Postural responses triggered by multidirectional leg lifts and surface tilts

The aim of the present study was to investigate the relationship between proactive and reactive components of postural control. We contrasted the kinematic and electromyographic (EMG) responses to multidirectional voluntary leg lifts with those elicited by unexpected surface tilts. In particular, we addressed the role of trunk stabilization following either a voluntary or forced weight shift from double to single limb support. Nine young female subjects stood with a standing posture of 45° toe-out and their arms abducted to shoulder level. On the experimenters signal, subjects either (1) lifted one leg as fast as possible in one of six directions (R/L side, R/L diagonal front, R/L diagonal back) to a height of 45° or (2) maintained standing as the support surface tilted at a rate of 53°/s to a height of 10° in one of six directions (R/L-up, R/L diagonal toes-up, R/L diagonal toes-down). For both tasks, our results showed that the center of pressure (COP) displacement began before or in conjunction with displacement of the center of mass (COM), after which the COP oscillated about the horizontal projection of the COM. In addition, the muscles were recruited in a distal-to-proximal sequence, either in anticipation of the voluntary leg lift or in response to the sudden surface tilt. Thus, the COP was being used dynamically to control displacement of the COM. The axial postural strategy comprising head, trunk, and pelvis movements was quantified by means of principal component analysis. More than 95% of the variance in the data could be described by the first two eigenvectors, which revealed specific coordination patterns dominated by pelvis rotation in one direction and head/trunk rotation in the opposite direction. Unexpected surface tilting elicited an automatic response strategy that focused on controlling the orientation of the head and trunk with respect to the vertical gravity vector while trunk verticality was compromised for movement generation and the recovery of postural equilibrium during leg lifting. In conclusion, regardless of the type (voluntary versus involuntary) or direction of perturbation, the strategy employed by the central nervous system to control the body COM displacement concerns mainly trunk stabilization.     

Neck,trunk and limb muscle responses during postural perturbations in humans

Summary This study examined the EMG onsets of leg, trunk, and neck muscles in 10 standing human subjects in response to support surface anterior and posterior translations, and to plantar and dorsiflexion rotations. The objective of the study was to test the hypothesis that the responses radiating upward from distal leg muscles represent part of a large ascending synergy encompassing axial muscles along the entire length of the body. If these responses are not ascending, then the muscles of the neck, and possibly the trunk, can be independently activated by vestibular, proprioceptive or visual inputs. We analysed the timing of postural muscle responses within and between body segments in order to determine whether they maintained a consistent temporal relationship under translational and rotational platform movement paradigms. Our results did not strongly support an ascending pattern of activation in all directions of platform perturbation. Temporal differences between activation patterns to platform perturbations in the forward or backward directions were revealed. In response to posterior platform translations we observed an ascending pattern of muscle responses along the extensor surface of the body. In addition, responses elicited in the neck flexor and abdominal muscles occurred as early as those of the stretched ankle muscles. This pattern of upward radiation from stretched ankle muscles was not as clear for anterior platform displacements, where early neck flexor muscle responses were observed during the ascending sequence on the flexor surface of the body. Platform rotations caused fewer responses in the neck and upper trunk muscles than translations, and all muscle responses occurred simultaneously rather than sequentially. Probable differences in the stimulation of vestibular and neck proprioceptive inputs and the mechanical demands of the rotation and translation paradigms are discussed.     

Effect of stance width on multidirectional postural responses

The effect of stance width on postural responses to 12 different directions of surface translations was examined. Postural responses were characterized by recording 11 lower limb and trunk muscles, body kinematics, and forces exerted under each foot of 7 healthy subjects while they were subjected to horizontal surface translations in 12 different, randomly presented directions. A quasi-static approach of force analysis was done, examining force integrals in three different epochs (background, passive, and active periods). The latency and amplitude of muscle responses were quantified for each direction, and muscle tuning curves were used to determine the spatial activation patterns for each muscle. The results demonstrate that the horizontal force constraint exerted at the ground was lessened in the wide, compared with narrow, stance for humans, a similar finding to that reported by Macpherson for cats. Despite more trunk displacement in narrow stance, there were no significant changes in body center of mass (CoM) displacement due to large changes in center of pressure (CoP), especially in response to lateral translations. Electromyographic (EMG) magnitude decreased for all directions in wide stance, particularly for the more proximal muscles, whereas latencies remained the same from narrow to wide stance. Equilibrium control in narrow stance was more of an active postural strategy that included regulating the loading/unloading of the limbs and the direction of horizontal force vectors. In wide stance, equilibrium control relied more on an increase in passive stiffness resulting from changes in limb geometry. The selective latency modulation of the proximal muscles with translation direction suggests that the trunk was being actively controlled in all directions. The similar EMG latencies for both narrow and wide stance, with modulation of only the muscle activation magnitude as stance width changed, suggest that the same postural synergy was only slightly modified for a change in stance width. Nevertheless, the magnitude of the trunk displacement, as well as of CoP displacement, was modified based on the degree of passive stiffness in the musculoskeletal system, which increased with stance width. The change from a more passive to an active horizontal force constraint, to larger EMG magnitudes especially in the trunk muscles and larger trunk and CoP excursions in narrow stance are consistent with a more effortful response for equilibrium control in narrow stance to perturbations in all directions.     

