Human Postural Control.

From ancient Greece to nowadays, research on posture control was guided and shaped by many concepts. Equilibrium control is often considered part of postural control. However, two different levels have become increasingly apparent in the postural control system, one level sets a distribution of tonic muscle activity ("posture") and the other is assigned to compensate for internal or external perturbations ("equilibrium"). While the two levels are inherently interrelated, both neurophysiological and functional considerations point toward distinct neuromuscular underpinnings. Disturbances of muscle tone may in turn affect movement performance. The unique structure, specialization and properties of skeletal muscles should also be taken into account for understanding important peripheral contributors to postural regulation. Here, we will consider the neuromechanical basis of habitual posture and various concepts that were rather influential in many experimental studies and mathematical models of human posture control.

Tonic and Rhythmic Spinal Activity Underlying Locomotion.

In recent years, many researches put significant efforts into understanding and assessing the functional state of the spinal locomotor circuits in humans. Various techniques have been developed to stimulate the spinal cord circuitries, which may include both diffuse and quite specific tuning effects. Overall, the findings indicate that tonic and rhythmic spinal activity control are not separate phenomena but are closely integrated to properly initiate and sustain stepping. The spinal cord does not simply transmit information to and from the brain. Its physiologic state determines reflex, postural and locomotor control and, therefore, may affect the recovery of the locomotor function in individuals with spinal cord and brain injuries. This review summarizes studies that examine the rhythmogenesis capacity of cervical and lumbosacral neuronal circuitries in humans and its importance in developing central pattern generator-modulating therapies.

Tapping into rhythm generation circuitry in humans during simulated weightlessness conditions.

An ability to produce rhythmic activity is ubiquitous for locomotor pattern generation and modulation. The role that the rhythmogenesis capacity of the spinal cord plays in injured populations has become an area of interest and systematic investigation among researchers in recent years, despite its importance being long recognized by neurophysiologists and clinicians. Given that each individual interneuron, as a rule, receives a broad convergence of various supraspinal and sensory inputs and may contribute to a vast repertoire of motor actions, the importance of assessing the functional state of the spinal locomotor circuits becomes increasingly evident. Air-stepping can be used as a unique and important model for investigating human rhythmogenesis since its manifestation is largely facilitated by a reduction of external resistance. This article aims to provide a review on current issues related to the "locomotor" state and interactions between spinal and supraspinal influences on the central pattern generator (CPG) circuitry in humans, which may be important for developing gait rehabilitation strategies in individuals with spinal cord and brain injuries.

Lighten Up: Specific Postural Instructions Affect Axial Rigidity and Step Initiation in Patients With Parkinson's Disease.

BACKGROUND: Parkinson's disease (PD) is associated with stooped postural alignment, increased postural sway, and reduced mobility. The Alexander Technique (AT) is a mindfulness-based approach to improving posture and mobility by reducing muscular interference while maintaining upward intentions. Evidence suggests that AT can reduce disability associated with PD, but a mechanism for this effect has not yet been established. OBJECTIVE: We investigated whether AT-based instructions reduce axial rigidity and increase upright postural alignment, and whether these instructions have different effects on postural alignment, axial rigidity, postural sway, and mobility than effort-based instructions regarding posture. METHOD: Twenty subjects with PD practiced 2 sets of instructions and then attempted to implement both approaches (as well as a relaxed control condition) during quiet standing and step initiation. The "Lighten Up" instructions relied on AT principles of reducing excess tension while encouraging length. The "Pull Up" instructions relied on popular concepts of effortful posture correction. We measured kinematics, resistance to axial rotation, and ground reaction forces. RESULTS: Both sets of experimental instructions led to increases in upright postural alignment relative to the control condition. Only the Lighten Up instructions led to reduced postural sway, reduced axial postural tone, greater modifiability of tone, and a smoother center of pressure trajectory during step initiation, possibly indicating greater movement efficiency. CONCLUSION: Mindful movement approaches such as AT may benefit balance and mobility in subjects with PD by acutely facilitating increased upright postural alignment while decreasing rigidity.

Light and heavy touch reduces postural sway and modifies axial tone in Parkinson's disease.

