Increased dynamic regulation of postural tone through Alexander Technique training.

Gurfinkel and colleagues (2006) recently found that healthy adults dynamically modulate postural muscle tone in the body axis during anti-gravity postural maintenance and that this modulation is inversely correlated with axial stiffness. Our objective in the present study was to investigate whether dynamic modulation of axial postural tone can change through training. We examined whether teachers of the Alexander Technique (AT), who undergo "long-term" (3-year) training, have greater modulation of axial postural tone than matched control subjects. In addition, we performed a longitudinal study on the effect of "short-term" (10-week) AT training on the axial postural tone of individuals with low back pain (LBP), since short term AT training has previously been shown to reduce LBP. Axial postural tone was quantified by measuring the resistance of the neck, trunk and hips to small (±10°), slow (1°/s) torsional rotation during stance. Modulation of tone was determined by the torsional resistance to rotation (peak-to-peak, phase-advance, and variability of torque) and axial muscle activity (EMG). Peak-to-peak torque was lower (∼50%), while phase-advance and cycle-to-cycle variability were enhanced for AT teachers compared to matched control subjects at all levels of the axis. In addition, LBP subjects decreased trunk and hip stiffness following short-term AT training compared to a control intervention. While changes in static levels of postural tone may have contributed to the reduced stiffness observed with the AT, our results suggest that dynamic modulation of postural tone can be enhanced through long-term training in the AT, which may constitute an important direction for therapeutic intervention.

Haptic touch reduces sway by increasing axial tone.

It is unclear how haptic touch with a stable surface reduces postural sway. We hypothesized that haptic input enhances postural stability due to alterations in axial postural tone. We measured the influence of heavy and light touch (LT) of the hands on a stable bar on axial postural tone and postural sway during stance in 14 healthy adults. A unique "Twister" device measured hip torque by fixing the upper body in space while oscillating the surface in yaw ±10 at 1 deg/s. Subjects were tested while: (1) standing quietly with their arms at their sides, (2) lightly touching a rigid bar in front of them and (3) firmly gripping the bar. Horizontal and vertical sway was not restricted by the device's yaw fixation, therefore, the subjects remained in a state of active postural control during the three touch conditions. Haptic touch significantly increased hip postural tone by 44% during light touch, from 2.5±0.9 to 3.6±1.0 Nm (P=0.005), and by 40% during firm grip to 3.5±0.8 Nm (P=0.005). Increases in hip postural tone were associated with a reduction in postural sway (r=-0.55, P=0.001). This is the first study showing that axial postural tone can be modified by remote somatosensory input and provides a potential explanation for how light touch improves postural stability. Changes in subjects' perception from trunk to surface rotation when changing from no touch (NT) to haptic touch, suggests that the CNS changes from using a global, to a local, trunk reference frame for control of posture during touch. The increase of hip postural tone during touching and gripping can be explained as a suppression of hip muscle shortening reactions that normally assist axial rotation.

Axial kinesthesia is impaired in Parkinson's disease: effects of levodopa.

