Effects of lateral perturbations and changing stance conditions on anticipatory postural adjustment

The study investigates the role of lateral muscles and changing stance conditions in anticipatory postural adjustments (APAs). Subjects stood laterally to an aluminum pendulum released by an experimenter and were required to stop it with their right or left hand. Stance conditions were manipulated by having the subjects stand in the following positions: on a single limb (SS), with feet together (narrow base of support, NB), and with feet shoulder width apart (regular base of support, RB). Bilateral EMG activity of dorsal, ventral, and lateral trunk and leg muscles and ground reaction forces were recorded and quantified within the time intervals typical of APAs. Anticipatory postural adjustments were seen in all experimental conditions, and their magnitudes depended on the stance and the side of perturbation. Accordingly, APAs in lateral muscles increased on the side of perturbation in SS condition, while simultaneous activation of dorsal muscles occurred on the contralateral side. Smaller APAs were seen in lateral muscles in conditions with a wider base of support (NB, RB) and APAs in dorsal muscles were smaller in NB – in comparison to RB – stance. The results of the present study provide new data on the role of lateral, ventral, and dorsal muscles in anticipatory postural control when dealing with lateral perturbations in conditions of postural instability.     

Effects of amplitude and predictability of perturbations to the arm on anticipatory and reactionary muscle responses to maintain balance

Disturbances to balance arising from forces applied to the upper limb have received relatively little attention compared to disturbances arising from support surface perturbations. In this study we applied fast ramp perturbations to the hand in anterior, posterior, medial and lateral directions. The effects of perturbation predictability and amplitude on the postural response of upper limb, trunk and lower limb muscles were investigated. Perturbations were applied either in blocks of constant amplitude and direction (predictable) or with direction and amplitude varying randomly (random) from trial to trial. The spatial-temporal patterns of anticipatory muscle activation under the predictable condition and the reactionary responses following the perturbation under both conditions were similarly organized. The size of the response increased systematically with the perturbation magnitude for both anticipatory and reactionary changes in muscle activation. However, the slope of the relation between perturbation amplitude and the magnitude of the change in muscle activation was greater when perturbations were predictable than when they were randomly selected. The timing of both the anticipatory and reactionary increases in muscle activation was invariant across perturbation amplitudes. The characteristics of the reactionary responses have a similar organization to the long latency muscle responses to support surface perturbations.     

The role of anticipatory postural adjustments in compensatory control of posture: 1. Electromyographic analysis

Anticipatory (APAs) and compensatory (CPAs) postural adjustments are the two principal mechanisms that the central nervous system uses to maintain equilibrium while standing. We studied the role of APAs in compensatory postural adjustments. Eight subjects were exposed to external predictable and unpredictable perturbations induced at the shoulder level, while standing with eyes open and closed. Electrical activity of leg and trunk muscles was recorded and analyzed during four epochs representing the time duration typical for anticipatory and compensatory postural control. No anticipatory activity of the trunk and leg muscles was seen in the case of unpredictable perturbations; instead, significant compensatory activation of muscles was observed. When the perturbations were predictable, strong anticipatory activation was seen in all the muscles: such APAs were associated with significantly smaller compensatory activity of muscles and COP displacements after the perturbations.The outcome of the study highlights the importance of APAs in control of posture and points out the existence of a relationship between the anticipatory and the compensatory components of postural control. It also suggests a possibility to enhance balance control by improving the APAs responses during external perturbations.     

The effect of the amplitude of motor action on anticipatory postural adjustments.

The purpose of this study was to determine whether changes in the amplitude of a motor action triggering the same perturbation affect anticipatory postural adjustments (APAs). Healthy subjects performed releases of the same load with shoulder abduction movements of different amplitudes. Changes in the electrical activity of trunk and leg muscles, as well as displacements of the center of pressure were recorded. Generally, there were no differences in anticipatory activity of muscles and displacements of the center of pressure between series of load releases induced by motor actions of different amplitudes. We suggest that the CNS arranges APAs based on the magnitude of the perturbation if the same muscle groups generate motor actions of different amplitudes.     

