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Separability of EEG signals recorded during right and left motor imagery using adaptive autoregressive parameters.

Electroencephalogram (EEG) recordings during right and left motor imagery can be used to move a cursor to a target on a computer screen. Such an EEG-based brain-computer interface (BCI) can provide a new communication channel to replace an impaired motor function. It can be used by, e.g., patients with amyotrophic lateral sclerosis (ALS) to develop a simple binary response in order to reply to specific questions. Four subjects participated in a series of on-line sessions with an EEG-based cursor control. The EEG was recorded from electrodes overlying sensory-motor areas during left and right motor imagery. The EEG signals were analyzed in subject-specific frequency bands and classified on-line by a neural network. The network output was used as a feedback signal. The on-line error (100%-perfect classification) was between 10.0 and 38.1%. In addition, the single-trial data were also analyzed off-line by using an adaptive autoregressive (AAR) model of order 6. With a linear discriminant analysis the estimated parameters for left and right motor imagery were separated. The error rate obtained varied between 5.8 and 32.8% and was, on average, better than the on-line results. By using the AAR-model for on-line classification an improvement in the error rate can be expected, however, with a classification delay around 1 s.

Adult↗

The patterns of control signals underlying elbow joint movements in humans.

Position, velocity, flexor and extensor electromyographic (EMG) activity of fast, moderate and slow elbow movements to a target were recorded and simulated using a model in which reciprocal and co-activation central commands, proprioceptive feedback and mechanical properties of muscles were considered. Two hypotheses concerning the pattern of shift in the equilibrium point (EP) underlying the movements were tested. First, the nervous system specifies a constant rate of EP shift to produce movement and encodes displacement by the duration of the shift (ramp-shaped pattern). Second, in fast movements, the EP rapidly shifts towards the future final position but then shifts back and forth eventually reaching the final EP (N-shaped pattern). The ramp pattern was consistent with kinematic and EMG experimental data regardless of movement speed. In contrast, the N-shaped pattern was incompatible with the kinematic characteristics of fast movements.

Elbow Joint↗

Intrinsic dynamics and mechanosensory modulation of non-nutritive sucking in human infants.

The human infant is endowed with a complex mechanism for the ingestion of nutrients that becomes functional in late fetal life. The sucking motor pattern is generally accepted to be under the control of pattern generating circuitry located in the brainstem reticular formation. Systems under the control of a central pattern generator (CPG) may use afferent feedback to allow for changing environmental conditions. Although it is clear that afferent pathways serving the orofacial region become responsive to mechanical stimulation early in fetal life, little is known about the integration of afferent information into the suck CPG. The actifier, a device for the mechanical stimulation of intraoral and perioral tissues, was designed and used to investigate the response properties of the human infant suck CPG to patterned mechanical stimulation. Sinusoid and square waveform stimuli elicited responses including modulation of jaw kinematics and synchronization [entrainment] of non-nutritive suck motor patterns to the mechanical stimulus. These data provide evidence that the suck CPG is responsive to mechanical stimulation of perioral and intraoral soft tissues.

Afferent Pathways↗

Modification of human gastric acid secretion with operant-conditioning procedures.

In an attempt to control gastric acid secretion with operant-conditioning techniques, four normal women were given visual feedback on gastric pH plus money reinforcers. When money was made dependent on increased secretion in a differential-reinforcement-of-high-rates schedule, the rate of secretion of three of the four subjects increased to three times baseline. When money was then made dependent on decreased secretion in a differential-reinforcement-of-other-behaviors schedule, the rate of secretion of these three subjects returned to baseline levels. Heart rate, respiratory frequency, abdominal electromyographic activity, and stomach motility (measured by the electrogastrogram method) were not consistently correlated with acid secretion across subjects, although individual subjects showed substantial correlations between acid secretion and one or more other physiological response.

Adult↗

Subject expectancy effects in frontal EMG conditioning.