BACKGROUND: Light touch with a stable object reduces postural sway by increasing axial postural tone in healthy subjects. However, it is unknown whether subjects with Parkinson's disease (PD), who have more postural sway and higher axial postural tone than healthy subjects, can benefit from haptic touch. OBJECTIVE: To investigate the effect of light and heavy touch on postural stability and hip tone in subjects with PD. METHODS: Fourteen subjects with mid-stage PD and 14 healthy control subjects were evaluated during quiet standing with eyes closed with their arms (a) crossed, (b) lightly touching a fixed rigid bar in front of them, and (c) firmly gripping the bar. Postural sway was measured with a forceplate, and axial hip tone was quantified using a unique device that measures the resistance of the hips to yaw rotation while maintaining active stance. RESULTS: Subjects with PD significantly decreased their postural sway with light or heavy touch (P < .001 vs arms crossed), similarly as control subjects. Without touch, hip tone was larger in PD subjects. With touch, however, tone values were similar in both groups. This change in hip tone with touch was highly correlated with the initial amount of tone (PD, r = -.72 to -.95; controls, r = -.74 to -.85). CONCLUSIONS: The authors showed, for the first time, that subjects with PD benefit from touch similarly to control subjects and that despite higher axial postural tone, PD subjects are able to modulate their tone with touch. Future studies should investigate the complex relationship between touch and postural tone.

Prolonged weight-shift and altered spinal coordination during sit-to-stand in practitioners of the Alexander Technique.

The Alexander Technique (AT) is used to improve postural and movement coordination and has been reported to be clinically beneficial, however its effect on movement coordination is not well-characterized. In this study we examined the sit-to-stand (STS) movement by comparing coordination (phasing, weight-shift and spinal movement) between AT teachers (n=15) and matched control subjects (n=14). We found AT teachers had a longer weight-shift (p<0.001) and shorter momentum transfer phase (p=0.01), than control subjects. AT teachers also increased vertical foot force monotonically, rather than unweighting the feet prior to seat-off, suggesting they generate less forward momentum with hip flexors. The prolonged weight-shift of AT teachers occurred over a greater range of trunk inclination, such that their weight shifted continuously onto the feet while bringing the body mass forward. Finally, AT teachers had greatly reduced spinal bending during STS (cervical, p<0.001; thoracic, p<0.001; lumbar, p<0.05). We hypothesize that the low hip joint stiffness and adaptive axial postural tone previously reported in AT teachers underlies this novel "continuous" STS strategy by facilitating eccentric contractions during weight-shift.

Contributions of skin and muscle afferent input to movement sense in the human hand.

In the stationary hand, static joint-position sense originates from multimodal somatosensory input (e.g., joint, skin, and muscle). In the moving hand, however, it is uncertain how movement sense arises from these different submodalities of proprioceptors. In contrast to static-position sense, movement sense includes multiple parameters such as motion detection, direction, joint angle, and velocity. Because movement sense is both multimodal and multiparametric, it is not known how different movement parameters are represented by different afferent submodalities. In theory, each submodality could redundantly represent all movement parameters, or, alternatively, different afferent submodalities could be tuned to distinctly different movement parameters. The study described in this paper investigated how skin input and muscle input each contributes to movement sense of the hand, in particular, to the movement parameters dynamic position and velocity. Healthy adult subjects were instructed to indicate with the left hand when they sensed the unseen fingers of the right hand being passively flexed at the metacarpophalangeal (MCP) joint through a previously learned target angle. The experimental approach was to suppress input from skin and/or muscle: skin input by anesthetizing the hand, and muscle input by unexpectedly extending the wrist to prevent MCP flexion from stretching the finger extensor muscle. Input from joint afferents was assumed not to play a significant role because the task was carried out with the MCP joints near their neutral positions. We found that, during passive finger movement near the neutral position in healthy adult humans, both skin and muscle receptors contribute to movement sense but qualitatively differently. Whereas skin input contributes to both dynamic position and velocity sense, muscle input may contribute only to velocity sense.

Progression of forearm intravenous regional anesthesia with ropivacaine.

