occur rapidly on changing haptic conditions. Here, we describe the effects and time-course of stabilization produced by a haptic cue derived from a walking cane. We intended to confirm that cane use reduces body sway, to evaluate the effect of vision on stabilization by a cane, and to estimate the delay of the changes in body sway after addition and withdrawal of haptic input. Seventeen healthy young subjects stood in tandem position on a force platform, with eyes closed or open (EO). They gently lowered the cane onto and lifted it from a second force platform. Sixty trials per direction of haptic shift (Touch → NoTouch, T-NT; NoTouch → Touch, NT-T) and visual condition (EC-EO) were acquired. Traces of Center of foot Pressure (CoP) and the force exerted by cane were filtered, rectified, and averaged. The position in space of a reflective marker positioned on the cane tip was also acquired by an optoelectronic device. Cross-correlation (CC) analysis was performed between traces of cane tip and CoP displacement. Latencies of changes in CoP oscillation in the frontal plane EC following the T-NT and NT-T haptic shift were statistically estimated. The CoP oscillations were larger in EC than EO under both T and NT (p < 0.001) and larger during NT than T conditions (p < 0.001). Haptic-induced effect under EC (Romberg quotient NT/T ~ 1.2) was less effective than that of vision under NT condition (EC/EO ~ 1.5) (p < 0.001). With EO cane had little effect. Cane displacement lagged CoP displacement under both EC and EO. Latencies to changes in CoP oscillations were longer after addition (NT-T, about 1.6 s) than withdrawal (T-NT, about 0.9 s) of haptic input (p < 0.001). These latencies were similar to those occurring on fingertip touch, as previously shown. Overall, data speak in favor of substantial equivalence of the haptic information derived from both "direct" fingertip contact and "indirect" contact with the floor mediated by the cane. Cane, finger and visual inputs would be similarly integrated in the same neural centers for balance control. Haptic input from a walking aid and its processing time should be considered when designing prostheses for locomotion.

Calibration of the Leg Muscle Responses Elicited by Predictable Perturbations of Stance and the Effect of Vision.

Motor adaptation due to task practice implies a gradual shift from deliberate control of behavior to automatic processing, which is less resource- and effort-demanding. This is true both for deliberate aiming movements and for more stereotyped movements such as locomotion and equilibrium maintenance. Balance control under persisting critical conditions would require large conscious and motor effort in the absence of gradual modification of the behavior. We defined time-course of kinematic and muscle features of the process of adaptation to repeated, predictable perturbations of balance eliciting both reflex and anticipatory responses. Fifty-nine sinusoidal (10 cm, 0.6 Hz) platform displacement cycles were administered to 10 subjects eyes-closed (EC) and eyes-open (EO). Head and Center of Mass (CoM) position, ankle angle and Tibialis Anterior (TA) and Soleus (Sol) EMG were assessed. EMG bursts were classified as reflex or anticipatory based on the relationship between burst amplitude and ankle angular velocity. Muscle activity decreased over time, to a much larger extent for TA than Sol. The attenuation was larger for the reflex than the anticipatory responses. Regardless of muscle activity attenuation, latency of muscle bursts and peak-to-peak CoM displacement did not change across perturbation cycles. Vision more than doubled speed and the amount of EMG adaptation particularly for TA activity, rapidly enhanced body segment coordination, and crucially reduced head displacement. The findings give new insight on the mode of amplitude- and time-modulation of motor output during adaptation in a balancing task, advocate a protocol for assessing flexibility of balance strategies, and provide a reference for addressing balance problems in patients with movement disorders.

Stepping in Place While Voluntarily Turning Around Produces a Long-Lasting Posteffect Consisting in Inadvertent Turning While Stepping Eyes Closed.

Training subjects to step in place on a rotating platform while maintaining a fixed body orientation in space produces a posteffect consisting in inadvertent turning around while stepping in place eyes closed (podokinetic after-rotation, PKAR). We tested the hypothesis that voluntary turning around while stepping in place also produces a posteffect similar to PKAR. Sixteen subjects performed 12 min of voluntary turning while stepping around their vertical axis eyes closed and 12 min of stepping in place eyes open on the center of a platform rotating at 60°/s (pretests). Then, subjects continued stepping in place eyes closed for at least 10 min (posteffect). We recorded the positions of markers fixed to head, shoulder, and feet. The posteffect of voluntary turning shared all features of PKAR. Time decay of angular velocity, stepping cadence, head acceleration, and ratio of angular velocity after to angular velocity before were similar between both protocols. Both postrotations took place inadvertently. The posteffects are possibly dependent on the repeated voluntary contraction of leg and foot intrarotating pelvic muscles that rotate the trunk over the stance foot, a synergy common to both protocols. We propose that stepping in place and voluntary turning can be a scheme ancillary to the rotating platform for training body segment coordination in patients with impairment of turning synergies of various origin.

