Reversals of anticipatory postural adjustments during voluntary sway in humans

We describe reversals of anticipatory postural adjustments (APAs) with the phase of a voluntary cyclic whole-body sway movement. Subjects ( n = 9) held a standard load in extended arms and released it by a bilateral shoulder abduction motion in a self-paced manner at different phases of the sway. The load release task was also performed during quiet stance in three positions: in the middle of the sway range and close to its extreme forward and backward positions. Larger APAs were seen during the sway task as compared to quiet stance. Although the direction of postural perturbation associated with the load release was always the same, the direction of the APAs in the leg muscles reversed when the subjects were close to the extreme forward position as compared to the APAs in other phases and during quiet stance. The trunk muscles showed smaller APA modulation at the extreme positions but larger modulation when passing through the middle position, depending on the direction of sway, forward or backward. The phenomenon of APA reversals emphasizes the important role of safety in the generation of postural adjustments associated with voluntary movements. Based on these findings, APAs could be defined as changes in the activity of postural muscles associated with a predictable perturbation that act to provide maximal safety of the postural task component.     

Role of lateral muscles and body orientation in feedforward postural control

The study investigates the role of lateral muscles and body orientation in anticipatory postural adjustments (APAs). Subjects stood in front of an aluminum pendulum and were required to stop it with their right or left hand. An experimenter released the pendulum inducing similar body perturbations in all experimental series. The perturbation directions were manipulated by having the subjects standing on the force platform with different body orientations in relation to the pendulum movements. Consequently, perturbations were induced in sagittal, oblique, and frontal planes. Ground reaction forces and bilateral EMG activity of dorsal, ventral, and lateral trunk and leg muscles were recorded and quantified within the time intervals typical of APAs. Anticipatory postural adjustments were seen in all experimental conditions; their magnitudes depended on the body orientation in relation to the direction of perturbation. When the perturbation was produced in the lateral and oblique planes, APAs in the gluteus medius muscles were greater on the side opposite to the side of perturbation. Conversely, simultaneous anticipatory activation of the external obliques, rectus abdominis, and erector spinae muscles was observed on the side of perturbation when it was induced in the lateral plane. The results of the present study provide additional information on the directional specificity of anticipatory activation of ventral and dorsal muscles. The findings provide new data on the role of lateral muscles in feedforward postural control and stress the importance of taking into consideration their role in the control of upright posture.     

Anticipatory postural adjustments in individuals with multiple sclerosis

Individuals with multiple sclerosis (MS) frequently exhibit difficulties in balance maintenance. It is known that anticipatory postural adjustments (APAs) play an important role in postural control. However, no information exists on how people living with MS utilize APAs for control of posture. A group of individuals with MS and a group of healthy control subjects performed rapid arm flexion and extension movements while standing on a force platform. Electromyographic (EMG) activity of six trunk and leg muscles and displacement of center of pressure (COP) were recorded and quantified within the time intervals typical of APAs. Individuals with MS demonstrated diminished ability to produce directional specific patterns of anticipatory EMGs as compared to control subjects. In addition, individuals with MS demonstrated smaller magnitudes of anticipatory muscle activation. This was associated with larger displacements of the COP during the balance restoration phase. These results suggest the importance of anticipatory postural control in maintenance of vertical posture in individuals with MS. The outcome of the study could be used while developing rehabilitation strategies focused on balance restoration in individuals with MS.     

