Double-Step Paradigm in Microgravity: Preservation of Sensorimotor Flexibility in Altered Gravitational Force Field.

The way we can correct our ongoing movements to sudden and unforeseen perturbations is key to our ability to rapidly adjust our behavior to novel environmental demands. Referred to as sensorimotor flexibility, this ability can be assessed by the double-step paradigm in which participants must correct their ongoing arm movements to reach targets that unexpectedly change location (i.e., target jump). While this type of corrections has been demonstrated in normogravity in the extent of reasonable spatiotemporal constraints underpinning the target jumps, less is known about sensorimotor flexibility in altered gravitational force fields. We thus aimed to assess sensorimotor flexibility by comparing online arm pointing corrections observed during microgravity episodes of parabolic flights with normogravity standards. Seven participants were asked to point as fast and as accurately as possible toward one of two visual targets with their right index finger. The targets were aligned vertically in the mid-sagittal plane and were separated by 10 cm. In 20% of the trials, the initially illuminated lower target was switched off at movement onset while the upper target was concomitantly switched on prompting participants to change the trajectory of their ongoing movements. Results showed that, both in normogravity and microgravity, participants successfully performed the pointing task including when the target jumped unexpectedly (i.e., comparable success rate). Most importantly, no significant difference was found in target jump trials regarding arm kinematics between both gravitational environments, neither in terms of peak velocity, relative deceleration duration, peak acceleration or time to peak acceleration. Using inverse dynamics based on experimental and anthropometrical data, we demonstrated that the shoulder torques for accelerating and decelerating the vertical arm movements substantially differed between microgravity and normogravity. Our data therefore highlight the capacity of the central nervous system to perform very fast neuromuscular adjustments that are adapted to the gravitational constraints. We discuss our findings by considering the contribution of feedforward and feedback mechanisms in the online control of arm pointing movements.

Effect of gravity-like torque on goal-directed arm movements in microgravity.

Gravitational force level is well-known to influence arm motor control. Specifically, hyper- or microgravity environments drastically change pointing accuracy and kinematics, particularly during initial exposure. These modifications are thought to partly reflect impairment in arm position sense. Here we investigated whether applying normogravitational constraints at joint level during microgravity episodes of parabolic flights could restore movement accuracy equivalent to that observed on Earth. Subjects with eyes closed performed arm reaching movements toward predefined sagittal angular positions in four environment conditions: normogravity, hypergravity, microgravity, and microgravity with elastic bands attached to the arm to mimic gravity-like torque at the shoulder joint. We found that subjects overshot and undershot the target orientations in hypergravity and microgravity, respectively, relative to a normogravity baseline. Strikingly, adding gravity-like torque prior to and during movements performed in microgravity allowed subjects to be as accurate as in normogravity. In the former condition, arm movement kinematics, as notably illustrated by the relative time to peak velocity, were also unchanged relative to normogravity, whereas significant modifications were found in hyper- and microgravity. Overall, these results suggest that arm motor planning and control are tuned with respect to gravitational information issued from joint torque, which presumably enhances arm position sense and activates internal models optimally adapted to the gravitoinertial environment.

Online control of anticipated postural adjustments in step initiation: evidence from behavioral and computational approaches.

Anticipatory postural adjustments (APAs) prior to step execution are thought to be immutable once released. Here we challenge this assumption by testing whether APAs can be modified online if a body perturbation occurs during execution. Two directions of perturbation (resisting and assisting) relative to the body weight transfer were used during the execution of APAs. We found that APAs are modified online (increase in both ground pressure and muscle activity) to compensate for resisting perturbations. The outcomes of a biomechanical model confirmed that the early changes in the APAs resulted from an active control of the APAs and were not merely mechanical consequences of the perturbation. However, no modification of the initial feedforward command was observed for assisting perturbations. The motor command changes for the resisting perturbation may originate from the mismatch between passively originated forces and those actively specified by the central command when acting in the opposite direction. The absence of a mismatch in the assisting perturbation might explain why the central nervous system was not prompted to modify the APAs in this condition.

Can prepared anticipatory postural adjustments be updated by proprioception?

