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PubMed2004

Neural coupling between upper and lower limbs during recumbent stepping.

During gait rehabilitation, therapists or robotic devices often supply physical assistance to a patient's lower limbs to aid stepping. The expensive equipment and intensive manual labor required for these therapies limit their availability to patients. One alternative solution is to design devices where patients could use their upper limbs to provide physical assistance to their lower limbs (i.e., self-assistance). To explore potential neural effects of coupling upper and lower limbs, we investigated neuromuscular recruitment during self-driven and externally driven lower limb motion. Healthy subjects exercised on a recumbent stepper using different combinations of upper and lower limb exertions. The recumbent stepper mechanically coupled the upper and lower limbs, allowing users to drive the stepping motion with upper and/or lower limbs. We instructed subjects to step with 1) active upper and lower limbs at an easy resistance level (active arms and legs); 2) active upper limbs and relaxed lower limbs at easy, medium, and hard resistance levels (self-driven); and 3) relaxed upper and lower limbs while another person drove the stepping motion (externally driven). We recorded surface electromyography (EMG) from six lower limb muscles. Self-driven EMG amplitudes were always higher than externally driven EMG amplitudes (P < 0.05). As resistance and upper limb exertion increased, self-driven EMG amplitudes also increased. EMG bursts during self-driven and active arms and legs stepping occurred at similar times. These results indicate that active upper limb movement increases neuromuscular activation of the lower limbs during cyclic stepping motions. Neurologically impaired humans that actively engage their upper limbs during gait rehabilitation may increase neuromuscular activation and enhance activity-dependent plasticity.

Adult↗
PubMed2004

A new interpretation of spontaneous sway measures based on a simple model of human postural control.

This study shows that center-of-pressure (COP) traces that closely resemble physiologically measured COP functions can be produced by an appropriate selection of model parameters in a simple feedback model of the human postural control system. Variations in the values of stiffness, damping, time delay, and noise level determine the values of 15 sway measures commonly used to characterize spontaneous sway. Results from model simulations indicate that there is a high degree of correlation among these sway measures, and the measures cluster into three different groups. Only two principal components accounted for about 92% of the variation among the different sway measures analyzed. This model can be used to formulate hypotheses regarding the cause of postural control deficits reported in the literature. This is accomplished using a multidimensional optimization procedure to estimate model parameters from a diverse set of spontaneous sway measures. These model parameters describe physiologically meaningful features of the postural control system as opposed to conventional sway measures that provide only a parametric description of sway. To show the application of this method, we applied it to published data of spontaneous sway from elderly subjects and contrasted it to the data of young healthy subjects. We found that modest increases in stiffness and damping and a fairly large increase in noise level with aging could account for the variety of sway measures reported in the literature for elderly subjects.

Age Factors↗
PubMed2002

Modifications of vestibular responses of individual reticulospinal neurons in lamprey caused by unilateral labyrinthectomy.

A postural control system in the lamprey is driven by vestibular input and maintains the dorsal-side-up orientation of the animal during swimming. After a unilateral labyrinthectomy (UL), the lamprey continuously rolls toward the damaged side. Normally, a recovery of postural equilibrium ("vestibular compensation") takes about 1 mo. However, illumination of the eye contralateral to UL results in an immediate and reversible restoration of equilibrium. Here we used eye illumination as a tool to examine a functional recovery of the postural network. Important elements of this network are the reticulospinal (RS) neurons, which are driven by vestibular input and transmit commands for postural corrections to the spinal cord. In this study, we characterized modifications of the vestibular responses in individual RS neurons caused by UL and the effect exerted on these responses by eye illumination. The activity of RS neurons was recorded from their axons in the spinal cord by chronically implanted electrodes, and spikes in individual axons were extracted from the population activity signals. The same neurons were recorded both before and after UL. Vestibular stimulation (rotation in the roll plane through 360 degrees ) and eye illumination were performed in quiescent animals. It was found that the vestibular responses on the UL-side changed only slightly, whereas the responses on the opposite side disappeared almost completely. This asymmetry in the bilateral activity of RS neurons is the most likely cause for the loss of equilibrium in UL animals. Illumination of the eye contralateral to UL resulted, first, in a restoration of vestibular responses in the neurons inactivated by UL and in an appearance of vestibular responses in some other neurons that did not respond to vestibular input before UL. These responses had directional sensitivity and zones of spatial sensitivity similar to those observed before UL. However, their magnitude was smaller than before UL. Second, the eye illumination caused a reduction of the magnitude of vestibular responses on the UL side. These two factors tend to restore symmetry in bilateral activity of RS neurons, which is the most likely cause for the recovery of equilibrium in the swimming UL lamprey. Results of this study are discussed in relation to the model of the roll control system proposed in our previous studies as well as in relation to the vestibular compensation.