Deficits in task-specific modulation of anticipatory postural adjustments in individuals with spastic diplegic cerebral palsy

We examined whether individuals with spastic diplegic cerebral palsy (SDCP) have the ability to utilize lower leg muscles in anticipatory postural adjustments (APAs) associated with voluntary arm movement while standing, as well as the ability to modulate APAs with changes in the degree of postural perturbation caused by arm movement. Seven individuals with spastic diplegia (SDCP group, 12-22 yr of age) and seven age- and sex-matched individuals without disability (control group) participated in this study. Participants flexed both shoulders and lifted a load under two different load conditions, during which electromyographic activities of focal and postural muscles were recorded. Although the timing of anticipatory activation of the erector spinae and medial hamstring (MH) muscles was similar in the two participant groups, that of the gastrocnemius (GcM) muscle was significantly later in the SDCP group than in the control group. An increase in anticipatory postural muscle activity with an increase in load was observed in MH and GcM in the control group but not in GcM in the SDCP group. The degree of modulation in MH was significantly smaller in the SDCP group than in the control group. An additional experiment confirmed that these differences in APAs between the two participant groups were unlikely to be attributable to their differences in initial standing posture before load lift. The present findings suggest that lower leg muscles play a minor role in APAs in individuals with spastic diplegia. In addition, it is likely that these individuals have difficulty modulating anticipatory postural muscle activity with changes in the degree of postural perturbation.     

Anticipatory EMG patterns associated with preferred and non-preferred arm pointing movements

Rapid arm movements generate balance perturbations, which are anticipated and counteracted by postural adjustments. The effect of lateral preference on the control of the postural preparation to maximal velocity upperlimb pointing movements was investigated in right-handers. The muscular activities characterizing the postural adjustments were compared for preferred and non-preferred upper-limb movements. Movements were performed in two sitting conditions differing by their stability ("full on seat" and "edge of seat"). The electromyographic activity of the arm-movement prime mover and of several trunk and hip muscles involved in postural control was recorded with surface electrodes. Results indicate the presence of a reproducible pattern of anticipatory postural adjustments (APAs) involving trunk and hip muscles preceding the activation of the prime mover. In the "full on seat" condition, APAs started earlier and movement velocity was higher for preferred than for non-preferred arm movements. In the "edge of seat" condition, maximal velocity of movement did not differ significantly between both sides, but a higher excitation level of postural muscles was required to achieve this similar performance when the non-preferred upper limb was used. These findings are consistent with the hypothesis that handedness involves differences in the postural control associated with upperlimb movements, in other words that lateral preference is associated with a postural laterality.     

The role of action in postural preparation for loading and unloading in standing subjects

The main purpose of the present study has been to find an answer to the question: Can the subject generate anticipatory postural adjustments (APAs) when a predictable postural perturbation occurs in the absence of a voluntary action? Answering this question would allow us to distinguish between two competing hypotheses on the relation between APAs and voluntary movements. One hypothesis considers both APAsigma and voluntary "focal" movements different peripheral patterns associated with a single control process, while the alternative hypothesis considers them outcomes of two parallel control processes. Healthy subjects performed series of loading and unloading trials that included: (1) catching a falling load onto another load held in extended hands; (2) catching a falling load onto a tray attached to the trunk; (3) allowing a falling load to hit another load out of the extended hands, causing an unloading; and (4) releasing a load held in extended hands by a voluntary shoulder movement. In series 1, precautions were taken to avoid possible small hand movements prior to the impact of the falling load. Available visual information on the trajectory of the falling load was manipulated. In all conditions, except when the subject's eyes were closed, APAs were seen with patterns that were adequate for counteracting expected perturbations. Quantitative electromyographic indices of APAs depended on the availability of visual information and particular methods of introducing postural perturbations despite the fact that the magnitude of the perturbation was always the same. Our findings support a hypothesis that control processes resulting in APAs can be different from control processes associated with focal voluntary movements.     

