Roles of dynamic and stationary components of a motor command in establishing the equilibrium state at simplest targeted movements

We studied in humans interrelations between the kinematic characteristics of targeted movements of the arm and current levels of EMG of the muscles providing these movements; the movements were relatively slow, and the attained joint angle was held for a time. The EMG level was considered a correlate of the level of integral motor commands (efferent activity of the respective motoneuronal pools). Application of low-amplitude non-inertial loadings, directed against the forces developed by one or another muscle group, allowed us to provide realization of targeted movements exclusively by the activity of this muscle group, without Involvement of the antagonists. It was demonstrated that the target equilibrium joint angle is reached synchronously with the dynamic phase of EMG activity, before the latter reaches a stationary level. The structure of the dynamic EMG phase itself is complex; in most cases it is split into several components. The dependence between the joint angle and amplitude of the EMG stationary phase is rather complex and variable, and usually it is difficult to predict the characteristics of this phase based on simple biomechanical considerations. There are proofs that at the performance of the studied movements and maintaining a target position there are some components in the mechanical muscle activity, which are not controlled by the motor commands. Thus, the stationary level of a motor command represents only one of several factors responsible for attaining and maintaining a target equilibrium position. Establishing this position is provided, first of all, by interaction of dynamic components of the motor commands to different muscles. Our results show that the attempts to interpret the processes of control of targeted movements on the basis of modifications of the equilibrium point hypothesis are inadequate; these data are in better compliance with the concept of impulse-temporal control; at their interpretation it is also necessary to take more thoroughly into account nonlinear properties of the muscle reactions.     

On the errors in assessment of severity of involuntary movements using surface EMG.

Surface electromyogram (SEMG) is a useful tool to depict involuntary movements, but evaluation of the intensity of such movements with SEMG over multiple recording instances requires awareness of factors influencing quantified SEMG signals. We investigated the differences in the amplitude of SEMGs due to electrode displacement in isometric voluntary contraction in the upper arm, forearm and lower leg in 8 healthy men. The SEMGs of gross muscle activity simultaneously recorded with 4 electrode pairs from the agonist and antagonist sides in 3 displacement conditions with respect to parallel position, interelectrode distance, and rotation were compared. The amount of EMG integration (equivalent to the average SEMG amplitude) of each electrode pair was compared to the reference electrode pair with interelectrode distance of 40 mm placed on the center of the tested muscles. The average EMG difference ratios ranged 1.1-2.2%/mm in parallel shift, 1.0-1.9%/mm in distance shift, and 0.3-0.6%/degree in rotation shift. Displacement error of electrodes in separate recording instances should be reduced using anatomical landmarks, when SEMG is applied as a quantitative method to evaluate change in the states of involuntary movements.     

An equilibrium-point model of electromyographic patterns during single-joint movements based on experimentally reconstructed control signals

The purpose of this work has been to develop a model of electromyographic (EMG) patterns during single-joint movements based on a version of the equilibrium-point hypothesis, a method for experimental reconstruction of the joint compliant characteristics, the dual-strategy hypothesis, and a kinematic model of movement trajectory. EMG patterns are considered emergent properties of hypothetical control patterns that are equally affected by the control signals and peripheral feedback reflecting actual movement trajectory. A computer model generated the EMG patterns based on simulated movement kinematics and hypothetical control signals derived from the reconstructed joint compliant characteristics. The model predictions have been compared to published recordings of movement kinematics and EMG patterns in a variety of movement conditions, including movements over different distances, at different speeds, against different-known inertial loads, and in conditions of possible unexpected decrease in the inertial load. Changes in task parameters within the model led to simulated EMG patterns qualitatively similar to the experimentally recorded EMG patterns. The model's predictive power compares it favourably to the existing models of the EMG patterns.     

