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Activity of thoracic and lumbar epaxial extensors during postural responses in the cat.

This study examined the role of trunk extensor muscles in the thoracic and lumbar regions during postural adjustments in the freely standing cat. The epaxial extensor muscles participate in the rapid postural responses evoked by horizontal translation of the support surface. The muscles segregate into two regional groups separated by a short transition zone, according to the spatial pattern of the electromyographic (EMG) responses. The upper thoracic muscles (T5-9) respond best to posteriorly directed translations, whereas the lumbar muscles (T13 to L7) respond best to anterior translations. The transition group muscles (T10-12) respond to almost all translations. Muscles group according to vertebral level rather than muscle species. The upper thoracic muscles change little in their response with changes in stance distance (fore-hindpaw separation) and may act to stabilize the intervertebral angles of the thoracic curvature. Activity in the lumbar muscles increases along with upward rotation of the pelvis (iliac crest) as stance distance decreases. Lumbar muscles appear to stabilize the pelvis with respect to the lumbar vertebrae (L7-sacral joint). The transition zone muscles display a change in spatial tuning with stance distance, responding to many directions of translation at short distances and focusing to respond best to contralateral translations at the long stance distance.

Animals↗

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.

Adult↗

Spatial-frequency-related efficacy of visual stabilisation of posture.

The present study investigates the efficacy of visual stabilisation of posture for different spatial frequencies of a visual stimulus. Circular sine wave gratings were used to analyse the correlation between perception of motion in depth and stabilisation of fore-aft sway by the mechanism of detecting changes in target size. Body sway was recorded by a force-measuring platform (series A) and, in addition, by simultaneous tracking of infrared markers fixed to the subject's body (series B). Mean velocity and amplitude (RMS) of body sway were calculated in both sagittal (a-p) and lateral (l-r) planes. Sagittal sway was of least magnitude when viewing contrast gratings with lowest thresholds, whereas higher thresholds resulted in increasing sway parameters. As intended by the design of the stimuli, sagittal sway was correlated closer with the stabilising effect exerted by the different stimuli than was lateral sway. Sway velocity was reduced more efficiently, however, with a lower correlation with the psychophysical transfer function, than was RMS sway. Since sway velocity measured by the platform is suggested to depend to a greater extent on dynamic muscle forces generated at each individual body site the results indicate that visual information can be used to reduce and thereby optimise dynamic muscle action (sway velocity) even though static body sway is either not or less reduced. A comparable economisation of sway velocity but not of RMS sway was also seen at the end of posture investigations, indicative of positive training effects.

Adult↗

Effect of head position on postural orientation and equilibrium.

This study examined (1) how changes in head position affect postural orientation variables during stance and (2) whether changes in head position affect the rapid postural response to linear translation of the support surface in the horizontal plane. Cats were trained to stand quietly on a moveable platform and to maintain five different head positions: center, left, right, up, and down. For each head position, stance was perturbed by translating the support surface linearly in 16 different directions in the horizontal plane. Postural equilibrium responses were quantified in terms of the ground reaction forces, kinematics, dynamics (net joint torques), body center of mass, and electromyographic (EMG) responses of selected limb and trunk muscles. A change in head position involved rotation of not only the neck but also the scapulae and anterior trunk. Tonic EMG levels were modulated in several forelimb and scapular muscles but not hindlimb muscles. Finally, large changes in head orientation in both horizontal and vertical planes did not hamper the ability of cats to maintain postural equilibrium during linear translation of the support surface. The trajectory of the body's center of mass was the same, regardless of head position. The main change was observed in joint torques at the forelimbs evoked by the perturbation. Evoked EMG responses of forelimb and scapular muscles were modulated in terms of magnitude but not spatial tuning. Hindlimb responses were unchanged. Thus, the spatial and temporal pattern of the automatic postural response was unchanged and only amplitudes of evoked activity were modulated by head position.

Animals↗

Effect of altered sensory conditions on multivariate descriptors of human postural sway.

