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The role of the neck and trunk in facilitating head stability during walking.

An apparent goal of the human postural system is to maintain head stability during walking. Although much is known about sensory-motor stabilising mechanisms associated with the head and neck, less is known about how the postural system attenuates motion between the trunk and neck segments in order to regulate head motion. Therefore the purpose of this study was to determine the role that the neck and the trunk play in stabilising the head at a range of walking speeds. Eight healthy male subjects (age: 23+/-4 years) performed self-selected slow, preferred, and fast walking speed trials along a 30 m walkway. Four custom-designed wireless triaxial accelerometers were attached to the head, upper trunk, lower trunk, and shank of each subject to measure vertical (VT), anterior-posterior (AP), and mediolateral (ML) accelerations. Acceleration data were examined in each direction using RMS, power spectral, harmonic, and regularity measures. Signal regularity was increased from the lower to upper trunk for all walking speeds and directions with the exception of the slow speed in the AP direction. Evidence from analysis of power spectral and amplitude characteristics of acceleration signals was suggestive that accelerations are also attenuated from the lower to upper trunk by dynamics of the intervening trunk segment. Differences in selected power spectral and amplitude characteristics between the accelerations of the upper trunk and head due to the intervening neck segment were only detected in the AP direction at preferred and fast walking speeds. Overall the findings of the present study suggest that the trunk segment plays a critical role in regulating gait-related oscillations in all directions. Only accelerations in the direction of travel at preferred and fast speeds required additional control from the neck segment in order to enhance head stability during walking.

Acceleration↗

Impact of subthalamic nucleus stimulation on the initiation of gait in Parkinson's disease.

The effects of subthalamic nucleus (STN) stimulation on the anticipatory postural actions associated with the initiation of gait were studied in ten patients with idiopathic Parkinson's disease undergoing therapeutic deep brain stimulation. Kinematic, dynamic and electromyographic analysis was performed before and while subjects were starting gait in response to an external cue. Effects of STN stimulation on the standing posture preceding the go signal included significant improvement of the vertical alignment of the trunk and shank, decrease of the hip joint moment, backward shift of the center of pressure (CoP) and reduction of abnormal tonic and/or rhythmic activity in the thigh and leg muscles. Responses to bilateral STN stimulation were more consistent than those evoked by unilateral stimulation. Moreover, comparison between postural changes induced by STN stimulation applied prior to the gait initiation cue and during simple quiet standing revealed more significant responses in the former condition. Effects on the actual gait initiation process included shortening of the imbalance phase, larger backward/lateral displacement of CoP and more physiological expression of the underlying anticipatory muscular synergy. Additional changes were shortening of the unloading phase, shortening of the first-swing phase and increase in the length of the first step. Results demonstrate substantial influence of STN stimulation on functionally basic motor control mechanisms. In particular, the evidence of more significant responses upon attention-demanding conditions and the remarkable effects on postural programmes sub-serving feed-forward regulation of the onset of complex multijoint movements, suggests a consistent action on postural sub-systems relying on cognitive data processing and internal models of body mechanics.

Aged↗

Kinematic synergy adaptation to an unstable support surface and equilibrium maintenance during forward trunk movement.

The aim of this investigation was to study the adaptation to an unstable support surface of kinematic synergy responsible for equilibrium control during upper trunk movements. Eight adult subjects were asked to bend their upper trunk forward to an angle of 35 degrees and then to hold the final position for 3 s, first in a standard condition, with two feet on the ground and the second, on a rocking platform swinging in the sagittal plane. The movement characteristics (duration, amplitude, and mean angular velocity of the trunk), the time course of the antero-posterior center of mass (CM) shift during the movement, and the EMG pattern of the main muscles involved in the movement were studied under the two experimental conditions. Kinematic synergy was quantified by performing a principal component analysis on the hip, knee, and ankle angle changes occurring during the movement. The results indicate that (1) the CM shift from the very onset of the movement remains controlled during performance of the forward trunk movement when the equilibrium constraints were increased; (2) the principal component analysis of the hip, knee, and ankle angle changes occurring during the movement showed a transition from one principal component (PC(1)) in the standard condition to two components in the rocking platform condition; (3) the greatest contribution of PC(1) (weight coefficients) was located at the hip level in both the standard and rocking platform conditions, while the greatest contribution of PC(2) in the rocking platform condition was located at the ankle level; and (4) the EMG pattern underlying kinematic synergy is modified. It is concluded that a simple adaptation of kinematic synergy by changing the weight coefficients of each pair of joints participating in the movement is no longer sufficient when the equilibrium constraints increase and, rather, disturbs equilibrium. The CNS has to provide two parallel controls, one to perform the trunk movement and the other to preserve equilibrium.