Temporal features of postural adaptation strategy to prolonged and repeatable balance perturbation.

Aim of this study was to get insight into the features of the postural adaptation process, occurring during a continuous 3-min and 0.6Hz horizontal sinusoidal oscillation of the body support base. We hypothesized an ongoing temporal organization of the balancing strategy that gradually becomes fine-tuned and more coordinated with the platform movement. The trial was divided into oscillation cycles and for each cycle: leg muscles activity and temporal relationship between Centre of Mass and Centre of Pressure A-P position were analyzed. The results of each cycle were grouped in time-windows of 10 successive cycles (time windows of 16.6s). Muscle activity was initially prominent and diminished progressively. The major burst of Tibialis Anterior (TA) muscle always occurred at the same time instant of the platform oscillation cycle, in advance with respect to the platform posterior turning point. This burst produced a body forward rotation that was delayed throughout the task. During prolonged and repeatable balance perturbation, an ongoing postural adaptation process occurs. When the effects of the perturbation become predictable, the CNS scales the level of muscle activity to counteracting the destabilizing effects of the perturbations. Furthermore, the CNS tunes the kinematics and the kinetic responses optimally by slightly delaying the onset of the body forward rotation, maintaining unchanged the time-pattern of postural muscle activation.

Processing time of addition or withdrawal of single or combined balance-stabilizing haptic and visual information.

We investigated the integration time of haptic and visual input and their interaction during stance stabilization. Eleven subjects performed four tandem-stance conditions (60 trials each). Vision, touch, and both vision and touch were added and withdrawn. Furthermore, vision was replaced with touch and vice versa. Body sway, tibialis anterior, and peroneus longus activity were measured. Following addition or withdrawal of vision or touch, an integration time period elapsed before the earliest changes in sway were observed. Thereafter, sway varied exponentially to a new steady-state while reweighting occurred. Latencies of sway changes on sensory addition ranged from 0.6 to 1.5 s across subjects, consistently longer for touch than vision, and were regularly preceded by changes in muscle activity. Addition of vision and touch simultaneously shortened the latencies with respect to vision or touch separately, suggesting cooperation between sensory modalities. Latencies following withdrawal of vision or touch or both simultaneously were shorter than following addition. When vision was replaced with touch or vice versa, adding one modality did not interfere with the effect of withdrawal of the other, suggesting that integration of withdrawal and addition were performed in parallel. The time course of the reweighting process to reach the new steady-state was also shorter on withdrawal than addition. The effects of different sensory inputs on posture stabilization illustrate the operation of a time-consuming, possibly supraspinal process that integrates and fuses modalities for accurate balance control. This study also shows the facilitatory interaction of visual and haptic inputs in integration and reweighting of stance-stabilizing inputs.

Rapid processing of haptic cues for postural control in blind subjects.

OBJECTIVES: Vision and touch rapidly lead to postural stabilization in sighted subjects. Is touch-induced stabilization more rapid in blind than in sighted subjects, owing to cross-modal reorganization of function in the blind? METHODS: We estimated the time-period elapsing from onset of availability of haptic support to onset of lateral stabilization in a group of early- and late-onset blinds. Eleven blind (age 39.4 years±11.7SD) and eleven sighted subjects (age 30.0 years±10.0SD), standing eyes closed with feet in tandem position, touched a pad with their index finger and withdrew the finger from the pad in sequence. EMG of postural muscles and displacement of centre of foot pressure were recorded. The task was repeated fifty times, to allow statistical evaluation of the latency of EMG and sway changes following the haptic shift. RESULTS: Steady-state sway (with or without contact with pad, no haptic shift) did not differ between blind and sighted. On adding the haptic stimulus, EMG and sway diminished in both groups, but at an earlier latency (by about 0.5 s) in the blinds (p <0.01). Latencies were still shorter in the early-than late-blinds. When the haptic stimulus was withdrawn, both groups increased EMG and sway at equally short delays. CONCLUSIONS: Blinds are rapid in implementing adaptive postural modifications when granted an external haptic reference. Fast processing of the stabilizing haptic spatial-orientation cues may be favoured by cortical plasticity in blinds. SIGNIFICANCE: These findings add new information to the field of sensory-guided dynamic control of equilibrium in man.

Leg muscle activity during tandem stance and the control of body balance in the frontal plane.