Stepping over an obstacle is preceded by a center of pressure (CoP) shift, termed anticipatory postural adjustments (APAs). It provides an acceleration of the center of mass forward and laterally prior to step initiation. The APAs are characterized in the lateral direction by a force exerted by the moving leg onto the ground, followed by an unloading of the stepping leg and completed by an adjustment corresponding to a slow CoP shift toward the supporting foot. While the importance of sensory information in the setting of the APAs is undisputed, it is currently unknown whether sensory information can also be used online to modify the feedforward command of the APAs. The purpose of this study was to investigate how the CNS modulates the APAs when a modification of proprioceptive information (Ia) occurs before or during the initiation of the stepping movement. We used the vibration of ankle muscles acting in the lateral direction to induce modification of the afferent inflow. Subjects learned to step over an obstacle, eyes closed, in synchrony to a tone signal. When vibration was applied during the initiation of the APAs, no change in the early APAs was observed except in the case of a cutaneous stimulation (low frequency vibration); it is thus possible that the CNS relies less on proprioceptive information during this early phase. Only the final adjustment of the CoP seems to take into account the biased proprioceptive information. When vibration was applied well before the APAs onset, a postural reaction toward the side of the vibration was produced. When subjects voluntarily initiated a step after the postural reaction, the thrust amplitude was set according to the direction of the postural reaction. This suggests that the planned motor command of the APAs can be updated online before they are triggered.

Behavioral outcomes following below-knee amputation in the coordination between balance and leg movement.

Lateral leg movement is accompanied by opposite movements of the supporting leg and trunk segments. This kinematic synergy shifts the center of mass (CM) towards the supporting foot and stabilizes its final position, while the leg movement is being performed. The aim of the present study was to provide insight in the behavioral substitution process responsible for the performance of this kinematic synergy. The kinematic synergy was assessed by the principal component analysis (PCA) applied to both hip joints and supporting ankle joint. Patients after unilateral below-knee amputation and control subjects were asked to perform a lateral leg raising. The first principal component (PC(1)) accounted for more than 99% of the total angular variance for all subjects (amputees and controls). PC(1) thus well represents the possibility to describe this complex multi-joint movement as a one degree of freedom movement with fixed ratios between joint angular time course. In control subjects, the time covariation between joints changes holds during all phases of the leg movement (postural phase, ascending and braking phases). In amputees, PC(1) score decreased during the ascending phase of the movement (i.e. when the body weight transfer is completed, while the movement is initiated). We conclude that a feedback mechanism is involved and discuss the hypothesis that this inter-joint coordination in amputees results from a failure in the pre-setting of the inter-joint coupling.

Sensori-motor adaptation to knee osteoarthritis during stepping-down before and after total knee replacement.

BACKGROUND: Stepping-down is preceded by a shift of the center of mass towards the supporting side and forward. The ability to control both balance and lower limb movement was investigated in knee osteoarthritis patients before and after surgery. It was hypothesized that pain rather than knee joint mobility affects the coordination between balance and movement control. METHODS: The experiment was performed with 25 adult individuals. Eleven were osteoarthritic patients with damage restricted to one lower limb (8 right leg and 3 left leg). Subjects were recruited within two weeks before total knee replacement by the same orthopedic surgeon using the same prosthesis and technics of surgery. Osteoarthritic patients were tested before total knee replacement (pre-surgery session) and then, 9 of the 11 patients were tested one year after the surgery when re-educative training was completed (post-surgery session). 14 adult individuals (men: n = 7 and women: n = 7) were tested as the control group. RESULTS: The way in which the center of mass shift forward and toward the supporting side is initiated (timing and amplitude) did not vary within patients before and after surgery. In addition knee joint range of motion of the leading leg remained close to normal before and after surgery. However, the relative timing between both postural and movement phases was modified for the osteoarthritis supporting leg (unusual strategy for stepping-down) before surgery. The "coordinated" control of balance and movement turned to be a "sequential" mode of control; once the body weight transfer has been completed, the movement onset is triggered. This strategy could be aimed at shortening the duration-time supporting on the painful limb. However no such compensatory response was observed. CONCLUSION: The change in the strategy used when supporting on the arthritis and painful limb could result from the action of nociceptors that lead to increased proprioceptor thresholds, thus gating the proprioceptive inputs that may be the critical afferents in controlling the timing of the coordination between balance and movement initiation control.

