Postural preparation to making a step: is there a 'motor program' for postural preparation?

We tested the hypothesis that a sequence of mechanical events occurs preceding a step that scales in time and magnitude as a whole in a task-specific manner, and is a reflection of a "motor program." Young subjects made a step under three speed instructions and four tasks: stepping straight ahead, down a stair, up a stair, and over an obstacle. Larger center-of-pressure (COP) and force adjustments in the anterior-posterior direction and smaller COP and force adjustments in the mediolateral direction were seen during stepping forward and down a stair, as compared with the tasks of stepping up a stair and over an obstacle. These differences were accentuated during stepping under the simple reaction time instruction. These results speak against the hypothesis of a single motor program that would underlie postural preparation to stepping. They are more compatible with the reference configuration hypothesis of whole-body actions.     

Changes in postural sway and its fractions in conditions of postural instability

We investigated changes in postural sway and its fractions associated with manipulations of the dimensions of the support area. Nine healthy adults stood as quietly as possible, with their eyes open, on a force plate as well as on 5 boards with reduced support area. The center of pressure (COP) trajectory was computed and decomposed into rambling (Rm) and trembling (Tr) trajectories. Sway components were quantified using RMS (root mean square) value, average velocity, and sway area. During standing on the force plate, the RMS was larger for the anterior-posterior (AP) sway components than for the mediolateral (ML) components. During standing on boards with reduced support area, sway increased in both directions. The increase was more pronounced when standing on boards with a smaller support area. Changes in the larger dimension of the support area also affected sway, but not as much as changes in the smaller dimension. ML instability had larger effects on indices of sway compared to AP instability. The average velocity of Rm was larger while the average velocity of Tr was smaller in the AP direction vs. the ML direction. The findings can be interpreted within the hypothesis of an active search function of postural sway. During standing on boards with reduced support area, increased sway may by itself lead to loss of balance. The findings also corroborate the hypothesis of Duarte and Zatsiorsky that Rm and Tr reveal different postural control mechanisms.     

Learning Upright Standing on a Multiaxial Balance Board

Upright stance on a balance board is a skill requiring complex rearrangement of the postural control. Despite the large use of these boards in training the standing posture, a comprehensive analysis of the learning process underlying the control of these devices is lacking. In this paper learning to maintain a stable stance on a multiaxial oscillating board was studied by analyzing performance changes over short and long periods. Healthy participants were asked to keep the board orientation as horizontal as possible for 20 sec, performing two sessions of 8 trials separated by 15-min pause. Memory consolidation was tested one week later. Amplitude and variability of the oscillations around horizontal plane and area and sway path of the board displacement decreased rapidly over the first session. The performance was stable during the second session, and retained after 1 week. A similar behavior was observed in the anterior-posterior and medial-lateral directions for amplitude and variability parameters, with less stable balance in the anterior-posterior direction. Approximate entropy and mean power frequency, assessing temporal dynamics and frequency content of oscillations, changed only in the anterior-posterior direction during the retention test. Overall, the ability to stand on a balance board is rapidly acquired, and retained for long time. The asymmetric stability between anterior-posterior and medial-lateral directions replicates a structure observed in other standing stances, suggesting a possible transfer from previous postural experiences. Conversely, changes in the temporal dynamics and the frequency content could be associated with new postural strategies developed later during memory consolidation.     

Center of pressure and center of mass behavior during gait initiation on inclined surfaces: A statistical parametric mapping analysis

This study analyzed gait initiation (GI) on inclined surfaces with 68 young adult subjects of both sexes. Ground reaction forces and moments were collected using two AMTI force platforms, of which one was in a horizontal position and the other was inclined by 8% in relation to the horizontal plane. Departing from a standing position, each participant executed three trials in the following conditions: horizontal position (HOR), inclined position at ankle dorsi-flexion (UP), and inclined position at ankle plantar-flexion (DOWN). Statistical parametric mapping analysis was performed over the entire center of pressure (COP) and center of mass (COM) time series. COP excursion did not show significant differences in the medial-lateral (ML) direction in both inclined conditions, but it was greater in the anterior-posterior (AP) direction for both inclined conditions. COP velocities are smaller in discrete portions of GI for the UP and DOWN conditions. COM displacement was greater in the ML direction during anticipatory postural adjustments (APA) in the UP condition, and COM moves faster in the ML direction during APA in the UP condition but slower at the end of GI for both the UP and the DOWN conditions. The COP-COM vector showed a greater angle in the DOWN condition. We observed changes for COP and COM in GI in both the UP and the DOWN conditions, with the latter showing changes for a great extent of the task. Both the UP and the DOWN conditions showed increased COM displacement and velocity. The predominant characteristic during GI on inclined surfaces, including APA, appears to be the displacement of the COM.     