Integration of sensory and motor inputs has been shown to be impaired in appendicular muscles and joints of Parkinson's disease (PD) patients. As PD advances, axial symptoms such as gait and balance impairments appear, which often progresses to complete inability stand or walk unaided. The current study evaluates kinesthesia in the axial musculature of PD patients during active postural control to determine whether impairments similar to those found in the appendages are also present in the hip and trunk. Using axial twisting, we quantified the detection threshold and directional accuracy of the hip relative to the feet (i.e. Hip Kinesthesia) and the hip relative to the shoulders (i.e. Trunk Kinesthesia). The relation of kinesthetic threshold to disease progression as measured by UPDRS and the effect of levodopa treatment on kinesthesia were assessed in 12 PD compared to age-matched controls. Subjects stood unaided while passively twisted at a very low constant rotational velocity (1 degrees /s). The results showed that accuracy in determining the direction of axial twisting was reduced in PD relative to healthy control subjects in the hip (PD-ON: 81%; PD-OFF: 91%; CTL=96%) and trunk (PD-ON: 81%; PD-OFF: 88%; CTL=95%). Thresholds for perception of axial twisting were increased when PD subjects were ON levodopa versus OFF in both the hip (p<0.01) and the trunk (p<0.05). The magnitude of decrease in sensitivity due to being ON levodopa was significantly correlated with the increase in UPDRS motor scores (Hip: r=0.90, p<0.01 and Trunk: r=0.60, p<0.05). This effect was not significantly correlated with equivalent levodopa dosage. PD subjects with disease onset on the left side of their body showed significantly higher axial thresholds than subjects with right PD onset (p<0.05). In conclusion, deficits in axial kinesthesia seem to contribute to the functional impairments of posture and locomotion in PD. Although levodopa has been shown to improve appendicular kinesthesia, we observed the opposite in the body axis. These findings underscore the dissociable neurophysiological circuits and dopaminergic pathways that are known to innervate these functionally distinct muscle groups.

Axial hypertonicity in Parkinson's disease: direct measurements of trunk and hip torque.

A cardinal feature of Parkinson's disease (PD) is muscle hypertonicity, i.e. rigidity. Little is known about the axial tone in PD or the relation of hypertonia to functional impairment. We quantified axial rigidity to assess its relation to motor symptoms as measured by UPDRS and determine whether rigidity is affected by levodopa treatment. Axial rigidity was measured in 12 PD and 14 age-matched controls by directly measuring torsional resistance of the longitudinal axis to twisting (+/-10 degrees ). Feet were rotated relative to fixed hips (Hip Tone) or feet and hips were rotated relative to fixed shoulders (Trunk Tone). To assess tonic activity only, low constant velocity rotation (1 degrees /s) and low acceleration (<12 degrees /s(2)) were used to avoid eliciting phasic sensorimotor responses. Subjects stood during testing without changing body orientation relative to gravity. Body parts fixed against rotation could translate laterally within the boundaries of normal postural sway, but could not rotate. PD OFF-medication had higher axial rigidity (p<0.05) in hips (5.07 N m) and trunk (5.30 N m) than controls (3.51 N m and 4.46 N m, respectively), which did not change with levodopa (p>0.10). Hip-to-trunk torque ratio was greater in PD than controls (p<0.05) and unchanged by levodopa (p=0.28). UPDRS scores were significantly correlated with hip rigidity for PD OFF-medication (r values=0.73, p<0.05). Torsional resistance to clockwise versus counter-clockwise axial rotation was more asymmetrical in PD than controls (p<0.05), however, there was no correspondence between direction of axial asymmetry and side of disease onset. In conclusion, these findings concerning hypertonicity may underlie functional impairments of posture and locomotion in PD. The absence of a levodopa effect on axial tone suggests that axial and appendicular tones are controlled by separate neural circuits.

Interaction of involuntary post-contraction activity with locomotor movements.

Involuntary post-contraction muscle activity may occur after performing a strong long-lasting (about 30 s) isometric muscle contraction (Kohnstamm phenomenon). Here we examined how this putative excitatory state may interact with a locomotor movement. The subjects stood upright and were asked to oppose a rotational force applied to the pelvis for about 30 s either in the clockwise or in the counterclockwise direction. After that, they were asked to perform various motor tasks with the eyes closed. During quiet standing, we observed an involuntary post-contraction torsion of the trunk. During walking, the post-contraction facilitatory effect of body torsion was not overridden by the voluntary activity, but instead significantly influenced the forward locomotor program such that subjects walked along a curved trajectory in the direction of the preceding torsion. In contrast, we did not observe any rotational component when subjects were asked to step in place. We conclude that the post-contraction rotational aftereffect does not transfer to just any motor task but apparently manifests itself in those movements that incorporate the activated axial muscle synergy or rotational component. We argue that central excitability changes following the voluntary effort may contribute to the phenomenon and highlight the role of tonic influences in fine-tuning of the spinal cord.

