Direct measurement of human ankle stiffness during quiet standing: the intrinsic mechanical stiffness is insufficient for stability

During quiet standing the human 'inverted pendulum' sways irregularly. In previous work where subjects balanced a real inverted pendulum, we investigated what contribution the intrinsic mechanical ankle stiffness makes to achieve stability. Using the results of a plausible model, we suggested that intrinsic ankle stiffness is inadequate for providing stability. Here, using a piezo-electric translator we applied small, unobtrusive mechanical perturbations to the foot while the subject was standing freely. These short duration perturbations had a similar size and velocity to movements which occur naturally during quiet standing, and they produced no evidence of any stretch reflex response in soleus, or gastrocnemius. Direct measurement confirms our earlier conclusion; intrinsic ankle stiffness is not quite sufficient to stabilise the body or pendulum. On average the directly determined intrinsic stiffness is 91 ± 23 % (mean ± s.d. ) of that necessary to provide minimal stabilisation. The stiffness was substantially constant, increasing only slightly with ankle torque. This stiffness cannot be neurally regulated in quiet standing. Thus we attribute this stiffness to the foot, Achilles' tendon and aponeurosis rather than the activated calf muscle fibres. Our measurements suggest that the triceps surae muscles maintain balance via a spring-like element which is itself too compliant to guarantee stability. The implication is that the brain cannot set ankle stiffness and then ignore the control task because additional modulation of torque is required to maintain balance. We suggest that the triceps surae muscles maintain balance by predictively controlling the proximal offset of the spring-like element in a ballistic-like manner.     

The passive, human calf muscles in relation to standing: the non-linear decrease from short range to long range stiffness

During human standing, tonic ankle extensor torque is required to support the centre of mass (CoM) forward of the ankles, and dynamic torque modulation is required to maintain unstable balance. Passive mechanisms contribute to both but the extent is controversial. Some groups have revealed a substantial intrinsic stiffness (65–90%) normalized to load stiffness, ' mgh '. Others regard their methodology as unsuitable for the low-frequency conditions of quiet standing and believe the passive contribution to be small (10–15%). Here we applied low-frequency ankle rotations to upright subjects who were supported at the waist allowing the leg muscles to be passive and we report normalized stiffness. The passive calf muscles provided: (i) an extensor torque capable of sustaining unstable balance without tonic activity at a mean CoM–ankle angle of 1.6 deg, (ii) a long range stiffness of 13 ± 2% and (iii) a short range (< 0.2 deg) stiffness of 67 ± 8%. Chordal ankle stiffness, derived from the torque versus angle relationship for 7 deg rotations, shows a non-linear decrease (stiffness α rotation^{−0.33±0.04}) from 101 ± 9% to 19 ± 5% for rotations of 0.03–7 deg, respectively. Thus, passive stiffness is well adapted for the continuum of postural and movement activity and has a substantial postural role eliminating the need for continuous muscle activity and increasing the unstable time constant of the human inverted pendulum. Ignoring the non-linear dependence of passive stiffness on sway size could lead to serious misinterpretation of experiments using perturbations and sensory manipulations such as eye closure, sway referencing and altered support surfaces.     

The frequency of human, manual adjustments in balancing an inverted pendulum is constrained by intrinsic physiological factors

