Coordination between posture and movement in a bimanual load lifting task: putative role of a medial frontal region including the supplementary motor area

Summary The aim of the present experimental series was to investigate the role of the medial frontal region including the supplementary motor area in the coordination between posture and movement in a bimanual load lifting task. The seated subject was instructed to maintain in a horizontal position one forearm (postural arm) which was loaded with a 1 kg weight. The unloading was performed either by the experimenter (imposed unloading) or by a voluntary movement of the other arm (voluntary unloading). In normal individuals, with the voluntary unloading, the movement control was accompanied by an anticipatory adjustment of the postural forearm flexor activity, which resulted in the maintenance of the forearm position despite the unloading. The anticipatory postural adjustments were impaired in 4 out of 5 patients with unilateral lesion of the SMA region; the defect was observed mainly when the postural forearm was contralateral to the lesion. No change in the anticipatory postural adjustment was observed in one patient with complete callosal section. This finding indicates that the coordination between the posture and movement in this task is not organized through callosal fibers linking the cortices on both sides but rather at a subcortical level. The anticipatory postural adjustments were abolished in two patients with spastic hemiparesis when the postural forearm was the spastic arm. It is suggested that the SMA region contralateral to the postural forearm, together with other premotor or motor areas, may select the circuits responsible for the phasic postural adjustments which are necessary to ensure postural maintenance, whereas the motor cortex contralateral to the voluntary movement controls both the movement and, via collaterals, the preselected circuits responsible for the associated postural adjustments.     

Acquisition of anticipatory postural adjustments in a bimanual load-lifting task: normal and pathological aspects

Anticipatory adjustments of forearm posture are associated with a voluntary load-lifting movement in bimanual load-lifting tasks. Three aspects of these adjustments are analyzed: their goal, their central organization, and their acquisition. The goal of the anticipatory adjustment in this task is to minimize the perturbation of forearm posture that occurs during unloading. The central organization is based on two parallel controls responsible, respectively, for the lifting movement of the moving forearm and the anticipatory postural adjustment of the postural forearm, their coordination depending on a central timing signal. The acquisition of the anticipatory postural adjustment was tested using a paradigm where the voluntary movement performed by one hand triggered, via an electronic switch, the load release of the postural forearm. It was achieved after 40–60 trials and was not graded as a function of the voluntary movement parameters, but of the disturbance of the postural arm about to occur. The learned anticipation was not transferred when, after a first acquisition session with one forearm as the postural forearm, a second learning session was performed with the other forearm as the postural forearm. The acquisition was tested in Parkinsonian and in hemiparetic patients with capsular lesion. The highest acquisition deficit was observed in hemiparetic patients, when the contralateral forearm was the postural forearm; the deficit was less important when the ipsilateral arm was postural. Surprisingly, the anticipatory postural adjustments in hemiparetic patients were rather well preserved when the natural load-lifting task was tested. These results suggest that the basal-ganglia SMA circuit and M1 premotor areas are important in the acquisition process. Received: 20 October 1998 / Accepted: 24 March 1999     

Building anticipatory postural adjustment during childhood: a kinematic and electromyographic analysis of unloading in children from 4 to 8 years of age

The present study addressed the question of how do anticipatory postural adjustments (APA) develop during childhood, in the range from 4 to 8 years, during a bimanual load-lifting task. This task required maintaining the stabilisation of the forearm position despite imposed or voluntary unloading of the forearm. Elbow angle and multiple surface EMG were recorded on the child postural forearm supporting a load. During voluntary unloading, the elbow flexion of the postural forearm was calculated as a percentage of the elbow flexion measured during the imposed situation. Improvement of the forearm stabilisation was observed mainly between 4--6 and 7--8 years of age, but the oldest children did not reach the adult level of stabilisation of the postural forearm. Moreover, a clear developmental sequence in the acquisition of APA was reported: first the selection of an efficient EMG pattern underlying the forearm stabilisation, and second the mastering of timing adjustments. In fact, regression of the co-contraction pattern was observed with age, together with selection of the adult-like reciprocal pattern. Mastering of the timing adjustments of the reciprocal pattern, characterised in adults by a well-synchronised co-ordination between onset of the flexor muscle contraction of the manual arm and onset of the flexor muscle inhibition of the postural forearm, progressively improves during development. Moreover, these results suggest that the internal representation of the consequences of unloading on the forearm stabilisation, underlying anticipatory function during a bimanual co-ordination task, slowly build up during childhood.     