Postural adjustments in arm and leg muscles associated with isodirectional and antidirectional coupling of upper limb movements in the horizontal plane

The hypothesis that anticipatory postural adjustments (APAs) may concur in generating the directional preference experienced during limb coupled movements was tested by measuring the electromyographic and mechanic postural actions elicited when moving: (1) one single arm/hand and, (2) both limbs, iso- or antidirectionally coupled. During fast adduction of the right arm in the horizontal plane (prime mover, pectoralis Major, rPM) APAs were recorded in the contralateral lPM as well as in the right ischiocruralis (rIC) muscle. This last action was associated to a transient increase of Tz (torque around body vertical axis) in the direction opposite to arm rotation. Both the APAs in rIC and the Tz changes nearly doubled in size when arms were coupled isodirectionally (adduction of one arm and abduction on the other) while they vanished when both arms were simultaneously adducted (antidirectional coupling). Conformably, during rhythmic arm oscillations APAs and Tz were cyclically modulated when movements were isodirectional, the modulation amplitude being strongly enhanced by increasing the movement frequency. When oscillations were antidirectional neither APAs nor Tz changes were observed, even if frequency was incremented. The postural actions linked to unidirectional or cyclic movements of the hand were affected by either coupling or frequency in the same way as arm movements, albeit much smaller in size. In conclusion, during antidirectional movements APAs in prime movers are synergic with voluntary activation and no postural engagement is requested to leg muscles. Conversely, during isodirectional movements, APAs in prime movers conflict with the voluntary commands and a strong, frequency-dependent, postural effort is required to leg muscles. How these factors may co-operate in determining the preference for antidirectional coupling is discussed.     

Walking delays anticipatory postural adjustments but not reaction times in a choice reaction task

During standing, anticipatory postural adjustments (APAs) and focal movements are delayed while performing a choice reaction task, compared with a simple reaction task. We hypothesized that APAs and focal movements of a choice reaction task would be similarly delayed during walking. Furthermore, reaction times are delayed during walking compared with standing. We further hypothesized that APAs and focal movements would be delayed during walking, compared with standing, for both simple and choice reaction tasks. Subjects either walked or stood on a treadmill while holding on to stable handles. They were asked to push or pull on the handles in response to a visual cue. Muscle activity was recorded from muscles of the leg (APA) and arm (RT). Our results were in agreement with previous work showing APA onset was delayed in the choice reaction task compared with the simple reaction task. In addition, the interval between the onset of APA and focal movement activity increased with choice reaction tasks. The task of walking did not delay the onset of focal movement for either the simple or choice reaction tasks. Walking did delay the onset of the APA, but only during choice reaction tasks. The results suggest the added demand of walking does not significantly modify the control of focal arm movements. However, additional attentional demands while walking may compromise anticipatory postural control.     

Modulation of anticipatory postural adjustments associated with unloading perturbation: effect of characteristics of a motor action

The purpose of the study was to determine whether characteristics of a motor action affect anticipatory postural adjustments (APAs). Standing subjects held a load between their hands with arms extended in front of their body. Next, subjects performed bilateral shoulder abduction movements (motor action) of three amplitudes at three instructed speeds. This motor action led to the release of the same load, inducing unloading perturbation in the sagittal plane. Electromyographic activities were recorded for the leg and trunk muscles. A change in the background muscle activity in these muscles was observed prior to the unloading perturbation and was quantified as APAs. APAs were dependant on instructed speed of the motor action; larger APA activities were observed in the leg and trunk muscles with a faster speed instruction. Meanwhile, the modulation of APAs was not observed by altering the movement amplitude. Moreover, experiments showed that motor action itself without a load release did not generate APA activity. Therefore, we concluded that the central nervous system selects information within a motor action (i.e., speed instruction) to approximate the magnitude of the forthcoming perturbation and modulate APAs, even when the unloading perturbation was unchanged.     