The influence of subject expectancies on performance has been examined extensively within the framework of behavior therapy but has received little attention within the context of EMG training. This study assessed the effects of subject expectancy on acquisition during frontal EMG conditioning. Prior to conditioning, subjects were either given no expectancy or instructed that learning to control the feedback stimulus would be either easy (high expectancy) or difficult (low expectancy). Subjects in the three expectancy conditions then underwent 20 min of contingent reinforcement for frontal EMG decreases. Three similar groups of no, low, or high expectancy subjects received noncontingent reinforcement. Differential EMG behavior was exhibited between the two no-expectancy groups, with the contingent group reliably lower in EMG activity than that noncontingent. However, differences were not observed between contingent and noncontingent subjects given either low or high expectancy sets. None of the groups could be differentiated on the basis of subjective variables such as anxiety, relaxation, or frustration. These findings suggest that subject expectancies, either positive or negative, interfere with the acquisition of conditioned EMG behavior.

Adolescent↗

Simultaneous monitoring of force, acceleration and electromyogram during computer-controlled infusion of atracurium in sheep.

Five sheep were anaesthetised with sodium thiopentone, nitrous oxide and oxygen, and the lungs ventilated artificially. A computer-controlled infusion of atracurium employing feedback from isometric force measurement was set to maintain 15, 50 and 85% depth of neuromuscular block. Each was held constant for 30 minutes, while isometric force and electromyogram were measured on the left forelimb and acceleration on the right. The acceleration transducer records at the three levels of block were practically identical with those from force measurements: mean differences (SD) between simultaneous measurements being 3.6 (5.4)%. EMG value was 10.1 (12.0)% different from force measurement. Correlation between simultaneous force and either acceleration or EMG was excellent (r = 0.986 and 0.938 respectively). In clinical practice all three techniques measure block comparably. In the sheep forelimb, EMG was a less reliable method of measuring block.

Animals↗

Early corrective reactions of the leg to perturbations at the torso during walking in humans.

The contribution of afferent feedback to the regulation of locomotion in humans is not well understood. Animal experiments have suggested that loading of the leg during the stance phase may enhance the magnitude of extensor burst activity and delay the onset of swing phase. The aim of the present study was to determine whether transient loading of the leg at the end of stance would enhance extensor-muscle activity and delay the onset of swing in walking humans. To test this hypothesis, we applied loads to the hips of subjects so that the load was applied along the long axis of the leg at the end of stance (down-back unsupported, DBU). This resulted in an unexpectedly complex reaction characterised by rapid co-contraction of antagonist pairs of muscles around the ankle and knee and a prolongation of the stance phase. We speculated that the complexity of the reaction was, in part, due to a disturbance in equilibrium. To address this possibility, two additional perturbation paradigms were tested: (1) subjects held a rail during the loading paradigm (down-back supported, DBS), or (2) subjects received only a posteriorly directed perturbation of the hips, which added no additional load to the leg (backward unsupported, BU). As predicted, the DBS perturbation resulted in an enhancement of the ongoing soleus-muscle activity, and the unexpected tibialis anterior burst that was observed during the DBU paradigm was absent. Allowing the subjects to hold a rail substantially reduced the change in the timing of the step cycle observed in the DBU paradigm. The BU perturbation prolonged the stance phase duration and, as expected, resulted in a burst of activity in tibialis activity. This was usually accompanied by a reduction in the ongoing soleus activity. Two important conclusions are drawn from the present study. First, loading of the leg at the end of stance phase enhances the ongoing extensor-muscle activity. We suggest that afferent feedback responding to the increase load supported by the leg leads to rapid enhancement of the active extensor muscles to compensate for the increased load and prevent collapse of the leg. Interestingly, the duration of the stance phase was only marginally increased when loading was applied without a postural disturbance (DBS). Second, posterior perturbation of the centre of mass at the end of stance phase evokes an "automatic postural response" in tibialis anterior. Of particular interest, this evoked postural response can occur simultaneously with an enhanced activation of soleus. This indicates that the DBU perturbation employed in this study elicited two responses, one to prevent the collapse of the leg and the other to stabilise the centre of mass.

Adult↗

Independence between the amount and structure of variability at low force levels.