Vestibular influences on human postural control in combinations of pitch and roll planes reveal differences in spatiotemporal processing

The present study examined the influence of bilateral peripheral vestibular loss (BVL) in humans on postural responses to multidirectional surface rotations in the pitch and roll planes. Specifically, we examined the effects of vestibular loss on the directional sensitivity, timing, and amplitude of early stretch, balance correcting, and stabilizing reactions in postural leg and trunk muscles as well as changes in ankle torque and trunk angular velocity following multidirectional rotational perturbations of the support surface. Fourteen normal healthy adults and five BVL patients stood on a dual axis rotating platform which rotated 7.5° at 50°/s through eight different directions of pitch and roll combinations separated by 45°. Directions were randomized within a series of 44 perturbation trials which were presented first with eyes open, followed by a second series of trials with eyes closed. Vestibular loss did not influence the range of activation or direction of maximum sensitivity for balance correcting responses (120–220 ms). Response onsets at approximately 120 ms were normal in tibialis anterior (TA), soleus (SOL), paraspinals (PARAS), or quadriceps muscles. Only SOL muscle activity demonstrated a 38- to 45-ms delay for combinations of forward (toe-down) and roll perturbations in BVL patients. The amplitude of balance correcting responses in leg muscles between 120 and 220 ms was, with one exception, severely reduced in BVL patients for eyes open and eyes closed conditions. SOL responses were decreased bilaterally for toe-up and toe-down perturbations, but more significantly reduced in the downhill (load-bearing) leg for combined roll and pitch perturbations. TA was significantly reduced bilaterally for toe-up perturbations, and in the downhill leg for backward roll perturbations. Forward perturbations, however, elicited significantly larger TA activity in BVL between 120 and 220 ms compared to normals, which would act to further destabilize the body. As a result of these changes in response amplitudes, BVL patients had reduced balance correcting ankle torque between 160 and 260 ms and increased torque between 280 and 380 ms compared to normals. There were no differences in the orientation of the resultant ankle torque vectors between BVL and normals, both of which were oriented primarily along the pitch plane. For combinations of backward (toe-up) and roll perturbations BVL patients had larger balance correcting and stabilizing reactions (between 350 and 700 ms) in PARAS than normals and these corresponded to excessive trunk pitch and roll velocities. During roll perturbations, trunk velocities in BVL subjects after 200 ms were directed along directions different from those of normals. Furthermore, roll instabilities appeared later than those of pitch particularly for backward roll perturbations. The results of the study show that combinations of roll and pitch surface rotations yield important spatiotemporal information, especially with respect to trunk response strategies changed by BVL which are not revealed by pitch plane perturbations alone. Our results indicate that vestibular influences are earlier for the pitch plane and are directed to leg muscles, whereas roll control is later and focused on trunk muscles. Electronic Publication     

The influence of postural threat on the cortical response to unpredictable and predictable postural perturbations

This study investigated the effects of postural threat on the cortical response associated with postural reactions to predictable and unpredictable perturbations to upright stance. Postural threat was manipulated by having individuals stand on an elevated surface to alter the context in which the postural task was performed. Ten healthy young adults experienced a series of predictable and unpredictable trunk perturbations when standing at ground level and at the edge of a platform located 3.2 m above the ground. Participants felt less confident, more fearful of falling, more anxious and less stable when standing at the high surface height. Unpredictable perturbations generated a large negative potential (N1) which was increased by 84% when standing at the high compared to low surface height. The magnitude of change in this potential was related to the magnitude of change in balance perceptions, such as confidence and fear. Predictable perturbations did not generate a N1 potential but instead produced an anticipation-related potential prior to the perturbation. This cortical activity observed in response to predictable perturbations was not influenced by postural threat. A large N1 potential was observed for a 'surprise' perturbation that followed a series of predictable perturbations. There was a trend for the amplitude of this potential to be increased when standing at the high compared to low surface height. The results of this study provide evidence for the modulating influence of psychological factors related to postural threat on the cortical activity associated with postural reactions to unpredictable perturbations.     