Two stages and three components of the postural preparation to action

Previous studies of postural preparation to action/perturbation have primarily focused on anticipatory postural adjustments (APAs), the changes in muscle activation levels resulting in the production of net forces and moments of force. We hypothesized that postural preparation to action consists of two stages: (1) Early postural adjustments (EPAs), seen a few hundred ms prior to an expected external perturbation and (2) APAs seen about 100 ms prior to the perturbation. We also hypothesized that each stage consists of three components, anticipatory synergy adjustments seen as changes in covariation of the magnitudes of commands to muscle groups (M-modes), changes in averaged across trials levels of muscle activation, and mechanical effects such as shifts of the center of pressure. Nine healthy participants were subjected to external perturbations created by a swinging pendulum while standing in a semi-squatting posture. Electrical activity of twelve trunk and leg muscles and displacements of the center of pressure were recorded and analyzed. Principal component analysis was used to identify four M-modes within the space of muscle activations using indices of integrated muscle activation. This analysis was performed twice, over two phases, 400–700 ms prior to the perturbation and over 200 ms just prior to the perturbation. Similar robust results were obtained using the data from both phases. An index of a multi-M-mode synergy stabilizing the center of pressure displacement was computed using the framework of the uncontrolled manifold hypothesis. The results showed high synergy indices during quiet stance. Each of the two stages started with a drop in the synergy index followed by a change in the averaged across trials activation levels in postural muscles. There was a very long electromechanical delay during the early postural adjustments and a much shorter delay during the APAs. Overall, the results support our main hypothesis on the two stages and three components of the postural preparation to action/perturbation. This is the first study to document anticipatory synergy adjustments in whole-body tasks. We interpret the results within the referent configuration hypothesis (an extension of the equilibrium-point hypothesis): The early postural adjustment is based primarily on changes in the coactivation command, while the APAs involve changes in the reciprocal command. The results fit an earlier hypothesis that whole-body movements are controlled by a neuromotor hierarchy where each level involves a few-to-many mappings organized to stabilize its overall output.     

The role of anticipatory postural adjustments (APAs) in interlimb coordination of coupled arm movements in the parasagittal plane: I. APAs associated with fast discrete flexion and extension movements of one arm or of both arms ISO- and ANTI-directionally coupled

Coupling stability during cyclic arm movements in the horizontal (transverse) plane is lower in ISO- than in ANTI-directional coupling. We proposed that such impairment arises from the interference exerted in ISO by the anticipatory postural adjustments (APAs) linked to the primary movements. To evaluate if a link between coupling stability and postural adjustments also exist for arm movements with different postural requirements, we focused on arm(s) flexion–extension in the parasagittal plane and started by analysing the APAs distribution in arm, trunk and leg muscles. Fast flexion and extension of the right arm elicited APAs in the left anterior and posterior deltoid that replicated the excitation–inhibition of the homologous prime movers; this pattern would favour ISO and contrast ANTI-coupled movements. Instead, in the left latissimus dorsi, APAs were opposite to the voluntary actions in the right latissimus dorsi, thus favouring ANTI coupling. Symmetrical APAs were also elicited in right and left erector spinae (rES, lES) and asymmetrical APAs in Ischiocruralis (rIC, lIC), while an antero-posterior force (Fy) and a moment about the vertical axis (Tz) were discharged to the ground. When fast discrete movements were ISO-coupled, APAs were symmetrical in trunk (rES, lES) and leg (rIC, lIC) muscles and a large Fy but no Tz was generated. In ANTI coupling, APAs in rES and lES remained symmetrical, whereas they became antisymmetrical in rIC and lIC. A large Tz and a small Fy were recorded. In conclusion, during parasagittal movements, APAs in are elicited in both ISO and ANTI coupling, at variance with horizontal movements where they are only present in ISO. This would suggest that the difference in coupling stability between the two modes is smaller (or even reversed) in parasagittal with respect to horizontal arm movements.     

Does the coordination between posture and movement during human whole-body reaching ensure center of mass stabilization?