Early postural adjustments in preparation to whole-body voluntary sway

We studied postural adjustments associated with a quick voluntary postural sway under two conditions, self-paced and simple reaction-time. Standing subjects were required to produce quick discrete shifts of the center of pressure (COP) forward. About 400-500 ms prior to the instructed COP shift, there were deviations of the COP in the opposite direction (backwards) accompanied by changes in the activation levels of several postural muscles. Under the reaction-time conditions, the timing of those early postural adjustments did not change (repeated measures MANOVA: p > 0.05) while its magnitude increased significantly (confirmed by repeated measures MANOVA: p < 0.05). These observations are opposite to those reported for anticipatory postural adjustments under simple reaction time conditions (a significant change in the timing without major changes in the magnitude). We conclude that there are two types of feed-forward postural adjustments. Early postural adjustments prepare the body for the planned action and/or expected perturbation. Some of these preparatory actions may be mechanically necessary. Later, anticipatory postural adjustments generate net forces and moments of force acting against those associated with the expected perturbation. Both types of adjustments fit well the referent configuration hypothesis, which offers a unified view on movement-posture control.     

The effects of early stages of aging on postural sway: A multiple domain balance assessment using a force platform

Technical advancements in instrumentation and analytical methods have improved the ability of assessing balance control. This study investigated the effects of early stages of aging on postural sway using traditional and contemporary postural indices from different domains. Eleven healthy young adults and fourteen healthy non-faller older adults performed two postural tasks: (a) functional limits of stability and (b) unperturbed bipedal stance for 120 s. Postural indices from spatial, temporal, frequency, and structural domains were extracted from the body’s center of pressure (COP) signals and its Rambling and Trembling components. Results revealed a preservation of functional limits of upright stability in older adults accompanied by larger, faster, and shakier body sway in both anterior-posterior and medio-lateral directions; increased medio-lateral sway frequency; increased irregularity of body sway pattern in time in both directions; and increased area, variability, velocity, and jerkiness of both rambling and trembling components of the COP displacement in the anterior-posterior direction (p < 0.02). Such changes might be interpreted as compensatory adjustments to the age-related decline of sensory, neural, and motor functions. In conclusion, balance assessment using postural indices from different domains extracted from the COP displacement was able to capture subtle effects of the natural process of aging on the mechanisms of postural control. Our findings suggest the use of such indices as potential markers for postural instability and fall risk in older adults.     

Biomechanical reorganisation of stepping initiation during acute dorsiflexor fatigue

During voluntary step initiation (SI), propulsive forces are generated during anticipatory postural adjustments (APA) which displace the centre-of-gravity (CoG) in the desired direction. These propulsive forces are implemented by ankle synergy, bilateral soleus inhibition followed by activation of tibialis anterior (TA). The aim of this study was to investigate the effect of fatigue applied to ankle dorsiflexors on APA associated with SI and on related motor performance. Eight young healthy participants initiated stepping before and after a protocol designed to generate fatigue in ankle dorsiflexors. Fatigue was induced by series of high-level isometric contractions performed until exhaustion. Results showed that, with fatigue, the level of TA activation during APA, anticipatory postural dynamics (backward centre-of-pressure displacement and forward CoG velocity) and related motor performance (peak of CoG velocity) were attenuated, while APA duration and total SI duration increased. These changes were interpreted as reflecting a protective strategy aiming to preserve the integrity of the fatigued muscles, rather than an impairment associated with muscle weakness.     