The progression of sensory blockade in the hand following a forearm Bier block with ropivacaine is currently unknown. The hands of 10 healthy adult human subjects were anesthetized with ropivacaine, and their sensitivities to cold and touch were tested until the completion of anesthesia. On average, insensitivity to cold occurred uniformly throughout the hand within 9 mins; however, touch sensation was not complete until approximately 20 mins after injection. The spread of anesthesia occurred in a semisystematic way, spreading proximally and distally from the site of injection (mid-dorsum of the hand), and, at a slower rate, from the dorsum of the hand to the palm.

Method to measure tone of axial and proximal muscle.

The control of tonic muscular activity remains poorly understood. While abnormal tone is commonly assessed clinically by measuring the passive resistance of relaxed limbs, no systems are available to study tonic muscle control in a natural, active state of antigravity support. We have developed a device (Twister) to study tonic regulation of axial and proximal muscles during active postural maintenance (i.e. postural tone). Twister rotates axial body regions relative to each other about the vertical axis during stance, so as to twist the neck, trunk or hip regions. This twisting imposes length changes on axial muscles without changing the body's relationship to gravity. Because Twister does not provide postural support, tone must be regulated to counteract gravitational torques. We quantify this tonic regulation by the restive torque to twisting, which reflects the state of all muscles undergoing length changes, as well as by electromyography of relevant muscles. Because tone is characterized by long-lasting low-level muscle activity, tonic control is studied with slow movements that produce "tonic" changes in muscle length, without evoking fast "phasic" responses. Twister can be reconfigured to study various aspects of muscle tone, such as co-contraction, tonic modulation to postural changes, tonic interactions across body segments, as well as perceptual thresholds to slow axial rotation. Twister can also be used to provide a quantitative measurement of the effects of disease on axial and proximal postural tone and assess the efficacy of intervention.

Reduced performance in balance, walking and turning tasks is associated with increased neck tone in Parkinson's disease.

Rigidity or hypertonicity is a cardinal symptom of Parkinson's disease (PD). We hypothesized that hypertonicity of the body axis affects functional performance of tasks involving balance, walking and turning. The magnitude of axial postural tone in the neck, trunk and hip segments of 15 subjects with PD (both ON and OFF levodopa) and 15 control subjects was quantified during unsupported standing in an axial twisting device in our laboratory as resistance to torsional rotation. Subjects also performed six functional tests (walking in a figure of eight [Figure of Eight], Timed Up and Go, Berg Balance Scale, supine rolling task [rollover], Functional Reach, and standing 360-deg turn-in-place) in the ON and OFF state. Results showed that PD subjects had increased tone throughout the axis compared to control subjects (p=0.008) and that this increase was most prominent in the neck. In PD subjects, axial tone was related to functional performance, but most strongly for tone at the neck and accounted for an especially large portion of the variability in the performance of the Figure of Eight test (r(OFF)=0.68 and r(ON)=0.74, p<0.05) and the Rollover test (r(OFF)=0.67 and r(ON)=0.55, p<0.05). Our results suggest that neck tone plays a significant role in functional mobility and that abnormally high postural tone may be an important contributor to balance and mobility disorders in individuals with PD.

Parkinson's disease shows perceptuomotor asymmetry unrelated to motor symptoms.

Evidence of neglect symptoms in Parkinson's disease (PD) have been reported during visuoperceptual tasks and linked to side of disease onset. The goal of this study was to determine if in PD perceptual asymmetry is also evident in perceptuomotor tasks without visual input. The task was to point to a remembered straight-ahead (SA) target in peripersonal space. During baseline pointing, a bias left of SA was evident in PD patients and right of SA in healthy controls. To evaluate whether this was linked to a proprioceptive bias in PD, pointing during axial twisting of the trunk was tested. Axial rotation (+/-15 degrees , 1 degrees s(-1)) of the lower-body about shoulders fixed against rotation induced a non-veridical perception of upper-body rotation and lower-body stationarity. Pointing endpoints were shifted right of the actual SA during clockwise (CW) lower-body rotation and left of SA during counter-clockwise (CC) rotation, despite the fact that the shoulders and head were not rotated. In PD patients, endpoints relative to SA were shifted less during CW than CC rotation of the lower-body, whereas controls showed symmetrical pointing. Levodopa did not significantly change this bias. Both hands were tested in each subject and bias appeared regardless of hand used. Neither disease progression nor side of disease onset was linked to the direction or size of pointing bias. These findings suggest that PD manifests a contraction of left external hemispace relative to right hemispace, which affects generation and execution of motor commands throughout disease progression.

