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A multisensory posture control model of human upright stance.

We present a multisensory postural control model based on experiments where the balance in normal subjects and vestibular loss patients was perturbed by application of external torque produced by force-controlled pull stimuli. The stimuli were applied while subjects stood on a stationary or body-sway-referenced motion platform with eyes closed and auditory cues masked. Excursions of the center of mass (COM) and the center of pressure (COP) were analyzed using a systems analysis approach. The results were compared to an 'inverted pendulum' model of posture control. The model receives input from four sensors: ankle proprioceptors, semicircular canals, otoliths, and plantar pressure sensors (somatosensory graviceptors). Sensor fusion mechanisms are used to yield separate internal representations of foot support motion, gravity, and external torque (pull). These representations are fed as global set point signals into a local control loop based on ankle proprioceptive negative feedback. This set point control upgrades the proprioceptive body-on-foot (support) stabilization into a body-in-space control which compensates for support tilt, gravity, and contact forces. This compensation occurs even when the stimuli are combined or a voluntary lean is superimposed. Model simulations paralleled our experimental findings.

Adult↗

Multisensory control of human upright stance.

The interaction of different orientation senses contributing to posture control is not well understood. We therefore performed experiments in which we measured the postural responses of normal subjects and vestibular loss patients during perturbation of their stance. Subjects stood on a motion platform with their eyes closed and auditory cues masked. The perturbing stimuli consisted of either platform tilts or external torque produced by force-controlled pull of the subjects' body on a stationary platform. Furthermore, we presented trials in which these two stimuli were applied when the platform was body-sway referenced (i.e., coupled 1:1 to body position, by which ankle joint proprioceptive feedback is essentially removed). We analyzed subjects' postural responses, i.e., the excursions of their center of mass (COM) and center of pressure (COP), using a systems analysis approach. We found gain and phase of the responses to vary as a function of stimulus frequency and in relation to the absence versus presence of vestibular and proprioceptive cues. In addition, gain depended on stimulus amplitude, reflecting a non-linearity in the control. The experimental results were compared to simulation results obtained from an 'inverted pendulum' model of posture control. In the model, sensor fusion mechanisms yield internal estimates of the external stimuli, i.e., of the external torque (pull), the platform tilt and gravity. These estimates are derived from three sensor systems: ankle proprioceptors, vestibular sensors and plantar pressure sensors (somatosensory graviceptors). They are fed as global set point signals into a local control loop of the ankle joints, which is based on proprioceptive negative feedback. This local loop stabilizes the body-on-foot support, while the set point signals upgrade the loop into a body-in-space control. Amplitude non-linearity was implemented in the model in the form of central threshold mechanisms. In model simulations that combined sensor fusion and thresholds, an automatic context-specific sensory re-weighting across stimulus conditions occurred. Model parameters were identified using an optimization procedure. Results suggested that in the sway-referenced condition normal subjects altered their postural strategy by strongly weighting feedback from plantar somatosensory force sensors. Taking this strategy change into account, the model's simulation results well paralleled all experimental results across all conditions tested.

Adult↗

Age-related differences in body segmental movement during perturbed stance in humans.

OBJECTIVE: The temporal and spatial features of body segmental movements of the head, trunk, thigh, shank and foot in response to a fast speed postural disturbance were compared among four age groups. It was hypothesized that the upper body movement was minimized in the healthy young adults, but was augmented in the older adults. DESIGN: Repeated measures design with 12 experimental conditions. BACKGROUND: It is not well understood yet why the elderly fall more than the young. In studies of locomotion and active balance tasks, the head position has been suggested as a reference frame for controlling dynamic equilibrium. This study was aimed at examining: (1) whether or not this reference frame would exist during involuntary balance tasks, which are most likely related to accidental falls in the elderly; and (2) how would the stability of this reference frame change with age. METHODS: Thirty-six healthy subjects of four age groups (20-70 + yr of age) participated in this study. Postural perturbation was introduced by a moving platform. The sagittal plane kinematics of five body segments was directly measured using Integrated Kinematic Sensors. RESULTS: Compared to the lower limb movement, the upper body movement, especially the head, was minimal (less than 7 degrees ) in the young adults (age group 1), and was largest (about 14 degrees ) in the older subjects (age groups 2, 3 and 4). Moreover, the maximum range of motion of the older adults was found to be positively correlated not only to the height of the segment, but also to the timings of the movements, while the young adults showed no correlation between range of motion and the timings of the movement. CONCLUSIONS: The findings suggested that the young adults indeed stabilized their upper body as a reference frame for maintaining postural balance. In contrast, this reference frame was not stabilized well in the older adults. In compensation, the older adults adopted a more rigid movement strategy, compared to a more flexible movement strategy in the young. All these changes would lead the older subjects to be less tolerable to postural disturbances and more susceptible to falls than the young subjects.