Three components of postural control associated with pushing in symmetrical and asymmetrical stance

A number of occupational and leisure activities that involve pushing are performed in symmetrical or asymmetrical stance. The goal of this study was to investigate early postural adjustments (EPAs), anticipatory postural adjustments (APAs), and compensatory postural adjustments (CPAs) during pushing performed while standing. Ten healthy volunteers stood in symmetrical stance (with feet parallel) or in asymmetrical stance (staggered stance with one foot forward) and were instructed to use both hands to push forward the handle of a pendulum attached to the ceiling. Bilateral EMG activity of the trunk and leg muscles and the center of pressure (COP) displacements in the anterior–posterior (AP) and medial–lateral (ML) directions were recorded and analyzed during the EPAs, APAs, and CPAs. The EMG activity and the COP displacement were different between the symmetrical and asymmetrical stance conditions. The COP displacements in the ML direction were significantly larger in staggered stance than in symmetrical stance. In staggered stance, the EPAs and APAs in the thigh muscles of the backward leg were significantly larger, and the CPAs were smaller than in the forward leg. There was no difference in the EMG activity of the trunk muscles between the stance conditions. The study outcome confirmed the existence of the three components of postural control (EPAs, APAs, and CPAs) in pushing. Moreover, standing asymmetrically was associated with asymmetrical patterns of EMG activity in the lower extremities reflecting the stance-related postural control during pushing. The study outcome provides a basis for studying postural control during other daily activities involving pushing.     

Vestibular and somatosensory contributions to responses to head and body displacements in stance

The relative contribution of vestibular and somatosensory information to triggering postural responses to external body displacements may depend on the task and on the availability of sensory information in each system. To separate the contribution of vestibular and neck mechanisms to the stabilization of upright stance from that of lower body somatosensory mechanisms, responses to displacements of the head alone were compared with responses to displacements of the head and body, in both healthy subjects and in patients with profound bilateral vestibular loss. Head displacements were induced by translating two 1-kg weights suspended on either side of the head at the level of the mastoid bone, and body displacements were induced translating the support surface. Head displacements resulted in maximum forward and backward head accelerations similar to those resulting from body displacements, but were not accompanied by significant center of body mass, ankle, knee, or hip motions. We tested the effect of disrupting somatosensory information from the legs on postural responses to head or body displacements by sway-referencing the support surface. The subjects' eyes were closed during all testing to eliminate the effects of vision. Results showed that head displacements alone can trigger medium latency (48–84 ms) responses in the same leg and trunk muscles as body displacements. Nevertheless, it is unlikely that vestibular signals alone normally trigger directionally specific postural responses to support surface translations in standing humans because: (1) initial head accelerations resulting from body and head displacements were in opposite directions, but were associated with activation of the same leg and trunk postural muscles; (2) muscle responses to displacements of the head alone were only one third of the amplitude of responses to body displacements with equivalent maximum head accelerations; and (3) patients with profound bilateral vestibular loss showed patterns and latencies of leg and trunk muscle responses to body displacements similar to those of healthy subjects. Altering somatosensory information, by sway-referencing the support surface, increased the amplitude of ankle muscle activation to head displacements and reduced the amplitude of ankle muscle activation to body displacements, suggesting context-specific reweighting of vestibular and somatosensory inputs for posture. In contrast to responses to body displacements, responses to direct head displacements appear to depend upon a vestibulospinal trigger, since trunk and leg muscle responses to head displacements were absent in patients who had lost vestibular function as adults. Patients who lost vestibular function as infants, however, had near normal trunk and leg response to head displacements, suggesting a substitution of upper trunk and neck somatosensory inputs for missing vestibular inputs during development.     