Standing on wedges modifies side-specific postural control in the presence of lateral external perturbations

Standing on wedges changes the position in the ankle joints and affects postural stability in the medial-lateral direction. The objective of the study was to investigate the role of wedges and external lateral perturbations on anticipatory (APA) and compensatory postural adjustments (CPA). Ten healthy young participants were exposed to perturbations applied to the lateral part of their right shoulder when standing on a planar surface, on a medial or lateral wedges. Bilateral electromyographic activity of dorsal and ventral postural muscles and the center of pressure (COP) displacement were recorded and analyzed during the APA and CPA phases. When exposed to the lateral perturbation, reciprocal activation of shank muscles was seen on the side of the perturbation while co-contraction of shank muscles was seen on the contralateral side during the APA and CPA phases. Standing on a wedge was associated with decreased magnitudes of co-contraction and reciprocal activation of shank muscles. The COP displacements were smaller in the APA phase and larger in the CPA phase while standing on wedges compared to standing on the planar surface. The outcome of the study provides a basis for future investigations of incorporating wedges in balance re-training paradigms for the elderly or individuals with neurological impairment.     

Effects of repeated ankle plantar-flexions on H-reflex and body sway during standing

The study investigated relations between effects of repeated ankle plantar-flexion movements exercise on the soleus Hoffmann (H) reflex and on postural body sway when maintaining upright stance. Ten young volunteers performed five sets of ankle plantar-flexions of both lower limbs. Assessment of the feet centre-of-pressure (COP) displacement and H-reflex tests were carried out in quiet stance before, during and after the exercise. H-max and M-max responses were obtained in 8 subjects and reported as the peak-to-peak amplitudes of the right soleus muscle electromyographic waves. Mean dispersion of COP along the antero-posterior direction increased significantly during the exercise; whilst the overall H-reflex response indicated a reduction without a concomitant modification in the M-max response. H-reflex responses, however, varied between participants during the first sets of exercise, showing two main trends of modulation: either depression or early facilitation followed by reduction of the H-reflex amplitude. The extent of reflex modulation in standing position was correlated to the concentric work performed during the exercise (r = 0.85; p < 0.01), but not to the antero-posterior COP dispersion. These results suggest that during a repeated ankle plantar-flexions exercise, modulation of the H-reflex measured in upright stance differs across individuals and is not related to changes of postural sway.     

Footwear and Foam Surface Alter Gait Initiation of Typical Subjects

Gait initiation is the task commonly used to investigate the anticipatory postural adjustments necessary to begin a new gait cycle from the standing position. In this study, we analyzed whether and how foot-floor interface characteristics influence the gait initiation process. For this purpose, 25 undergraduate students were evaluated while performing a gait initiation task in three experimental conditions: barefoot on a hard surface (barefoot condition), barefoot on a soft surface (foam condition), and shod on a hard surface (shod condition). Two force plates were used to acquire ground reaction forces and moments for each foot separately. A statistical parametric mapping (SPM) analysis was performed in COP time series. We compared the anterior-posterior (AP) and medial-lateral (ML) resultant center of pressure (COP) paths and average velocities, the force peaks under the right and left foot, and the COP integral x force impulse for three different phases: the anticipatory postural adjustment (APA) phase (Phase 1), the swing-foot unloading phase (Phase 2), and the support-foot unloading phase (Phase 3). In Phase 1, significantly smaller ML COP paths and velocities were found for the shod condition compared to the barefoot and foam conditions. Significantly smaller ML COP paths were also found in Phase 2 for the shod condition compared to the barefoot and foam conditions. In Phase 3, increased AP COP velocities were found for the shod condition compared to the barefoot and foam conditions. SPM analysis revealed significant differences for vector COP time series in the shod condition compared to the barefoot and foam conditions. The foam condition limited the impulse-generating capacity of COP shift and produced smaller ML force peaks, resulting in limitations to body-weight transfer from the swing to the support foot. The results suggest that footwear and a soft surface affect COP and impose certain features of gait initiation, especially in the ML direction of Phase 1.     