While standing naturally and when manually or pedally balancing an equivalent inverted pendulum, the load sways slowly (characteristic unidirectional duration ∼1 s) and the controller, calf muscles or hand, makes more frequent adjustments (characteristic unidirectional duration 400 ms). Here we test the hypothesis that these durations reflect load properties rather than some intrinsic property of the human neuromuscular system. Using a specialized set-up mechanically analogous to real standing, subjects manually balanced inverted pendulums with different moments of inertia through a compliant spring representing the Achilles tendon. The spring bias was controlled by a sensitive joystick via a servo motor and accurate visual feedback was provided on an oscilloscope. As moment of inertia decreased, inverted pendulum sway size increased and it became difficult to sustain successful balance. The mean duration of unidirectional balance adjustments did not change. Moreover, the mean duration of unidirectional inverted pendulum sway reduced only slightly, remaining around 1 s. The simplest explanation is that balance was maintained by a process of manual adjustments intrinsically limited to a mean frequency of two to three unidirectional adjustments per second corresponding to intermittent control observed in manual tracking experiments. Consequently the inverted pendulum sway duration, mechanically related to the bias duration, reflects an intrinsic constraint of the neuromuscular control system. Given the similar durations of sway and muscle adjustments observed in real standing, we postulate that the characteristic duration of unidirectional standing sway reflects intrinsic intermittent control rather than the inertial properties of the body.     

Human balancing of an inverted pendulum with a compliant linkage: neural control by anticipatory intermittent bias

These experiments were prompted by the recent discovery that the intrinsic stiffness of the ankle is inadequate to stabilise passively the body in standing. Our hope was that showing how a large inverted pendulum was manually balanced with low intrinsic stiffness would elucidate the active control of human standing. The results show that the pendulum can be satisfactorily stabilised when intrinsic stiffness is low. Analysis of sway size shows that intrinsic stiffness actually plays little part in stabilisation. The sway duration is also substantially independent of intrinsic stiffness. This suggests that the characteristic sway of the pendulum, rather than being dictated by stiffness and inertia, may result from the control pattern of hand movements. The key points revealed by these experiments are that with low intrinsic stiffness the hand provides pendulum stability by intermittently altering the bias of the spring and, on average, the hand moves in opposition to the load. The results lead to a new and testable hypothesis; namely that in standing, the calf muscle shortens as the body sways forward and lengthens as it sways backwards. These findings are difficult to reconcile with stretch reflex control of the pendulum and are of particular relevance to standing. They may also be relevant to postural maintenance in general whenever the CNS controls muscles which operate through compliant linkages. The results also suggest that in standing, rather than providing passive stability, the intrinsic stiffness acts as an energy efficient buffer which provides decoupling between muscle and body.     

Neuromusculoskeletal torque-generation process has a large destabilizing effect on the control mechanism of quiet standing

The delay of the sensory-motor feedback loop is a destabilizing factor within the neural control mechanism of quiet standing. The purposes of this study were 1) to experimentally identify the neuromusculoskeletal torque-generation process during standing posture and 2) to investigate the effect of the delay induced by this system on the control mechanism of balance during quiet standing. Ten healthy adults participated in this study. The ankle torque, ankle angle, and electromyograms from the right lower leg muscles were measured. A ground-fixed support device was used to support the subject at his/her knees, without changing the natural ankle angle during quiet standing. Each subject was asked to mimic the ankle torque fluctuation by exerting voluntary ankle extension while keeping the supported standing posture. Using the rectified soleus electromyogram as the input and the ankle torque as the output, a critically damped, second-order system (twitch contraction time of 0.152 +/- 0.027 s) successfully described the dynamics of the torque-generation process. According to the performed Bode analysis, the phase delay induced by this torque-generation process in the frequency region of spontaneous body sway during quiet standing was considerably large, corresponding to an effective time delay of about 200 to 380 ms. We compared the stability of the balance control system with and without the torque-generation process and demonstrated that a much smaller number of gain combinations can stabilize the model with the torque-generation process than without it. We concluded that the phase delay induced by the torque-generation process is a more destabilizing factor in the control mechanism of quiet standing than previously assumed, which restricts the control strategies that can stabilize the entire system.     