Blood pressure changes alter tracheobronchial cough: computational model of the respiratory-cough network and in vivo experiments in anesthetized cats

We tested the hypothesis, motivated in part by a coordinated computational cough network model, that alterations of mean systemic arterial blood pressure (BP) influence the excitability and motor pattern of cough. Model simulations predicted suppression of coughing by stimulation of arterial baroreceptors. In vivo experiments were conducted on anesthetized spontaneously breathing cats. Cough was elicited by mechanical stimulation of the intrathoracic airways. Electromyograms (EMG) of inspiratory parasternal, expiratory abdominal, laryngeal posterior cricoarytenoid (PCA), and thyroarytenoid muscles along with esophageal pressure (EP) and BP were recorded. Transiently elevated BP significantly reduced cough number, cough-related inspiratory, and expiratory amplitudes of EP, peak parasternal and abdominal EMG, and maximum of PCA EMG during the expulsive phase of cough, and prolonged the cough inspiratory and expiratory phases as well as cough cycle duration compared with control coughs. Latencies from the beginning of stimulation to the onset of cough-related diaphragm and abdominal activities were increased. Increases in BP also elicited bradycardia and isocapnic bradypnea. Reductions in BP increased cough number; elevated inspiratory EP amplitude and parasternal, abdominal, and inspiratory PCA EMG amplitudes; decreased total cough cycle duration; shortened the durations of the cough expiratory phase and cough-related abdominal discharge; and shortened cough latency compared with control coughs. Reduced BP also produced tachycardia, tachypnea, and hypocapnic hyperventilation. These effects of BP on coughing likely originate from interactions between barosensitive and respiratory brainstem neuronal networks, particularly by modulation of respiratory neurons within multiple respiration/cough-related brainstem areas by baroreceptor input.     

Effects of postural muscle fatigue on the relation between segmental posture and movement.

The purpose of this study was to examine whether fatigue of postural muscles might influence the coordination between segmental posture and movement. Seven healthy adults performed series of fifteen fast wrist flexions and extensions while being instructed to keep a dominant upper limb posture as constant as possible. These series of voluntary movements were performed before and after a fatiguing submaximal isometric elbow flexion, and also with or without the help of an elbow support. Surface EMG from muscles Delto?deus anterior, Biceps brachii, Triceps brachii, Flexor carpi ulnaris, Extensor carpi radialis were recorded simultaneously with wrist, elbow and shoulder accelerations and wrist and elbow displacements. Fatigue was evidenced by a shift of the elbow and shoulder muscles EMG spectra towards low frequencies. Kinematics of wrist movements and corresponding activations of wrist prime-movers, as well as the background of postural muscle activation before wrist movement were not modified. There were only slight changes in timing of postural muscle activations. These data indicate that postural fatigue induced by a low-level isometric contraction has no effect on voluntary movement and requires no dramatic adaptation in postural control.     

Anticipatory postural adjustments to arm movement reveal complex control of paraspinal muscles in the thorax

Anatomical and empirical data suggest that deep and superficial muscles may have different functions for thoracic spine control. This study investigated thoracic paraspinal muscle activity during anticipatory postural adjustments associated with arm movement. Electromyographic (EMG) recordings were made from the right deep (multifidus/rotatores) and superficial (longissimus) muscles at T5, T8, and T11 levels using fine-wire electrodes. Ten healthy participants performed fast unilateral and bilateral flexion and extension arm movements in response to a light. EMG amplitude was measured during 25 ms epochs for 150 ms before and 400 ms after deltoid EMG onset. During arm flexion movements, multifidus and longissimus had two bursts of activity, one burst prior to deltoid and a late burst. With arm extension both muscles were active in a single burst after deltoid onset. There was differential activity with respect to direction of trunk rotation induced by arm movement. Right longissimus was most active with left arm movements and right multifidus was most active with right arm movements. All levels of the thorax responded similarly. We suggest that although thoracic multifidus and longissimus function similarly to control sagittal plane perturbations, these muscles are differentially active with rotational forces on the trunk.     

Kinematic and EMG characteristics of simple shoulder movements with proprioception and visual feedback.

The objective of this study was to determine if simple, shoulder movements use the dual control hypothesis strategy, previously demonstrated with elbow movements, and to see if this strategy also applies in the absence of visual feedback. Twenty subjects were seated with their right arm abducted to 90 degrees and externally rotated in the scapular plane. Subjects internally rotated to a target position using a custom shoulder wheel at three different speeds with and without visual feedback. Kinematics were collected with a motion analysis system and electromyographic (EMG) recordings of the pectoralis major (PECT), infraspinatus (INFRA), anterior and posterior (ADELT, PDELT) deltoid muscles were used to evaluate muscle activity patterns during movements. Kinematics changed as movement speed increased with less accuracy (p<0.01). Greater EMG activity was observed in the PECT, PDELT, and INFRA with shorter durations for the ADELT, PDELT and INFRA. Movements with only kinesthetic feedback were less accurate (p<0.01) and performed faster (p<0.01) than movements with visual feedback. EMG activity suggests no major difference in CNS control strategies in movements with and without visual feedback. Greater resolution with visual feedback enables the implementation of a dual control strategy, allowing greater movement velocity while maintaining accuracy.     