Effect of slow, small movement on the vibration-evoked kinesthetic illusion.

The study reported in this paper investigated how vibration-evoked illusions of joint rotation are influenced by slow (0.3 degrees /s), small (2-4 degrees ) passive rotation of the joint. Normal human adults (n=15) matched the perceived position of the left ("reference") arm with the right ("matching") arm while vibration (50 pps, 0.5 mm) was applied for 30 s to the relaxed triceps brachii of the reference arm. Both arms were constrained to rotate horizontally at the elbow. Three experimental conditions were investigated: (1) vibration of the stationary reference arm, (2) slow, small passive extension or flexion of the reference arm during vibration, and (3) slow, small passive extension or flexion of the reference arm without vibration. Triceps brachii vibration at 50 pps induced an illusion of elbow flexion. The movement illusion began after several seconds, relatively fast to begin with and gradually slowing down to a stop. On average, triceps vibration produced illusory motion at an average latency of 6.3 s, amplitude of 9.7 degrees , velocity of 0.6 degrees /s, and duration of 16.4 s. During vibration, slow, small ( approximately 0.3 degrees /s, 1.3 degrees ) passive rotations of the joint dramatically enhanced, stopped, or reversed the direction of illusory movement, depending on the direction of the passive joint rotation. However, the subjects' perceptions of these passive elbow rotations were exaggerated: 2-3 times the size of the actual movement. In the absence of vibration, the subjects accurately reproduced these passive joint rotations. We discuss whether the exaggerated perception of slow, small movement during vibration is better explained by contributions of non muscle spindle Ia afferents or by changes in the mechanical transmission of vibration to the receptor.

The sit-up: complex kinematics and muscle activity in voluntary axial movement.

This paper describes the kinematics and muscle activity associated with the standard sit-up, as a first step in the investigation of complex motor coordination. Eight normal human subjects lay on a force table and performed at least 15 sit-ups, with the arms across the chest and the legs straight and unconstrained. Several subjects also performed sit-ups with an additional weight added to the head. Support surface forces were recorded to calculate the location of the center of pressure and center of gravity; conventional motion analysis was used to measure segmental positions; and surface EMG was recorded from eight muscles. While the sit-up consists of two serial components, 'trunk curling' and 'footward pelvic rotation', it can be further subdivided into five phases, based on the kinematics. Phases I and II comprise trunk curling. Phase I consists of neck and upper trunk flexion, and phase II consists of lumbar trunk lifting. Phase II corresponds to the point of peak muscle contraction and maximum postural instability, the 'critical point' of the sit-up. Phases III-V comprise footward pelvic rotation. Phase III begins with pelvic rotation towards the feet, phase IV with leg lowering, and phase V with contact between the legs and the support surface. The overall pattern of muscle activity was complex with times of EMG onset, peak activity, offset, and duration differing for different muscles. This complex pattern changed qualitatively from one phase to the next, suggesting that the roles of different muscles and, as a consequence, the overall form of coordination, change during the sit-up.

The effect of aging on dynamic position sense at the ankle.

The present study addressed whether dynamic position sense at the ankle--or sense of position and velocity during movement--shows a similar decline as a result of aging as previously described for static position sense and movement detection threshold. Additionally, the involvement of muscle spindle afferents in the possible age-related decline was studied. To assess dynamic position sense, blindfolded subjects had to open the hand briskly when the right ankle was rotating passively through a prescribed target angle. To assess the involvement of muscle spindles, the effect of tibialis anterior vibration was studied. The results showed that aging lead to a significant increase in deviation from the target angle at hand opening as well as in variability of performance. Vibration resulted in larger undershoot errors in the elderly compared to the young adults, suggesting that the age-related decline in performance on the dynamic position sense task is not (solely) due to muscle spindle function changes. Alternatively, this degeneration might be due to altered input from other sources of proprioceptive input, such as skin receptors. The elderly subjects did show a beneficial effect of practice with the task, which may provide solid fundaments for rehabilitation.