Effect of strength and speed of torque development on balance recovery with the ankle strategy

In the event of an unexpected disturbance to balance, the ability to recover a stable upright stance should depend not only on the magnitude of torque that can be generated by contraction of muscles spanning the lower extremity joints but also on how quickly these torques can be developed. In the present study, we used a combination of experimental and mathematical models of balance recovery by sway (feet in place responses) to test this hypothesis. Twenty-three young subjects participated in experiments in which they were supported in an inclined standing position by a horizontal tether and instructed to recover balance by contracting only their ankle muscles. The maximum lean angle where they could recover balance without release of the tether (static recovery limit) averaged 14.9 +/- 1.4 degrees (mean +/- SD). The maximum initial lean angle where they could recover balance after the tether was unexpectedly released and the ankles were initially relaxed (dynamic recovery limit) averaged 5.9 +/- 1.1 degrees, or 60 +/- 11% smaller than the static recovery limit. Peak ankle torque did not differ significantly between the two conditions (and averaged 116 +/- 32 Nm), indicating the strong effect on recovery ability of latencies in the onset and subsequent rates of torque generation (which averaged 99 +/- 13 ms and 372 +/- 267 N. m/s, respectively). Additional experiments indicated that dynamic recovery limits increased 11 +/- 14% with increases in the baseline ankle torques prior to release (from an average value of 31 +/- 18 to 54 +/- 24 N. m). These trends are in agreement with predictions from a computer simulation based on an inverted pendulum model, which illustrate the specific combinations of baseline ankle torque, rate of torque generation, and peak ankle torque that are required to attain target recovery limits.     

Relative contribution of ankle and hip muscles in regulation of the human orthograde posture in a frontal plane

It was investigated, whether the postural regulation in the frontal plane takes place mainly at the hip or at the ankle level. The elimination of ankle torque was achieved by providing a point support in the frontal plane. (Two boards were attached to subject feet, below each board a metal pipe 2 cm diameter was fixed, so, the subject stood on contrivances resembling ‘skates’). The lateral displacements of breast and hip, the angle of ‘skates’ tilt and the characteristics of frontal stabilogram and electromyogram of two ankle muscles (m. peroneus and m. soleus) were compared in two situations: (1) during normal standing; (2) under the conditions of the exclusion of ankle torque from postural control. During normal standing the body behaved as two-link inverted pendulum. Transition from normal standing to standing on free ‘skates’ produced changes in the kinematics of body movement. Under the conditions of ankle torque exclusion (free ‘skates’) breast and hip of a subject moved in a frontal plane as a single unit (one-link inverted pendulum). During standing on free ‘skates’ the electromyographic activity of m. peroneus and m. soleus was the same as during normal standing (approximately 70–100 μV).     

Effect of the hip motion on the body kinematics in the sagittal plane during human quiet standing

Human quiet stance is often modeled as a single-link inverted pendulum pivoting only around the ankle joints in the sagittal plane. However, several recent studies have shown that movement around the hip joint cannot be negligible, and the body behaves like a double-link inverted pendulum. The purpose of this study was to examine how the hip motion affects the body kinematics in the sagittal plane during quiet standing. Ten healthy subjects were requested to keep a quiet stance for 30 s on a force platform. The angular displacements of the ankle and hip joints were measured using two highly sensitive CCD laser sensors. By taking the second derivative of the angular displacements, the angular accelerations of both joints were obtained. As for the angular displacements, there was no clear correlation between the ankle and hip joints. On the other hand, the angular accelerations of both joints were found to be modulated in a consistent anti-phase pattern. Then we estimated the anterior–posterior (A–P) acceleration of the center of mass (CoM) as a linear summation of the angular acceleration data. Simultaneously, we derived the actual CoM acceleration by dividing A–P share force by body mass. When we estimated CoM acceleration using only the angular acceleration of the ankle joint under the assumption that movement of the CoM is merely a scaled reflection of the motion of the ankle, it was largely overestimated as compared to the actual CoM acceleration. Whereas, when we take the angular acceleration of the hip joint into the calculation, it showed good coincidence with the actual CoM acceleration. These results indicate that the movement around the hip joint has a substantial effect on the body kinematics in the sagittal plane even during quiet standing.     