TMS-responses during anticipatory postural adjustment in bimanual unloading in humans

Transcranial magnetic brain stimulation (TMS) was used to assess the influence of the corticospinal system on motor output during forearm unloading in humans. Unloading was obtained either "passively" by the experimenter, or "actively" with the subjects' own contralateral arm. Anticipatory postural adjustments consisted of changes in the activity of a forearm flexor muscle prior to active unloading of the limb and acted to stabilize the forearm position. Motor evoked potentials (MEPs) were recorded in the forearm flexor at different times during active and passive unloading, static forearm loading, and during lifting of an equivalent weight by the contralateral arm while the ipsilateral forearm was statically loaded and held stationary. In active unloading, MEP amplitude decreased with the decrease of muscle activity. Passive unloading resulted in a similar decrease of MEP as with active unloading. During stationary forearm loading, the change in MEP corresponded to the degree of loading. If during static loading the contralateral arm has lifted a separate, equivalent weight, the amplitude of MEP decreased. A possible role of direct corticospinal volley and the motor command mediated by subcortical structures in anticipatory postural adjustments is discussed.     

Acquisition of co-ordination between posture and movement in a bimanual task

The acquisition of co-ordination between posture and movement was investigated in human subjects performing a load lifting task. Sitting subjects held their left (postural) forearm in a horizontal position while supporting a 1 kg load via an electromagnet. Perturbation of the postural forearm position consisted of the load release triggered either by the experimenter (control) or by the subject voluntarily moving the other arm. In the latter case, the movement involved the elbow joint (load lifting (A), isometric force change at the wrist level (B), elbow rotation (C) and pressing a button with the wrist (D] or the fingers (grip isometric force change). We recorded the maximal amplitude and maximal velocity of the rotation of the postural forearm, the EMG of the forearm flexors on both sides and the force exerted either by the load on the postural arm or by the isometric contraction of the moving arm. The maximal forearm angular velocity after unloading was known to be related to the level of muscle contraction before unloading. 1. In the control situation, repetition of the imposed unloading test resulted in a progressive reduction in the maximal forearm rotation without any decrease in the maximal velocity. The amplitude and duration of the unloading reflex were found to increase in parallel. These results suggest that an adaptive mechanism took place which increased the gain of the unloading reflex loop and reduced the mechanical effect of the perturbation. This mechanism was found to come into play not only in the control situation but also in other paradigms where the perturbation was expected by the subjects. 2. A decrease in both maximal amplitude and velocity of forearm rotation together with a weak "anticipatory" deactivation of the forearm postural flexors was observed when the unloading was caused by an elbow movement (situations A, B, C) which indicates that a feedforward postural control took place. An interlimb coordination was built up and stabilized after 40-60 trials. Pressing a button with the wrist (weak force and displacement) was a less effective means of inducing an anticipatory control of the flexors of the postural forearm, which indicates that the intensity of the central control plays a role in the building up of the coordination. 3. A distal grip action exerting either weak (100 g) or a high (1 kg) force was able to reduce the maximal amplitude of the forearm rotation, but not the maximal velocity, which indicates that an improved reflex action takes place, but not a feedforward anticipatory postural control.(ABSTRACT TRUNCATED AT 400 WORDS)     

Cerebellar involvement in anticipating the consequences of self-produced actions during bimanual movements

Anticipatory postural adjustments (APA) during bimanual actions can be observed when participants hold an object in one hand and then lift it with the other hand. The postural force used to hold the object is reduced in anticipation of unloading, indicating an accurate prediction of the change in load. We examined patients with unilateral or bilateral cerebellar damage as well as two individuals lacking the corpus callosum on the bimanual unloading task. The acallosal patients showed an intact APA, suggesting subcortical integration of motor signals for anticipatory adjustments during bimanual actions. Contrary to the hypothesis that the cerebellum is critical for predicting and compensating for the consequences of our actions, we found that the well-learned APA in this task was largely intact in cerebellar patients. However, cerebellar damage abolished short-term adaptation of the APA, and the patients were unable to acquire an APA in a similar but previously untrained situation. These results indicate that while over-learned anticipatory adjustments are preserved after cerebellar lesions, adaptation of this response and the acquisition of a novel coordination requires the cerebellum ipsilateral to the postural hand. Furthermore, this structure appears to be essential for the accurate timing of previously learned behaviors. The patients with cerebellar damage showed poorly timed adjustments with the APA beginning earlier than in healthy participants.     