Action Potentials↗
PubMed2006

Postural muscle tone in the body axis of healthy humans.

Across the entire human body, postural tone might play its most critical role in the body's axis because the axis joins the four limbs and head into a single functioning unit during complex motor tasks as well as in static postures. Although postural tone is commonly viewed as low-level, tonic motor activity, we hypothesized that postural tone is both tonically and dynamically regulated in the human axis even during quiet stance. Our results describe the vertical distribution of postural muscle tone in the neck, trunk, and hips of standing human adults. Each subject stood blindfolded on a platform that axially rotated the neck, trunk, or pelvis at 1 degrees /s and +/-10 degrees relative to the neutral position (i.e., facing forward). The measured resistance to axial rotation was highest in the trunk and lowest in the neck and was characterized by several nonlinear features including short-range stiffness and hysteresis. In half of the subjects, axial muscle activity was relatively constant during axial rotation, and in the other half, muscle activity was modulated by lengthening and shortening reactions, i.e., decreasing activity in lengthening muscles and increasing activity in shortening muscles, respectively. Axial resistance to rotation was reduced in subjects whose muscle activity was modulated. The results indicate that axial tone is modulated sensitively and dynamically, this control originates, at least in part, from tonic lengthening and shortening reactions, and a similar type of control appears to exist for postural tone in the proximal muscles of the arm.

Adult↗
PubMed2005

Two kinematic synergies in voluntary whole-body movements during standing.

We used a particular computational approach, the uncontrolled manifold hypothesis, to investigate joint angle covariation patterns during whole-body actions performed by standing persons. We hypothesized that two kinematic synergies accounted for the leg/trunk joint covariation across cycles during a rhythmic whole-body motion to stabilize two performance variables, the trunk orientation in the external space and the horizontal position of the center of mass (COM). Subjects stood on a force plate and performed whole-body rhythmic movements for 45 s under visual feedback on one of the four variables, the position of the center of pressure or the angle in one of the three joints (ankle, knee, or hip). The Fitts-like paradigm was used with two target amplitudes and six indices of difficulty (ID) for each of the four variables. This was done to explore the robustness of kinematic postural synergies. A speed-accuracy trade-off was observed in all feedback conditions such that the movement time scaled with ID and the scaling differed between the two movement amplitudes. Principal-component (PC) analysis showed the existence of a single PC in the joint space that accounted for over 95% of the joint angle variance. Analysis within the uncontrolled manifold hypothesis has shown that data distributions in the joint angle space were compatible with stabilization of both trunk orientation and COM location. We conclude that trunk orientation and the COM location are stabilized by co-varied changes of the major joint angles during whole-body movements. Despite the strong effects of movement amplitude and ID on performance, the structure of the joint variance showed only minor dependence on these task parameters. The two kinematic synergies (co-varied changes in the joint angles that stabilized the COM location and trunk orientation) have proven to be robust over a variety of tasks.

Adaptation, Physiological↗
PubMed2005

A role for hip position in initiating the swing-to-stance transition in walking cats.

In this investigation, we obtained data that support the hypothesis that afferent signals associated with hip flexion play a role in initiating the swing-to-stance transition of the hind legs in walking cats. Direct evidence came from observations in walking decerebrate cats. Assisting the flexion of the hip joint during swing advanced the onset of activity in ankle extensor muscles, and this advance was strongly correlated with a reduction in the duration of hip flexor muscle activity. The hip angle at the time of onset of the flexion to extension transition was similar during assisted and unassisted steps. Additional evidence for the hypothesis that sensory signals related to hip flexion are important in regulating the swing-to-stance transition came from four normal animals trained to walk in a variety of situations designed to alter the coordination of movements at the hip, knee, and ankle joints during the swing phase. Although there were exceptions in some tasks and preparations, the angle of the hip joint at the time of onset of extensor activity was generally less variable than that of the knee and ankle joints. We also found no clear relationships between the angle of the limb and body axes, or the length of the limb axis, and the time of onset of extensor activity. Finally, there were no indications that the stretching of ankle extensor muscles during swing was a factor in regulating the transition from swing-to-stance.

Animals↗
PubMed2003

Dynamic regulation of sensorimotor integration in human postural control.