Human postural sway results from frequent, ballistic bias impulses by soleus and gastrocnemius

It has been widely assumed for nearly a century, that postural muscles operate in a spring-like manner and that muscle length signals joint angle (the mechano-reflex mechanism). Here we employ automated analysis of ultrasound images to resolve calf muscle (soleus and gastrocnemius) length changes as small as 10 μm in standing subjects. Previously, we have used balancing of a real inverted pendulum to make predictions about human standing. Here we test and confirm these predictions on 10 subjects standing quietly. We show that on average the calf muscles are actively adjusted 2.6 times per second and 2.8 times per unidirectional sway of the body centre of mass (CoM). These alternating, small (30–300 µm) movements provide impulsive, ballistic regulation of CoM movement. The timing and pattern of these adjustments are consistent with multisensory integration of all information regarding motion of the CoM, pattern recognition, prediction and planning using internal models and are not consistent with control solely by local reflexes. Because the system is unstable, errors in stabilization provide a perturbation which grows into a sway which has to be reacted to and corrected. Sagittal sway results from this impulsive control of calf muscle activity rather than internal sources (e.g. the heart, breathing). This process is quite unlike the mechano-reflex paradigm. We suggest that standing is a skilled, trial and error activity that improves with experience and is automated (possibly by the cerebellum). These results complement and extend our recent demonstration that paradoxical muscle movements are the norm in human standing.     

Influence of local sensory afference in the calibration of human balance responses

Summary Peripheral sensory modulation of balance behavior may require a calibrated mechanism which would maintain upright standing by a feedback control of torque at the ankle joint. The calibration of human balance was studied using a systematic presentation of perturbation excursions and velocities in normal freely standing subjects. All perturbations (posterior movements of a force platform) induced a forward body sway and were presented by first increasing and then decreasing the magnitude of perturbation. In preselected conditions the stability of the ankle and hence the accuracy of surface orientation inputs was altered using a foam base placed under the subjects feet. Each subject pressed a hand held response key at the moment a postural disturbance was detected. The automatic neuromuscular response (ANR) was recorded from the gastrocnemius muscles bilaterally and the perturbation detection time (DT) was obtained from the onset of thenar muscle discharge. The major findings in this study were: (1) Conscious DT changed as a function of step variations in perturbation excursion and was disassociated from the ANR latency. The ANR latency remained essentially constant in all conditions and did not have any influence on the kinematics of body sway. (2) Normalized peak body sway decreased during unstable ankle conditions and the reduction of body sway could be attributed to an increase in the gain of the ANR across a 200 ms integration period. The ANR 200 ms amplitude also showed higher correlations with perturbation magnitude during unstable (versus stable) ankle conditions. (3) The 200 ms gastrocnemius amplitude was modulated by excursion and velocity of platform displacement but the amplitude integrated over 100 ms was dependent on only the velocity of perturbation. Our results indicate that balance is controlled by a centrally initiated postural response but regulated in amplitude by local sensory information. These results establish that the gain of the ANR is functional, peripherally driven, and occurs subconsciously to alter the kinematics of body sway.     

The hallucinogen 1-[2,5-dimethoxy-4-iodophenyl]-2-aminopropane (DOI) increases cortical extracellular glutamate levels in rats

We examined how young and older adults adapt their posture to static balance tasks of increasing difficulty. Participants stood barefoot on a force platform in normal quiet, Romberg-sharpened and one-legged stance. Center of pressure (CoP) variations, electromyographic (EMG) activity of ankle and hip muscles and kinematic data were recorded. Both groups increased postural sway as a result of narrowing the base of support. Greater CoP excursions, EMG activity and joint displacements were noted in old compared to younger adults. Older adults displayed increased hip movement accompanied by higher hip EMG activity, whereas no similar increase was noted in the younger group. It is concluded that older adults rely more on their hip muscles when responding to self induced perturbations introduced by increased task constraints during quiet standing.     