The purpose of the current experiment was to investigate the amount (standard deviation (S.D.) and coefficient of variation (CV)) and structure (approximate entropy (ApEn)) of force variability at very low force levels. Participants produced isometric force output of index finger abduction at five levels (0.4, 0.8, 1.0, 2.0, and 4.0 N) with high and low visual feedback gain. The findings showed that: subjects scaled their force output to the targets; S.D. increased non-linearly with force level and decreased with visual gain; and CV decreased with force level as well as visual gain. ApEn of the force output did not change as a function of force level, although the high gain increased ApEn in contrast to low gain. It is proposed that the recruitment of additional motor units at very low force levels does not significantly alter the structure of the force output, although it does increase the magnitude of force and its amount of variability. Overall, the findings provide evidence that the amount and structure of motor variability can be influenced by separate control processes at low force levels.

Adult↗

Influence of clenching level on intercuspal contact area in various regions of the dental arch.

The purpose of this study was to evaluate the relationship between the clenching level and the intercuspal contact area in different regions of the dental arch. Twenty-five healthy subjects with natural normal dentitions and good occlusal support performed clenching tasks in the intercuspal position at four different levels (10, 30, 70 and 100% levels of maximum voluntary contraction) through EMG visual feedback from bilateral masseter and anterior temporal muscles. Simultaneously, the occlusal contacts were recorded with a silicone occlusal contact checking material (Black Silicone, GC Dental Industrial Corp., Tokyo, Japan). The occlusal records were analysed by an image analyser. Every area of the thickness less than 50 microm was determined to be an occlusal contact area. The occlusal contact areas on the anterior teeth (incisors and canines), the premolars (first and second premolars) and the molars (first and second molars) were calculated separately. The posterior occlusal contact area increased with an increase in the clenching levels but that of the anterior did not. The results of this study indicate that the increase in clenching forces affects the anterior and posterior occlusal contact areas differently.

Adult↗

Predictions of the existence, frequency, and amplitude of physiological tremor in normal man based on measured frequency-response characteristics.

Mathematical models for 1) musculoskeletal dynamics, and 2) reflex feedback, based on the results of the authors' frequency-response measurements on normal adult male human subjects, are combined to produce a model for physiological tremor in such subjects. An analysis of this model shows that the system will be unstable to small disturbances (that is, tremor will occur) under certain conditions of external loading. Further, when the system is unstable, nonlinearities in the model produce responses in the form of limit cycles, and both the frequency and amplitude of the resulting tremor can be calculated. For constant loads applied through a constant compliance, the model predicts the onset of tremor at low loads, a maximum intensity of tremor at loads corresponding to 30-50 percent of maximum voluntary effort, and a decrease in the tremor amplitude at still higher loads.

Biomechanical Phenomena↗

Proprioceptive feedback in humans expresses motor invariants during writing.

Proprioceptive feedback from populations of muscle spindle afferents feeds the brain with information relating to the instantaneous velocity and direction of ongoing movements. In this paper, we investigate whether the invariant relationship between the velocity and curvature of a trajectory, i.e. the two-thirds power law, is reflected in this muscle spindle feedback. Sixty unitary muscle spindle afferents from six ankle muscle groups were recorded using intraneural microelectrodes during imposed "writing-like" movements. The movements had kinematic parameters obeying the two-thirds power law and were imposed so that the tip of the foot followed trajectories forming four different letters and six numbers. The responses of the muscle spindle afferent populations were analysed using the population vector model. The results demonstrate that the neuronal trajectories attained from populations of muscle spindles clearly depict the path and kinematic parameters and express the movement invariants, i.e. the trajectory segmentation into units of action and the two-thirds power law. The central vs peripheral origin of such constraints involved in the motor system is discussed.

Action Potentials↗

Force control of isometric elbow flexion with visual feedback in cold with and without shivering.