Coexistence of stability and mobility in postural control: evidence from postural compensation for respiration.

This study evaluated the extent to which movement of the lower limbs and pelvis may compensate for the disturbance to posture that results from respiratory movement of the thorax and abdomen. Motion of the neck, pelvis, leg and centre of pressure (COP) were recorded with high resolution in conjunction with electromyographic activity (EMG) of flexor and extensor muscles of the trunk and hip. Respiration was measured from ribcage motion. Subjects breathed quietly, and with increased volume due to hypercapnoea (as a result of breathing with increased dead-space) and a voluntary increase in respiration. Additional recordings were made during apnoea. The relationship between respiration and other parameters was measured from the correlation between data in the frequency domain (i.e. coherence) and from time-locked averages triggered from respiration. In quiet standing, small angular displacements ( approximately 0.5 degrees ) of the trunk and leg were identified in raw data. Correspondingly, there were peaks in the power spectra of the angular movements and EMG. While body movement and EMG were coherent with respiration (>0.5), the coherence between respiration and COP displacement was low (<0.2). The amplitude of movement and coherence was increased when respiration was increased. The present data suggest that the postural disturbance that results from respiratory movement is matched, at least partly, and counteracted by small angular displacements of the lower trunk and lower limbs. Thus, stability in quiet stance is dependent on movement of multiple body segments and control of equilibrium cannot be reduced to control of a single joint.

Position sense during imperceptibly slow movements.

Proprioceptive input provides the nervous system with information related to body position and movement. This study investigated how normal humans perceive information related to the position of a limb when it is either at rest or moving very slowly, below the threshold for movement perception. Each subject's left hand, hidden from view, was horizontally translated in the frontal plane such that joint rotation was largely isolated to the shoulder. The translation speed was too slow for the subjects to detect movement. Subjects indicated their perception of hand position at 1- or 2-min intervals by pointing with the right index finger to the perceived location of the tip of the left middle finger. The constant error (i.e., bias) and variable error (i.e., SD of mean constant error) of each pointing movement was quantified. Partway through the trial, the direction of hand movement was reversed. In two additional experimental conditions, the subjects pointed at 1- or 2-min intervals at the stationary left hand, either when they assumed the hand would be moved or when they knew it would not. During slow movement, errors in the frontal and sagittal planes were found to be independent, and therefore, data analysis focused on the frontal plane, as this was the plane relevant to the subjects' task. All subjects performed the task sufficiently well to demonstrate a clear perception of hand location during the slow movement. The accuracy of perception was better when the left (target) hand was in its ipsilateral hemifield and, correspondingly, when the right hand pointed to its contralateral hemifield. There was no significant difference in constant error when the hand moved slowly, although there was a slightly higher variable error during slow movement than when the hand was stationary. Based on the similarity of results in trials with very slow and no hand movement, it was concluded that position-sense percepts are more accurately distinguished by the speed of movement rather than whether movement is occurring or not.

The role of paraspinal muscle spindles in lumbosacral position sense in individuals with and without low back pain.