Anticipatory balance control is affected by loadless training experiences.

The main purpose of this study was to identify whether a lot of sports training had any effect on the balance control associated with a leg movement. The nature of the training experience was also an important concern and we chose subject who had undergone specific training experience in absence of equilibrium constraints. To this end a comparison between control (untrained) subjects, triathletes and swimmers was designed to establish whether a general training in sports (triathletes) or a specific loadless training (swimmers), leads to differences in the balance control. A leg movement is preceded by a shift of the center of mass (CM) towards the supporting side to maintain equilibrium and forward to create the condition for progression. To provide an acceleration of the CM sideward and forward, an initial displacement of the center of pressure (CP) towards the moving limb and in posterior direction was performed. Interestingly, the lateral pressure onto the ground was greater increased in swimmers in both leg raising and obstacle avoidance tasks compared to the control group and/or triathletes whereas the backward CP shift in all group was the same. The initial control of the CM shift is very different in swimmers compared to triathletes and controls. The increased lateral pressure onto the ground in swimmers may be a result of a prolonged training in water. This suggests that prolonged training in the absence of equilibrium constraints has more of an effect on balance control than a prolonged general training. In addition, the lack of differences in the backward CP shift suggests that M/L and A/P controls support two functional goals: equilibrium maintenance and movement initiation.

Is perception of upper body orientation based on the inertia tensor? Normogravity versus microgravity conditions.

During lateral leg raising, a synergistic inclination of the supporting leg and trunk in the opposite direction to the leg movement is performed in order to preserve equilibrium. As first hypothesized by Pagano and Turvey (J Exp Psychol Hum Percept Perform, 1995, 21:1070-1087), the perception of limb orientation could be based on the orientation of the limb's inertia tensor. The purpose of this study was thus to explore whether the final upper body orientation (trunk inclination relative to vertical) depends on changes in the trunk inertia tensor. We imposed a loading condition, with total mass of 4 kg added to the subject's trunk in either a symmetrical or asymmetrical configuration. This changed the orientation of the trunk inertia tensor while keeping the total trunk mass constant. In order to separate any effects of the inertia tensor from the effects of gravitational torque, the experiment was carried out in normo- and microgravity. The results indicated that in normogravity the same final upper body orientation was maintained irrespective of the loading condition. In microgravity, regardless of loading conditions the same (but different from the normogravity) orientation of the upper body was achieved through different joint organizations: two joints (the hip and ankle joints of the supporting leg) in the asymmetrical loading condition, and one (hip) in the symmetrical loading condition. In order to determine whether the different orientations of the inertia tensor were perceived during the movement, the interjoint coordination was quantified by performing a principal components analysis (PCA) on the supporting and moving hips and on the supporting ankle joints. It was expected that different loading conditions would modify the principal component of the PCA. In normogravity, asymmetrical loading decreased the coupling between joints, while in microgravity a strong coupling was preserved whatever the loading condition. It was concluded that the trunk inertia tensor did not play a role during the lateral leg raising task because in spite of the absence of gravitational torque the final upper body orientation and the interjoint coupling were not influenced.

[Biomechanical consequences of a knee osteoarthritis on the opposite lower limb]. / Conséquences biomécaniques d'une gonarthrose unilatérale sur le membre inférieur opposé

UNLABELLED: The aim of this work was to study the compensatory strategies built up by patients with unilateral knee arthritis during stair descent. These compensatory strategies might induce increased biomechanical constraints on the unaffected knee. METHOD: A kinetic and kinematic analysis was performed in 11 patients with unilateral knee arthritis and in 14 control subjects using an ELITE system and two force-plates. The peak of vertical ground reaction forces when landing on the reception force-plate, the time to reach the peak and the duration of the different phases of the movement were studied during stair descent. RESULTS: The peak of vertical ground reaction forces was more important when landing on the unaffected limb than when landing on the affected limb. The time to reach this peak was longer in patients than in controls no matter which side was supporting. The duration of the single support phase was longer on the unaffected limb than on the affected limb. DISCUSSION AND CONCLUSION: This work has shown that patients with unilateral knee arthritis develop new strategies during stair descent. These new strategies imply increased biomechanical constraints on the unaffected limb and might favor arthritis on the sound side. These results support the idea that rehabilitation protocols of patients with unilateral knee arthritis should also involve the unaffected limb.