The effect of short-term changes in the body mass on anticipatory postural adjustments

The purpose of the study was to investigate whether anticipatory postural adjustments (APAs) are modified with short-term changes in the body mass. Nine subjects were asked to catch a 2.2 kg load with their arms extended under conditions of no weight and when additional weights of 10 and 20% of the subject’s body weight (BW) were attached to single body locations or when 20 or 40% BW were attached evenly to two locations. Attaching weights was associated with an increase of the whole body mass, but also involved changes in the vertical position of the center of mass (COM). Electromyographic activity of leg and trunk muscles and ground reaction forces were recorded and quantified within the typical time intervals of APAs. APAs were influenced by the magnitude of the weight attached to the body: an increase in the body mass was associated with anticipatory co-activation of trunk and leg muscles. The level of this co-activation increased with an increase in the magnitude of weight added to the body. At the same time, APAs were affected by the changes in the vertical position of COM. These findings suggest that in the case of short-term changes in the body mass, the CNS might prioritize information regarding the magnitude and location of the additional weight added to the body and utilize a strategy of anticipatory co-activation of postural muscles directed at the stabilization of body segments.     

Control of reactive balance adjustments in perturbed human walking: roles of proximal and distal postural muscle activity

Studies on the proactive control of gait have shown that proximal (hip/trunk) muscles are the primary contributors to balance control, while studies on reactive balance control during perturbed gait, examining only activity in distal (leg/thigh) muscles, have shown that these muscles are important in compensating for a gait disturbance. This study tested the hypothesis that proximal muscles are also primary contributors to reactive balance control during perturbed gait. Thirty-three young adults participated in a study in which an anterior slip was simulated at heel strike by the forward displacement of a force plate on which they walked. Surface electromyographic data were recorded from bilateral leg, thigh, hip and trunk muscles. Kinematic data were collected on joint angle changes in response to the perturbation. The results did not support the hypothesis that the proximal muscles contribute significantly to balance control during perturbed gait. The proximal muscles did not demonstrate more consistent activation, earlier onset latency, longer burst duration or larger burst magnitude than distal muscles. Moreover, although proximal postural activity was often present in the first slip trial, it tended to adapt away in later trials. By contrast, the typical postural responses exhibited by young adults consisted of an early (90–140 ms), high-magnitude (4–9 times muscle activity during normal walking) and relatively long duration (70–200 ms) activation of bilateral anterior leg muscles as well as the anterior and posterior thigh muscles. Thus, postural activity from bilateral leg and thigh muscles and the coordination between the two lower extremities were the key to reactive balance control and were sufficient for regaining balance within one gait cycle. The adaptive attenuation of proximal postural activity over repeated trials suggests that the nervous system overcompensates for a novel balance threat in the first slip trial and fine-tunes its responses with experience. Received: 24 March 1997 /Accepted: 4 September 1997     

Effect of head position on postural orientation and equilibrium

This study examined (1) how changes in head position affect postural orientation variables during stance and (2) whether changes in head position affect the rapid postural response to linear translation of the support surface in the horizontal plane. Cats were trained to stand quietly on a moveable platform and to maintain five different head positions: center, left, right, up, and down. For each head position, stance was perturbed by translating the support surface linearly in 16 different directions in the horizontal plane. Postural equilibrium responses were quantified in terms of the ground reaction forces, kinematics, dynamics (net joint torques), body center of mass, and electromyographic (EMG) responses of selected limb and trunk muscles. A change in head position involved rotation of not only the neck but also the scapulae and anterior trunk. Tonic EMG levels were modulated in several forelimb and scapular muscles but not hindlimb muscles. Finally, large changes in head orientation in both horizontal and vertical planes did not hamper the ability of cats to maintain postural equilibrium during linear translation of the support surface. The trajectory of the body’s center of mass was the same, regardless of head position. The main change was observed in joint torques at the forelimbs evoked by the perturbation. Evoked EMG responses of forelimb and scapular muscles were modulated in terms of magnitude but not spatial tuning. Hindlimb responses were unchanged. Thus, the spatial and temporal pattern of the automatic postural response was unchanged and only amplitudes of evoked activity were modulated by head position. Received: 14 October 1997 / Accepted: 22 April 1998     

Deceleration affects anticipatory and reactive components of triggered postural responses