Journal Article↗

Different effect of height on latency of leg and foot short- and medium- latency EMG responses to perturbation of stance in humans.

In standing humans, platform perturbations evoked short- (SLR) and medium-latency responses (MLR) in soleus (Sol), tibialis anterior (TA) and flexor digitorum brevis (FDB) muscles. The latency of all responses significantly increased with subjects' height. The slope of the regression lines for the MLRs versus height was significantly steeper than that for the SLRs. The conduction velocity of the afferent fibres mediating the FDB MLR, calculated on the basis of the regression, was 17.5 m/s. These findings indicate that the MLRs are transmitted through afferent fibres slower than those mediating the SLRs. While the latter fibres are spindle group Ia afferents, the former are suggested to be group II fibres.

Adolescent↗

Influence of visual feedback on successive control mechanisms in upright quiet stance in humans assessed by fractional Brownian motion modelling.

An up-to-date way to model the centre of pressure (CP) trajectories may consist in using fractional Brownian motion (fBm). By doing so, one may note that standing still is in fact controlled by two separate and successive mechanisms. The point raised in this study concerns the nature of these control mechanisms and their level of interaction. Following this idea, visual feedback (VFB), which is known to affect postural control by significantly decreasing sway magnitudes, was used. Twelve healthy adults, instructed to stand as still as possible, were tested under this VFB protocol (via a PC screen). In order to model the CP trajectories as fBm, variograms (mean square distances, MSD, expressed as a function of increasing time intervals deltat) were bi-logarithmically plotted. The main visual effect of VFB on these variograms concerns longest latency scaling regimes which reveal less stochastic and consequently more accurate control (P < 0.05 and P < 0.01 for X and Y components, respectively). An increase in the MSD of the transition point, which corresponds to the switch between the two control mechanisms, is also noted (P < 0.05). Overall, evidence is provided from this data that long latency scaling regimes do operate through a feedback process. Interestingly, this improved determinism in feedback control in turn induces a similar effect on the control operating over the shortest deltat. Thus, by privileging a control strategy based on feedback mechanisms, VFB in turn would make the subjects quicker in their initial displacement in order to reach a position capable of initiating a feedback mechanism.

Adult↗

Evaluation of the lambda model for human postural control during ankle strategy.

An accurate modeling of human stance might be helpful in assessing postural deficit. The objective of this article is to validate a mathematical postural control model for quiet standing posture. The postural dynamics is modeled in the sagittal plane as an inverted pendulum with torque applied at the ankle joint. The torque control system is represented by the physiological lambda model. Two neurophysiological command variables of the central nervous system, designated lambda and micro, establish the dynamic threshold muscle at which motoneuron recruitment begins. Kinematic data and electromyographic signals were collected on four young males in order to measure small voluntary sway and quiet standing posture. Validation of the mathematical model was achieved through comparison of the experimental and simulated results. The mathematical model allows computation of the unmeasurable neurophysiological commands lambda and micro that control the equilibrium position and stability. Furthermore, with the model it is possible to conclude that low-amplitude body sway during quiet stance is commanded by the central nervous system.

Adult↗

Human postural response to lower leg muscle vibration of different duration.