Diabetic neuropathy and surface sway-referencing disrupt somatosensory information for postural stability in stance

In order to determine the type of somatosensory information for postural control that is most affected by neuropathy, we compared the relative effects of three methods of sway-referencing the surface in a group of subjects with profound loss of somatosensory function associated with sensory polyneuropathy from diabetes with age-matched control subjects. Sway-referencing disrupted somatosensory feedback for postural control by servo-controlling the dorsi- and plantar-flexion rotation of the support surface in proportion to anterior-posterior excursion of (1) ankle angle, (2) center of body mass (CoM) angle or (3) filtered center of pressure (CoP). Postural sway in subjects with somatosensory loss was significantly larger than normal on a firm surface but not on the sway-referenced surfaces, suggesting that sway-referencing disrupts somatosensory information for postural control already disrupted by neuropathy. Control subjects standing on any sway-referenced surface swayed significantly more than neuropathy subjects who stood on a firm surface, suggesting that sway-referencing disrupts more somatosensory information than disrupted by severe neuropathy. CoP sway-referencing was less sensitive than ankle or CoM sway-referencing for distinguishing postural sway in subjects with somatosensory loss from age-matched control subjects. Given that filtered CoP sway-referencing disrupts the ability to utilize somatosensory information related to surface reactive force to a greater extent than the other two methods of sway-referencing, then these results support the hypothesis that subjects with diabetic peripheral neuropathy have lost more CoP information, than ankle or CoM angle information, for controlling postural sway in stance.     

Diabetic neuropathy and surface sway-referencing disrupt somatosensory information for postural stability in stance

In order to determine the type of somatosensory information for postural control that is most affected by neuropathy, we compared the relative effects of three methods of sway-referencing the surface in a group of subjects with profound loss of somatosensory function associated with sensory polyneuropathy from diabetes with age-matched control subjects. Sway-referencing disrupted somatosensory feedback for postural control by servo-controlling the dorsi- and plantar-flexion rotation of the support surface in proportion to anterior-posterior excursion of (1) ankle angle, (2) center of body mass (CoM) angle or (3) filtered center of pressure (CoP). Postural sway in subjects with somatosensory loss was significantly larger than normal on a firm surface but not on the sway-referenced surfaces, suggesting that sway-referencing disrupts somatosensory information for postural control already disrupted by neuropathy. Control subjects standing on any sway-referenced surface swayed significantly more than neuropathy subjects who stood on a firm surface, suggesting that sway-referencing disrupts more somatosensory information than disrupted by severe neuropathy. CoP sway-referencing was less sensitive than ankle or CoM sway-referencing for distinguishing postural sway in subjects with somatosensory loss from age-matched control subjects. Given that filtered CoP sway-referencing disrupts the ability to utilize somatosensory information related to surface reactive force to a greater extent than the other two methods of sway-referencing, then these results support the hypothesis that subjects with diabetic peripheral neuropathy have lost more CoP information, than ankle or CoM angle information, for controlling postural sway in stance.     

How experienced alpine-skiers cope with restrictions of ankle degrees-of-freedom when wearing ski-boots in postural exercises

The present study investigates the mechanisms underlying changes in postural strategy that occur to compensate for mechanical ankle joint restrictions induced by wearing ski-boots during postural exercises. Fourteen experienced skiers were asked to stand as still as possible in a stable (STA) posture and in 2 postures with instability in the medio/lateral and antero/posterior (ML and AP postures) direction. Postural tasks were performed with eyes open or closed and while wearing or not wearing ski-boots. The electromyographic (EMG) activity of representative lower limb muscles and positions of centre-of-foot pressure (COP) were recorded and analyzed. Our results illustrated enhanced postural performances with ski-boots in the STA posture, whereas postural performances remained unchanged when wearing ski-boots in the ML and AP postures. Analysis of COP sways in the frequency domain did not illustrate any modification in the contribution of different neuronal loops when the study subjects wore ski-boots. EMG showed that the mechanical effects of wearing ski-boots were compensated by changes in postural strategy through the reorganization of muscle coordination, made possible by inherent redundancies in the human body. The preservation of postural performances, despite restrictions of ankle degrees-of-freedom induced by ski-boots, emphasizes the subjects’ capacity to exploit the additional support provided by ski-boots by adequately adjusting muscle coordination to control posture in different balance conditions.     