STUDY DESIGN: A two-group experimental design with repeated measures on one factor was used. OBJECTIVES: To investigate the role of paraspinal muscle spindles in lumbosacral position sense in individuals with and without low back pain. SUMMARY OF BACKGROUND DATA: Proprioceptive deficits have been identified in patients with low back pain. The underlying mechanisms, however, are not well documented. METHODS: Lumbosacral position sense was determined before, during, and after lumbar paraspinal muscle vibration in 23 young patients with low back pain and in 21 control subjects. Position sense was estimated by calculating the mean absolute error, constant error, and variable error between six criterion and reproduction sacral tilt angles. RESULTS: Repositioning accuracy was significantly lower in the patient group than in healthy individuals (absolute error difference between groups = 2.7 degrees, P < 0.0001). Multifidus muscle vibration induced a significant muscle-lengthening illusion that resulted in an undershooting of the target position in healthy individuals (constant error = -3.1 degrees, P < 0.0001). Conversely, the position sense scores of the patient group did not display an increase in negative directional error but a significant improvement in position sense during muscle vibration (P < 0.05). No significant differences in absolute error were found between the first and last trial in the healthy individuals (P >/= 0.05) and in the patient group (P > 0.05). CONCLUSIONS: Patients with low back pain have a less refined position sense than healthy individuals, possibly because of an altered paraspinal muscle spindle afference and central processing of this sensory input. Furthermore, muscle vibration can be an interesting expedient for improving proprioception and enhancing local muscle control.

Proprioceptive control of multijoint movement: unimanual circle drawing.

The present experiments addressed whether proprioception is used by the central nervous system (CNS) to control the spatial and temporal characteristics of unimanual circle drawing. Circle drawing is a multijoint movement, in which the muscles crossing the elbow and the shoulder are sequentially activated. The spatial and temporal characteristics of circle drawing depend on the precise coordination of these sequential activation patterns, and proprioception is ideally suited to support this coordination. Blindfolded human subjects produced a counterclockwise circular drawing motion (diameter = 16 cm) with the dominant arm at a repetition rate of 1/s. In some trials, 60-70 Hz vibration was applied to the tendons of the biceps brachii and/or the anterior deltoid. Spatial parameters measured from hand-movement data included the x- and y-axis diameters, circularity, and drift of the hand in the workspace. Vibration of either the biceps brachii or the anterior deltoid caused subjects to draw circles with decreased diameter, with changes in circularity, and with a systematic drift of the hand. These distortions to circle drawing by tendon vibration demonstrate that the CNS uses proprioceptive information to accomplish the spatial characteristics of this motor task. Simultaneous vibration of both muscles produced a drift that exceeded the individual vibration effects, which suggests that the CNS combined proprioceptive information related to elbow and shoulder rotation to control the movement of the hand. The temporal characteristics of circle drawing were quantified from joint angle data. While vibration did not significantly influence the relative phase between elbow and shoulder rotation, the variability of the phase relationship increased significantly, which suggests that proprioception contributes to phase stabilization. During circle drawing, elbow flexion-extension movements were produced with limited activation of the biceps. Nevertheless, biceps vibration distorted the circle metrics, suggesting that a muscle's significance as a sensory transducer is independent of its activity level.

Proprioceptive control of multijoint movement: bimanual circle drawing.

Proprioception is used by the central nervous system (CNS) in the control of the spatial and temporal characteristics of single joint and multiple joint movement. The present study addressed the role of proprioception in the control of bilateral cyclical movements of the limbs. Normal blindfolded human subjects drew circles simultaneously and symmetrically with the two arms (16 cm diameter, 1 /s) upon two digitizing tablets. In selected trials, vibration (60-70 Hz) was applied to the tendon of the biceps and/or anterior deltoid muscles of the dominant arm to distort the proprioceptive information from muscle spindle afferents. One goal of this study was to identify whether tendon vibration influenced the spatial characteristics of circles drawn by the vibrated, dominant arm and the non-vibrated, non-dominant arm. A second goal was to determine the effect of vibration on the temporal coupling between the two arms during circle drawing. The results revealed that tendon vibration affected the spatial characteristics of circles drawn by the vibrated arm in a manner similar to that previously found for unilateral circle drawing. During bimanual circle drawing, vibration had only a minimal effect on the spatial characteristics of the non-vibrated, non-dominant arm. Temporal interlimb coupling was quantified by the relative phasing between the arms. Without tendon vibration, the dominant arm led the non-dominant arm. Vibration of the dominant arm increased the average phase lead. In a first control experiment, vibration of the non-dominant arm decreased the phase lead of the dominant arm, or even reversed it to a non-dominant arm phase lead. In a second control experiment, the subjects performed the bimanual circle-drawing task with vision of only the vibrated arm, in which case there was no spatial distortion of the circles drawn by the vibrated arm, but the phase relation between the two arms was still shifted as if vision were completely unavailable. It was concluded that, in bimanual movements such as these, the spatial and temporal characteristics of movement are controlled independently. Whereas the spatial characteristics of hand movement seem to be controlled unilaterally, the temporal characteristics of interlimb coupling appear to be controlled by proprioceptive information from both limbs, possibly by a proprioceptive triggering mechanism.