Posturographic measures in healthy young adults during quiet sitting in comparison with quiet standing

Measures of postural steadiness – known as posturography – are commonly used for balance assessment during quiet standing. Although quiet sitting balance may be studied via posturography as well, this has not been done to date. As such, the purpose of this study was to characterize the posturography during quiet sitting in comparison with quiet standing and to provide a benchmark for future studies investigating differences in balance regulation and execution. Twelve young and healthy people agreed to quietly sit and stand on a force platform with their eyes open and closed. For each condition, one trial of 2 min was executed and the anterior–posterior, medial–lateral, and resultant distance fluctuations of the body's center of pressure (COP) were calculated. Finally, time-domain, frequency-domain, and stabilogram diffusion function (SDF) measures were identified and compared for all COP time series. The results consistently indicate that, for quiet sitting, the body sway size and velocity were smaller and the power-weighted average frequency larger than for quiet standing. Moreover, the SDF analysis revealed that quiet sitting shows fewer drifts over short time intervals, but also fewer controlled adjustments in the longer term to bring the system back to equilibrium. The observed differences can be partially explained by biomechanical and dynamic differences of the body portions that are in motion during quiet sitting and standing. The SDF analysis suggests, however, that also the balance control strategies are not identical. These findings may be especially useful for the assessment of sitting balance and the development of novel balance rehabilitation techniques and assistive devices.     

Paradoxical muscle movement in human standing

In human standing, gravity causes forward toppling about the ankle joint which is prevented by activity in the soleus and gastrocnemius muscles. It has long been assumed that when people sway forwards the calf muscles are stretched and conversely that they shorten with backward sway. Consequently, for many years, two explanations for standing stabilization have flourished. First, tonic muscle activity itself may generate adequate intrinsic ankle stiffness. Second, if intrinsic ankle stiffness is inadequate, the resistance to stretch of the calf muscles may be augmented by stretch reflexes or by central control. These explanations require that the passive tissue (Achilles' tendon, foot) transmitting the calf muscle tension is stiff. However, our recent measurements have indicated that this passive tissue is not stiff during standing. Accordingly, we predicted a counterintuitive mode of control where the muscles and body must, on average, move in opposite directions (paradoxical movements). Here we use dynamic ultrasound imaging in vivo with novel automated tracking of muscle length to test our hypothesis. We show that soleus and gastrocnemius do indeed move paradoxically, shortening when the body sways forward and lengthening when the body returns. This confirms that intrinsic ankle stiffness is too low to stabilize human standing. Moreover, it shows that the increase in active tension is associated with muscle shortening. This pattern cannot be produced by muscle stretch reflexes and can only arise from the anticipatory neural control of muscle length that is necessary for balance.     

A new paradigm for human stick balancing: a suspended not an inverted pendulum

We studied 14 skilled subjects balancing a stick (a television antenna, 52 cm, 34 g) on their middle fingertip. Comprehensive three-dimensional analyses revealed that the movement of the finger was 1.75 times that of the stick tip, such that the balanced stick behaved more like a normal noninverted pendulum than the inverted pendulum common to engineering models for stick balancing using motors. The average relation between the torque applied to the stick and its angle of deviation from the vertical was highly linear, consistent with simple harmonic motion. We observed clearly greater rotational movement of the stick in the anteroposterior plane than the mediolateral plane. Despite this magnitude difference, the duration of stick oscillatory cycles was very similar in both planes, again consistent with simple harmonic motion. The control parameter in balancing was the ratio of active torque applied to the stick relative to gravitational torque. It determined both the pivot point and oscillatory cycle period of the pendulum. The pivot point was located at the radius of gyration (about the centre of mass) of the stick from its centre of mass, showing that the subjects attuned to the gravitational dynamics and mass distribution of the stick. Hence, the key to controlling instability here was mastery of the physics of the unstable object. The radius of gyration may—similar to centre of mass—contribute to the kinesthesis of rotating limb segments and control of their gravitational dynamics.     