Adaptation, Physiological↗

Do you know where your arm is if you think your head has moved?

Reproduction of a previously presented elbow position is affected by changes in head position. As movement of the head is associated with local biomechanical changes, the aim of the present study was to determine if illusory changes in head position could induce similar effects on the reproduction of elbow position. Galvanic vestibular stimulation (GVS) was applied to healthy subjects in supine lying. The stimulus was applied during the presentation of an elbow position, which the subject then reproduced without stimulation. In the first study, 13 subjects received 1.5 mA stimuli, which caused postural sway in standing, confirming that the firing of vestibular afferents was affected, but no illusory changes in head position were reported. In the second study, 13 subjects received 2.0-3.0 mA GVS. Six out of 13 subjects reported consistent illusory changes in head position, away from the side of the anode. In these subjects, anode right stimulation induced illusory left lateral flexion and elbow joint position error towards extension (p=0.03), while anode left tended to have the opposite effect (p=0.16). The GVS had no effect on error in subjects who did not experience illusory head movement with either 1.5 mA stimulus (p=0.8) or 2.0-3.0 mA stimulus (p=0.7). This study demonstrates that the accuracy of elbow repositioning is affected by illusory changes in head position. These results support the hypothesis that the perceived position of proximal body segments is used in the planning and performance of accurate upper limb movements.

Adult↗

A cognitive intersensory interaction mechanism in human postural control.

Human control of upright body posture involves inputs from several senses (visual, vestibular, proprioceptive, somatosensory) and their central interactions. We recently studied visual effects on posture control and their intersensory interactions and found evidence for the existence of an indirect and presumably cognitive mode of interaction, in addition to a direct interaction (we found, e.g., that a 'virtual reality' visual stimulus has a weaker postural effect than a 'real world' scene, because of its illusory character). Here we focus on the presumed cognitive interaction mechanism. We report experiments in healthy subjects and vestibular loss patients. We investigated to what extent a postural response to lateral platform tilt is modulated by tilt of a visual scene in an orthogonal rotational plane (anterior-posterior, a-p, direction). The a-p visual stimulus did not evoke a lateral postural response on its own. But it enhanced the response to the lateral platform tilt (i.e., it increased the evoked body excursion). The effect was related to the velocity of the visual stimulus, showed a threshold at 0.31 degrees /s, and increased monotonically with increasing velocity. These characteristics were similar in normals and patients, but body excursions were larger in patients. In conclusion, the orthogonal stimulus arrangement in our experiments allowed us to selectively assess a cognitive intersensory interaction that upon co-planar stimulation tends to be merged with direct interaction. The observed threshold corresponds to the conscious perceptual detection threshold of the visual motion, which is clearly higher than the visual postural response threshold. This finding is in line with our notion of a cognitive phenomenon. We postulate that the cognitive mechanism in normals interferes with a central visual-vestibular interaction mechanism. This appears to be similar in vestibular loss patients, but patients use less effective somatosensory instead of vestibular anti-gravity mechanisms.

Adult↗

Localization of the subjective vertical during roll, pitch, and recumbent yaw body tilt.