Interaction between attention demanding motor and cognitive tasks and static postural stability.

BACKGROUND: Due to the often-reported decrease in postural stability in the elderly, it is important to understand factors that may contribute to reduced postural stability. It is possible that attention-demanding focal tasks performed concurrent with postural regulation influence postural stability. OBJECTIVE: This study utilized dual-task methodology to determine if motor or cognitive focal tasks interact with center of pressure (COP) excursion during static bipedal stance in healthy young and healthy elderly subjects (n = 18). METHODS: The cognitive task involved silently solving an orally-presented multi-step arithmetic problem over a 30-second period. The motor task was a 30-second bilateral static finger-thumb pinch task performed at 10% of maximal voluntary contraction with a pair of pinch-force transducers. Each focal task was performed separately, and in a condition in which both tasks were performed simultaneously. COP excursion was compared in quiet standing (no focal task) and during performance of the focal tasks with full vision and with vision occluded. RESULTS: Performance on the focal tasks was unaffected by increased postural demands during stance as compared to a seated baseline condition. This was the case for both age groups, and for the full vision and occluded vision conditions. Medio-lateral COP excursion was reduced over the quiet standing pretest condition when attentional focus was on the cognitive task, suggesting that COP was influenced centrally during cognition. In contrast, COP excursion increased over the quiet standing pretest condition when performing the motor focal task, suggesting a reduced ability to suppress sway when the motor system was concurrently occupied with a voluntary task that shared the same input-output resources. CONCLUSION: The ability to share attentional resources among focal and postural tasks was similar in healthy young and elderly subjects.

Equilibrium and movement control strategies in trans-tibial amputees.

This study was aimed at identifying changes in equilibrium and movement control strategies in trans-tibial amputees (TTA) related to both the biomechanical changes and the loss of afferent inflow. The coordinations between equilibrium and movement were studied in traumatical TTA and in controls during transition from bipedal to monopodal stance. TTA failed to perform the task in a high percentage of trials both when the sound and the prosthetic limb were supporting. Significant differences were also found between TTA and controls in the duration of the weight transfer phase, in the length of the initial centre of pressure (CP) displacement and in the electromyographic (EMG) patterns. Despite adaptive posturomotor control strategies, transition from bipedal to monopodal stance remains a difficult task to perform for TTA, both when the supporting limb is the affected one and when the sound one is. The results of this study are discussed with respect to the rehabilitation programme and the prosthesis design for transtibial amputees.

Body orientation and control of coordinated movements in microgravity.

The present paper focuses on the organization of posture and movement under normal and microgravity conditions. Two reference values subserving the control of erect posture and the performance of movements are analyzed. The first is 'geometrical' in nature and corresponds to the orientation of a body segment with respect to the external world. The second reference value, which involves the mass and inertia of the body segments, is the position of the centre of mass with respect to the foot support area. The reorganization of these parameters which occurs under microgravity is discussed in the framework of a hierarchical model of posture. Suggestions are made for training procedures which could be used to prevent loss of balance from occurring in astronauts on landing after long space flights.

Postural reorganization of weight-shifting in below-knee amputees during leg raising.

The position of the center of gravity (CG) is a reference value that is controlled by the nervous system during the performance of movements. In order to maintain equilibrium, leg movement is preceded by a shift of the CG towards the supporting side. This CG shift is initiated by an early displacement of the center of pressure (CP) towards the moving leg. This characteristic CP thrust partly results from the activity of a distal muscle in the leg to be moved: the gastrocnemius medialis (GM). The aim of this study was to determine how this weight-shifting is initiated when the distal muscles are missing, as in amputees, and to identify any change in the central command. Experiments were performed on ten subjects: five below-knee amputees with no pathology and five control subjects. While standing, the subjects were instructed to raise one leg laterally as fast as possible to an angle of 45 degrees and to maintain the final position. The same weight-shifting strategy was used by both groups, whereas local adaptations associated with the behavior occurred. When the GM is lacking, an early tensor-fasciae-latae (TFL) burst is observed just prior to and associated with the onset of the lateral CP change. This moving-leg abductor may be responsible for initiating the thrust at a proximal level when that leg is still on the ground. In addition, upon analyzing the lateral displacement of the CP, two modes of CP shift were detected. The first CP-shift mode has been previously described and the second mode (which we term here the pre-pushing mode) was used by both amputees and controls. The prepushing mode consisted of two thrusts: an early thrust onto the ground was exerted by the leg about to become the supporting leg followed by the previously described thrust exerted by the leg about to be raised. The early thrust, which could be exerted by either the sound or prosthetic leg, may have increased the efficiency of the second, classical thrust by initiating a swing.