Body lean response to bilateral vibrations of soleus muscles were investigated in order to understand the influence of proprioceptive input from lower leg in human stance control. Proprioceptive stimulation was applied to 17 healthy subjects by two vibrators placed on the soleus muscles. Frequency and amplitude of vibration were 60 Hz and 1 mm, respectively. Vibration was applied after a 30 s of baseline. The vibration duration of 10, 20, 30 s respectively was used with following 30 s rest. Subjects stood on the force platform with eyes closed. Postural responses were characterized by center of pressure (CoP) displacements in the anterior-posterior (AP) direction. The CoP-AP shifts as well as their amplitudes and velocities were analyzed before, during and after vibration. Vibration of soleus muscles gradually increased backward body tilts. There was a clear dependence of the magnitude of final CoP shift on the duration of vibration. The amplitude and velocity of body sway increased during vibration and amplitude was significantly modulated by duration of vibration as well. Comparison of amplitude and velocity of body sway before and after vibration showed significant post-effects. Presented findings showed that somatosensory stimulation has a long-term, direction-specific influence on the control of postural orientation during stance. Further, the proprioceptive input altered by soleus muscles vibration showed significant changes in postural equilibrium during period of vibration with interesting post-effects also.

Adult↗

Lateral roentgenography in the diagnosis and treatment of dysplasia/dislocation of the hip.

The author emphasizes the importance of true lateral and multiple-position roentgenograms in providing a more comprehensive demonstration of the pathology, the attainment of reduction and the surgical alterations which are necessary to maintain a congruous reduction. A simple convenient technique for lateral roentgenography of the hip in children is presented. A series of cases with roentgenograms is also presented to demonstrate the value of the lateral roentgenograms. The horizontal lateral roentgenogram simulates the position of the upright human stance, and thus, depicts malocclusion, which may be seen when anteversion is present. Even though the anterior view depicts sphericity and apparent coverage, the lateral roentgenogram may show malocclusion which may lead to degenerative changes later in life. It is the author's belief that the treatment of children's hip conditions will show improved long-term results if predicated on the comprehensive x-ray examination outlined in this article.

Child↗

Dynamics of human postural transitions.

In the present study, the authors examined transitions between postural coordination modes involved in human stance. The analysis was motivated by dynamical theories of pattern formation, in which coordination modes and transitions between modes are emergent, self-organized properties of the dynamics of animal-environment systems. In 2 experiments, standing participants tracked a moving target with the head. Results are consistent with the hypothesis that changes in body coordination follow typical nonequilibrium phase transitions, exhibiting multistability, bifurcation, critical fluctuations, hysteresis, and critical slowing down. The findings suggest that posture may be organized in terms of dynamical principles and favor the existence of general and common principles governing pattern formation and flexibility in complex systems.

Adult↗

Modification of human postural response to leg muscle vibration by electrical vestibular stimulation.

In order to understand proprioceptive and vestibular contributions to human stance posture, the effect of electrical vestibular stimulation on body lean induced by leg muscle vibration was investigated. The magnitudes and directions of postural responses were registered as changes in the center of foot pressure (COP) with a force platform. Vestibular stimulation consisted of 1 mA, binaural, bipolar galvanic current and proprioceptive input from tibialis anterior or soleus muscles was altered vibratory stimulation. The body lean induced by combined vibratory and galvanic stimulation could be largely considered as a summation of responses evoked by the galvanic and vibratory stimulation alone. The results of the present study showed that both vestibular and proprioceptive signals play important roles in the estimation of internal representation of the body vertical.

Adult↗

The influence of artificially increased hip and trunk stiffness on balance control in man.