Localization of the subjective vertical during body tilt in pitch and in roll has been extensively studied because of the relevance of these axes for aviation and control of posture. Studies of yaw orientation relative to gravity are lacking. Our goal was to perform the first thorough evaluation of static orientation in recumbent yaw and to collect as efficiently as possible roll and pitch orientation data which would be consistent with the literature, using the same technique as our yaw tests. This would create the first comprehensive, coherent data set for all three axes suitable for quantitative tri-dimensional modeling of spatial orientation. We tested localization of the vertical for subjects tilted in pitch (-100 degrees to +130 degrees ), in roll (-90 degrees to +90 degrees ), and in yaw while recumbent (-80 degrees to +80 degrees ). We had subjects point a gravity-neutral probe to the gravitational vertical (haptically indicated vertical) and report verbally their perceived tilt. Subjects underestimated their body tilts in recumbent yaw and pitch and overestimated their tilts in roll. The haptic settings for pitch and roll were consistent with data in the literature obtained with haptic and visual indications. Our data constitute the first tri-dimensional assessment of the subjective vertical using a common measurement procedure and provide the basis for the tri-axial modeling of vestibular function presented in the companion paper.

Adult↗

The influence of natural body sway on neuromuscular responses to an unpredictable surface translation.

Previous research has shown that the postural configuration adopted by a subject, such as active leaning, influences the postural response to an unpredictable support surface translation. While those studies have examined large differences in postural conditions, it is of additional interest to examine the effects of naturally occurring changes in standing posture. Thus, it was hypothesized that the normal postural sway observed during quiet standing would affect the responses to an unpredictable support surface translation. Seventeen young adults stood quietly on a moveable platform and were perturbed in either the forward or backward direction when the location of the center of pressure (COP) was either 1.5 standard deviations anterior or posterior to the mean baseline COP signal. Postural responses, in the form of electromyographic (EMG) latencies and amplitudes, were recorded from lower limb and trunk muscles. When the location of the COP at the time of the translation was in the opposite, as compared to the same, direction as the upcoming translation, there was a significantly earlier onset of the antagonists (10-23%, i.e. 15-45 ms) and a greater EMG amplitude (14-39%) in four of the six recorded muscles. Stepping responses were most frequently observed during trials where the position of the COP was opposite to the direction of the translation. The results support the hypothesis that postural responses to unpredictable support surface translations are influenced by the normal movements of postural sway. The results may help to explain the large variability of postural responses found between past studies.

Adult↗

Muscle synergies during voluntary body sway: combining across-trials and within-a-trial analyses.

We investigated co-varied changes in muscle activity during voluntary sway tasks that required a quick shift of the center of pressure (COP). We hypothesized that multi-muscle synergies (defined as task-specific covariation of elemental variables, muscle modes) stabilize a COP location in the anterior-posterior direction prior to a voluntary COP shift and that during the shift the synergies would weaken. Standing subjects performed two tasks, a cyclic COP shift over a range corresponding to 80% of the maximal amplitude of voluntary COP shift at 1 Hz and a unidirectional quick COP shift over the same nominal amplitude. The cyclic sway task was used to define muscle modes (M-modes, leg and trunk muscle groups with parallel scaling of muscle activation level within a group) and the relations between small changes in the magnitudes of M-modes [in the principal component analysis (PCA), the M-mode magnitudes are equivalent to PC scores] and COP shifts. A novel approach was used involving PCA applied to indices of muscle integrated activity measured both within a trial and across trials. The unidirectional sway task was performed in a self-paced (SP) manner and under a typical simple reaction time (RT) instruction. M-modes were also defined along trials at those tasks; they have been shown to be similar across tasks. Integrated indices of muscle activity in the SP-sway and RT-sway tasks were transformed into the M-modes. Variance in the M-mode space was partitioned into two components, one that did not affect the average value of COP shift (V (UCM)) and the other that did (V (ORT)). An index (DeltaV) corresponding to the normalized difference between V (UCM) and V (ORT) was computed. During steady-state posture, DeltaV was positive corresponding to most M-mode variance lying in a sub-space corresponding to a stable COP location across trials. Positive DeltaV values have been interpreted as reflecting a multi-M-mode synergy stabilizing the COP location. The magnitude of DeltaV was larger in SP trials than in RT trials. During voluntary COP shifts, the DeltaV magnitude dropped to zero or even became negative. We conclude that M-mode synergies stabilize COP location during quiet standing, while these synergies weaken or disappear during fast voluntary COP shifts. Under RT conditions, the COP stabilizing synergies were weaker supposedly to facilitate a quick COP shift without time for preparation. The suggested method of M-mode identification may potentially be applied to analysis of postural synergies in persons with impaired postural control such as elderly persons, persons with atypical development, or in the course of rehabilitation after an injury.