Trunk muscle contributions of to L4–5 joint rotational stiffness following sudden trunk lateral bend perturbations

The purpose of this research was to investigate the contributions of individual muscles to joint rotational stiffness and total joint rotational stiffness about the lumbar spine’s L_4–5 joint prior to, and following, sudden dynamic lateral perturbations to the trunk. Kinematic and surface EMG data were collected while subjects maintained a kneeling posture on a robotic platform, while restrained so that motions caused by the perturbation were transferred to the pelvis, causing motion of the trunk and head. The robotic platform caused sudden inertial trunk lateral perturbations to the right or left, with or without timing and direction knowledge. An EMG-driven model of the lumbar spine was used to calculate the muscle forces and contributions to joint rotational stiffness during the perturbations. Data showed 95% and 106% increases in total joint rotational stiffness_, about the lateral bend and axial twist axes, when subjects had knowledge of the timing of the perturbation. Also, the contralateral muscles exhibited a significantly larger total joint rotational stiffness about the lateral bend axis, and earlier surface EMG responses, than the ipsilateral muscles. The results indicate that, when the timing of the perturbation was unknown, subjects relied more on delayed muscle forces following the perturbation to stiffen the L_4–5 joint.     

The Temporal Structure of Vertical Arm Movements

The present study investigates how the CNS deals with the omnipresent force of gravity during arm motor planning. Previous studies have reported direction-dependent kinematic differences in the vertical plane; notably, acceleration duration was greater during a downward than an upward arm movement. Although the analysis of acceleration and deceleration phases has permitted to explore the integration of gravity force, further investigation is necessary to conclude whether feedforward or feedback control processes are at the origin of this incorporation. We considered that a more detailed analysis of the temporal features of vertical arm movements could provide additional information about gravity force integration into the motor planning. Eight subjects performed single joint vertical arm movements (45° rotation around the shoulder joint) in two opposite directions (upwards and downwards) and at three different speeds (slow, natural and fast). We calculated different parameters of hand acceleration profiles: movement duration (MD), duration to peak acceleration (D PA), duration from peak acceleration to peak velocity (D PA-PV), duration from peak velocity to peak deceleration (D PV-PD), duration from peak deceleration to the movement end (D PD-End), acceleration duration (AD), deceleration duration (DD), peak acceleration (PA), peak velocity (PV), and peak deceleration (PD). While movement durations and amplitudes were similar for upward and downward movements, the temporal structure of acceleration profiles differed between the two directions. More specifically, subjects performed upward movements faster than downward movements; these direction-dependent asymmetries appeared early in the movement (i.e., before PA) and lasted until the moment of PD. Additionally, PA and PV were greater for upward than downward movements. Movement speed also changed the temporal structure of acceleration profiles. The effect of speed and direction on the form of acceleration profiles is consistent with the premise that the CNS optimises motor commands with respect to both gravitational and inertial constraints.     

Anticipatory postural adjustments to arm movement reveal complex control of paraspinal muscles in the thorax

Anatomical and empirical data suggest that deep and superficial muscles may have different functions for thoracic spine control. This study investigated thoracic paraspinal muscle activity during anticipatory postural adjustments associated with arm movement. Electromyographic (EMG) recordings were made from the right deep (multifidus/rotatores) and superficial (longissimus) muscles at T5, T8, and T11 levels using fine-wire electrodes. Ten healthy participants performed fast unilateral and bilateral flexion and extension arm movements in response to a light. EMG amplitude was measured during 25 ms epochs for 150 ms before and 400 ms after deltoid EMG onset. During arm flexion movements, multifidus and longissimus had two bursts of activity, one burst prior to deltoid and a late burst. With arm extension both muscles were active in a single burst after deltoid onset. There was differential activity with respect to direction of trunk rotation induced by arm movement. Right longissimus was most active with left arm movements and right multifidus was most active with right arm movements. All levels of the thorax responded similarly. We suggest that although thoracic multifidus and longissimus function similarly to control sagittal plane perturbations, these muscles are differentially active with rotational forces on the trunk.     