Lightweight corsets were used to produce mid-body stiffening, rendering the hip and trunk joints practically inflexible. To examine the effect of this artificially increased stiffness on balance control, we perturbed the upright stance of young subjects (20-34 years of age) while they wore one of two types of corset or no corset at all. One type, the "half-corset", only increased hip stiffness, and the other, the "full-corset", increased stiffness of the hips and trunk. The perturbations consisted of combined roll and pitch rotations of the support surface (7.5 deg, 60 deg/s) in one of six different directions. Outcome measures were biomechanical responses of the legs, trunk, arms and head, and electromyographic (EMG) responses from leg, trunk, and upper arm muscles. With the full-corset, a decrease in forward stabilising trunk pitch rotation compared to the no-corset condition occurred for backward pitch tilts of the support surface. In contrast, the half-corset condition yielded increased forward trunk motion. Trunk backward pitch motion after forwards support-surface perturbations was the same for all corset conditions. Ankle torques and lower leg angle changes in the pitch direction were decreased for both corset conditions for forward pitch tilts of the support-surface but unaltered for backward tilts. Changes in trunk roll motion with increased stiffness were profound. After onset of a roll support-surface perturbation, the trunk rolled in the opposite direction to the support-surface tilt for the no-corset and half-corset conditions, but in the same direction as the tilt for the full-corset condition. Initial head roll angular accelerations (at 100 ms) were larger for the full-corset condition but in the same direction (opposite platform tilt) for all conditions. Arm roll movements were initially in the same direction as trunk movements, and were followed by large compensatory arm movements only for the full-corset condition. Leg muscle (soleus, peroneus longus, but not tibialis anterior) balance-correcting responses were reduced for roll and pitch tilts under both corset conditions. Responses in paraspinals were also reduced. These results indicate that young healthy normals cannot rapidly modify movement strategies sufficiently to account for changes in link flexibility following increases in hip and trunk stiffness. The changes in leg and trunk muscle responses failed to achieve a normal roll or pitch trunk end position at 700 ms (except for forward tilt rotations), even though head accelerations and trunk joint proprioception seemed to provide information on changed trunk movement profiles over the first 300 ms following the perturbation. The major adaptation to stiffness involved increased use of arm movements to regain stability. The major differences in trunk motion for the no-corset, half-corset and full-corset conditions support the concept of a multi-link pendulum with different control dynamics in the pitch and roll planes as a model of human stance. Stiffening of the hip and trunk increases the likelihood of a loss of balance laterally and/or backwards. Thus, these results may have implications for the elderly and others, with and without disease states, who stiffen for a variety of reasons.

Abdomen↗

The role of cutaneous receptors in the foot.

Cutaneous receptors in the foot sole appear to contribute to the control of human stance and locomotion. Two approaches were undertaken to establish the characteristics of the receptors in the sole. Psychophysical vibrotactile thresholds (range 25-400 Hz) were determined across the unloaded sole in young and elderly subjects. Thresholds were lower in the ball and arch of the sole, than in the heel and toe regions. Elderly subjects demonstrated significantly elevated thresholds for high-frequency vibration. Secondly, microneurographic recordings were made from skin afferents of the unloaded sole in young subjects. Results indicated that while similar types of cutaneous receptors exist in the sole of the foot and hand, there appear to be differences in receptor density and distribution. Our results demonstrate that cutaneous afferent inputs from the foot sole provide useful information for the control of posture and locomotion.

Adult↗

Control of the body vertical by vestibular and proprioceptive inputs.

The study examines the influence of vestibular and leg proprioceptive cues on the maintenance of the body vertical in human stance. Vestibular body orientation cues were changed by applying bipolar currents to both mastoid bones (cosine-bell wave form of 3.3 s duration, 1 mA current intensity). Proprioceptive input was modified by vibrating the tibialis anterior muscle (at f = 90 Hz, step of 5 s duration and 1 mm amplitude). Furthermore, the vestibular stimulus was paired with the muscle vibration using three different temporal relationships between the stimuli. Body lean responses were analyzed in terms of sway trajectories of the center of foot pressure on the body support surface (horizontal plane). With the anode on the right mastoid, vestibular body lean response was essentially straight towards the right side, and with the anode on left mastoid towards the left side. Vibration of right tibialis anterior muscle induced an almost straight body lean forward and to the right. Upon combined stimulation, responses with complex trajectory resulted, which depended on the stimulus interval. These responses reflected a superposition of the individual vestibular and proprioceptive effects. The results show that the body vertical is under the continuous control of leg proprioceptive and vestibular inputs, which sum linearly. We present a concept according to which these inputs are used for establishing a reference system for the control of the body vertical.