Adult↗

Multisensory information for postural control: sway-referencing gain shapes center of pressure variability and temporal dynamics.

The authors investigated the multisensory control of posture by altering sensory information across the visual and somatosensory systems. The support surface and visual surround were sway-referenced to anterior/posterior center of mass sway and the gain between postural sway and degree of sway referencing was manipulated (gain settings were 0.2, 1.0, and 1.8). These alterations in the sensory environment lead to observed changes in the temporal structure of the center of pressure (COP) trajectories. COP path length increased across gain settings while COP coefficient of variation decreased. The COP became increasingly more deterministic across more challenging sensory organization test (SOT) conditions and with increasing gain, and more nonstationary across more challenging SOT conditions and when the support surface was sway-referenced using a 1.8 gain setting. These findings indicate that changes in the responsiveness of the support surface and/or visual surround within each of the sway-referenced SOT conditions had functional consequences for the control of posture as evidenced by the variations in postural sway dynamics.

Adolescent↗

Visual motion combined with base of support width reveals variable field dependency in healthy young adults.

We previously reported responses to induced postural instability in young healthy individuals viewing visual motion with a narrow (25 degrees in both directions) and wide (90 degrees and 55 degrees in the horizontal and vertical directions) field of view (FOV) as they stood on different sized blocks. Visual motion was achieved using an immersive virtual environment that moved realistically with head motion (natural motion) and translated sinusoidally at 0.1 Hz in the fore-aft direction (augmented motion). We observed that a subset of the subjects (steppers) could not maintain continuous stance on the smallest block when the virtual environment was in motion. We completed a posteriori analyses on the postural responses of the steppers and non-steppers that may inform us about the mechanisms underlying these differences in stability. We found that when viewing augmented motion with a wide FOV, there was a greater effect on the head and whole body center of mass and ankle angle root mean square (RMS) values of the steppers than of the non-steppers. FFT analyses revealed greater power at the frequency of the visual stimulus in the steppers compared to the non-steppers. Whole body COM time lags relative to the augmented visual scene revealed that the time-delay between the scene and the COM was significantly increased in the steppers. The increased responsiveness to visual information suggests a greater visual field-dependency of the steppers and suggests that the thresholds for shifting from a reliance on visual information to somatosensory information can differ even within a healthy population.

Adult↗

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.

Adaptation, Physiological↗

Early development of postural adjustments in standing with and without support.

This study investigates the early development of postural adjustments during external perturbations in two different standing positions: standing with support and standing without support. The aim of the study was to assess a group of 13 infants four times during the period in life when independent standing is achieved; at 8, 10, 12 and 14 months. However, longitudinal data could be achieved only in four infants. Muscle activations of the neck, hip and ankle were recorded using surface electromyography. Based on earlier studies and controversies, three main issues were addressed: (1) Is direction specificity present before independent standing is established? (2) How do postural adjustments change with increasing age (8-14 months)? (3) Are postural adjustments task-specific in the young child? The results showed that our small sample of infants aged 8 and 10 months, who were not yet able to stand independently, exhibited direction-specific postural adjustments both during standing with and without support, though not consistently during all trials and at all body levels. Therefore, we argue that direction specificity might constitute a prerequisite for the development of independent standing. We also found that the development of postural adjustments in standing with support resembles that of sitting, i.e. great variation in the postural adjustments at early age, and fine-tuning to the situation with increasing age and experience. This, we find that this is in agreement with the proposal that postural control develops through a selection process of the most suitable postural adjustments for the situation from a repertoire of direction-specific postural adjustments. The development of postural adjustments during standing without support is discussed. Additionally, differences in response rates were noted between the two standing positions, indicating that even before independent standing is established, sophisticated sensorimotor integration enables task-specific postural adjustments.