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.     

Modifications of anticipatory postural adjustments in a rock climbing task: the effect of supporting wall inclination.

The aim of this study was to analyse the influence of initial postural constraint on the realisation of a leg release in a rock climbing task. Two conditions were tested: a vertical posture and an overhanging posture. The overhanging posture was characterised by a large sustentation base, which enhanced the mechanical possibilities of the system. Subjects had to release their right foot in both postural conditions. In the vertical posture, movement's effectuation was associated with anticipatory postural adjustments (APAs). In the overhanging posture, the movement was performed without APAs. The results indicated that APAs were modulated according to the possibilities of force creation of the system. Hence, the disappearance of APAs in the overhanging posture was explained by the efficiency of the system to create the impulse necessary to perform the task.     

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.     

Anticipatory postural adjustments in children with hemiplegia and diplegia

Anticipatory postural adjustments (APAs) play an important role in the performance of many activities requiring the maintenance of standing posture. However, little is known about if and how children with cerebral palsy (CP) generate APAs. Two groups of children with CP (hemiplegia and diplegia) and a group of children with typical motor development performed arm flexion and extension movements while standing on a force platform. Electromyographic activity of six trunk and leg muscles and displacement of center of pressure (COP) were recorded. Children with CP were able to generate anticipatory postural adjustments and produce directionally specific APAs and COP displacements similar to those described in adults and typically developing children. However, children with diplegia were unable to generate APAs of the same magnitude as children with typical development and hemiplegia and had higher baseline muscle activity prior to movement. In children with diplegia, COP was posteriorly displaced and peak acceleration was smaller during bilateral extension compared to children with hemiplegia. The outcomes of the study highlight the role of APAs in the control of posture of children with CP and point out the similarities and differences in anticipatory control in children with diplegia and hemiplegia. These differences may foster ideas for treatment strategies to enhance APAs in children with CP.     

Force-Controlled Balance Perturbations Associated with Falls in Older People: A Prospective Cohort Study

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

Effect of exercise-induced fatigue on postural control of the knee

Muscle fatigue is associated with reduced power output and work capacity of the skeletal muscle. Fatigue-induced impairments in muscle function are believed to be a potential cause of increased injury rates during the latter stages of athletic competition and often occur during unexpected perturbations. However the effect of fatigue on functionally relevant, full body destabilizing perturbations has not been investigated. This study examines the effect of muscle fatigue on the activation of the quadriceps and hamstrings to fast, full body perturbations evoked by a moveable platform. Surface electromyographic (EMG) signals were recorded from the knee extensor (vastus medialis, rectus femoris, and vastus lateralis) and flexor muscles (biceps femoris and semitendinosus) of the right leg in nine healthy men during full body perturbations performed at baseline and immediately following high intensity exercise performed on a bicycle ergometer. In each condition, participants stood on a moveable platform during which 16 randomized postural perturbations (eight repetitions of two perturbation types: 8 cm forward slides, 8 cm backward slides) with varying inter-perturbation time intervals were performed over a period of 2-3 min. Maximal voluntary knee extension force was measured before and after the high intensity exercise protocol to confirm the presence of fatigue. Immediately after exercise, the maximal force decreased by 63% and 66% for knee extensors and flexors, respectively (P<0.0001). During the post-exercise postural perturbations, the EMG average rectified value (ARV) was significantly lower than the baseline condition for both the knee extensors (average across all muscles; baseline: 19.7±25.4μV, post exercise: 16.2±19.4 μV) and flexors (baseline: 24.3±20.9 μV, post exercise: 13.8±11.0 μV) (both P<0.05). Moreover the EMG onset was significantly delayed for both the knee extensors (baseline: 132.7±32.9 ms, post exercise: 170.8±22.9 ms) and flexors (baseline: 139.1±38.8 ms, post exercise: 179.3±50.9 ms) (both P<0.05). A significant correlation (R(2)=0.53; P<0.05) was identified between the percent reduction of knee extension MVC and the percent change in onset time of the knee extensors post exercise. This study shows that muscle fatigue induces a reduction and delay in the activation of both the quadriceps and hamstring muscles in response to rapid destabilizing perturbations potentially reducing the stability around the knee.     