Upright stance in humans is inherently unstable, requiring corrective action based on spatial-orientation information from sensory systems. One might logically predict that environments providing access to accurate orientation information from multiple sensory systems would facilitate postural stability. However, we show that, after a period in which access to accurate sensory information was reduced, the restoration of accurate information disrupted postural stability. In eyes-closed trials, proprioceptive information was altered by rotating the support surface in proportion to body sway (support surface "sway-referencing"). When the support surface returned to a level orientation, most subjects developed a transient 1-Hz body sway oscillation that differed significantly from the low-amplitude body sway typically observed during quiet stance. Additional experiments showed further enhancement of the 1-Hz oscillation when the surface transitioned from a sway-referenced to a reverse sway-referenced motion. Oscillatory behavior declined with repetition of trials, suggesting a learning effect. A simple negative feedback-control model of the postural control system predicted the occurrence of this 1-Hz oscillation in conditions where too much corrective torque is generated in proportion to body sway. Model simulations were used to distinguish between two alternative explanations for the excessive corrective torque generation. Simulation results favor an explanation based on the dynamic reweighting of sensory contributions to postural control rather than a load-compensation mechanism that scales torque in proportion to a fixed combination of sensory-orientation information.

Adaptation, Physiological↗
PubMed2005

Impairment and recovery of postural control in rabbits with spinal cord lesions.

The aim of this study was to characterize impairment and subsequent recovery of postural control after spinal cord injuries. Experiments were carried out on rabbits with three types of lesion--a dorsal (D), lateral (L), or ventral (V) hemisection (HS) at T(12) level. The animals were maintaining equilibrium on a platform periodically tilted in the frontal plane. We assessed the postural limb/trunk configuration from video recordings and postural reflexes in the hindquarters from kinematical and electromyographic (EMG) recordings. We found that for a few days after DHS or LHS, the animals were not able to maintain the dorsal-side-up position of their hindquarters. This ability was then gradually restored, and the dynamic postural reflexes reached the prelesion value within 2-3 wk. By contrast, a VHS almost completely abolished postural reflexes, and they did not recover for > or =7 wk. The DHS, LHS, and VHS caused immediate and slowly compensated changes in the postural limb/trunk configuration as well as gradually developing changes. After DHS, both hind limbs were placed in an abnormal rostral and medial position. After LHS, the limb on the undamaged side was turned inward and occurred at the abnormal medial position; LHS also caused a gradually developing twisting of the caudal trunk. VHS caused gradually developing extension of the ankle and knee joints. These findings show that ventral spinal pathways are of crucial importance for postural control. When a part of these pathways is spared, postural reflexes can be restored rapidly, but not the postural limb/trunk configuration. Spinal and supraspinal mechanisms responsible for postural deficits and their compensation are discussed.

Animals↗
PubMed2002

Strategies for dynamic stability during locomotion on a slippery surface: effects of prior experience and knowledge.

Falls due to slips are prevalent in everyday life. The purpose of this study was to determine the reactive recovery responses used to maintain dynamic stability during an unexpected slip, establish the time course of response adaptation to repeated slip perturbations, and distinguish the proactive strategies for negotiating a slippery surface. Twelve young adults participated in the study in which a slip was generated following foot contact on a set of steel free-wheeling rollers. Surface electromyographic (EMG) data were collected from rectus femoris, biceps femoris, tibialis anterior, and the medial head of gastrocnemius on the perturbed limb. Whole body kinematics were recorded using an optical imaging system: from this the center of mass, foot angle, and medial-lateral stability margins were determined. In addition, braking/loading and accelerating/unloading impulses while in contact with the rollers and the rate of loading the rollers were determined from ground reaction forces. Results demonstrate that the reactive recovery response to the first slip consisted of a rapid onset of a flexor synergy (146-199 ms), a large arm elevation strategy, and a modified swing limb trajectory. With repeated exposure to the slip perturbation, the CNS rapidly adapts within one slip trial through global changes. These changes include the attenuation of muscle response magnitude, reduced braking impulse, landing more flat-footed, and elevating the center of mass. Individuals implement a "surfing strategy" while on the rollers when knowledge of the surface condition was available before hand. Furthermore, knowledge of a slip results in a reduced braking impulse and rate of loading, a shift in medial-lateral center of mass closer to the support limb at foot contact on the rollers and a more flat foot landing. In conclusion, prior experience with the perturbations allows subsequent modification and knowledge of the surface condition results in proactive adjustments to safely traverse the slippery surface.

Adaptation, Physiological↗
PubMed2006

Context-dependent modulation of interlimb cutaneous reflexes in arm muscles as a function of stability threat during walking.