Multivariate descriptors of sway were used to test whether altered sensory conditions result not only in changes in amount of sway but also in postural coordination. Eigenvalues and directions of eigenvectors of the covariance of shnk and hip angles were used as a set of multivariate descriptors. These quantities were measured in 14 healthy adult subjects performing the Sensory Organization test, which disrupts visual and somatosensory information used for spatial orientation. Multivariate analysis of variance and discriminant analysis showed that resulting sway changes were at least bivariate in character, with visual and somatosensory conditions producing distinct changes in postural coordination. The most significant changes were found when somatosensory information was disrupted by sway-referencing of the support surface (P = 3.2 x 10(-10)). The resulting covariance measurements showed that subjects not only swayed more but also used increased hip motion analogous to the hip strategy. Disruption of vision, by either closing the eyes or sway-referencing the visual surround, also resulted in altered sway (P = 1.7 x 10(-10)), with proportionately more motion of the center of mass than with platform sway-referencing. As shown by discriminant analysis, an optimal univariate measure could explain at most 90% of the behavior due to altered sensory conditions. The remaining 10%, while smaller, are highly significant changes in posture control that depend on sensory conditions. The results imply that normal postural coordination of the trunk and legs requires both somatosensory and visual information and that each sensory modality makes a unique contribution to posture control. Descending postural commands are multivariate in nature, and the motion at each joint is affected uniquely by input from multiple sensors.

Adult↗

Gaze stabilization during dynamic posturography in normal and vestibulopathic humans.

Dynamic posturography by measurement of center of pressure (COP) is a widely employed technique for evaluating the vestibular system. However, the relationship of COP motion to vestibulo-ocular reflex (VOR) function and image stability on the retina has not been determined previously. To assess these relationships, we report gaze, head, and trunk stability during dynamic posturography in 11 normal volunteers, 7 subjects with unilateral vestibular lesions, and 3 subjects with bilateral vestibular lesions. Posturographic tasks consisted of standing still and standing on a platform that was sliding (0.2 Hz), tilting (0.1 Hz), or covered with a foam cushion 6 cm thick while tilting (0.1 Hz). Each perturbation was imposed in the anterior-posterior and repeated in the medial-lateral direction, in both light and darkness. Subjects viewed (or in darkness remembered) a target located 50, 100, or 500 cm distant. COP, angular eye position, and angular and linear orbit and trunk positions were measured using magnetic search coils and flux gate magnetometer sensors. With the target visible, the velocity of image motion on the retina was on average always less than 1 degree/s, well within the range consistent with high visual acuity. In darkness, gaze velocity increased for normal and vestibulopathic subjects. During tilt, vestibulopathic subjects had a significantly greater gaze velocity than controls. Gain of the angular VOR (eye velocity/head velocity) was significantly lower in darkness than in light and in vestibulopathic as compared to control subjects. Gain of the VOR was significantly correlated with gaze instability, but variation in VOR gain accounted for only 20-40% of the variance. In darkness, the velocity of the COP was significantly greater in vestibulopathic than control subjects for every condition tested. In light, this difference was small and often not significant. Although spectral analysis of the COP indicated frequencies above 1 Hz that were not observed in motion of the trunk and orbit, root mean square (RMS) velocities of the trunk and orbit in the horizontal plane were higher in darkness and in vestibulopathic subjects, mirroring COP findings. Only in vestibulopathic subjects tested in darkness was there a correlation between COP velocity and gaze velocity; COP velocity was otherwise uncorrelated with gaze. Gaze velocity was greater with near than with distant targets. Vertical VOR gain was higher with near targets. No other significant effects of target distance were found. Head movement strategy, VOR gain, and COP were all unaffected by target proximity. These data show that gaze velocity measurements during dynamic posturography in darkness are sensitive to vestibular loss. With a visible target, both COP and gaze stability of vestibulopathic subjects are difficult to distinguish from normal. During visual feedback, it is likely that image stabilization over the range of frequencies tested is achieved through better head stability and through visual tracking, allowing vestibulopathic subjects to maintain adequate visual acuity.

Adult↗

Forearm postural control during unloading: anticipatory changes in elbow stiffness.