Effect of local and general fatiguing exercises on disturbed and static postural control

This study compared the effect of local and general fatiguing exercise on disturbed and static postural control performances. Surface electromyography and center of pressure signals were respectively recorded during self-initiated perturbation test and static postural stability test from 7 young male subjects. Local fatiguing exercise was performed using intermittent isometric knee extensions at the level of 40% of maximal voluntary torques. General fatiguing exercise was implemented with rowing ergometer at a speed of 200 ± 5 m/min. Results of disturbed postural tests showed no significant change of anticipatory postural adjustment (APAs) organizations in individual muscles following both fatiguing exercises, but observed larger APAs coactivations in trunk and dorsal muscle pairs following local than general fatiguing exercise, and larger compensatory postural adjustments (CPAs) coactivation in dorsal muscle pair after both fatiguing exercises. In addition, the results of static postural tests indicated efficient static postural stability accompanying the down-weighting of visual input and the up-weighting of vestibular/somatosensory component following both fatiguing exercises. These findings evidenced a general compensation in the central nervous system in response to the neuromuscular deficiencies induced by local fatiguing exercise and put forward the function of sensory recalibration in maintaining postural stability under fatigue conditions.     

The role of proprioceptive information in programming of anticipatory postural components of voluntary movements

Anticipatory components of the EMG activity of the postural muscles during a voluntary movement were analyzed to find out how the CNS regulates these components in response to changes in the movement parameters and what information is used for programming these components. The fast lift of an arm in an erect posture was used as a model. The parameters of the movement were modified by varying weights held in the hand (0.5, 1.0, and 1.5 kg) and the preliminary information on these weights: lifting the hand holding a weight and lifting an unknown weight from a support in the absence of information on its value or after receiving verbal information on it. Our experiments showed that the program of maintaining an erect posture while performing a fast voluntary lift of the arm involves anticipatory adjustments of postural muscles (the soleus muscle, biceps muscle of the thigh, and sacrospinal muscle) using information on the parameters of the movement to be performed. For all these muscles, the anticipation time did not depend on either experimental conditions or the velocity of lifting the arm. The duration of the activity and its amount had different dependences on the lifted weight. The parameters of inhibition of the soleus muscle did not depend on the lifted weight, the activity of the biceps muscle of the thigh was mainly regulated by varying its amplitude, and the regulation of the sacrospinal muscle involved both amplitude and duration changes. It was shown that the adjustment of anticipatory movement components can be only based on proprioceptive rather than verbal (conscious) information.     

Anticipatory postural adjustments in the shoulder girdle in the reach movement performed in standing by post-stroke subjects

Abstract

After a stroke in middle cerebral artery territory, there is a high probability of dysfunction of the ventromedial pathways, mainly related with postural control mechanisms such as the anticipatory postural adjustments (APAs). According to neuroanatomical knowledge, these pathways have a predominant ipsilesional disposition, which justifies a bilateral postural control dysfunction, often neglected in rehabilitation. In order to assess this bilateral postural control dysfunction, electromyography activity was assessed in eight post-stroke and 10 healthy individuals in the anterior deltoids, the superior and lower trapezius, and the latissimus dorsi as they reached for a bottle with both upper limbs separately at a self-selected velocity and fast velocity while standing associated with trunk kinematics analysis. Through this analysis it was possible to compare the timing of APAs in scapular muscles between sides in post-stroke and with healthy individuals, and to verify if there is a relation between the timing and the displacement of the trunk in the temporal window of the APAs. Indeed, post-stroke individuals show a delayed activation of APAs on scapular girdle muscles on both ipsilesional and contralesional sides, which were not reflected in the trunk displacement.     