Representation of wrist joint kinematics by the ensemble of muscle spindles from synergistic muscles.

Proprioceptive information about movement is transmitted to the central nervous system by a variety of receptor types, which are widely distributed among the muscles, joints, and skin. Muscle spindles are known to be an important and reliable source of information for the perception of movement kinematics. Previous studies that focused on the characteristics of single muscle spindle firing patterns have left the impression that each receptor fires in relation to a number of kinematic variables, leaving the following question unanswered: what role is played by the ensemble of muscle spindles within the same muscle or within synergistic muscles? The study described in this paper addressed whether the perception of joint position and velocity is based on the net input of muscle spindles residing in all synergistic muscles crossing a joint. Normal human adults performed a motor coordination task that required perception of joint velocity and dynamic position at the wrist. The task was to open the left hand briskly as the right wrist was passively rotated in the flexion direction through a prescribed target angle. In randomly occurring trials, the tendons to three muscles [extensor carpi radialis (ECR), extensor carpi ulnaris (ECU), and extensor digitorum (ED)] were vibrated either individually or in different combinations during the performance of the motor task. Tendon vibration is known to distort muscle spindle firing patterns, and consequently, kinesthesia. By comparing performance errors with and without tendon vibration, the relative influences of muscle spindles residing in ECR, ECU, and ED were quantified. Vibration of the individual ECR, ECU, or ED tendons produced systematic undershoot errors in performance, consistent with the misperception of wrist velocity and dynamic position. Performance errors were larger when combinations of, rather than individual, muscle tendons were vibrated. The error resulting from simultaneous vibration of ECR and ECU was roughly equal to the sum of the errors produced by vibration of the individual tendons. These effects of vibrating synergistic tendons at the wrist suggest that kinesthesia is derived from the integrated input of muscle spindles from all synergistic muscles.

Proprioceptive consequences of tendon vibration during movement.