Development of coordinated movement in chicks: II. Temporal analysis of hindlimb muscle synergies at embryonic day 10 in embryos with spinal gap transections.

Spinal neural circuits can recruit muscles to produce organized patterns of activity early in embryonic development. In a previous study, using multichannel electromyographic (EMG) recordings, we characterized burst parameters for these patterns in the legs of chick embryos during spontaneous motility in ovo at embryonic days (E) 9 and E10 (Bradley and Bekoff, 1990). Results of the study suggested both neural and biomechanical factors play an important role in the development of coordinated limb movements. In this study, to explore the contribution of descending neural inputs to the control of leg movements during motility, we applied similar methods to characterize motor patterns produced by the spinal cord in the absence of descending inputs. Thoracic spinal gap transections were performed at E2 and EMG patterns were recorded at E10. Several EMG features for chronic spinal embryos were similar to those for normal embryos and demonstrate that lumbar spinal circuits can be correctly assembled to control limb movements in the absence of connectivity with more rostral neural structures during early differentiation processes. However, certain aspects of the EMG patterns in chronic spinal embryos were different from patterns in normal embryos and provide support for conclusions drawn earlier by Oppenheim (1975). Specifically, our data support the view that propriospinal and/or supraspinal inputs function to regulate the timing of cyclic limb movements controlled by spinal neural circuits. Finally, we consider the possible long-term effects of chronic spinal gap transections as compared to acute spinal transections on the development of motility.     

Predictions of mechanical output of the human M. triceps surae on the basis of electromyographic signals: the role of stimulation dynamics

In order to assess the significance of the dynamics of neural control signals for the rise time of muscle moment, simulations of isometric and dynamic plantar flexion contractions were performed using electromyographic signals (EMG signals) of m. triceps surae as input. When excitation dynamics of the muscle model was optimized for an M-wave of the medial head of m. gastrocnemius (GM), the model was able to make reasonable predictions of the rise time of muscle moment during voluntary isometric plantar flexion contractions on the basis of voluntary GM EMG signals. The rise time of muscle moment in the model was for the greater part determined by the amplitude of the first EMG burst. For dynamic jumplike movements of the ankle joint, however, no relationship between rise time of muscle moment in the experiment and muscle moment predicted by the model on the basis of GM EMG signals was found. Since rise time of muscle moment varied over a small range for this movement, it cannot be completely excluded that stimulation dynamics plays a role in control of these simple single-joint movements.     

Oxygen consumption of active rainbow trout, Salmo gairdneri Richardson, derived from electromyograms obtained by radiotelemetry

A radiotelemetry apparatus is described for sensing and transmitting electromyograms (EMGs) from free-swimming fish. EMGs are recorded from the epaxial muscles of adult rainbow trout during periods in spontaneous (= routine) activity, and forced-swim, respirometers. When such EMG records are integrated, subjected to spectral analysis, and computer-averaged, the EMG values (in μV) are highly correlated with the fish oxygen consumption during the activity periods. However, there is a marked difference between the regression slopes for oxygen v . EMG value for the data from the spontaneous, and forced-swim, respirometers; the former slope is the steeper. The probable explanation of this phenomenon is that whereas in forced swims the epaxial myomeres are responsible for most of the activity of the fish, in spontaneous activity other muscle systems (e.g. of the lateral, dorsal and ventral fins) come to account for a greater relative proportion of body movement. The difference in slope, although great, is evidently a regular phenomenon. The shift from one regression to the other occurs at a fairly precise epaxial EMG value ( c . 5 μV). This suggests that the laboratory calibration of EMG value in terms of oxygen consumption can be utilized in the wild so that EMG records from free-swimming fish, fitted with telemetry packages can be used to deduce oxygen consumption attributable to activity. It also appears that such records can be used as a guide to the type of activity of the fish, i.e. desultory movements or free cruising.     