Understanding the physiological and psychological factors that contribute to healthy and pathological balance control in man has been made difficult by the confounding effects of the perturbations used to test balance reactions. The present study examined how postural responses were influenced by the acceleration–deceleration interval of an unexpected horizontal translation. Twelve adult males maintained balance during unexpected forward and backward surface translations with two different acceleration–deceleration intervals and presentation orders (serial or random). “SHORT” perturbations consisted of an initial acceleration (peak acceleration 1.3 m s⁻²; duration 300 ms) followed 100 ms later by a deceleration. “LONG” perturbations had the same acceleration as SHORT perturbations, followed by a 2-s interval of constant velocity before deceleration. Surface and intra-muscular electromyography (EMG) from the leg, trunk, and shoulder muscles were recorded along with motion and force plate data. LONG perturbations induced larger trunk displacements compared to SHORT perturbations when presented randomly and larger EMG responses in proximal and distal muscles during later (500–800 ms) response intervals. During SHORT perturbations, activity in some antagonist muscles was found to be associated with deceleration and not the initial acceleration of the support surface. When predictable, SHORT perturbations facilitated the use of anticipatory mechanisms to attenuate early (100–400 ms) EMG response amplitudes, ankle torque change and trunk displacement. In contrast, LONG perturbations, without an early deceleration effect, did not facilitate anticipatory changes when presented in a predictable order. Therefore, perturbations with a short acceleration–deceleration interval can influence triggered postural responses through reactive effects and, when predictable with repeated exposure, through anticipatory mechanisms.     

The effect of aging on anticipatory postural control

The aim of the study was to investigate the differences in anticipatory postural adjustments (APAs) between young and older adults and its effect on subsequent control of posture. Ten healthy older adults and thirteen healthy young adults were exposed to predictable external perturbations using the pendulum impact paradigm. Electromyographic activity of the trunk and leg muscles, the center of pressure (COP), and center of mass (COM) displacements in the anterior–posterior direction were recorded and analyzed during the anticipatory and compensatory postural adjustments (CPAs) phases of postural control. The effect of aging was seen as delayed anticipatory muscle activity and larger compensatory muscle responses in older adults as compared to young adults. Moreover, in spite of such larger reactive responses, older adults were still more unstable, exhibiting larger COP and COM peak displacements after the perturbation than young adults when exposed to similar postural disturbances. Nonetheless, while APAs are impaired in older adults, the ability to recruit muscles anticipatorily is largely preserved; however, due to their smaller magnitudes and delayed onsets, it is likely that their effectiveness in reducing the magnitude of CPAs is smaller. The outcome of the study lends support toward investigating the ways of improving anticipatory postural control in people with balance impairments due to aging or neurological disorders.     

Directional sensitivity of stretch reflexes and balance corrections for normal subjects in the roll and pitch planes

A large body of evidence has been collected which describes the response parameters associated with automatic balance corrections in man to perturbations in the pitch plane. However, perturbations to human stance can be expected from multiple directions. The purpose of the present study was to describe the directional sensitivities of muscle responses re-establishing disturbed stance equilibrium in normal subjects. The contributions of stretch reflex and automatic balance-correcting responses to balance control, and concomitant biomechanical reactions, were examined for combinations of pitch and roll perturbations of the support surface. More specifically, muscle responses, initial head accelerations and trunk velocities were analyzed with the intention of identifying possible origins of directionally specific triggering signals and to examine how sensory information is used to modulate triggered balance corrections with respect to direction. Fourteen healthy adults were required to stand on a dual-axis rotating platform capable of delivering rotational perturbations with constant amplitude (7.5°) and velocity (50°/s) through multiple directions in the pitch and roll planes. Each subject was randomly presented with 44 support surface rotations through 16 different directions separated by 22.5° first under eyes-open, and then, for a second identical set of rotations, under eyes-closed conditions. Bilateral muscle activities from tibialis anterior, soleus, lateral quadriceps and paraspinals were recorded, averaged across direction, and areas calculated over intervals with significant bursts of activity. Trunk angular velocity and ankle torque data were averaged over intervals corresponding to significant biomechanical events. Stretch reflex (intervals of 40–100, 80–120 ms) and automatic balance-correcting responses (120–220, 240–340 ms) in the same muscle were sensitive to distinctly different directions. The directions of the maximum amplitude of balance-correcting activity in leg muscles were oriented along the pitch plane, approximately 180° from the maximum amplitude of stretch responses. Ankle torques for almost all perturbation directions were also aligned along the pitch plane. Stretch reflexes in paraspinal muscles were tuned along the 45° plane but at 90° to automatic balance corrections and 180° to unloading responses in the same muscle. Stretch reflex onsets in paraspinal muscles were observed at 60 ms, as early as those of soleus muscles. In contrast, unloading reflexes in released paraspinal muscles were observed at 40 ms for perturbations which caused roll of the trunk towards the recorded muscle. Onsets of trunk roll velocities were earlier and more rapid than those observed for pitch velocities. Trunk pitch occurred for pure roll directions but not vice versa. When considered together, early stretch and unloading of paraspinals, and concomitant roll and pitch velocities of the trunk requiring a roll-and-pitch-based hip torque strategy, bring into question previous hypotheses of an ankle-based trigger signal or ankle-based movement strategies for postural balance reactions. These findings are compatible with the hypothesis that stretch-, force- and joint-related proprioceptive receptors at the level of the trunk provide a directionally sensitive triggering mechanism underlying a, minimally, two-stage (pitch-based leg and pitch-and-roll-based trunk) balance-correcting strategy. Accelerometer recordings from the head identified large vertical linear accelerations only for pitch movements and angular roll accelerations only during roll perturbations with latencies as early as 15 ms. Thus, it appears that balance corrections in leg and trunk muscles may receive strong, receptor-dependent (otolith or vertical canal) and directionally sensitive amplitude-modulating input from vestibulospinal signals. Received: 4 September 1998 / Accepted: 30 April 1999     