1. Previous studies have used tendon vibration to investigate kinesthetic illusions in the isometric limb and end point control in the moving limb. These previous studies have shown that vibration distorts the perceptions of static joint angle and movement and causes systematic errors in the end point of movement. In this paper we describe the effects of tendon vibration during movement while human subjects performed a proprioceptively coordinated motor task. In an earlier study we showed that the CNS coordinates this motor task-a movement sequence-with proprioceptive information related to the dynamic position and velocity of the limb. 2. When performing this movement sequence, each subject sat at a table and opened the right hand as the right elbow was passively rotated in the extension direction through a prescribed target angle. Vision of the arm was prevented, and the movement velocity was changed randomly from trial to trial, leaving proprioception as the only useful source of kinematic information with which to perform the task. 3. In randomly occurring trials, vibration was applied to the tendon of the biceps brachii, a muscle that lengthens during elbow extension. In some experiments the timing of tendon vibration was varied with respect to the onset of elbow rotation, and in other experiments the frequency of vibration was varied. In each experiment we compared the accuracy of the subject's response (i.e., the elbow angle at which the subject opened the hand) in trials with tendon vibration with the accuracy in trials without tendon vibration. 4. The effect of tendon vibration depended on the frequency of vibration. When the biceps tendon was vibrated at 20 Hz, subjects opened the hand after the elbow passed through the target angle ("overshooting"). Overshooting is consistent with an underestimate of the actual displacement or velocity of the elbow. Vibration at 30 Hz had little or no effect on the elbow angle at hand opening. Vibration at 40 Hz caused subjects to open the hand before the elbow reached the target angle ("undershooting"). Undershooting is consistent with an overestimate of the actual displacement or velocity of the elbow. The size of the error depended on the velocity of the passively imposed elbow rotation. 5. The effect of tendon vibration also depended on the timing of vibration. If 40-Hz vibration began at the onset of movement, the subject undershot the target. If 40-Hz vibration started 5 s before movement onset and continued throughout the movement, the undershoot error increased in magnitude. However, if 40-Hz vibration started 5 s before movement onset and then stopped at movement onset, the subject overshot the target. When vibration was shut off during movement, a transition occurred from an over-shooting error to an undershooting error at a time that depended on the velocity of elbow rotation. 6. In a separate experiment, subjects were instructed to match either the perceived dynamic position or the perceived velocity of rotation imposed on the right elbow by actively rotating the left elbow. In both matching tasks, tendon vibration produced oppositely directed errors depending on the frequency of vibration. Vibration at 20 Hz produced a perception of decreased elbow velocity and a bias in dynamic position in the flexion direction, and vibration at 40 Hz produced the opposite perceptions. 7. We conclude that muscle spindle afferents, which are activated by tendon vibration, are an important source of the dynamic position and velocity information that the CNS uses to coordinate this movement sequence task. The observed effects of vibration timing and frequency suggest that perceptual changes evoked by vibration cannot be explained by the simple summation of sensory input evoked by movement and by vibration. Rather, the bias in perception produced by vibration appears to be related to the difference between vibration- and movement-evoked activity in muscle spindle afferents.

Proprioceptive coordination of discrete movement sequences: mechanism and generality.

A "discrete" movement sequence is defined as a movement with a single goal that involves a series of overlapping joint rotations. Reaching-and-grasping and throwing are examples of discrete movement sequences. The central nervous system (CNS) can use reafferent proprioceptive information from one joint rotation in a sequence to coordinate subsequent rotations at other joints. The experiments reported in this paper demonstrate how the human CNS uses proprioceptive information to coordinate discrete movement sequences. We examined the mechanism (at an information processing level) underlying proprioceptive coordination and the generality (i.e., the boundary conditions) of these mechanisms as they apply to everyday movement sequences. Adult human subjects performed a discrete movement sequence that resembles backhand throwing: elbow extension followed by hand opening. The task was to open the hand as the elbow passed through a prescribed "target" angle. We eliminated visual information and made the arrival time at the target angle unpredictable so that the available kinematic information was provided exclusively by proprioception. The subjects were capable of performing this motor task with a high degree of precision, thereby demonstrating that the nervous system can use proprioceptive input to coordinate discrete movement sequences. Our data indicate that precise coordination is achieved by extracting kinematic information related to both the velocity of elbow rotation as well as the elbow position during movement (i.e., "dynamic position"). Dynamic position information appears to be encoded as both absolute joint angle and angular distance, although more precisely as angular distance. Although our experiments were conducted under rather restrictive laboratory conditions, this mechanism of motor coordination might also apply to everyday movement. Our results suggest that this mechanism could be employed for passive as well as active movement sequences, with and without opposing loads; it could exert its influence in discrete movement sequences as brief as 210 ms or as long as 1.5 s; and it does not involve any significant degree of learning (this proprioceptive mechanism appears to be readily available for use on the first attempt of a novel motor task).

Fusimotor reflexes in relaxed forearm muscles produced by cutaneous afferents from the human hand.