Slow dynamics of postural sway are in the feedback loop

Postural sway is considered to have two fundamental stochastic components, a slow nonoscillatory component and a faster damped-oscillatory component. The slow component has been shown to account for the majority of sway variance during quiet stance. Postural control is generally viewed as a feedback loop in which sway is detected by sensory systems and appropriate motor commands are generated to stabilize the body's orientation. Whereas the mechanistic source for the damped-oscillatory sway component is most likely feedback control of an inverted pendulum, the underlying basis for the slow component is less clear. We investigated whether the slow process was inside or outside the feedback loop by providing standing subjects with sum-of-sines visual motion. Linear stochastic models were fit to the experimental sway trajectories to determine the stochastic structure of sway as well as the transfer function from visual motion to sway. The results supported a fifth-order stochastic model, consisting of a slow process and two damped-oscillatory components. Importantly, the slow process was determined to be inside the feedback loop. This supports the hypothesis that the slow component is due to errors in state estimation because state estimation is inside the feedback loop rather than a moving reference point or an exploratory process outside the feedback loop.     

Larger center of pressure minus center of gravity in the elderly induces larger body acceleration during quiet standing

When an inverted pendulum approximates quiet standing, it is assumed that the distance between the center of pressure and the vertical projection of the center of mass on the ground (COP–COG) reflects the relationship between the controlling and controlled variables of the balance control mechanism, and that the center of mass acceleration (ACC) is proportional to COP–COG. As aging affects the control mechanism of balance during quiet standing, COP–COG must be influenced by aging and, as a result, ACC is influenced by aging as well. The purpose of this study was to test the hypotheses that aging results in an increased COP–COG amplitude and, as a consequence, that ACC becomes larger in the elderly than the young. Fifteen elderly and 11 young subjects stood quietly on a force platform with their eyes open or closed. We found that (1) the standard deviations of COP–COG and ACC were larger in the elderly than in the young, irrespective of the eye condition; (2) COP–COG is proportional to ACC in both age groups, i.e., the inverted pendulum assumption holds true for quiet standing. The results suggest that a change in the control strategy that is due to aging causes a larger COP–COG in the elderly and, as a consequence, that ACC becomes larger as well.     

Visual-vestibular interactions in postural control during the execution of a dynamic task

The purpose of this experiment was to determine the interaction between visual and vestibular information during the transition from quiet standing to the completion of a forward step. Six subjects were asked to take one step forward at the sound of an audio tone, with their eyes open or closed, and terminate the step in a standing position. During stimulation trials, galvanic vestibular stimulation (GVS) was delivered 1500 ms before the auditory cue. GVS was delivered at an intensity three-fold that of each subject's quiet stance threshold with either stimulus right, left or no stimulation. Force data were collected from three forceplates for the calculation of centre of pressure (CoP), and kinematic data were used to calculate centre of mass (CoM) and body trajectories. In quiet stance all subjects responded to the GVS perturbation by demonstrating upper body segment roll and whole body sway towards the anode electrode. Unexpectedly, in the presence of vision during quiet stance, the upper body roll response was not attenuated, even though the CoP sway patterns were reduced when vision was available. During the initiation phase of the step, despite ongoing GVS stimulation, there were no significant effects seen in CoM, CoP or upper body roll responses. During step execution, however, both CoM displacement and upper body roll demonstrated significant effects and both responses were significantly reduced when subjects' eyes were open. Analysis of the medio-lateral CoP integrals also indicated a strong stimulation effect between conditions late in the execution phase, which were largely attenuated with vision. The results suggest that the importance of visual and vestibular information varies depending on the phase of the task. In addition, the different integration between visual and vestibular input during quiet standing suggests a dual role for vestibular information. We propose that vestibular information in quiet standing has a role in maintaining whole body postural stability, as well as playing an integral role in the alignment of the body segments in preparation for proper movement execution. Vision was demonstrated to differentially attenuate these responses based on the phase of the task. Thus, visual and vestibular information appear to be integrated differently across the different phases of a forward-stepping task.     