Ocular and stabilization feedback: an evaluation of two EMG biofeedback control procedures

This study evaluated the adequacy of two novel EMG biofeedback control procedures. During a single training session, 36 subjects received either contingent EMG feedback from the frontal region (Veridical), contingent feedback for vertical eye movements (Ocular), or a feedback condition where the signal increased with deviations in any direction from baseline EMG levels (Stabilization). The results supported the use of Ocular but not Stabilization feedback as a control procedure in frontalis EMG biofeedback studies. Ocular feedback did not produce reductions in frontalis EMG but did lead to changes in subjective measures of nonspecific treatment effects that were at least comparable to those obtained with Veridical feedback. Stabilization subjects produced small but significant reductions in EMG, felt the most bored as a result of their feedback training, and were the most likely to rate themselves as having received false feedback. The implications of attribution theory and multiprocess relaxation theory for the evaluation of nonspecific treatment effects are discussed.     

Development of coordinated movement in chicks: II. Temporal analysis of hindlimb muscle synergies at embryonic day 10 in embryos with spinal gap transections

Spinal neural circuits can recruit muscles to produce organized patterns of activity early in embryonic development. In a previous study, using multichannel electromyographic (EMG) recordings, we characterized burst parameters for these patterns in the legs of chick embryos during spontaneous motility in ovo at embryonic days (E) 9 and E10 (Bradley and Bekoff, 1990). Results of the study suggested both neural and biomechanical factors play an important role in the development of coordinated limb movements. In this study, to explore the contribution of descending neural inputs to the control of leg movements during motility, we applied similar methods to characterize motor patterns produced by the spinal cord in the absence of descending inputs. Thoracic spinal gap transections were performed at E2 and EMG patterns were recorded at E10. Several EMG features for chronic spinal embryos were similar to those for normal embryos and demonstrate that lumbar spinal circuits can be correctly assembled to control limb movements in the absence of connectivity with more rostral neural structures during early differentiation processes. However, certain aspects of the EMG patterns in chronic spinal embryos were different from patterns in normal embryos and provide support for conclusions drawn earlier by Oppenheim (1975). Specifically, our data support the view that propriospinal and/or supraspinal inputs function to regulate the timing of cyclic limb movements controlled by spinal neural circuits. Finally, we consider the possible long-term effects of chronic spinal gap transections as compared to acute spinal transections on the development of motility. © 1992 John Wiley & Sons, Inc.     

Estimation of dynamic joint torques and trajectory formation from surface electromyography signals using a neural network model

In this study, human arm movement was re-constructed from electromyography (EMG) signals using a forward dynamics model acquired by an artificial neural network within a modular architecture. Dynamic joint torques at the elbow and shoulder were estimated for movements in the horizontal plane from the surface EMG signals of 10 flexor and extensor muscles. Using only the initial conditions of the arm and the EMG time course as input, the network reliably reconstructed a variety of movement trajectories. The results demonstrate that posture maintenance and multijoint movements, entailing complex via-point specification and co-contraction of muscles, can be accurately computed from multiple surface EMG signals. In addition to the model's empirical uses, such as calculation of arm stiffness during motion, it allows evaluation of hypothesized computational mechanisms of the central nervous system such as virtual trajectory control and optimal trajectory planning.     

Further evaluation of an EMG technique for assessment of the deep cervical flexor muscles

A novel surface electromyographic (EMG) technique was recently described for the detection of deep cervical flexor muscle activity. Further investigation of this technique is warranted to ensure EMG activity from neighbouring muscles is not markedly influencing the signals recorded. This study compared deep cervical flexor (DCF) muscle activity with the activity of surrounding neck and jaw muscles during various anatomical movements of the neck and jaw in 10 volunteer subjects. DCF EMG activity was recorded with custom electrodes inserted via the nose and fixed by suction to the posterior mucosa of the oropharynx. Surface electrodes were placed over the sternocleidomastoid, anterior scalene, masseter and suprahyoid muscles. Positioned in supine, subjects performed isometric cranio-cervical flexion, cervical flexion, right and left cervical rotation, jaw clench and resisted jaw opening. Across all movements examined, EMG amplitude of the DCF muscles was greatest during neck movements that would require activity of the DCF muscles, particularly during cranio-cervical flexion, their primary anatomical action. The actions of jaw clench and resisted jaw opening demonstrated significantly less DCF EMG activity than the cranio-cervical flexion action (p < 0.05). Across all other movements, the neighbouring neck and jaw muscles demonstrated greatest EMG amplitude during their respective primary anatomical actions, which occurred in the absence of increased EMG amplitude recorded from the DCF muscles. The finding of substantial EMG activity of the DCF muscles only during neck actions that would require their activity, particularly cranio-cervical flexion, and not during actions involving the jaw, provide further assurance that the majority of myoelectric signals detected from the nasopharyngeal electrode are from the DCF muscles.     