OBJECTIVES: We investigated the pattern of activity of the tibialis anterior (TA), soleus (SOL) and peroneus longus (PER) muscles of both legs during tandem stance, in order to highlight their respective role in maintaining balance. METHODS: Twelve young healthy subjects were asked to stand with feet in line for successive 15s-epochs, on a dynamometric platform with (EO) and without (EC) vision. EMG was recorded from the six muscles simultaneously. Collected signals were displacement of the centre of feet pressure (CoP) and EMG. Variables calculated for each recorded epoch were mean level, variability and distribution between legs of EMG, and cross-correlation between EMG and CoP traces and between EMG of homonymous muscles. RESULTS: CoP motion was larger along the medio-lateral (M-L) than antero-posterior (A-P) axis, and larger with EC than EO particularly in the M-L axis. Muscle activity was larger in the rear than in the front leg, as expected, except for PER. Activity increased with the increase in M-L CoP oscillations, except for SOL, which was tonically active, both legs, regardless of the amplitude of the oscillations. Manipulating vision had no effect on the variability of the EMG for equal mean levels of activity, for any muscle. Cross-correlation between EMG of rear leg muscles and M-L CoP sway gave higher coefficients for TA and PER than SOL, and appropriate time-delays between TA or PER and CoP motion, indicating a role of these muscles in the control of M-L sway. Except for the tonically active SOL, the homonymous muscles of the two legs were active out-of-phase, indicating a mutual push-pull action of the pairs. This was confirmed by the reciprocal activation of TA and PER of the same leg. CONCLUSIONS: Overall, in spite of a large inter-trial and inter-subject variability, the neural command to the leg muscles during tandem stance implies a task-sharing rule, whereby SOL keeps the body upright while the reciprocal PER and TA activities produce the alternate impulses necessary for body stabilization in the frontal plane. SIGNIFICANCE: Knowledge of the normal mode of control of balance in frontal plane can foster new investigation in both posture and gait control, in addition to offering tools for understanding balance problems of elderly persons and patients at risk of fall.

Adaptation to continuous perturbation of balance: progressive reduction of postural muscle activity with invariant or increasing oscillations of the center of mass depending on perturbation frequency and vision conditions.

We investigated the adaptation of balancing behavior during a continuous, predictable perturbation of stance consisting of 3-min backward and forward horizontal sinusoidal oscillations of the support base. Two visual conditions (eyes-open, EO; eyes-closed, EC) and two oscillation frequencies (LF, 0.2 Hz; HF, 0.6 Hz) were used. Center of Mass (CoM) and Center of Pressure (CoP) oscillations and EMG of Soleus (Sol) and Tibialis Anterior (TA) were recorded. The time course of each variable was estimated through an exponential model. An adaptation index allowed comparison of the degree of adaptation of different variables. Muscle activity pattern was initially prominent under the more challenging conditions (HF, EC and EO; LF, EC) and diminished progressively to reach a steady state. At HF, the behavior of CoM and CoP was almost invariant. The time-constant of EMG adaptation was shorter for TA than for Sol. With EC, the adaptation index showed a larger decay in the TA than Sol activity at the end of the balancing trial, pointing to a different role of the two muscles in the adaptation process. At LF, CoM and CoP oscillations increased during the balancing trial to match the platform translations. This occurred regardless of the different EMG patterns under EO and EC. Contrary to CoM and CoP, the adaptation of the muscle activities had a similar time-course at both HF and LF, in spite of the two frequencies implying a different number of oscillation cycles. During adaptation, under critical balancing conditions (HF), postural muscle activity is tuned to that sufficient for keeping CoM within narrow limits. On the contrary, at LF, when vision permits, a similar decreasing pattern of muscle activity parallels a progressive increase in CoM oscillation amplitude, and the adaptive balancing behavior shifts from the initially reactive behavior to one of passive riding the platform. Adaptive balance control would rely on on-line computation of risk of falling and sensory inflow, while minimizing balance challenge and muscle effort. The results from this study contribute to the understanding of plasticity of the balance control mechanisms under posture-challenging conditions.

Sensori-motor integration during stance: time adaptation of control mechanisms on adding or removing vision.

Sudden addition or removal of visual information can be particularly critical to balance control. The promptness of adaptation of stance control mechanisms is quantified by the latency at which body oscillation and postural muscle activity vary after a shift in visual condition. In the present study, volunteers stood on a force platform with feet parallel or in tandem. Shifts in visual condition were produced by electronic spectacles. Ground reaction force (center of foot pressure, CoP) and EMG of leg postural muscles were acquired, and latency of CoP and EMG changes estimated by t-tests on the averaged traces. Time-to-reach steady-state was estimated by means of an exponential model. On allowing or occluding vision, decrements and increments in CoP position and oscillation occurred within about 2s. These were preceded by changes in muscle activity, regardless of visual-shift direction, foot position or front or rear leg in tandem. These time intervals were longer than simple reaction-time responses. The time course of recovery to steady-state was about 3s, shorter for oscillation than position. The capacity of modifying balance control at very short intervals both during quiet standing and under more critical balance conditions speaks in favor of a necessary coupling between vision, postural reference, and postural muscle activity, and of the swiftness of this sensory reweighing process.