The whole-body center of mass (CoM) has been classically regarded as the stabilized reference value for human voluntary movements executed upon a fixed base of support. Axial synergies (opposing displacements of head and trunk with hip segments) are believed to minimize antero-posterior (A/P) CoM displacements during forward trunk movements. It is also widely accepted that anticipatory postural adjustments (APAs) create forces of inertia that counteract disturbances arising from the moving segment(s). In the present study, we investigated CoM stabilization by axial synergies and APAs during a whole-body reaching task. Subjects reached towards an object placed on the ground in front of them in their sagittal plane using a strategy of coordinated trunk, knee, and hip flexion. The reaching task imposed constraints on arm-trajectory formation and equilibrium maintenance. To manipulate equilibrium constraints, differing conditions of distance and speed were imposed. The comparison of distance conditions suggested that axial synergies were not entirely devoted to CoM stabilization: backward A/P hip displacements reduced as head and trunk forward A/P displacements increased. Analysis of upper- and lower-body centers of mass in relation to the CoM also showed no strict minimization of A/P CoM displacements. Mechanical analysis of the effects of APAs revealed that, rather than acting to stabilize the CoM, APAs created necessary conditions for forward CoM displacement within the base of support in each condition. The results have implications for the CoM as the primary stabilized reference for posture and movement coordination during whole-body reaching and for the central control of posture and voluntary movement. Received: 14 July 1998 / Accepted: 23 May 1999     

Anticipatory postural adjustments in arm muscles associated with movements of the contralateral limb and their possible role in interlimb coordination

While sitting on a turnable stool, with both shoulders flexed at 90° or, alternatively, with arms parallel to the trunk and the elbows flexed at 90°—the hands being semisupine—subjects performed unidirectional and cyclic movements on the horizontal plane of the right arm (adduction–abduction) or hand (flexion–extension). The left arm was still, in a position symmetrical to that of the right limb and with the hand contacting a fixed support by the palmar or dorsal surface. During both unidirectional and cyclic arm or hand movements, activation of the prime mover muscles (right Pectoralis Major for arm adduction and Infraspinatus for abduction; right Flexor Carpi Radialis and Extensor Carpi Radialis for the hand movements) was accompanied by activation of the homologous muscles of the contralateral arm and inhibition of antagonists. The contralateral activities (1) regularly preceded the burst in the movement prime movers and (2) were organised in fixation chains that, exerting forces on the hand fixed support, will counterbalance the rotatory action exerted on the trunk by the primary movement. Based on these features, these activities may be classified as anticipatory postural adjustments (APAs). The observed APAs distribution is such as to favour the preferential (mirror symmetrical) coupling of upper limb movements on the horizontal plane. The possible role of these APAs in determining the different constraints experienced when performing mirror symmetrical versus isodirectional coupling is discussed.     

Coordination between postural and movement controls: effect of changes in body mass distribution on postural and focal component characteristics

Whole-body reaching movements are accomplished through a combination of anticipatory postural adjustments (APAs) and focal movements. Two different modes of central organization is usually proposed for this coordination: first, a single-process control, where the APAs and the focal movements would share a common command; second, where the APAs and the focal movements would be independently controlled through parallel commands (dual-process control). Here, we investigated which one of these modes of control could better explain the coordination between the trunk and the upper limb for standing subjects reaching for a target located beyond arm’s length. This was done evaluating the effect of changing the APAs settings on the arm movement. The APAs modification was achieved by shifting the subject’s centre of mass prior to the focal movement onset; this was done by adding an asymmetric load on either side of the head (a control condition with the load fixed centrally at the top of the head was also performed). As it changed the body mass distribution, the muscular torques and the orientation of the head inertia tensor, it is assumed that the addition of the asymmetric load led to a change in the APAs. Analyses indeed showed that both the initial head and trunk displacement towards the supporting side (during the unloading of the moving leg) were smaller when the load was fixed on the side of the supporting leg than when it was fixed on the side of the moving leg. However, changing the initial conditions, and therefore the APAs settings, had no significant effect on the path and kinematics of the focal hand movement. Therefore, subjects cancelled out the effect of the trunk motion on the hand-in-space motion through compensatory arm movements. These results support the dual-process control hypothesis for the postural and the focal components.