Cutaneous reflexes evoked in the muscles of the arms with electrical stimulation of nerves of the foot ("interlimb reflexes") are observed during walking. These reflexes have been suggested to coordinate the actions of the legs and arms when walking is disturbed. Recently, we showed that cutaneous reflexes evoked in the leg muscles after stimulation at the foot are modulated according to the level of postural threat during walking. We hypothesized that the amplitude of interlimb cutaneous reflexes would similarly be modulated when subjects walk in unstable environments. Subjects walked on a treadmill under four walking conditions: 1) normal; 2) normal with unpredictable anterior-posterior (AP) perturbations; 3) arms crossed; and 4) arms crossed with unpredictable AP perturbations. Interlimb reflexes evoked from electrical stimulation of the right superficial peroneal or sural nerves were recorded bilaterally, at four points of the step cycle. These reflexes were compared between conditions in which the arms were moving in a similar manner: 1) normal versus AP walking and 2) arms crossed versus arms crossed with AP perturbations. Differences in reflex amplitudes between arms-crossed conditions were observed in most upper limb muscles when subjects were perturbed while walking compared with undisturbed walking. This effect was less apparent when the arms were swinging freely. The results indicate that the strength of interlimb connections is influenced by the level of postural threat (i.e., the context of the behavior), thereby suggesting that these reflexes serve a functional link between the legs and arms during locomotion.

Adult↗
PubMed2006

Stimulus-dependent changes in the vestibular contribution to human postural control.

Humans maintain stable stance in a wide variety of environments. This robust behavior is thought to involve sensory reweighting whereby the nervous system adjusts the relative contribution of sensory sources used to control stance depending on environmental conditions. Based on prior experimental and modeling results, we developed a specific quantitative representation of a sensory reweighting hypothesis that predicts that a given reduction in the contribution from one sensory system will be accompanied by a corresponding increase in the contribution from different sensory systems. The goal of this study was to test this sensory-reweighting hypothesis using measures that quantitatively assess the relative contributions of the proprioceptive and graviceptive (vestibular) systems to postural control during eyes-closed stance in different test conditions. Medial/lateral body sway was evoked by side-to-side rotation of the support surface (SS) while simultaneously delivering a pulsed galvanic vestibular stimulus (GVS) through electrodes behind the ears. A model-based interpretation of sway evoked by SS rotations provided estimates of the proprioceptive weighting factor, Wp, and showed that Wp declined with increasing SS amplitude. If the sensory-reweighting hypothesis is true, then the decline in Wp should be accompanied by a corresponding increase in Wp, the graviceptive weighting factor, and responses to the GVS should increase in proportion to the value of Wp derived from responses to SS rotations. Results were consistent with the predictions of the proposed sensory-reweighting hypothesis. GVS-evoked sway increased with increasing SS amplitude, and Wp measures derived from responses to GVS and from responses to SS rotations were highly correlated.

Adult↗
PubMed2003

Thresholds for inducing protective stepping responses to external perturbations of human standing.

Standing subjects were unexpectedly pulled forward to identify a threshold boundary that evokes stepping in terms of the size of the pull relative to the base of support (BoS). Performances in a range of sensorimotor tests were correlated with the threshold boundary parameters. Younger and older subjects were studied to identify age-related changes in stepping and the threshold boundaries. The threshold boundary had a forward limit (T(L)) that, when crossed, always made subjects step no matter how slowly they were pulled. As velocity increased, the threshold position that produced a step shifted nearer to the ankles. Eventually a pull velocity was reached above which velocity had no further effect and a position threshold (T(H)) was identified behind which subjects never stepped. Thus the position threshold boundary for stepping is a posterior-going sigmoidal function of perturbation velocity. Older subjects stepped more than the young (69% vs. 40% of trials). For the older subjects, T(L) (91% vs. 107% BoS) and T(H) (59% vs. 72% BoS) were closer to the ankles, and the transition between T(L) and T(H) occurred at lower velocities (96% vs. 121% BoS.s(-1)). Across the entire study population many sensorimotor factors were associated with T(L) and T(H). However, these associations were not present when age was removed as a factor. Thus, although the older subjects use protective stepping more often, this cannot be attributed directly to the sensorimotor factors tested here. It can be explained by stepping as a triggered response to the perturbation event rather than later sensory input about body movement.

Adult↗
PubMed2004

Controlling human upright posture: velocity information is more accurate than position or acceleration.