BACKGROUND: Exposure to cold impairs manual performance through effects on muscle tissue and control mechanisms. The purpose of this study was to assess the influence of increased muscle tone and shivering on ability to maintain required force during isometric flexion over a wide range of effort levels. METHODS: Lightly clad male subjects (n = 6) were exposed to thermoneutral air (TN, 27 degrees C) for 30 min, or to cold air (CA, 10 degrees C) for 30 min followed by a cold drink (1 L, 8 degrees C) to cause vigorous shivering (SH). At the end of each condition, subjects performed isometric elbow flexion at 10, 20, 40, and 80% of individual maximal voluntary contraction (MVC) for 10 s each, using digital visual feedback to control the force. We analyzed mean force output (F), the coefficient of force variation (FCv), information transmission (F/SD), and the coefficient of force auto-correlation (Ra), and the averaged electromyogram (aEMG) from elbow flexors, elbow extensor, and pectoral girdle muscles. RESULTS: Compared with TN, CA with increased muscle tone raised the aEMG by 5-30% but did not impair any of the force characteristics. In SH, F was not affected, while FCv and Ra were significantly increased at 10% MVC, while aEMG increased by 30-400% depending on the specific muscle and MVC level. CONCLUSION: Neither thermoregulatory muscle tone nor shivering influenced the control of force output during isometric elbow flexion, except that at the lowest MVC (10%) the force output was more variable during shivering.

Adult↗

Human balancing of an inverted pendulum: is sway size controlled by ankle impedance?

Using the ankle musculature, subjects balanced a large inverted pendulum. The equilibrium of the pendulum is unstable and quasi-regular sway was observed like that in quiet standing. Two main questions were addressed. Can subjects systematically change sway size in response to instruction and availability of visual feedback? If so, do subjects decrease sway size by increasing ankle impedance or by some alternative mechanism? The position of the pendulum, the torque generated at each ankle and the soleus and tibialis anterior EMG were recorded. Results showed that subjects could significantly reduce the mean sway size of the pendulum by giving full attention to that goal. With visual feedback sway size could be minimised significantly more than without visual feedback. In changing sway size, the frequency of the sways was not changed. Results also revealed that ankle impedance and muscle co-contraction were not significantly changed when the sway size was decreased. As the ankle impedance and sway frequency do not change when the sway size is decreased, this implies no change in ankle stiffness or viscosity. Increasing ankle impedance, stiffness or viscosity are not the only methods by which sway size could be reduced. A reduction in torque noise or torque inaccuracy via a predictive process which provides active damping could reduce sway size without changing ankle impedance and is plausible given the data. Such a strategy involving motion recognition and generation of an accurate motor response may require higher levels of control than changing ankle impedance by altering reflex or feedforward gain.

Adolescent↗

Voluntary activation of human motor axons in the absence of muscle afferent feedback. The control of the deafferented hand.

The ability to activate human motoneurons supplying individual intrinsic muscles of the hand was examined during acute deafferentation of the muscles. Tungsten microelectrodes were inserted percutaneously into motor fascicles of the ulnar nerve of 5 subjects, which was then blocked distally with local anaesthetic. In 4 subjects unitary action potentials were recorded from 16 motor axons, which were identified with respect to their target muscles. In the complete absence of muscle afferent feedback, subjects could voluntarily recruit motoneurons, grade their discharge and sustain a constant level of activity. Significant facilitation of motor efforts was provided by cutaneous feedback from the digits via the median nerve. During attempted maximal voluntary efforts the mean discharge frequencies of single motor axons were significantly lower than those of normally-innervated motor units. This finding suggests that peripheral afferents have a net facilitatory influence on motoneurons. However, during prolonged (20-30 s) maximal voluntary efforts the deafferented motoneurons did not display the progressive decline in discharge frequency shown by normally-innervated motor units during contractile fatigue, a finding consistent with two possible explanations: disfacilitation or reflex inhibition of the motoneuron pool by peripheral afferents. The results also indicate that the otherwise intact nervous system can perform some simple motor tasks with no proprioceptive input other than knowledge of the motor commands. Other factors may contribute to the poor motor performance reported for patients with severe sensory deficits.

Adult↗

Inhibition of respiratory activity during passive ventilation: a role for intercostal afferents?