Age Factors↗

Role of gravity-based information on the orientation and localization of the perceived body midline.

The present study focused on the influence of gravity-based information on the orientation and localization of the perceived body midline. The orientation was investigated by the rolling adjustment of a rod on the subjects' Z-axis and the localization by the horizontal adjustment of a visual dot as being straight ahead. Experiment 1 investigated the effect of the dissociation between the Z-axis and the direction of gravity by placing subjects in roll tilt and supine postures. In roll tilt, the perception of the body midline orientation was deviated in the direction of body tilt and the perception of its localization was deviated in the opposite direction. In the supine body orientation, estimates of the Z-axis and straight-ahead remained veridical as when the body was upright. Experiment 2 highlighted the relative importance of the otolithic and tactile information using diffuse pressure stimulation. The estimation of body midline orientation was modified contrarily to the estimation of its localization. Thus, subjects had no absolute representation of their egocentric space. The main hypothesis regarding the dissociation between the orientation and localization of the body midline may be related to a difference in the integration of sensory information. It can be suggested that the horizontal component of the vestibulo-ocular reflex (VOR) contributed to the perceived localization of the body midline, whereas its orientation was mainly influenced by tactile information.

Adult↗

Imagery of different sensory modalities: hypnotizability and body sway.

Postural control in subjects with high (Highs) and low (Lows) susceptibility to hypnosis is differentially affected by changes in visual and neck tactile/proprioceptive input. The aim of the present experiment was to investigate whether imagery of the visual and tactile sensory modalities also induces different modulation of postural control in Highs and Lows. Fourteen Highs and 16 Lows were included in the study; they were recorded while standing upright with eyes closed during visual and tactile imagery tasks and during mental computation. Their posture and movement were recorded with an Elite System and their experience was assessed after each task in a structured interview. Visual imagery was judged "easier" than tactile imagery by Lows, while Highs performed both tasks easily and judged the tactile imagery less effortful and more vivid than Lows. No difference was observed for the mental computation. The Highs' body sway was not affected by the cognitive tasks, while Lows showed a task-related modulation of body sway. The results are in line with the hypothesis of lower vulnerability of Highs to the effects of tasks interfering with postural control and of different sensory-motor integration in Highs and Lows.

Adult↗

The effects of instability and additional hand support on anticipatory postural adjustments in leg, trunk, and arm muscles during standing.

We investigated the role of additional perceptual information (finger touch) and additional mechanical support (hand grasp) on the anticipatory postural adjustments (APAs) associated with fast arm movements performed by standing subjects. The subjects performed fast, unilateral shoulder ante-flexion movements while standing on a stable force platform or on an unstable board with instability in a sagittal or in a frontal plane. Changes in the background activity of leg, trunk, and arm muscles and displacements of the center of pressure were quantified within time intervals typical for APAs. Leg and trunk muscles showed a significant drop in APAs with added finger touch and no further changes when the touch was substituted with hand grasp. Arm muscles showed no changes or a small drop in APAs with touch and a significant increase in APAs with grasp. These changes were seen during both stable and unstable standing. We conclude that APAs can show changes associated not only with mechanical aspects of a task, but also with its perceptual aspects. Based on the equilibrium-point hypothesis of motor control, an additional analysis of APAs was performed. With additional support, we observed a significant modulation of an index related to the co-activation of agonist-antagonist muscle pairs, while no changes in reciprocal activation were found. A similar analysis performed across all the leg and trunk muscles with respect to control of the center of mass lead to similar results. We conclude that the central nervous system seems to simplify adjustments of control patterns to changes in task parameters by modulation of only one of the two major central commands.

Adult↗

Initial evidence for the mixing and soft assembly of the ankle, suspensory, and hip muscle patterns.