Nutritional control of respiratory and other muscular activities in relation to plasma prostaglandin E in the fetal sheep

The effects of nutrient availability on fetal plasma prostaglandin E (PGE) concentrations, on fetal breathing movements and electromyographic (EMG) activities of fetal nuchal and forelimb muscles were investigated in pregnant ewes by varying dietary intake and by manipulation of fetal plasma glucose concentration. The incidence of fetal breathing movements (06.00-10.00 h) decreased with increasing gestational age while fetal arterial concentrations of plasma PGE increased significantly over the same period of gestation. Maternal fasting for 48 h reduced the incidence of fetal breathing movements and the amount of nuchal EMG activity (06.00-10.00 h) in animals older than 130 days but had no effect earlier in gestation. No changes in forelimb EMG activity were observed during fasting at any gestational age. Plasma PGE levels increased significantly during fasts begun both before and after 130 days of gestation. When data from fed and fasted states were combined for all fetuses, irrespective of gestational age, there was a significant inverse correlation between fetal breathing movements incidence and plasma PGE concentration in utero. This relationship was even more pronounced when the fetuses were considered individually. Insulin infusions induced hypoglycaemia, an increase in fetal plasma PGE concentration and a significant reduction in the incidence of fetal breathing movements at all ages. Glucose infusions of fetal breathing movements only after 130 days and had no effect on plasma PGE levels in utero at any gestational age. Neither insulin nor glucose infusions altered the EMG activities of the nuchal and forelimb muscles. The results show that glucose availability is an important factor in determining the incidence of fetal breathing movements in utero and indicate that nutritionally induced changes in fetal breathing movements are mediated in part by PGE. They also suggest that PGE is a physiological regulator of fetal breathing movements in the sheep during late gestation.     

Control of single-joint movements with a reversal.

We studied the kinematic and electromyographic (EMG) patterns during single-joint elbow flexion movements with a reversal and tested two hypotheses. First, that the amplitude of the second phase of the movement (M(2)) will be controlled by two different means, a drop in the second flexor burst for a small M(2) and an increase in the integral of the extensor burst for larger M(2). Second, based on the muscle stretch-shortening cycle (SSC), that movements reversing without a delay will show a larger extensor burst, as compared to movements that reverse after a delay. Changes in EMG patterns with M(2) amplitude supported the first hypothesis and could be interpreted within the framework of the equilibrium-point hypothesis. The observations also corroborate a hypothesis that discrete movements represent outcomes of an oscillatory control process stopped at a particular phase. In Experiment-2, even the shortest delay at the target led to a significantly larger extensor burst. However, there were no differences in the peak velocity of M2 with and without the delay. These observations do not support a major role of stretch reflexes in the SSC effects during such movements. However, they are compatible with the idea of peripheral factors, such as peripheral muscle and tendon elasticity, playing a major potentiating role in the SSC.     

Ocular and stabilization feedback: An evaluation of two EMG biofeedback control procedures

This study evaluated the adequacy of two novel EMG biofeedback control procedures. During a single training session, 36 subjects received either (1) contingent EMG feedback from the frontal region (Veridical), (2) contingent feedback for vertical eye movements (Ocular), or (3) a feedback condition where the signal increased with deviations in any direction from baseline EMG levels (Stabilization). The results supported the use of Ocular but not Stabilization feedback as a control procedure in frontalis EMG biofeedback studies. Ocular feedback did not produce reductions in frontalis EMG but did lead to changes in subjective measures of nonspecific treatment effects that were at least comparable to those obtained with Veridical feedback. Stabilization subjects produced small but significant reductions in EMG, felt the most bored as a result of their feedback training, and were the most likely to rate themselves as having received false feedback. The implications of attribution theory and multiprocess relaxation theory for the evaluation of nonspecific treatment effects are discussed.This research was supported in part by grants from the National Institutes of Health (AM31500) and the Robert Wood Johnson Foundation. Portions of this research were presented at the Sixth Annual Meeting of the Society of Behavioral Medicine, New Orleans, March 1985.     