Are human anticipatory postural adjustments affected by a modification of the initial position of the center of gravity?

During a lateral leg raising task, the position of the center of gravity (CG) in the horizontal plane shifts towards the supporting leg prior to the movement onset. The aim of this study was to explore whether the anticipatory postural adjustments were calibrated as a function of the initial horizontal location of the CG. Experiments were performed on 8 healthy subjects, with three initial positions of the CG (close to the supporting leg, between the two legs, close to the moving leg). Simultaneous kinematic, kinetic and electromyographic (EMG) data were recorded with the ELITE. system. The results show that the duration of the kinetic variables and EMG pattern are scaled as a function of the distance covered by the CG and constitute the means of modulating the CG shift. They suggest that the evaluation of the support conditions is necessary to calibrate the CG shift, this is done during the early phase of the postural adjustments.

Voluntary head stabilization in space during oscillatory trunk movements in the frontal plane performed in weightlessness.

The ability voluntarily to stabilize the head in space during lateral rhythmic oscillations (0.59+/-0.09 Hz) of the trunk has been investigated during microgravity (microG) and normal gravity (nG) conditions (parabolic flights). Five healthy young subjects, who gave informed consent, were examined. The movements were performed with eyes open or eyes closed, during phases of either microG or nG. The main result was that head orientation with respect to vertical may be stabilized about the roll axis under microG with, as well as without vision, despite the reduction in vestibular afferent and muscle proprioceptive inputs. Moreover, the absence of head stabilization about the yaw axis confirms that the degrees of freedom of the neck can be independently controlled, as was previously reported. These results seem to indicate that voluntary head stabilization does not depend crucially upon static vestibular afferents. Head stabilization in space may in fact be organized on the basis of either dynamic vestibular afferents or a short-term memorized postural body schema.

Is the regulation of the center of mass maintained during leg movement under microgravity conditions?

1. Investigations on stance regulation have already suggested that the body's center of mass is the variable controlled by the CNS to maintain equilibrium. The aim of this study was to determine how the center of mass of the body is regulated when leg movements are made under different gravitoinertial force conditions. 2. Kinematic and electromyographic (EMG) recordings were made during both straight-and-level flight (earth-normal gravity condition, nG) and periods of weightlessness in parabolic flight (microgravity condition, microG). The standing subjects were restrained to the floor (kept from floating away in microG) and were instructed to raise one leg laterally to an angle of 45 degrees as fast as possible. 3. Two modes of center of mass (CM) control were identified during leg movement in nG: a "shift mode" and a "stabilization mode." The shift mode served to transfer the CM toward the supporting side before the leg raising, and it preceded the phase of single limb support. The stabilization mode took place after the CM shift was completed and was aimed at stabilizing the CM during raising of the leg. In this phase, the movement of the raising leg is counterbalanced by a lateral inclination of the trunk in the opposite direction. As a consequence, CM position did not change with respect to the position reached before the leg raising, and its projection on the ground remained within the support area delineated by the stance foot. 4. Under microG, the CM position did not change before the leg raising. Moreover, gastrocnemius medialis activity observed in the moving leg under nG, preceding the initiation of the body weight transfer toward the supporting leg, was greatly reduced. While the leg is raising, the simultaneous and opposite lateral trunk movement was still present in microG. 5. Results suggest that the body weight transfer corresponding to the shift mode, might depend on the gravity constraints, whereas the stabilization mode, which remains unchanged in microG, might be a motor stereotype that does not depend on the gravity conditions.

Voluntary head stabilization in space during trunk movements in weightlessness.