The onset latencies of automatic muscle responses to backward translations of the support surface in standing subjects were assessed to test two longstanding hypotheses: that (1) the ankle, hip, and suspensory muscle patterns represent discrete synergies; and (2) the suspensory pattern cannot be mixed with the ankle and the hip patterns (based on theoretical modeling). Muscle responses were recorded at the leg (medial gastrocnemius and tibialis anterior), thigh (hamstring and quadriceps), lower trunk (paraspinals and abdominals), and the upper trapezius muscles. A principal factor analysis with orthogonal varimax rotation was conducted to describe the variation among the muscle onset latencies by means of the smallest possible set of newly defined variables, or factors. Muscles that were correlated with each other were grouped into a factor. The analysis produced three factors with eigenvalues > or =1, which accounted for 81% of the total variance. Based on the first muscle response in each factor, the muscle patterns resembled the hip, ankle, and suspensory synergies. Muscle responses within a synergy were activated before the completion of the muscle responses in the other synergies, indicating that the synergies, though mixed, are triggered separately. This is consistent with the hypothesis that the ankle, hip, and suspensory patterns represent discrete synergies. On the other hand, the results do not support the hypothesis that the three patterns cannot be combined.

Adult↗

Voluntary head stabilisation in space during oscillatory trunk movements in the frontal plane performed before, during and after a prolonged period of weightlessness.

The ability to voluntarily stabilise the head in space exhibited by two subjects during lateral rhythmic oscillations of the trunk has been investigated before, during and after a prolonged period of microgravity (microG) exposure. In flight acquisitions were performed onboard the Core Module of the Russian Space Station MIR as part of the T4 "Human Posture in Microgravity" experiment of the 179-days ESA-RKA mission EUROMIR-95. Data collection and kinematic analysis were performed by means of a space-qualified version of the automatic motion analyser ELITE. Head stabilisation in space strategy was estimated by means of the head anchoring index and cross-correlation analysis. Results show that head orientation may be well stabilised about the roll axis both with and without the presence of visual information. This was true despite the expected reduction in vestibular efficiency and muscular proprioception occurring in-flight. In one subject, however, vision was found to improve head stabilisation in space post-flight, presumably to recover from the postural deficiency induced by the long-term microG exposure. Head stability during trunk movements was achieved with either over-compensatory (out-of-phase), under-compensatory (in-phase) or mixed movement strategies, as was attested by the analysis of cross-correlation functions between head and shoulder movements. In weightlessness, vision occlusion seemed to influence the choice of the strategies to be used as well as the reduction of movement variability. The feedforward nature of compensatory head movements suggests that head stabilisation could be based in weightlessness on the internal postural body scheme, supposed to be adapted to the weightless environment within 5 months of microG exposure.

Adaptation, Biological↗

In a complex sequential movement, what component of the motor program is improved with intensive practice, sequence timing or ensemble motor learning?

We investigated whether the higher maximum speed of complex athletic movements attained by experts compared with novices might solely be ascribed to the acquisition of a more efficient timing of the initiation of the simple movements that comprise complex sporting gestures. Six novices in fencing and five experts in fencing performed three experimental series: 7 trials of touche, 7 trials of lunge (control series) and 50 trials of lunge + touche (test series), where the touche and lunge were initiated with different chronological sequences imposed by the experimenter. The lunge and the touche can be assimilated to a fast forward step and to a pointing task, respectively. We compared the maximum speed of touche between the two groups, recorded by an accelerometer fixed to the hand guard of the foil, and the speed of the centre of mass obtained from a force plate. The speed of the centre of mass was not statistically different between the two groups in the control and in the test series. The maximum speed of touche was not statistically different between the two groups in the control series. In contrast, in the test series, the maximum speed of touche was higher in the fencers' than in novices' groups, while the timing of initiation of the lunge and the touche was similar. The results of the test series show that the higher maximum speed of touche of fencers compared with novices cannot solely be ascribed to the acquisition of a more efficient initiation timing in motor programming. In a complex fencing gesture, the higher maximum speed of touche following intensive practice is discussed with regard to the inhibition of negative effects linked to the "refractory period" associated with the initiation of two closely spaced motor programs.

Adult↗