Reactive gait and postural adjustments following the first exposures to (un)expected stepdown

This study evaluated the reactive biomechanical strategies associated with both upper- and lower-body (lead and trail limbs) following the first exposures to (un)expected stepdown at comfortable (1.22 ± 0.08 m/s) and fast (1.71 ± 0.11 m/s) walking velocities. Eleven healthy adults completed 34 trails per walking velocity over an 8-m, custom-built track with two forceplates embedded in its center. For the expected stepdown, the track was lowered by 0-, −10- and −20-cm from the site of the second forceplate, whereas the unexpected stepdown was created by camouflaging the second forceplate (−10-cm). Two-way repeated-measurement ANOVAs detected no velocity-related effects of stepdown on kinematic and kinetic parameters during lead limb stance-phase, and on the trail limb stepping kinematics. However, analyses of significant interactions revealed greater peak flexion angles across the trunk and the trail limb joints (hip, knee and ankle) in unexpected versus expected stepdown conditions at a faster walking velocity. The −10-cm unexpected stepdown (main effect) had a greater influence on locomotor behavior compared to expected conditions due mainly to the absence of predictive adjustments, reflected by a significant decrease in peak knee flexion, contact time and vertical impulse during stance-phase. Walking faster (main effect) was associated with an increase in hip peak flexion and net anteroposterior impulse, and a decrease in contact time and vertical impulse during stepdown. The trail limb, in response, swung forward faster, generating a larger and faster recovery step. However, such reactive stepping following unexpected stepdown was yet a sparse compensation for an unstable body configuration, assessed by significantly smaller step width and anteroposterior margin-of-stability at foot-contact in the first-recovery-step compared with expected conditions. These findings depict the impact of the expectedness of stepdown onset on modulation of global dynamic postural control for a successful accommodation of (un)expected surface elevation changes in young, healthy adults.     

The effect of short-term changes in body mass distribution on feed-forward postural control

It was recently shown that short-term changes in the whole body mass and associated changes in the vertical position of the center of mass (COM) modify anticipatory postural adjustments (APAs) [Li X, Aruin AS. The effect of short-term changes in the body mass on anticipatory postural adjustments. Exp Brain Res 2007;181:333–46]. In this study, we investigated whether changes in the body mass distribution and related changes in the anterior–posterior COM position affect APA generation. Fourteen subjects were instructed to catch a 2.2 kg load with their arms extended while standing with no additional weight or while carrying a 9.08 kg weight. Adding weight to a backpack, front pack or belly pocket was associated with an increase of the whole body mass, but it also involved changes in the anterior–posterior (A/P) and vertical positions of the COM. Electromyographic activity of leg and trunk muscles, body kinematics, and ground reaction forces were recorded and quantified within the typical time intervals of APAs. APAs were modified in conditions with changed body mass distribution: increased magnitude of anticipatory EMG activity in leg and trunk muscles, as well as co-activation of leg muscles and decreased anticipatory displacement of the COM in the vertical direction, were seen in conditions with increased body mass. Changes in the COM position induced in both A/P and vertical directions were associated with increased anticipatory EMG activity. In addition, they were linked to a co-activation of muscles at the ankle joints and significant changes in the center of pressure (COP) position. Modifications of the COM position induced in the A/P direction were related to increased anticipatory EMG activity in the leg and trunk muscles. At the same time, no significant differences in anticipatory EMG activity or displacement of COP were observed when changes of COM position were induced in the vertical direction. The study outcome suggests that the CNS uses different strategies while generating APAs in conditions with changes in the COM position induced in the anterior–posterior and vertical directions.     

Synergistic co-activation in multi-directional postural control in humans

To examine the muscle synergies of multi-directional postural control, we calculated the target-directed variance fraction (η) and net action direction of each muscle using the electromyogram-weighted averaging (EWA) method. Subjects stood barefoot on a force platform and maintained their posture by producing a center of pressure (COP) in twelve target directions. Surface electromyograms were recorded from 6 right-sided muscles: tibialis anterior (TA), soleus (SOL), lateral gastrocnemius (LG), medial gastrocnemius (MG), fibularis longus (FL), and gluteus medius (GM). η was calculated from COP with duration of 20-s, during which the COP was relatively constant. The EWA method was applied to the EMG and the two COP components to estimate the net action direction of each muscle. The results showed that η values in all directions did not cross the 0.8 threshold. This suggests that human postural control is achieved by synergistic co-activation. The EWA revealed that the net action directions of TA, SOL, LG, MG, and GM were 277.6°, 71.1°, 87.7°, 94.0°, and 2.2°, respectively. This suggests that postural maintenance by muscle synergy can be attributed to the relevant muscles having various action directions. These results demonstrate that muscle synergies can be investigated using COP fluctuations.     