Adult↗

Simulating mechanical consequences of voluntary movement upon whole-body equilibrium: the arm-raising paradigm revisited.

Voluntary arm-raising movement performed during the upright human stance position imposes a perturbation to an already unstable bipedal posture characterised by a high body centre of mass (CoM). Inertial forces due to arm acceleration and displacement of the CoM of the arm which alters the CoM position of the whole body represent the two sources of disequilibrium. A current model of postural control explains equilibrium maintenance through the action of anticipatory postural adjustments (APAs) that would offset any destabilising effect of the voluntary movement. The purpose of this paper was to quantify, using computer simulation, the postural perturbation due to arm raising movement. The model incorporated four links, with shoulder, hip, knee and ankle joints constrained by linear viscoelastic elements. The input of the model was a torque applied at the shoulder joint. The simulation described mechanical consequences of the arm-raising movement for different initial conditions. The variables tested were arm inertia, the presence or not of gravity field, the initial standing position and arm movement direction. Simulations showed that the mechanical effect of arm-raising movement was mainly local, that is to say at the level of trunk and lower limbs and produced a slight forward displacement of the CoM (1.5 mm). Backward arm-raising movement had the same effect on the CoM displacement as the forward arm-raising movement. When the mass of the arm was increased, trunk rotation increased producing a CoM displacement in the opposite direction when compared to arm movement performed without load. Postural disturbance was minimised for an initial standing posture with the CoM vertical projection corresponding to the ankle joint axis of rotation. When the model was reduced to two degrees of freedom (ankle and shoulder joints only) the postural perturbation due to arm-raising movement increased compared to the four-joints model. On the basis of these results the classical assumption that APAs stabilise the CoM is challenged.

Arm↗

Characterisation of the quadriceps stretch reflex during the transition from swing to stance phase of human walking.

The main objective of this study was to characterize the stretch reflex response of the human thigh muscles to an unexpected knee flexion at the transition from stance to swing during walking. Eleven healthy subjects walked on a treadmill at their preferred speed. Reliable and constant knee flexions (6-12 degrees amplitude, 230-350 degrees /s velocity, 220 ms duration) were applied during the late swing and early stance phase of human walking by rotating the knee joint with a specifically designed portable stretch apparatus affixed to the left knee. Responses from rectus femoris (RF), vastus lateralis (VL), vastus medialis (VM), biceps femoris (BF), medial hamstrings (MH) and medial gastrocnemius (GM) were recorded via bipolar surface electromyograms (EMG). The onset of the response in the RF, VL and VM, remained stable and independent of the time in the step cycle when the stretch was applied. Across all subjects the response onset (mean +/- SD) occurred at 23+/-1, 24+/-1 and 23+/-1 ms for RF, VL and VM, respectively. The duration of the initial response was 90-110 ms, at which time the EMG signal returned towards baseline levels. Three reflex response windows, labelled the short latency reflex (SLR), the medium latency reflex (MLR) and the late latency reflex response (LLR), were analysed. The medium and late reflex responses of all knee extensors increased significantly ( p=0.008) as the gait cycle progressed from swing to stance. This was not related to the background EMG activity. In contrast, during standing at extensor EMG levels similar to those attained during walking the reflex responses were dependent on background EMG. During walking, LLR amplitudes expressed as a function of the background activity were on average two to three times greater than SLR and MLR reflex amplitudes. Distinct differences in SLR and LLR amplitude were observed for RF, VL and VM but not in the MLR amplitude. This may be related to the different pathways mediating the SLR, MLR and LLR components of the stretch response. As for the knee extensor antagonists, they exhibited a response to the stretch of the quadriceps at latencies short enough to be monosynaptic. This is in agreement with the suggestion by Eccles and Lundberg (1958) that there may be functional excitatory connections between the knee extensors and flexors in mammals.

Adult↗