Control of roll and pitch motion during multi-directional balance perturbations

Does the central nervous system (CNS) independently control roll and pitch movements of the human body during balance corrections? To help provide an answer to this question, we perturbed the balance of 16 young healthy subjects using multi-directional rotations of the support surface. All rotations had pitch and roll components, for which either the roll (DR) or the pitch (DP) component were delayed by 150 ms or not at all (ND). The outcome measures were the biomechanical responses of the body and surface EMG activity of several muscles. Across all perturbation directions, DR caused equally delayed shifts (150 ms) in peak lateral centre of mass (COM) velocity. Across directions, DP did not cause equally delayed shifts in anterior–posterior COM velocity. After 300 ms however, the vector direction of COM velocity was similar to the ND directions. Trunk, arm and knee joint rotations followed this roll compared to pitch pattern, but were different from ND rotation synergies after 300 ms, suggesting an intersegmental compensation for the delay effects. Balance correcting responses of muscles demonstrated both roll and pitch directed components regardless of axial alignment. We categorised muscles into three groups: pitch oriented, roll oriented and mixed based on their responses to DR and DP. Lower leg muscles were pitch oriented, trunk muscles were roll oriented, and knee and arm muscles were mixed. The results of this study suggest that roll, but not pitch components, of balance correcting movement strategies and muscle synergies are separately programmed by the CNS. Reliance on differentially activated arm and knee muscles to correct roll perturbations reveals a dependence of the pitch response on that of roll, possibly due to biomechanical constraints, and accounts for the failure of DP to be transmitted equally in time across all limbs segments. Thus it appears the CNS preferentially programs the roll response of the body and then adjusts the pitch response accordingly.     

Is lower leg proprioception essential for triggering human automatic postural responses?

It is unknown to what extent automatic postural responses are triggered by lower leg proprioception. This issue was addressed by studying postural control in five carefully selected patients with subtle diabetic polyneuropathy (restricted to the lower legs) and 15 healthy subjects. All patients had bilaterally absent Achilles tendon reflexes and weak or absent patella tendon reflexes, but muscle strength was fully preserved. Subjects were tested while standing on a supporting, movable force-plate. The contribution of lower leg proprioception to automatic postural responses was investigated by randomly exposing the subjects to either a 4 degrees 'toe-up' rotational perturbation ('normal ankle input'), a simultaneous 4-cm rearward translation and 4 degrees toe-up rotation ('enhanced ankle input'), or a simultaneous 4-cm rearward translation and 4 degrees 'toe-down' rotation ('nulled ankle input'). We recorded surface EMG (stretch reflexes and balance-correcting responses) from leg and trunk muscles, ankle torque and angular velocities of the upper and lower legs and trunk. We argued that automatic postural responses that have abnormally small amplitudes in patients and are modulated in controls with the velocity of different types of ankle rotations must receive a major input from lower leg proprioception. Conversely, automatic postural responses that are weakly modified in amplitude or onset by different ankle perturbations and are present despite nulled ankle inputs and, finally, are unaffected in patients with distal polyneuropathy must be triggered or modulated by inputs other than from lower leg proprioception. Normal postural synergies and strategies were maintained in patients, although within a given synergy the timing and amplitude of some automatic postural responses were abnormal. A few automatic postural responses appeared to be triggered or modulated by lower leg proprioception. Thus, early stretch reflexes in soleus and medial gastrocnemius were severely diminished in patients, while in controls these stretch reflexes were modulated by different ankle perturbations. Furthermore, balance-correcting responses in tibialis anterior were diminished and delayed in patients, while in controls these balance-correcting responses were modulated by different ankle perturbations. Other automatic postural responses were apparently not triggered or modulated by lower leg proprioception, but likely received a major input from more proximal sensory systems. Thus, in both groups prominent balance-correcting responses were present in several muscles (soleus, gastrocnemius, quadriceps, paraspinals and trapezius) during the 'nulled ankle input' condition, where ankle position was stabilised over the first 250 ms. During the 'enhanced ankle input' condition, where prominent ankle dorsiflexion occurred during the first 200 ms, amplitudes of balance-correcting responses were only marginally weaker in patients than in controls. We analysed body segment displacements to unveil the potential nature of proximal triggers for automatic postural responses. As opposed to the 'inverted pendulum' concept of postural control, early movement occurred in the knees, hips and trunk well before the onset of automatic postural responses. For example, during the 'nulled ankle input' condition, the lower leg moved forward with early knee flexion, followed by knee extension. The trunk extended backwards at 80 ms, which was followed by forward flexion. The absent stretch reflex and weaker balance-correcting responses in patients produced changed trunk velocity profiles (mainly a reduced initial backward motion of the trunk), but lower-body segment movements showed no consistent differences between the two groups. Considering these body segment displacements, any automatic postural response with an onset within the first 200 ms could well be triggered by receptors located at the knee, hip or trunk. (ABSTRACT TRUNCATED)     