In this study, the equilibrium-point hypothesis of muscle-torque generation is used to evaluate the changes in central control parameters in the process of postural-maintenance learning. Muscle torque is described by a linear spring equation with modifiable stiffness, viscosity, and equilibrium angle. The stiffness is considered to be the estimation of the central command for antagonist-muscle coactivation and the equilibrium angle to be the estimation of the reciprocal command for a shift of invariant characteristics of the joint. In the experiments, a load applied to the forearm was released. The subjects were instructed to maintain their forearm in the initial horizontal position. Five sessions of approximately twenty trials each were carried out by eight subjects. During two "control" series, the load release was triggered by the experimenter. During three "learning" series, the load supported by one forearm was released by the subject's other hand. The elbow-joint angle, the angular acceleration, and the external load on the postural forearm were recorded. These recordings as well as anthropometric forearm characteristics were used to calculate the elbow-joint torque (which we called "experimental"). Linear regression analysis was performed to evaluate the equilibrium angle, joint stiffness, and viscosity at each trial. The "theoretical" torque was calculated using a linear spring equation with the found parameters. The good agreement observed between experimental and theoretical joint-torque time courses, apart from the very early period following unloading, argues in favor of the idea that the movement was mainly performed under a constant central command presetting the joint stiffness and the equilibrium angle. An overall increase in the stiffness occurred simultaneously with a decrease in the equilibrium angle during the "learning" series in all the subjects. This suggests that subjects learn to compensate for the disturbing effects of unloading by increasing the joint stiffness. The mechanism possibly responsible for the presetting of the central control parameters is discussed.

Cognition↗

The effects of stochastic galvanic vestibular stimulation on human postural sway.

Galvanic vestibular stimulation serves to modulate the continuous firing level of the peripheral vestibular afferents. It has been shown that the application of sinusoidally varying, bipolar galvanic currents to the vestibular system can lead to sinusoidally varying postural sway. Our objective was to test the hypothesis that stochastic galvanic vestibular stimulation can lead to coherent stochastic postural sway. Bipolar binaural stochastic galvanic vestibular stimulation was applied to nine healthy young subjects. Three different stochastic vestibular stimulation signals, each with a different frequency content (0-1 Hz, 1-2 Hz, and 0-2 Hz), were used. The stimulation level (range 0.4-1.5 mA, peak to peak) was determined on an individual basis. Twenty 60-s trials were conducted on each subject - 15 stimulation trials (5 trials with each stimulation signal) and 5 control (no stimulation) trials. During the trials, subjects stood in a relaxed, upright position with their head facing forward. Postural sway was evaluated by using a force platform to measure the displacements of the center of pressure (COP) under each subject's feet. Cross-spectral measures were used to quantify the relationship between the applied stimulus and the resulting COP time series. We found significant coherency between the stochastic vestibular stimulation signal and the resulting mediolateral COP time series in the majority of trials in 8 of the 9 subjects tested. The coherency results for each stimulation signal were reproducible from trial to trial, and the highest degree of coherency was found for the 1- to 2-Hz stochastic vestibular stimulation signal. In general, for the nine subjects tested, we did not find consistent significant coherency between the stochastic vestibular stimulation signals and the anteroposterior COP time series. This work demonstrates that, in subjects who are facing forward, bipolar binaural stochastic galvanic stimulation of the vestibular system leads to coherent stochastic mediolateral postural sway, but it does not lead to coherent stochastic anteroposterior postural sway. Our finding that the coherency was highest for the 1- to 2-Hz stochastic vestibular stimulation signal may be due to the intrinsic dynamics of the quasi-static postural control system. In particular, it may result from the effects of the vestibular stimulus simply being superimposed upon the quiet-standing COP displacements. By utilizing stochastic stimulation signals, we ensured that the subjects could not predict a change in the vestibular stimulus. Thus, our findings indicate that subjects can act as "responders" to galvanic vestibular stimulation.

Adolescent↗

Time-dependent influence of sensorimotor set on automatic responses in perturbed stance.