Balance control under different passive contributions of the ankle extensors: quiet standing on inclined surfaces

Human bipedal stance is often modeled as a single inverted pendulum that pivots at the ankle joints in the sagittal plane. Because the center of body mass is usually maintained in front of the ankle joints, ankle extensor torque is continuously required to prevent the body from falling. During quiet standing, both passive and active mechanisms contribute to generate the ankle extensor torque counteracting gravity. This study aimed to investigate the active stabilization mechanism in more detail. Eight healthy subjects were requested to stand quietly on three different surfaces of 1) toes-up, 2) level, and 3) toes-down. Surface electromyogram (EMG) was recorded from the medial head of the gastrocnemius (MG), soleus (SOL), and tibialis anterior muscles. Inclination angle of the body was simultaneously measured. As a result, we found that EMG activities of MG and SOL were lowest during the toes-up standing and highest during the toes-down, indicating that increased (decreased) passive contribution required less (more) extensor torque generated by active muscle contraction. Frequency domain analysis also revealed that sway-related modulation of the ankle extensor activity (0.12–4.03 Hz) was unchanged among the three foot inclinations. On the other hand, isometric contraction strength of these muscles increased as the slope declined (toes-up < level < toes-down). These results support the idea that by regulating the isometric contraction strength, the CNS maintains a constant level of muscle tone and resultant ankle stiffness irrespective of the passive contribution. Such control scheme would be crucial when we consider the low bandwidth of the intermittent controller.     

Effects of 20-day bed rest with and without strength training on postural sway during quiet standing

Aim: To examine the effect of unweighting as a possible contributory factor to a reduced calf muscle volume on postural sway during quiet standing, changes in postural sway following bed rest with or without strength training were investigated. Methods: Twelve young men participated in a 20‐day bed‐rest study. Subjects were divided into a non‐training group (BR‐Con) and a strength training group (BR‐Tr). For the BR‐Tr group, training was comprised of dynamic calf‐raise and leg‐press exercises to maintain the muscle volume of the plantar flexors. Before and after bed rest, subjects maintained quiet standing in a barefoot position on a force platform with their eyes open or closed. During the quiet stance, foot centre‐of‐pressure (CoP) and the mean velocity of CoP was calculated. Muscle volume of the plantar flexors was computed using axial magnetic resonance images of the leg. Results: After the bed‐rest period, the muscle volume decreased in the BR‐Con group but not in the BR‐Tr group. The mean velocity of CoP as an assessment of postural sway, however, increased in both groups. These results indicate that the strength training during bed rest cannot counteract the increase in postural sway. Conclusion: We concluded that postural sway increases following 20 days of bed rest despite maintenance of the muscle volume of plantar flexors as the main working muscles for the human postural standing.     

Body oscillations in balancing due to segmental stretch reflex activity

Summary While subjects balanced on a seesaw consisting of a platform with a curved base, the antero-posterior sway of head and body as well as changes in the angle of the ankle joint were recorded and analysed for their frequency power spectrum. The EMG of leg muscles and the position of the resultant force exerted by the seesaw on a force-measuring platform were simultaneously registered and analysed. Balancing oscillations of 4–5 Hz were observed under this condition. They were accompanied by short, reciprocally organized bursts of EMG activity in the leg muscles. When stimulating the tibialis nerves to produce a displacement, the delay until the counterbalancing EMG activity started (about 40 ms) was in the time range of a fast-conducting segmental reflex. After partial ischaemic blocking of group I afferents from the leg muscles or fixation of the ankle joints, the predominant sway frequency was lacking, bursts of EMG activity became longer and stronger, and body balance was more unstable. Altering the height of the seesaw showed that a threshold change in the ankle angle was the determining factor in the production of spinal stretch reflex activity for fast regulation of balance.This work was supported by the Deutsche Forschungsgemeinschaft (SFB 70 — Hirnforschung und Sinnesphysiologie)