The purpose of this study was to determine whether receptors from the rib cage are primarily responsible for inhibitory feedback of inspiratory muscle activity during mechanical ventilation. Seven quadriplegics with C5-C6 lesions were compared to 6 normals during mechanical ventilation. All subjects were mechanically hyperventilated with a nasal mask to suppress intrinsic inspiratory muscle activity. End-tidal partial pressure of carbon dioxide (PETCO2) was increased by either adding CO2 (FICO2) or decreasing tidal volume (VT) until reoccurrence of inspiratory activity, defined as the recruitment threshold (PCO2RT). The difference between PCO2RT and eupneic PETCO2 indicated the presence and magnitude of volume-related inhibition of inspiratory muscle activity during mechanical ventilation. Substantial inhibition of inspiratory activity was observed in both quadriplegics and normals. We conclude that afferent information from the rib cage is not obligatory for the mediation of volume-related inhibition of inspiratory muscle activity during mechanical ventilation.

Adult↗

Is a stimulus conveying task-relevant information a sufficient condition to elicit a stimulus-preceding negativity?

Movement-preceding and stimulus-preceding negativities were recorded when a movement was followed by one of three informative visual stimuli. The meaning of the visual stimulus alternated between (a) conveying a task-relevant instruction about a subsequent time production task and (b) providing feedback (knowledge of results) about performance on the current time production task. In a control condition, premovement and postmovement scalp potentials were recorded when subjects made the same movements but in a voluntary, self-paced manner. Under all conditions, movements were preceded by a movement-preceding negativity, and neither the amplitude nor the lateral asymmetry of this negativity was affected by the subsequent presentation of either kind of informative stimulus. When the movement was followed by a stimulus conveying knowledge of results, the negativity in the postmovement epoch was enhanced, but this enhancement was not evident in epochs preceding instruction stimuli. We conclude that not all task relevant stimuli elicit a stimulus-preceding negativity, and we provide a functional interpretation of this negativity in terms of emotional anticipation and the contingency of the stimulus on a previous event.

Adult↗

Activity patterns of the masticatory muscles during feedback-controlled simulated clenching activities.

In 10 normal subjects, the electromyographic (EMG) activities of the masseter, anterior and posterior temporalis, medial pterygoid, inferior lateral pterygoid, as well as the anterior digastric, were bilaterally recorded during clenching with various resulting force vectors. The intraoral force transfer was achieved with a three-component force transducer. The direction and magnitude of the force vectors were controlled by visual feedback. The goal of the study was to investigate how various clenching directions at constant magnitude of force influence the EMG activity in the masticatory muscles during gradual increase of the horizontal force component. Depending on the force direction and the individual muscle, an increase or decrease of activity was observed during clenching with increasing horizontal force components at constant magnitude of force. The inferior lateral pterygoid exhibited the highest activation (about 80% of maximal voluntary clenching) of all masticatory muscles. The medial pterygoid showed the greatest range of variation in activation behavior, and it was the most active muscle in relation to all clenching directions. In conclusion, the results show that with growing horizontal force components at constant magnitude of force, all muscles demonstrated an increase or decrease of activity in several clenching directions.

Adult↗

Isovelocity investigation of the lengthening behaviour of the erector spinae muscles.

The purpose of this study was to investigate the force-velocity (F/v) relationship for the erector spinae muscles in submaximal activation movements, with particular attention to their response during lengthening movements and at lower shortening contraction velocities. Dynamic models that predict lower back muscle forces require reasonable representations of the modulating effect of instantaneous velocity. Ten males were observed performing trunk flexion and extension in the sagittal plane under constant load. Contraction velocities were measured as the first derivative from a devise sensitive to changes in spine curvature, and controlled by a visual feedback system while a constant load was applied through a chest harness. The erector spinae exhibited a yielding phenomenon which causes an abrupt drop in force during constant velocity stretching under constant, submaximal, stimulation. The findings were consistent with previous isovelocity muscle lengthening experiments. Yielding appeared dependent on the level of load/activation supporting the theory of a "state-variable" F/v relationship. The eccentric behaviour of the lower erectors (L3) seemed independent of velocity and length, while that of the upper erectors (T9) showed a dependence on length. At lower concentric velocities, concavity in torque-velocity curves was noted after a "threshold" velocity. The findings of this study strongly reinforce the notion that the F/v length relationship is not a continuous hyperbolic relationship during muscle shortening and that the commonly modelled force augmentation effect of lengthening is incorrect, at least for submaximal activation of the extensors of the lower back.

Adult↗