These experiments tested the hypothesis that the ability to change sensorimotor set quickly for automatic responses depends on the time interval between successive surface perturbations. Sensorimotor set refers to the influence of prior experience or context on the state of the sensorimotor system. Sensorimotor set for postural responses was influenced by first giving subjects a block of identical backward translations of the support surface, causing forward sway and automatic gastrocnemius responses. The ability to change set quickly was inferred by measuring the suppression of the stretched antagonist gastrocnemius responses to toes-up rotations causing backward sway, following the translations. Responses were examined under short (10-14 s) and long (19-24 s) inter-trial intervals in young healthy subjects. The results showed that subjects in the long-interval group changed set immediately by suppressing gastrocnemius to 51% of translation responses within the first rotation and continued to suppress them over succeeding rotations. In contrast, subjects in the short-interval group did not change set immediately, but required two or more rotations to suppress gastrocnemius responses. By the last rotation, the short-interval group suppressed gastrocnemius responses to 33%, similar to the long-interval group of 29%. Associated surface plantarflexor torque resulting from these responses showed similar results. When rotation and translation perturbations alternated, however, the short-interval group was not able to suppress gastrocnemius responses to rotations as much as the long-interval group, although they did suppress more than in the first rotation trial after a series of translations. Set for automatic responses appears to linger, from one trial to the next. Specifically, sensorimotor set is more difficult to change when surface perturbations are given in close succession, making it appear as if set has become progressively stronger. A strong set does not mean that responses become larger over consecutive trials. Rather, it is inferred by the extent of difficulty in changing a response when it is appropriate to do so. These results suggest that the ability to change sensorimotor set quickly is sensitive to whether the change is required after a long or a short series of a prior different response, which in turn depends on the time interval between successive trials. Different rate of gastrocnemius suppression to toes-up rotation of the support surface have been reported in previous studies. This may be partially explained by different inter-trial time intervals demonstrated in this study.

Adult↗

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.

Adult↗

Long-term adaptation of postural control in microgravity.

Orbital microgravity represents a unique environment, which allows the isolation of variables assumed to be involved in the mechanism of body positioning in space. In this context, the alignment of the trunk axis along allocentric references and the positioning of the body center of mass inside the supporting base compete for the role of the primary-controlled variable when assuming erect posture. This paper reports the quantitative evaluation of the postural strategies exhibited by two subjects with feet fixed to the floor of the space module along a 4-month period of exposure to microgravity. With respect to previous findings in parabolic flights and short term space missions, the analysis focused on long-term process of sensorimotor adaptation to weightlessness. Results show that while trunk-axis orientation is preserved and used as a stable postural frame of reference, the positioning of the body center of mass appears to be significantly biased backward and turns out to be involved in a long-term process of adaptation throughout the entire flight towards the re-emergence of a typically terrestrial postural regulation compatible with equilibrium.

Adult↗

Kinematic synergies and equilibrium control during trunk movement under loaded and unloaded conditions.

The aim of the present investigation was to study the adaptation of the kinematic synergy responsible for equilibrium control during upper trunk movements to a 10-kg load added to the subject's shoulders. Five 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 without any load and then with a 10-kg load fixed to their shoulders. The final anteroposterior CM positions 400 ms after the movement offset, the time course of the anteroposterior center of mass (CM) shift during the movement, the EMG pattern of the main muscles involved in the movement and the initial CP shift were studied under both unloaded and loaded conditions. The kinematic synergy was quantified by performing a principal components analysis on the hip, knee and ankle angle changes occurring during the movement. The results indicate that: (1) the final anteroposterior position of the CM changed little if at all in the presence of the additional load, and that the anteroposterior CM shift was minimized throughout the duration of the movement; (2) the kinematic synergy was still characterized, in the presence of the additional load, by a strong coupling between the angle changes, as indicated by the fact that the first principal component (PC1) accounted for more than 99% of the hip, knee and ankle joint movements. A change was observed, however, in the ratio between the angles: the ankle extension increased, thus compensating for the additional theoretical forward CM shift that the additional load could be expected to cause; (3) the lack of change in the initial backward CP shift observed under loaded condition as well as the lack of change of the initial agonist EMG bursts suggest that the initial feedforward control of the kinematic synergy was not affected in the presence of the additional load. An increase in the antagonist bursts, presumably reflecting an adaptation of the kinematic synergy, was observed during the late phase of the movement; and (4) it is concluded that the adaptation of the kinematic synergy to the load was due to a specific change in the feedback control during the braking phase of the movement which presumably increases the ankle joint extension and consequently causes an increased backward shift of the hip which compensates for the forward shift due to the load.

Adult↗

Tactile impoverishment and sensorimotor restriction deteriorate the forepaw cutaneous map in the primary somatosensory cortex of adult rats.