Perturbed step initiation in cerebellar subjects: 2. Modification of anticipatory postural adjustments

Although ataxias of stance and gait are frequent manifestations of cerebellar disease, the number of human studies examining stance or gait in cerebellar subjects is limited. In the present study, we examined whether anticipatory postural adjustments were impaired in cerebellar subjects during perturbed and unperturbed step initiation. The first aim was to show possible abnormalities in timing, force and kinematic parameters of anticipatory postural adjustments in unperturbed stepping in cerebellar subjects. Second, we examined the ability of cerebellar subjects to modify anticipatory postural adjustments associated with step initiation in response to a backward translation. Finally, we asked whether cerebellar subjects (and controls) make use of predictive knowledge of perturbation amplitude in perturbed stepping. Only few abnormalities of anticipatory postural adjustments were found in cerebellar subjects compared to controls. Both in the unperturbed and perturbed step conditions, force production as well as step length and step velocity were reduced in cerebellar subjects compared to controls, suggesting compensatory slowing. Cerebellar subjects also appeared to be less able to use predictive information of perturbation amplitude to scale anticipatory postural adjustments than control subjects. Nevertheless, in unperturbed steps, temporal parameters of anticipatory postural adjustments were preserved in cerebellar subjects. When subjects voluntarily initiated a step in response to the surface translation, both control and cerebellar subjects adapted by executing the anticipatory postural adjustments for step more rapidly. Furthermore, both control and cerebellar subjects were able to use online information regarding perturbation amplitude to scale parameters of step initiation in perturbed stepping. Overall, our findings suggest that the cerebellum is neither critical for the basic motor program underlying unperturbed step initiation nor for many adaptive changes occurring during perturbed step initiation. Like its role in predictive scaling of automatic postural responses to external perturbations, the cerebellum appears to be important for predictive adaptation of anticipatory postural adjustments during step initiation.     

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.     

Anticipatory control related to the upward propulsive force during the rising on tiptoe from an upright standing position

The purpose of the present investigation was to explore whether and how the amplitude and/or duration parameters of anticipatory postural adjustments (APAs) affect the upward propulsive force (FZamp) during rising on tiptoes from upright standing position. Seven subjects initiated the movement of rising on tiptoe from a strictly regulated initial body position at various velocities at the subjects own spontaneous pace. In this experiment, electromyographic (EMG) activities of the muscles in the tibialis anterior (TA) and soleus, the goniogram in the ankle joint, and the anteroposterior and vertical ground reaction forces were simultaneously recorded. The average amplitude and the duration of the EMG activities and ground reaction force were measured, and the displacement of the center of body mass and the center of foot pressure (CP) were calculated and reported as the mean value plus its standard deviation. FZamp correlated positively and significantly with the maximum backward CP shift, the average amplitude of the backward loading and the average EMG amplitude in the TA in the anticipation phase, whereas FZamp did not correlate positively or significantly with the anticipatory duration of the backward CP shift of the backward loading and of the EMG activity in the TA. These results suggest that the amplitude parameters of the APAs associated with rising on tiptoes are dependent on the upward propulsive force, and that APAs may be directly involved not only with compensation of postural disturbance but also with increasing performance of the focal movement.     

Target size modifies anticipatory postural adjustments and subsequent elementary arm pointing

We herein studied whether arm-pointing movements from an upright posture (i.e. Belenkii’s paradigm) toward various targets demanding a low degree of precision could influence associated anticipatory postural adjustments (APAs) and subsequent arm movements. Six subjects were asked to use their right arm to point (without finger touch) to targets of 2, 4 and 8 cm in diameter (respectively, D2, D4 and D8). APAs were measured by recording the electromyographic activity of the right anterior deltoid and biceps femoris, as well as the kinematics of the right arm. Longer APA durations and lower values for the ratio between acceleration duration and total duration of the focal movement were observed for D4 compared to D2 and D8, whereas precision was constant across all three targets. Thus, the medium target seemed to engender an optimum motor strategy for accomplishing the accuracy and velocity requirements of the task. These results emphasize that subjects build perceptual representations of their environment as well as representations of the actions to be produced. We suggest that, even in this simple movement traditionally studied from a biomechanical perspective, APAs function not only to compensate for perturbations to equilibrium, but also reflect a cognitive representation of the precision requirements of the task.     