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.

Scaling and non-scaling of muscle activity, kinematics, and dynamics in sit-ups with different degrees of difficulty.

The purpose of this study was to investigate how the CNS adjusts motor patterns for variants of a complex axial movement-the sit-up. Adjustments were induced by changing the support surface contact and mass distribution of the body. Healthy adults performed straight-legged sit-ups, 3 s in duration, with support added to or removed from the lumbar trunk, or with mass added to the head or to the legs. Each of these interventions either increased or decreased the difficulty of the task. The study addressed the extent to which changes in sit-up difficulty are compensated by scaling of muscle activity, kinematics, and dynamics versus the extent to which they are compensated by changing discretely the motor pattern. The analysis of muscle activity, kinematics, and dynamics focused on the first 30-40% of the sit-up-the trunk flexion phase-since this is the most critical part of the movement. Our results demonstrate that, in some respects, sit-up kinematics and dynamics scaled with difficulty, but in other respects, they did not. Muscle activity also scaled, in many respects, but in more difficult sit-ups, abdominal flexor activity decreased instead of increased. Non-scaling changes in these parameters suggest that complex movements, such as the sit-up, may require discrete changes in motor pattern in order to deal with large loads, which challenge the available leverage.

Motor coordination can be fully understood only by studying complex movements.

In this chapter, we use the sit-up to illustrate the complexity of coordination in movements that involve many muscles, joints, degrees of freedom, and high levels of muscle activity. Complex movements often involve the body axis. In addition to the intentional, focal part of any voluntary movement, complex movements also include "associated movements" that are not consciously controlled, but are necessary for the movement to succeed. Some associated movements serve a purpose, and others may not. During sitting up, the leg-lift is a purposive associated movement, whereas three-joint flexion is a non-purposive associated movement. The control of complex movements is also likely to be complex and, we argue, is hierarchically controlled. Associated movements may, themselves, be hierarchically organized and triggered by lower brain structures, local changes in neuronal excitability, and sensory feedback. Complex movements typically involve a high level of mobility. Because this mobility can lead to instability, anticipatory postural adjustments, a type of purposive associated movement, are commonly used to regulate posture. Thus, a number of important aspects of motor coordination can only be revealed by the study of complex movements.

Position sensitivity of human muscle spindles: single afferent and population representations.

The representation of joint position at rest and during movement was investigated in 44 muscle spindle primary afferents originating from the extensor carpi radialis brevis (ECRb) and extensor digitorum (ED) of normal human subjects. Position sensitivity was estimated for each afferent, and 43 of 44 were position sensitive. In each trial, six sequential ramp-and-hold movements (2-6 degrees, 2 degrees/s, total 24 degrees) flexed the relaxed wrist, beginning from the angle at which the afferent was just recruited. Joint position was represented by three specific features of afferent firing patterns: the steady-state firing rate during the 4-s hold period between ramps, the initial burst at the beginning of each ramp, and the ramp increase in firing rate later in the movement. The position sensitivity of the initial burst (1.27 +/- 0.90 pps/degree, mean +/- SD) was several times higher than that of the hold period (0.40 +/- 0.30 pps/degree) and not different from that of the ramp increase in firing rate (1.36 +/- 0.68 pps/degree). The wrist position sensitivities of ECRb and ED afferents were equivalent, as were their recruitment angles and angular ranges of position sensitivity. Muscle spindle afferents, both individually and as a population, were shown to represent static joint position via the hold rate and the initial burst. Afferents were recruited over the entire 110 degree range of wrist positions investigated; however, the angular range over which each feature represented joint position was extremely limited (approximately 15 degrees). The population response, based on the summed activity of the 43 afferents, was monotonically related to joint position, and it was strongly influenced by the afferent recruitment pattern, but less so by the position sensitivities of the individual afferents.