Reaction to the perturbation of the vertical posture under different conditions of standing and in the case of additional support

The reaction of equilibrium restoration in response to the perturbation of the vertical posture of a subject standing on a stable or unstable support was studied. Perturbation was induced by a sudden forward or backward shift of the support surface. In some of the experiments, the subject was holding onto a handrail suspended on a long soft belt with a fixed upper end. The results of the study showed that the reaction to support movement depended on the direction of the perturbation. The soleus muscle was activated first upon a backward movement, and the anterior tibial muscle was activated first upon a forward movement, with a latency of about 50 ms. Within 30–70 ms, bursts of activity were also detected in the respective antagonist muscles. Sudden movement of the platform caused bursts of impulses in the arm muscles even in the absence of contact with the handrail. These impulse bursts had a longer latency (80–130 ms) and probably played an auxiliary role in the process of the restoration of balance. In the case of standing on an unstable support, the latency of leg muscle activation increased. When a subject was holding onto a handrail, the intensity of impulse bursts decreased in the leg muscles and increased in the arm muscles, while the latent periods of the bursts in the arm muscles decreased. This effect proved to be still more pronounced in the case of balance maintenance on an unstable support. Thus, the change in the response to external perturbations during maintenance of the vertical posture on an unstable support demonstrates that an additional contact of the hand changes the adjustment of the posture control system.     

Work overload induced changes in fast and slow skeletal muscle myosin heavy chain gene expression

Work induced hypertrophy of the slow postural soleus and the fast phasic plantaris muscles was produced by tenotomy of the synergistic gastrocnemius muscle. Increases in weight of both muscles were associated with proportionately even larger increases in total RNA and mRNA levels. Alterations in levels of specific myosin heavy chain (MHC) isoform mRNAs were measured using the slot blot procedure with radioactively labelled oligonucleotides as probes. Type 1 MHC gene expression was unaffected in both muscles by work overload, whereas type 2a was deinduced in the soleus and type 2b was deinduced in the plantaris. The neonatal MHC gene was transiently reinduced in the plantaris.     

Biomechanical capabilities influence postural control strategies in the cat hindlimb

During postural responses to perturbations, horizontal plane forces generated by the cat hindlimb are stereotypically directed either towards or away from the animal's center of mass, independent of perturbation direction. We used a static, three-dimensional musculoskeletal model of the hindlimb to investigate possible biomechanical determinants of this "force constraint strategy." We hypothesized that directions in which the hindlimb can produce large forces are preferentially used in postural control. We computed feasible force sets (FFSs) based on hindlimb configurations of three cats during postural equilibrium tasks and compared them to horizontal plane postural force directions. The grand mean FFS was bimodal, with maxima near the posterior-anterior axis (-86+/-8 degrees and 71+/-4 degrees ), and minima near the medial-lateral axis (177+/-8 degrees and 8+/-8 degrees ). Experimental postural force directions clustered near both maxima; there were no medial postural forces near the absolute minimum. However, the medians of the anterior and posterior postural force direction histograms in the right hindlimb were rotated counter-clockwise from the FFS maxima (p<0.05; Wilcoxon signed-rank test). Because the posterior-anterior alignment of the FFS is consistent with a hindlimb structure optimized for locomotion, we conclude that the biomechanical capabilities of the hindlimb strongly influence, but do not uniquely determine the force directions observed in the force constraint strategy. Forces used in postural control may reflect a balance between a neural preference for using forces in the directions of large feasible forces and other criteria, such as the stabilization of the center of mass, and muscular coordination strategies.