Analyzing gastrocnemius EMG-activity and sway data from quiet and perturbed standing.

In an experiment, we combined force plate measurements and surface EMG in studying quiet and perturbed standing, involving MS (Multiple sclerosis) and controls. The aim of this paper is to report the results thus obtained on the relation between filtered gastrocnemius (GA) EMG and the anterior-posterior center-of-pressure (A/P COP) coordinate. The main finding is the good correspondence between A/P COP and the filtered GA EMG in the low frequency range. The EMG envelope was calculated using a zero-lag filter. Combining this with time shifts around 250-350 ms produced a high correlation (85.5+/-8.4%) between the GA-EMG envelope and the A/P COP. This EMG-COP relation was closest when using a low cut-off frequency value around 1 Hz in calculating the EMG envelope. Based on this filtering procedure we estimated the average EMG-COP time shift to be 283+/-43 ms between the GA-EMG envelope and A/P COP (which "lags" behind EMG envelope). This shift is consistent with the 1 Hz cut-off and phase shift produced by a corresponding critically damped second-order filter, and is about twice the corresponding twitch time. These results suggest that GA is to a large extent responsible for the phasic control of the anterior-posterior balance during quiet standing. A small difference (p<0.03) was found between mean time shift thus obtained for controls (n=4) and MS (n=6) while sway area showed a major difference (p<0.01). The paper also compares three alternative filters for numerical calculation of the EMG-envelope.     

Electromyographic and kinematic studies of tail movements in dogs during treadmill locomotion

Electromyographic (EMG) activities of three tail muscles, the extensor caudae lateralis (ECL), abductor caudae externus (ACE), and flexor caudae longus (FCL), were recorded bilaterally in seven adult dogs during walking, trotting, and galloping on a treadmill. Each dog's movements were recorded with a 16 mm high-speed camera system, and angular movements of the tail were analyzed. During walking and trotting, reciprocal EMG bursts were observed between right and left tail muscles and corresponded with lateral movements of the tail. The tonic discharges that were observed in ECL and FCL seemed to maintain the position of the tail. During galloping, synchronized EMG activity of all tail muscles produced reactive torques to counter those generated by cyclic limb movements and kept the tail in a stable position. These results suggest that tail movements are important in maintaining body balance during locomotion in the dog. © 1993 Wiley-Liss, Inc.     

Activity and functions of the human gluteal muscles in walking,running, sprinting,and climbing

It has been suggested that the uniquely large gluteus maximus (GMAX) muscles were an important adaptation during hominin evolution based on numerous anatomical differences between humans and extant apes. GMAX electromyographic (EMG) signals have been quantified for numerous individual movements, but not across the range of locomotor gaits and speeds for the same subjects. Thus, comparing relative EMG amplitudes between these activities has not been possible. We assessed the EMG activity of the gluteal muscles during walking, running, sprinting, and climbing. To gain further insight into the function of the gluteal muscles during locomotion, we measured muscle activity during walking and running with external devices that increased or decreased the need to control either forward or backward trunk pitch. We hypothesized that 1) GMAX EMG activity would be greatest during sprinting and climbing and 2) GMAX EMG activity would be modulated in response to altered forward trunk pitch demands during running. We found that GMAX activity in running was greater than walking and similar to climbing. However, the activity during sprinting was much greater than during running. Further, only the inferior portion of the GMAX had a significant change with altered trunk pitch demands, suggesting that the hip extensors have a limited contribution to the control of trunk pitch movements during running. Overall, our data suggest that the large size of the GMAX reflects its multifaceted role during rapid and powerful movements rather than as a specific adaptation for a single submaximal task such as endurance running. Am J Phys Anthropol 153:124–131, 2014. © 2013 Wiley Periodicals, Inc.