We investigated the effects of sensory deprivation on the forepaw representation in the primary somatosensory cortex (SI) in the adult rat. Cortical maps were constructed from high-resolution multiunit recordings of the response of layer IV neurons to somatosensory stimuli. The main features of the forepaw representation were described in terms of areal extent and topography of the cortical map, and sensory submodality, size, and location of the receptive field (RF) of small clusters of the cortical neurons. After being weaned, two groups of Long-Evans rats were housed in a standard (SE) or impoverished (IE) environment for 65-115 days. A third group of SE rats was subjected to severe sensorimotor restriction (SR) of one forepaw for 7 days or 14 days, by using a one-sleeved cast. A concomitant effect of unilateral forelimb immobilization was a forced use of the nonrestricted forelimb in postural balance. The maps of both forepaws were derived 24 h after the cast was removed and the animal was allowed normal limb use. In a fourth group, SE rats experienced a 7-day immobilization followed by symmetrical limb use for 7 days before we mapped the hemisphere contralateral to the casted limb. For the SE and IE rats, the total areal extent of the cutaneous forepaw representation was similar, but IE rats exhibited a significant expansion of cortical islets serving high-threshold, presumably noncutaneous inputs, which were included in the cutaneous maps. In addition, SI neurons of IE rats had greatly enlarged glabrous, but not hairy, skin RFs. For the SR rats, the areal extent of the cutaneous map of the casted forepaw decreased by about 50%, after both 7- and 14-day forelimb immobilization. Large cortical sectors presumed to be formerly activated by cutaneous inputs were driven by high-threshold inputs that disrupted the somatotopic representation of the forepaw skin surfaces. These "emergent" representational sectors were topographically organized. By contrast, the areal extent and topography of the non-casted forepaw representation did not differ from those of SE rats. The size of glabrous RFs on the casted forepaw was similar to that of SE rats. On the contrary, glabrous RFs on the noncasted forepaw of SR rats were larger than those on their casted forepaw. The size of hairy RFs was not altered by the forelimb restriction. Interestingly, alteration of the somatotopic features of the casted forepaw map persisted after 7 days of symmetric use of the forelimbs. The present study demonstrates that continuous sensory experience is needed for the organizational features of SI maps to be maintained.

Adrenal Glands↗

Online steering: coordination and control of body center of mass, head and body reorientation.

Steering is an integral component of adaptive locomotor behavior. Along with reorientation of gaze and body in the direction of intended travel, body center of mass must be controlled in the mediolateral plane. In this study we examine how these subtasks are sequenced when steering is planned early or initiated under time constraints. Whole body kinematics were monitored as individuals were required to change their direction of travel by varying amounts when visually cued either at the beginning of the walk or one stride before. The analyses focused on the transition stride from one travel direction to another. Timing of changes (with respect to first right foot contact) in trunk roll angle, head and trunk yaw angle, and right foot displacement in the mediolateral plane were analyzed. The magnitude of these measures along with right and left foot placement at the beginning and right foot placement at the end of the transition stride were also analyzed. The results show the CNS uses two mechanisms, foot placement and trunk roll motion (piking action about the hip joint in the frontal plane), to move the center of mass towards the new direction of travel in the transition stride, preferring to use the first option when planning can be done early. Control of body center of mass precedes all other changes and is followed by initiation of head reorientation. Only then is the rest of the body reorientation initiated.

Adult↗

Cross-axis adaptation of the translational vestibulo-ocular reflex.

The adaptive plasticity of the translational vestibulo-ocular reflex (VOR) was investigated in rhesus monkeys after 2-h exposure to either vertical or torsional optic flow stimulation accompanied by lateral translation stimuli (0.5 Hz). Because of the inherent ambiguity in the otolith system for the detection of gravitoinertial accelerations, we hypothesized that cross-axis adaptation of the translational VOR during lateral motion would be preferentially selective for a torsional optic flow stimulus that would mimic a roll tilt movement. However, we found that both vertical and torsional adaptation was possible. Furthermore, there was no significant preference for whether the torsional adaptation was in phase or out of phase with the apparent tilt induced by the motion stimulus. These results suggest that, at least at 0.5 Hz, there seems to be no preferential, visually induced adaptive capacity of the otolith system for tilt/translation reinterpretation during motion. Like the rotational VOR, translational VOR appears to exhibit a general form of cross-axis adaptation that operates for different directions of optic flow stimulation.