1. This study was designed to determine whether cutaneous receptors in the hand exert reflex effects on fusimotor neurones innervating relaxed muscles. Recordings were made from fifty-four muscle spindle afferents in the radial nerve while the arm was held relaxed in a supporting frame. Cutaneous afferents were activated by trains of stimuli at non-noxious levels to the superficial radial nerve or to the palmar surface of the fingers. 2. For the population of muscle spindle afferents, the mean discharge rate was 7.1 +/- 6.4 Hz (range 0-24 Hz). Thirty-three per cent had no background discharge, and this occurred significantly more often in finger extensors than wrist extensors. 3. Trains of cutaneous stimuli produced no change in the discharge rates of the majority of spindle endings irrespective of whether the spindle afferent had a background discharge or was given one by muscle stretch. However, with two of forty afferents, the stimuli produced an increase in discharge at latencies of 135 and 155 ms. 4. With a further fourteen muscle spindle endings, the dynamic responses to stretch were measured 100-400 ms after the trains of cutaneous stimuli. For four spindle afferents there was a statistically significant change in the dynamic response to stretch occurring at conditioned-stretch intervals of 100-200 ms. For two afferents the dynamic response decreased by 17 and 26% and for two others it increased by about 24 and 37%. 5. While these results support the view that the level of background fusimotor drive is low in the relaxed state, they suggest that there is some dynamic fusimotor drive to completely relaxed muscles operating on the human hand, and that this drive can be altered reflexly by cutaneous afferent inputs from the hand.

The measurement of tremor using simple laser systems.

Quantification of movement pathologies is an important challenge of the clinical and research laboratories today. Basically, two problems must be addressed. The first one is to find the appropriate technology; the second is to develop adequate measures from the raw data which will best discriminate between health and pathology. In this paper, we propose a simple method to record and analyse tremor and other microdisplacements of the upper extremities based on the recording of position by laser analog sensors. Any microcomputer equipped to perform analog-digital conversion is compatible for use with this system. The performance of the laser system is examined and compared with the performance of accelerometers. Finally, data recorded with a laser analog sensor from a patient with Parkinson's disease and a control subject are presented. This new laser-based quantitative method may prove to be an important tool in early and differential diagnosis of neurodegenerative diseases of the central nervous system.

Proprioceptive coordination of movement sequences: discrimination of joint angle versus angular distance.

1. The purpose of these experiments was to determine the accuracy with which human subjects could discriminate proprioceptive (nonretinotopic) targets during movement. The targets were located at either a specified angle in joint space, or a specified angular distance from an initial joint angle. 2. In these experiments the right elbows of normal human subjects were passively extended from either predictable or unpredictable starting angles. The subjects were instructed to open the right hand to indicate that the elbow was passing through a target joint angle or a target angular distance. The subjects were not given visual information about the location of the elbow, so they had to rely on proprioceptive input to perform this task. The target (criterion joint angle or angular distance) was learned by the use of proprioception during 8-15 practice trials. 3. Four experiments were conducted. In three experiments the target was located at a constant joint angle, and in the fourth experiment the target was located at a constant angular distance from the starting angle. The starting angle in all four experiments was pseudorandomly varied from trial to trial. 4. On the basis of an analysis of constant errors, subjects were more accurate at discriminating angular distance than joint angle. The slope of the relationship between the starting position and the constant errors was dictated by the task requirement. 5. In the distance discrimination experiment, when the starting angle was more flexed than the intermediate (i.e., central) position, the subjects slightly overshot the target distance. Conversely, when the starting angle was more extended than the intermediate position, the subjects slightly undershot the target distance. 6. In the joint angle discrimination experiments, the opposite results were obtained. Subjects overshot the target when the starting position of the elbow was more extended than the intermediate starting position, and they undershot the target when the starting position was more flexed than the intermediate starting position. The amplitude of these systematic errors increased when the subjects were unaware that the initial angle of their elbow was variable. 7. It is concluded that, in kinesthetic tasks of this type, the discrimination of angular distance is more accurate than the discrimination of joint angle. We hypothesize that the nervous system extracts kinematic information related to both joint angle and angular distance from proprioceptors, and that the encoding and or decoding of angular distance is more accurate than that of absolute joint angle.(ABSTRACT TRUNCATED AT 400 WORDS)