Brain motor system function in a patient with complete spinal cord injury following extensive brain–computer interface training

Anticipatory postural adjustment (APA) during bimanual action is observed when participants hold an object in one hand and then lift that object with the other hand. The decrease in activity of a forearm flexor muscle prior to an active forearm unloading acts to stabilize the forearm position. Recent studies have investigated the influence of the corticospinal system on muscle activity during APA through transcranial magnetic stimulation. It was shown that at different times during APA, the amplitude of motor-evoked potentials in the forearm flexors decreased in conjunction with the decrease of muscle activity. If the unloading is triggered via an electromagnet by lifting an equal weight by the other arm, the anticipatory postural adjustment is learned through the repetition of unloading (three series of 20 trials). Using the transcranial magnetic stimulation, we examined changes in the motor-evoked potential in the forearm flexors before and after APA learning. Motor-evoked potential amplitude did not significantly change as forearm flexor activity decreased. The motor-evoked potential/background electromyogram ratio, however, increased in the final learning session in comparison to the initial learning session and stationary loading. The present results corroborate a hypothesis on the fundamental role of the motor cortex in the suppression of synergies that interfere with the execution of the new coordination in the process of motor learning.     

Scaling anticipatory postural adjustments dependent on confidence of load estimation in a bi-manual whole-body lifting task

Anticipatory control of motor output enables fast and fluent execution of movement. This applies also to motor tasks in which the performance of movement brings about a disturbance to balance that is not completely predictable. For example, in bi-manual lifting the pick-up of a load causes a forward shift of the centre of mass with consequent disturbance of posture. Anticipatory postural adjustments are scaled to the expected magnitude of the perturbation and are initiated well before the availability of sensory information characterising the full nature of the postural disturbance. However, when the postural disturbance unexpectedly changes, the anticipatory adjustment of joint torques is not equilibrated and may result in a disturbance to balance. In a previous study, it was demonstrated that apart from anticipatory postural adjustments, corrective responses after load pick-up are used to further compensate the postural disturbance. In this study it was examined whether the central nervous system (CNS) assembles a strategy that incorporates both anticipatory control and corrective responses, in which the magnitude of the anticipatory postural adjustments depends on the perceived level of predictability of the postural disturbance. Subjects performed series of lifts in which the magnitude of the load was never revealed to the subject. Two boxes equal in size and colour, but different in mass (6 and 16 kg), were used. Differences in expectation were created by several lifts with the 16-kg load before the 6-kg box was presented. It was observed that the number of strong corrective responses (stepping) varied with the number of 16-kg trials that formed the prior experience when the final 6-kg trial was presented. The follow-up question was whether control relied more on anticipation in the stepping trials, compared with trials in which such gross signs of imbalance were absent. In this study it was shown that subjects when stepping (i) exhibited differential anticipatory postural adjustments in comparison with 6-kg trials in which expectation was not shaped by preceding 16-kg trials, and (ii) scaled the anticipatory postural adjustments similar to those preceding lift-off of the 16-kg trial preceding it. These findings emphasise the programmed nature of the anticipatory postural adjustments and the ability of the CNS to selectively tune the anticipatory postural adjustments to stored information gained during the previous lift(s). Received: 30 October 1995 / Accepted: 10 September 1997     

Development of anticipatory postural adjustments in a bimanual load-lifting task in children

Anticipatory postural adjustments (APA) are needed to perform a movement without perturbing posture. We investigated the development of APA in 3- to 4-year-old children during a bimanual load-lifting task. The task required maintaining a stable elbow position despite imposed or voluntary unloading of the forearm. Although children can compensate the consequences of unloading by using APA, their performance did not reach an adults' level. In addition, children showed high intra-individual variability in the voluntary situation, revealed by the coexistence of both adult-like and immature patterns in kinematic and electromyographic data. In conclusion, the present study reports that APA, associated with a bimanual load-lifting task, are still being set up in 3- to 4-year-old-children. The intra-individual variability should decrease with age and be associated with a progressive mastering of the timing parameters characterizing APA.     