The problem of how the nervous system fuses sensory information from multiple modalities for upright stance control remains largely unsolved. It is well established that the visual, vestibular, and somatosensory modalities provide position and rate (e.g., velocity, acceleration) information for estimation of body dynamics. However, it is unknown whether any particular property dominates when multisensory information is fused. Our recent stochastic analysis of postural sway during quiet stance suggested that sensory input provides more accurate information about the body's velocity than its position or acceleration. Here we tested this prediction by degrading major sources of velocity information through removal/attenuation of sensory information from vision and proprioception. Experimental measures of postural sway were compared with model predictions to determine whether sway behavior was indicative of a deficit in velocity information rather than position or acceleration information. Subjects stood with eyes closed on a support surface that was 1) fixed, 2) foam, or 3) sway-referenced. Six measures characterizing the stochastic structure of postural sway behaved in a manner consistent with model predictions of degraded velocity information. Results were inconsistent with the effect of degrading only position or acceleration information. These findings support the hypothesis that velocity information is the most accurate form of sensory information used to stabilize posture during quiet stance. Our results are consistent with the assumption that changes in sway behavior resulting from commonly used experimental manipulations (e.g., foam, sway-referencing, eyes closed) are primarily attributed to loss of accurate velocity information.

Acceleration↗
PubMed2005

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.

Adult↗
PubMed2005

Synchronization of motor units in human soleus muscle during standing postural tasks.

During standing posture, the soleus muscles acts to control sway in the anteroposterior (AP) direction. The soleus muscles bilaterally share a common function during standing tasks. We sought to determine whether common descending inputs, as evidenced by the synchronization of bilateral motor unit pairs, were employed as a strategy to control this common function. Single motor units were recorded from the soleus muscles in subjects who stood on adjacent force platforms for 5 min with their eyes open or closed. While standing with the eyes open, only 4/39 bilateral motor unit pairs showed significant synchronization. Similarly, only 3/36 motor unit pairs were significantly synchronized during the eyes closed task. The low incidence of synchronization was observed despite a high correlation in the amount of sway in the AP direction between legs in both the eyes open and eyes closed tasks (rho = 0.80 and rho = 0.83, respectively). When the extent of synchronization was assessed between pairs of motor units within the same leg with the eyes open, 10/12 pairs were synchronized. Furthermore, when pairs of soleus motor units were recorded both bilaterally and unilaterally during voluntary isometric ankle plantarflexion, only 4/30 bilateral pairs showed significant synchronization, whereas 19/24 unilateral pairs had significant synchronization. In this study, there was little evidence of the existence of synchronization between bilateral soleus motor unit pairs in either postural tasks or voluntary isometric contractions. In cases in which bilateral synchronization was observed, it was considerably weaker than the synchronization of motor units within a single soleus muscle. The results of this study reveal that it is rather uncommon for bilateral soleus motoneurons to receive common descending synaptic inputs, whereas two motoneurons within a single soleus muscle do.

Action Potentials↗
PubMed1979

Statoacoustic properties of utricular afferents.

1. We classified the utricular afferents on the basis of their spontaneous acitivity and responses to tilts and vibrations. 2. Type I afferents fire spontaneously in a regular pattern; their responses to tilts consist of a phasic-tonic change in firing rate. They may respond to vibrations by increasing or decreasing their rate and show no adaptation. 3. The spontaneous activity and the responses to tilts of type II are similar to those observed in type I afferents. The differences become apparent when the preparation is subjected to a vibrational stimulus, since type II neurons increase their firing rate regardless of the stimulus frequency and show adaptation. 4. Type III neurons have no spontaneous activity. They respond to tilts by firing during the transition from one position to the other. They respond to a vibrational stimulus with maintained firing and show no adaptation. 5. We studied the dynamic responses of each type of neuron. We used sensitivity curves for the study of type III afferents and proposed a statistical method to define gain curves for the study of the other types. 6. The gain curves generated by type I neurons reach their maximum at frequencies of stimulation close to the spontaneous rate of firing. 7. In the gain curves of type II afferents the maximum corresponds to frequencies higher than their spontaneous activity. 8. Sensitivity curves and gain curves give similar results for type III fibers. The sensitivity curves of these afferents were classified into four subtypes. 9. We studied the responses of the three types of afferents to bursts of sinusoidal vibrations. 10. We concluded that the properties of types I and II fibers are fit to carry information about movements and position of the head, but also transmit acoustical information. Type III fibers are more adapted to provide information about acoustical stimuli, but can also convey information about head movements.

Acoustic Stimulation↗