Ankle intrinsic stiffness changes with postural sway

In standing, the human body is inherently unstable and its stabilization requires constant regulation of ankle torque, generated by a combination of ankle intrinsic properties, peripheral reflexes, and central contributions. Ankle intrinsic stiffness, which quantifies the joint intrinsic properties, has been usually assumed constant in standing; however, there is strong evidence that it is highly dependent on the joint torque, which changes significantly with sway in stance. In this study, we examined how ankle intrinsic stiffness changes with postural sway during standing. Ten subjects stood on a standing apparatus, while subjected to pulse perturbations of ankle position. The mean torque of a short period before the start of each pulse was used as a measure of background torque. Responses with similar background torques were grouped together and used to estimate the parameters of an intrinsic stiffness model. Stiffness estimates were normalized to the critical stiffness and the background torque was transformed to the center of pressure location. We found that in most subjects, the normalized stiffness increased linearly with the movement of center of pressure towards the toes, with an average slope of 2.11 ± 0.80 1/m·rad. This modulation of ankle intrinsic stiffness seems functionally appropriate, since the intrinsic stiffness increases quickly, as the center of pressure moves toward the toes and the limits of stability. These large changes of ankle intrinsic stiffness with postural sway must be incorporated in any model of stance control.     

Comparison of Leg Muscle Responses to Transcranial Magnetic Stimulation upon Standing on Stable and Unstable Supports

The role of supraspinal structures in postural adjustment upon standing on stable and unstable supports was studied in healthy individuals. For this purpose, transcranial magnetic stimulation (TMS) of the motor cortex was used in the region of leg representation. The subject stood with the eyes closed on a firm floor or on an unstable support in the form of a paperweight (20 cm in height, with a base radius of 32 cm) with mobility in the sagittal direction. Electric responses of four muscles—the soleus muscle, the anterior tibial muscle, the femoral biceps muscle, and the femoral rectus muscle—were recorded. It was shown that, in all the muscles, the response to TMS upon standing on an unstable support increased by 1.8–2.7 times as compared with the response upon standing on a firm floor. Since the increase in the tonic activity of the muscles studied was statistically insignificant upon switching over from standing on a firm floor to standing on an unstable support, it is hypothesized that the increase in the amplitude of muscle responses is connected with an increased activity of the supraspinal structures or with an increase in the effectiveness of corticospinal connections. The results are discussed from the point of view of the role of the motor cortex in maintaining balance on an unstable support.     

The relationship between dynamic balancing ability and posture-related modulation of the soleus H-reflex

Soleus H-reflex reveals down modulation with increased postural difficulty. Role of this posture-related reflex modulation is thought to shift movement control toward higher motor centers in order to facilitate more precise postural control. Present study hypothesized that the ability to modulate H-reflex is related to one’s ability to dynamically balance while in an unstable posture. This study examined the relationship between dynamic balancing ability and soleus H-reflex posture-related modulation. Thirty healthy adults participated. The soleus maximal H-reflex (Hmax), motor response (Mmax), and background EMG activity (bEMG) were obtained during three postural conditions: prone, open-legged standing, and closed-legged standing. Hmax/Mmax ratios were normalized via the corresponding bEMG in order to remove the effects of background muscle activity from the obtained H-reflex. Reflex modulation was calculated as the ratio of the normalized Hmax/Mmax ratios in one postural condition to another posture in a more difficult condition. Dynamic balancing ability was assessed by testing stability while standing on a wobble board. A significant negative correlation was observed between balancing scores and reflex modulation from open-legged standing to closed-legged standing. This suggests that the ability to modulate monosynaptic stretch reflex excitability in response to a changing posture is a significant factor for dynamic balancing.