Anticipatory postural adjustment before bimanual unloading reactions: The role of the motor cortex in motor learning

The role of the motor cortex in forming a learned coordination (stabilization of the forearm on unloading) was studied in humans. Subjects maintained a 1-kg weight with the right (postural) forearm, the weight being attached via an electromagnet. Unloading of the postural arm was initiated by the subjects by lifting a similar load with the left arm. In control experiments, lifting of the load did not lead to unloading of the postural arm. Changes in motor cortex excitability were studied by transcranial magnetic stimulation applied to the representation area of the right biceps muscle in the motor cortex at the beginning and end of the experiments. Repeated unloading tests showed progressive decreases in the amplitude of the movement of the unloaded forearm, which were accompanied by increases in the anticipatory inhibition of the electromyogram of the biceps muscle of the unloaded arm (learning). Muscle responses to transcranial magnetic stimulation during the learning process showed no significant changes. Analysis of normalized muscle responses to transcranial magnetic stimulation (response/baseline) showed that these increased at the end of training and reached a significantly higher level than seen at the beginning of training. These results lead to the conclusion that the motor cortex plays a fundamental role in inhibiting synergies and coordinations which would interfere with the formation of the new coordination during motor learning. __________ Translated from Zhurnal Vysshei Nervnoi Deyatel’nosti imeni I. P. Pavlova, Vol. 56, No. 5, pp. 603–610, September–October, 2006.     

Developmental sequence in the acquisition of anticipation during a new co-ordination in a bimanual load-lifting task in children

The aim of this experiment was to test the capacity of children to develop anticipation at various stages of their growth. We studied the learning process of an artificial co-ordination by means of a procedure of double unloading in two groups of children, and a group of adults. Elbow-joint angle measurement was used to assess for the improvement of the forearm stabilization through the six learning sessions. The main results report a developmental sequence of the acquisition of anticipatory postural adjustments (APA) similar to the one established for the ontogenesis of APA. A difference in learning dynamics, but also in the final level of performance, is reported between children aged 5–6 and children aged 7–8 years. The improvement occurring at the age of 7 years probably indicates maturation of internal representations, favourable to the faster setting up of operational anticipated control.     

The build-up of anticipatory behaviour An analysis of the development of gait initiation in children

This study analyses the anticipatory postural adjustments during the gait initiation process in children aged 2.5, 4, 6 and 8 years. In adults, anticipation during gait initiation includes a shift in the centre of foot pressure (CP) both backwards and towards the stepping foot. Backward displacement and the duration of the anticipation phase covary with the gait progression velocity reached by the subject at the end of the first step. In the present study, the children walked on a force plate that allowed us to calculate the acceleration of the centre of mass and the displacements of the CP. The results showed three main characteristics of the development of anticipatory behaviour: (1) The occurrence of anticipatory displacements of the CP increased progressively with age. Systematic backward anticipation was found for all children except one of the youngest, whereas the lateral displacement was systematically observed later, in the 6-year group; (2) the amplitude of the spatial parameters showed a significant increase with age; (3) contrary to the adult, the amplitude of the backward shift did not covary with the forthcoming velocity in the youngest groups. This covariation became significant at 6 years and remained significant at 8 years. The results showed that even if anticipatory behaviour was present in 2.5-year-old children it is only later that the child is able of more accurate tuning of feedforward control, probably due to better control of the overall postural adjustments. Received: 2 April 1997 / Accepted: 4 November 1997     

The role of motor action in anticipatory postural adjustments studied with self-induced and externally triggered perturbations

This study investigated the relation between the magnitude of a motor action triggering a postural perturbation and the magnitude of anticipatory postural adjustments. Subjects stood on a force platform and held, in extended arms, a balloon with a 2.2-kg load suspended on a rigid cord. In different series, unloadings were induced by fast bilateral shoulder abduction movements, by popping the balloon with a tack taped to the subject's right middle finger, or by the experimenter popping the balloon. Anticipatory postural adjustments were seen during all self-initiated unloadings as changes in the level of activation of postural muscles and in displacements of the center of pressure. However, absolute values of these changes were significantly smaller in the series with balloon popping as compared to the series with shoulder abductions. Such reactions were absent when the unloading was triggered by the experimenter. We conclude that a self-triggered perturbation is always associated with anticipatory postural adjustments, while the magnitude of the adjustments may be scaled with respect to the magnitude of a motor action used to induce the perturbation.     