Coexistence of stability and mobility in postural control: evidence from postural compensation for respiration

This study evaluated the extent to which movement of the lower limbs and pelvis may compensate for the disturbance to posture that results from respiratory movement of the thorax and abdomen. Motion of the neck, pelvis, leg and centre of pressure (COP) were recorded with high resolution in conjunction with electromyographic activity (EMG) of flexor and extensor muscles of the trunk and hip. Respiration was measured from ribcage motion. Subjects breathed quietly, and with increased volume due to hypercapnoea (as a result of breathing with increased dead-space) and a voluntary increase in respiration. Additional recordings were made during apnoea. The relationship between respiration and other parameters was measured from the correlation between data in the frequency domain (i.e. coherence) and from time-locked averages triggered from respiration. In quiet standing, small angular displacements ( approximately 0.5 degrees ) of the trunk and leg were identified in raw data. Correspondingly, there were peaks in the power spectra of the angular movements and EMG. While body movement and EMG were coherent with respiration (>0.5), the coherence between respiration and COP displacement was low (<0.2). The amplitude of movement and coherence was increased when respiration was increased. The present data suggest that the postural disturbance that results from respiratory movement is matched, at least partly, and counteracted by small angular displacements of the lower trunk and lower limbs. Thus, stability in quiet stance is dependent on movement of multiple body segments and control of equilibrium cannot be reduced to control of a single joint.     

Practice modifies the developing automatic postural response

 The purpose of this study was to examine effects of experience with a postural task on components of the automatic postural response including: (1) probability of activation of functionally appropriate postural muscles; (2) number of functionally appropriate postural muscles activated; and (3) onset latencies of functionally appropriate postural muscles in infants. Infants (n=15; age 36–48 weeks old) able to pull themselves into a standing position but not able to walk independently were tested using a postural task requiring the infant to stand and balance, with support, following a forward or backward movement of the support surface (platform perturbation). Infants were tested twice at 5-day intervals. One-half of the infants, the training group, were given intense platform perturbation training on the days between test sessions. Infants in the second group were also brought into the laboratory on the days between test sessions but were not exposed to platform perturbations during those days. Electromyograms of six leg and trunk muscles were recorded during test sessions to provide muscle onset latencies, probability of muscle activation data, and the number of postural muscles activated following a perturbation. Training infants demonstrated significant increases in probability of activating functionally appropriate muscles with tibialis anterior, quadriceps, and abdominal muscles activated in response to backward sway and gastrocnemius muscle in response to forward sway. The number of functionally appropriate postural muscles activated in a single trial also increased in the training group. There were no significant changes in mean postural muscle onset latencies or number of trials with antagonist muscle coactivation for either training or control groups. These findings suggest that during development selective parameters of the automatic postural response are affected by experience with the postural task. Received: 30 November 1996 / Accepted: 12 September 1996     

Central programming of postural movements: adaptation to altered support-surface configurations