Adaptation, Physiological↗

Postural reflexes evoked by tapping forehead and chest.

We investigated whether a tap with a reflex hammer to the forehead can elicit responses in the leg muscles and whether vestibular stimulation is the crucial prerequisite for eliciting these responses. We also measured the postural changes caused by the tap and by the compensatory, presumably reflex-like reactions of the subject. Tap-evoked activity of leg muscles was easily elicited during upright stance in normal subjects and was also seen in two subjects without vestibular function. The pattern of muscle activation clearly showed a counteraction to the tap-evoked perturbation of stance. Taps applied to the chest elicited similar reflexes. Since these two conditions imply a different activation of the vestibular apparatus, the vestibular input alone cannot account for the observed leg muscle reflexes. We suggest that multisensory reflex pathways that integrate vestibular and proprioceptive inputs account for these reflexes.

Adult↗

Vestibulospinal reflexes: quantitative effects of sensory feedback and postural task.

Vestibulospinal and vestibulocollic reflexes evoked by galvanic stimulation were studied in 20 normal volunteers. In an initial "baseline" study, subjects stood unsupported on a flat surface and a narrow base with their eyes shut and with their heads rotated to the left. The effects of vision, external support and increasing stance width were examined both individually and in combination. In a second series, the effects of the same factors were examined while subjects stood on a compliant surface. Short latency (SL) and medium latency (ML) vestibulospinal reflex responses were evoked using 4 mA/20 ms galvanic vestibular stimulation (GVS) and measured from the right soleus muscle. Vestibulocollic reflexes to short duration galvanic stimulation (4 mA/2 ms) were also measured under similar conditions. Both SL and ML vestibulospinal reflexes decreased significantly and to a similar degree with vision, external support and increasing stance width on a flat surface. On the compliant surface, stance width did not result in a decrease in the ML reflex. Reflex amplitudes further decreased in a non-linear fashion with each additional sensory modality ("factor") that was made available; the degree of attenuation due to the addition of a second and third factor closely approximated the product of the effect of each factor in isolation. Standing on a compliant surface resulted in enhancement of average SL and ML reflexes under all conditions. Vestibulocollic reflexes in contrast showed no significant modulation with vision, external support and stance width. The dissociation between vestibulocollic and vestibulospinal reflexes indicates that the modulation of vestibulospinal reflexes with task occurs proximal to the primary vestibular afferents. Vestibulospinal reflexes were largest when subjects stood on a narrow base, on a compliant surface, deprived of vision and external support, consistent with the importance of vestibular function under these conditions. Although attenuated, vestibulospinal reflexes were preserved in most subjects even when vision and external support were available and a wider stance width was adopted. The combination of different factors caused a multiplicative attenuation of the initial response.

Adult↗

Postural coordination patterns associated with the swinging frequency of arms.

Voluntary arm movements frequently perturb body equilibrium in an upright posture. The motions of leg joints need to be coordinated according to the properties of voluntary arm movements in order to maintain body equilibrium, and this may cause a change in postural pattern. The purpose of this study was to determine whether the kinematic pattern generation of upright posture is influenced by a change in the swinging frequency of arm movements and whether the pattern generation is correlated with a change in joint torque about the shoulder joint. Four male subjects in an upright posture were instructed to swing their arms at seven different frequencies, determined by the maximum swinging frequency of each subject (35%max, 40-60%max, 65%max). Segment rotations around the shoulder, hip, and ankle joints were analyzed at kinematic and kinetic levels. The results of kinematic analysis indicated that tight coupling between motions of the shoulder and hip joints was generated in lower-frequency trials (under 40-45%max), whereas tight coupling between motions of the shoulder and ankle joints was generated in higher frequency trials (more than 40-45%max). Furthermore, the results of kinetic analysis revealed that changes in the joint torque patterns about the shoulder and hip joints occurred in trials at 40-45%max. The mean value of 40-45%max was close to the eigenfrequency of each subject's arm. We concluded that (1) postural patterns associated with a gradual change in the swinging frequency of the arms can be divided into two coordination modes (a hip-shoulder in-phase mode and an ankle-shoulder in-phase mode), and (2) these two patterns may be divided by the eigenfrequency of the arm.

Adult↗