Feedforward postural muscle modes and multi-mode coordination in mild cerebellar ataxia

The purpose of this study was to investigate postural muscle synergies (M-modes) and quantitative multi-mode coordination to ensure reproducible center of pressure (COP) in anterior–posterior trajectories associated with voluntary-induced perturbations in patients with mild cerebellar ataxia. We applied the framework of the uncontrolled manifold hypothesis for the patients with ataxia. Nine patients diagnosed with spinocerebellar degeneration (SCD) and nine healthy adults stood on a force plate performed the voluntary unloading task. Ground reaction forces and surface electromyogram signals of ten trunk and leg muscles were recorded. Total variance of the first three principal components in the SCD group was similar to the control group. The co-contraction M-modes, uniting muscle pairs with opposing actions at major leg joints, were observed more frequently in the SCD group than in the control group during anticipatory postural adjustments. The quantitative multi-mode coordinations to ensure stable COP trajectories prior to and after motor actions were smaller in the SCD group than in the control group. We conclude that individuals with mild cerebellar ataxia organize feedforward muscle modes and show more co-contraction modes and impaired coordination during feedback and feedforward postural control.     

Coordination between posture and movement in a bimanual load-lifting task: is there a transfer?

The present experimental series was designed to test the possibility that an anticipatory postural adjustment learned during the performance of a bimanual load lifting task may be transferred between the upper extremities. Eight seated subjects were asked to maintain horizontally one forearm (postural arm) loaded with a 1kg load, which was fixed to the arm by means of an electromagnet. The unloading was triggered either by the experimenter pressing a switch (control) or by the subjects making a voluntary movement with their other arm (moving arm). In the latter case, the subject lifted a 1-kg load resting on a force platform with the moving hand, and the switching off was triggered when the force level reached a threshold of 0.5 kg. The maximum amplitude (MA) and the maximum velocity (MV) of the postural forearm elbow joint rotation occuring after the unloading were measured at each trial. The learning process was estimated by performing a regression analysis on each series of trials, using an exponential model, and from the intercept of the regression curve with the ordinate. 1. During the original learning session (three series of 20 trials), a decrease in MA and MV was found to occur both within the series and between the series during a session. 2. After the initial learning session, the sides of the postural and moving arm were interchanged to test whether any transfer had occurred. The first series of trials in the second session (transfer) and the last series of trials in the original learning session were compared and found to be significantly different in terms of the intercept (seven subjects in the case of MA, five subjects in the case of MV) and the slope (five subjects), indicating a lack of transfer. 3. The data recorded during the second transfer learning session indicated that learning occurred in all eight subjects in the case of MA and in six subjects in the case of MV. It was observed that the original learning session did not facilitate the second one. 4. The lack of transfer of the anticipatory postural adjustment observed in this task is discussed with reference to the data in the literature.     

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.     

Feedforward versus feedback control in children and adults subjected to a postural disturbance

 Any action performed by standing subjects is generally accompanied by compensatory postural activities, which reduce or abolish the postural disturbance generated by the movements and keep the subjects’ center of gravity within the supporting base. These postural activities are triggered by either anticipatory and/or feedback-based control processes, depending on the information available and on the behavioral context. To investigate the respective involvement of these two components in postural control during development, we studied the extent to which the postural equilibrium of children (3- to 10-year-olds) and adults was disturbed by the same physical event, an unloading, depending on whether it was initiated by the subject or externally imposed. The subjects were standing on a force platform with their eyes closed, holding a load (5% of their own body weight) in their hands, with arms vertical and forearms horizontal. Two conditions were applied: (1) the subjects voluntarily released the load and (2) the load was unpredictably removed. The unloading resulted in a backward movement of the center of pressure, which was smaller with self-initiated than imposed disturbances in all age groups. This difference varied depending mainly on the age-related changes in the relative amplitude of the self-initiated disturbance, which decreased between 3- to 5-, and 6- to 8-year-olds (who showed no marked postural instability after self-initiated unloading), and increased again in the two older groups (9- to 10-year-olds and adults), in which it also became more consistent . It was concluded that feedforward control becomes more efficient as children grow up, but that its relative contribution to postural control does not show a monotonic pattern of development. Received: 20 December 1997 / Accepted: 9 October 1998     