We studied the extent to which automatic postural actions in standing human subjects are organized by a limited repertoire of central motor programs. Subjects stood on support surfaces of various lengths, which forced them to adopt different postural movement strategies to compensate for the same external perturbations. We assessed whether a continuum or a limited set of muscle activation patterns was used to produce different movement patterns and the extent to which movement patterns were influenced by prior experience. Exposing subjects standing on a normal support surface to brief forward and backward horizontal surface perturbations elicited relatively stereotyped patterns of leg and trunk muscle activation with 73- to 110-ms latencies. Activity began in the ankle joint muscles and then radiated in sequence to thigh and then trunk muscles on the same dorsal or ventral aspect of the body. This activation pattern exerted compensatory torques about the ankle joints, which restored equilibrium by moving the body center of mass forward or backward. This pattern has been termed the ankle strategy because it restores equilibrium by moving the body primarily around the ankle joints. To successfully maintain balance while standing on a support surface short in relation to foot length, subjects activated leg and trunk muscles at similar latencies but organized the activity differently. The trunk and thigh muscles antagonistic to those used in the ankle strategy were activated in the opposite proximal-to-distal sequence, whereas the ankle muscles were generally unresponsive. This activation pattern produced a compensatory horizontal shear force against the support surface but little, if any, ankle torque. This pattern has been termed the hip strategy, because the resulting motion is focused primarily about the hip joints. Exposing subjects to horizontal surface perturbations while standing on support surfaces intermediate in length between the shortest and longest elicited more complex postural movements and associated muscle activation patterns that resembled ankle and hip strategies combined in different temporal relations. These complex postural movements were executed with combinations of torque and horizontal shear forces and motions of ankle and hip joints. During the first 5-20 practice trials immediately following changes from one support surface length to another, response latencies were unchanged. The activation patterns, however, were complex and resembled the patterns observed during well-practiced stance on surfaces of intermediate lengths.(ABSTRACT TRUNCATED AT 400 WORDS)     

Influence of fear of falling on anticipatory postural control of medio-lateral stability during rapid leg flexion

During leg flexion from erect posture, postural stability is organized in advance during “anticipatory postural adjustments” (APA). During these APA, inertial forces are generated that propel the centre of gravity (CoG) laterally towards stance leg side. This study examined how fear of falling (FoF) may influence this anticipatory postural control of medio-lateral (ML) stability. Ten young healthy participants performed a series of leg flexions at maximal velocity from low and high surface heights (6 and 66 cm above ground, respectively). In this latter condition with increased FoF, stance foot was placed at the lateral edge of the support surface to induce maximal postural threat. Results showed that the amplitude of ML inertial forces generated during APA decreased with FoF; this decrease was compensated by an increase in APA duration so that the CoG position at time of swing foot-off was located further towards stance leg side. With these changes in ML APA, the CoG was propelled in the same final (unipodal) position above stance foot as in condition with low FoF. These results contrast with those obtained in the literature during quiet standing which showed that FoF did not have any influence on the ML component of postural control. It is proposed that ML APA are modified with increased FoF, in such a way that the risk of a sideway fall induced by the large CoG motion is attenuated.     

Automated postural responses are modified in a functional manner by instruction

The restoration of upright balance after a perturbation relies on highly automated and, to a large extent, stereotyped postural responses. Although these responses occur before voluntary control comes into play, previous research has shown that they can be functionally modulated on the basis of cognitive set (experience, advanced warning, instruction, etc.). It is still unknown, however, how the central nervous system deals with situations in which the postural response is not necessarily helpful in the execution of a task. In the present study, the effects of instruction on automated postural responses in neck, trunk, shoulder, and leg muscles were investigated when people were either instructed to recover balance after being released from an inclined standing posture [balance recovery (BR) trials], or not to recover at all and fall onto a safety mattress in the most comfortable way [fall (F) trials], in both backward and leftward directions. Participants were highly successful in following the instructions, consistently exhibiting stepping responses for balance recovery in BR trials, and suppressing stepping in the F trials. Yet EMG recordings revealed similar postural responses with onset latencies between 70 and 130 ms in both BR and F trials, with slightly delayed responses in F trials. In contrast, very pronounced and early differences were observed between BR and F trials in response amplitudes, which were generally much higher in BR than in F trials, but with clear differentiation between muscles and perturbation directions. These results indicate that a balance perturbation always elicits a postural response, irrespective of the task demands. However, when a specific balance recovery response is not desired after a perturbation, postural responses can be selectively downregulated and integrated into the motor output in a functional and goal-oriented way.