External postural perturbations induce multiple anticipatory postural adjustments when subjects cannot pre-select their stepping foot

Previous research on human balance recovery suggests that, prior to an externally triggered postural perturbation, healthy subjects can pre-select their postural response based on the environmental context, but it is unclear whether this pre-selection includes the selection of a stepping leg when performing compensatory steps. We sought to determine how pre-selecting a stepping limb affects the compensatory steps and stability of young, healthy subjects when responding to postural perturbations. Nine healthy subjects (24–37 years of age) stepped in response to backward translations of a platform under their feet when, prior to the perturbations, the subjects either knew whether they were to step with their left or right leg to a visual target (the Predictable condition) or did not know whether to step with their left or right leg until one of two targets appeared at perturbation onset (the Unpredictable condition). The Unpredictable condition also included randomly inserted trials of toes-up rotations and backward translations without targets (catch trials). The results showed that, in the Predictable condition, the subjects consistently exhibited one anticipatory postural adjustment (APA; a lateral weight shift toward the stance limb) before stepping accurately to the target with the correct leg. In the Unpredictable condition, the subjects either (1) exhibited multiple APAs, late step onsets, and forward center-of-mass (CoM) displacements that were farther beyond their base of support, or (2) exhibited an early step with only one APA and kept their CoM closer to the base of support, but also stepped more often with the incorrect leg. Thus, when the subjects had to select a stepping leg at perturbation onset, they either became more unstable and used multiple APAs to delay stepping in order to provide enough time to select the correct stepping leg, or they stepped earlier to remain stable but often stepped with the incorrect leg. In addition, responses to catch trials in the Unpredictable condition included distorted step placements that resembled steps to anticipated targets, despite allowing the subjects to step with a leg of their choice and to a location of their choice. Lastly, the subjects’ voluntary stepping latencies to visual targets presented without perturbations were twice as long as their stepping latencies to the backward platform translations. Therefore, healthy subjects appear to pre-select their stepping limb, even when the perturbation characteristics are unpredictable, because relying on visual input provided at perturbation onset requires a delayed response that leads to greater instability.     

Anticipatory postural adjustments during self-paced and reaction-time movements

We studied the changes in the anticipatory postural adjustments (APAs), associated with dropping a load from extended arms and during fast bilateral shoulder flexion movements, when movements were performed in a self-paced manner and under a simple reaction-time instruction. The latter instruction applied time pressure and did not allow the regular pattern of APAs to be used. In particular, the following questions were asked: (1) are there changes in the relative timing of APAs under the reaction time condition; (2) are changes in the relative timing of APAs associated with changes in APAs themselves; (3) can different postural strategies be used to maintain stability under self-paced and reaction time conditions; and (4) are changes in APAs related to actual reaction time or to a change in the instruction? In particular, under reaction-time conditions, APAs occurred later in time, typically simultaneously with the initiation of the focal movement. Additional changes in electromyographic (EMG) patterns in postural muscles included an increase in the amplitude of EMG bursts and “speeding-up” some of the tri-phasic patterns in postural dorsal-ventral muscle pairs. This was accompanied by a smaller early shift of the center of pressure followed by its more rapid delayed displacement. There was considerable variability in the changes of EMG and dynamic characteristics across subjects. Some of the changes in the EMG patterns in postural muscles depended on actual reaction time, while others were related to a change in the instruction and occurred even if actual reaction times were long enough to allow for the typical self-paced APA patterns to occur. These findings can be interpreted as supporting the parallel control hypothesis for the focal movement and postural adjustments. Alternatively, they can be interpreted within a framework that implies the generation of a single control function, which is transformed into two components, one directed at the focal muscles/joints and the other directed at postural muscles/joints.