The ability to voluntarily stabilize the head in space during lateral rhythmic oscillations of the trunk has been investigated during parabolic flights. Five healthy young subjects, who gave informed consent, were examined. The movements were performed with eyes open or eyes closed, either during phases of microgravity or phases of normal gravity. The main result to emerge from this study is that the head may be stabilized in space about the roll axis under microgravity conditions with, as well as without vision, despite the reduction of the vestibular afferent and the muscle proprioceptive inputs. Moreover, the absence of head stabilization about the yaw axis confirms that the degrees of freedom of the neck can be independently controlled, as it was previously shown. These results seem to indicate that voluntary head stabilization does not depend crucially upon static vestibular afferents. Head stabilization in space may be in fact organized on the basis of either dynamic vestibular afferents or a postural body scheme.

Body orientation and regulation of the center of gravity during movement under water.

Professional divers were instructed to adopt a vertical posture under water with their feet fixed to the ground and to perform a fast forward or backward upper trunk bending movement in response to a tone. Kinematic and EMG analyses were performed. It was first noted that the divers adopted a forward inclined, erect posture, suggesting that the verticality was misevaluated, although the effects of gravity were still exerted on the otoliths. Second, the upper trunk movements were still accompanied by opposite movements of lower segments and, as a result, the center of gravity displacement was still minimized, although not so accurately as on the ground. The EMG pattern consisting of early activation of a set of trunk, thigh, and shank muscles continued to occur under water. These results suggest that "axial synergies" associated with upper trunk movements are learned motor habits that regulate the center of gravity position regardless of the equilibrium constraints.

Is the trunk a reference frame for calculating leg position?

Naive subjects and dancers were instructed to raise a leg laterally toward 45 degrees. The final position reached by the leg by each group of subjects was quite different: 48 degrees in dancers, i.e. close to the required value, and 56 degrees in the naive subjects. The reason for this difference was investigated. During the body weight transfer toward the supporting side prior to the leg movement, naive subjects inclined both leg and trunk laterally, whereas the dancers' trunk remained vertical. It was observed that in naive subjects the trunk inclination and the overestimation of the final leg position were closely correlated. The results suggest that in both naive subjects and dancers, the trunk axis serves as a reference value for calculating the leg position.

Coordination between equilibrium and head-trunk orientation during leg movement: a new strategy build up by training.

1. During unilateral leg movements performed while standing, it is necessary to displace the center of gravity toward the other leg to maintain equilibrium. In addition, the orientation of particular segments, such as the head and trunk, which are used as reference values for organizing the motor act, needs to be preserved. The aim of the present study was to investigate the coordination between movement, equilibrium, and local posture. 2. Experiments were carried out on standing subjects who were instructed to raise one leg laterally to an angle of 45 degrees in response to a light. Two sources of light placed in front of the subject indicated the side on which the movement was to be performed. Three main aspects of the posturokinetic sequence were investigated in two populations, naive subjects and dancers: 1) The body weight transfer toward the supporting leg was found to have two components: first, a "ballistic" one, initiated by a thrust exerted by the moving leg; and second, an "adjustment" component during which the displacement of the center of gravity (CG) reaches a final position (steady state). An early burst in the gastrocnemius medialis of the moving leg often precedes the onset of the center of pressure change. Two differences between naive subjects and dancers were observed: first, the new CG position was almost reached in one step very near to the end of the ballistic component and required only a short adjustment in dancers, whereas in naive subjects it was reached in two steps, including a much longer adjustment component. Second, the dancers were able to minimize the CG displacement toward the supporting side; this might be because they form a better internal representation of the biomechanical limits of stability because of their long training. 2) The onset of the lateral displacement of the malleolus marker of the moving leg always occurred when the body weight had almost completed its transfer to above the support foot. This shows that the positioning of the CG in a new position compatible with equilibrium maintenance was a prerequisite for the leg movement to be performed. The relative timing of events during the posturokinetic sequence was fairly fixed in the dancers, whereas it varied from one trial to another in the naive subjects. 3) The coordination between movement, equilibrium, and head-trunk orientation involves two control strategies. An "inclination" strategy was used by the naive subjects; this consisted of an external rotation of the supporting leg around the anteroposterior ankle joint axis. A counter-rotation at the neck level ensured the stability of the interorbital line in the horizontal plane.(ABSTRACT TRUNCATED AT 400 WORDS)