Anticipatory adjustments of multi-finger synergies in preparation for self-triggered perturbations

We studied changes in multi-finger synergies associated with predictable and unpredictable force perturbations applied to a finger during a multi-finger constant total force production task. The main hypothesis was that indices of multi-finger synergies can show anticipatory changes in preparation for a predictable perturbation. Subjects sat in a chair and pressed on force sensors with the four fingers of the right hand. The task was to produce a constant level of total force. The fingers acted against loads that produced upward directed forces. The loads (applied either to the index or to the ring finger) could be disengaged either by the subject or by the experimenter. An index of finger co-variation, ΔV was computed across sets of 12 trials at each time sample and for all tasks separately. During steady-state force production, all subjects showed positive ΔV values corresponding to strong negative covariation among finger forces interpreted as a force-stabilizing synergy. Prior to self-triggered unloading, subjects showed an anticipatory drop in ΔV that started 100–125 ms prior to the unloading time. Such early changes were absent in trials with experimenter-triggered unloading. After an unloading, subjects changed forces of both perturbed and unperturbed fingers and reached a new sharing pattern of the total force. In experimenter-triggered conditions, changes in the forces of unperturbed fingers could be seen as early as 120 ms following an unloading. The index ΔV dropped following a perturbation and then recovered; the recovery occurred faster in self-triggered conditions. We conclude that humans can use feed-forward changes in multi-finger synergies (anticipatory synergy adjustments) in anticipation of a predictable perturbation. These changes may help avoid prolonged weakening of a multi-digit force-stabilizing synergy. We discuss a possibility that anticipatory postural adjustments may represent a particular case of the phenomenon of anticipatory synergy adjustments and suggest a hierarchical control scheme that incorporates a possibility of independent control over the output of a multi-element system and covariation patterns among outputs of its elements.     

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.     

Influence of central set on anticipatory and triggered grip-force adjustments

The effects of predictability of load magnitude on anticipatory and triggered grip-force adjustments were studied as nine normal subjects used a precision grip to lift, hold, and replace an instrumented test object. Experience with a predictable stimulus has been shown to enhance magnitude scaling of triggered postural responses to different amplitudes of perturbations. However, this phenomenon, known as a central-set effect, has not been tested systematically for grip-force responses in the hand. In our study, predictability was manipulated by applying load perturbations of different magnitudes to the test object under conditions in which the upcoming load magnitude was presented repeatedly or under conditions in which the load magnitudes were presented randomly, each with two different pre-load grip conditions (unconstrained and constrained). In constrained conditions, initial grip forces were maintained near the minimum level necessary to prevent pre-loaded object slippage, while in unconstrained conditions, no initial grip force restrictions were imposed. The effect of predictable (blocked) and unpredictable (random) load presentations on scaling of anticipatory and triggered grip responses was tested by comparing the slopes of linear regressions between the imposed load and grip response magnitude. Anticipatory and triggered grip force responses were scaled to load magnitude in all conditions. However, regardless of pre-load grip force constraint, the gains (slopes) of grip responses relative to load magnitudes were greater when the magnitude of the upcoming load was predictable than when the load increase was unpredictable. In addition, a central-set effect was evidenced by the fewer number of drop trials in the predictable relative to unpredictable load conditions. Pre-load grip forces showed the greatest set effects. However, grip responses showed larger set effects, based on prediction, when pre-load grip force was constrained to lower levels. These results suggest that anticipatory processes pertaining to load magnitude permit the response gain of both voluntary and triggered rapid grip force adjustments to be set, at least partially, prior to perturbation onset. Comparison of anticipatory set effects for reactive torque and lower extremity EMG postural responses triggered by surface translation perturbations suggests a more general rule governing anticipatory processes.