Elastic energy and muscle potentiation,a continuing discussion: A reply to Hof and van den Berg

In a previous study (van Ingen Schenau 1984) a number of arguments were formulated to show that the role of storage and re-utilisation of elastic energy in human movements is often overestimated. One of the used arguments concerned the estimation of the amount of stored elastic energy on the basis of knee extension and plantar flexion torques in combination with knee flexion and dorsiflexion angles during running. In their comment to this argument, Hof and van den Berg (1986) show convincingly that one should take into account the internal stretch distance of the elastic component rather than those flexion angles. In the present paper the further arguments of Hof and van den Berg concerning the compliance of human muscle and tendons are discussed. Based on the other arguments previously published and on a new point concerning the limited capacity for muscles to add external work to the elastic energy which is released during concentric contraction, it is concluded that all main arguments against the significance of the storage and re-utilisation of elastic energy still hold.     

Mechanical output about the ankle in countermovement jumps and jumps with extended knee

In this study the mechanical output (e.g., force, contraction velocity, instantaneous power) about the ankle was measured during a jump with and without occurrence of transportation of power and pre-stretch potentiation. To examine this, a model of the m. triceps surae was used. Eleven subjects performed a maximal one-legged countermovement jump (CMJ) and a maximal one-legged jump with extended knee (EKJ). Ground reaction forces, cinematographic data and electromyograms of m. triceps surae were recorded. The power output, defined as the product of moment and angular velocity, and work done about the ankle were higher during CMJ (1404 W, 130 J) than during EKJ (852 W, 76 J). The peak moments were the same during both jumps. The model of the m. triceps surae showed that the power delivered by m. triceps surae was higher during CMJ than during EKJ, as a result of catapult-action of m. gastrocnemius tendon. The difference in work done is explained, in addition to transportation of energy, by a greater contraction range of m. soleus during CMJ. There is no reason to assume that pre-stretch potentiation plays a role in the difference in mechanical output. The results show that the mechanical output of muscles in complex movements is strongly dependent on the dynamics of the movement, and not only on its contractile capacity.     

Effect of taping and its conditions on electromyographic responses of knee extensor muscles

IntroductionThe present study investigated the effect of stretchable characteristics of elastic therapeutic tape and its elongation on surface electromyography (EMG) of knee extensor muscles during knee extension movements.MethodsNine healthy men performed knee extension movement with the application of normal elastic tape or highly stretchable tape and without the tapes (control). Tapes were applied on the anterior thigh to cross the knee joint with no elongation and elongation of 50 and 75% of the maximum stretchability. Surface EMG was recorded from the vastus lateralis (VL) muscle and proximal (RFp) and distal (RFd) sites of the rectus femoris muscle.ResultsUnder the no-elongation conditions, decreases in the surface EMG amplitude of the VL and RFd muscles were observed with normal tape during the isometric contraction phase and with highly stretchable tape during isometric and eccentric contraction phases, compared with the control (p < 0.05). There were no significant differences in surface EMG among the different elongation conditions in any muscles (p > 0.05).DiscussionThese results suggest that the stretchable characteristics of tapes change the effect of elastic tape application on neuromuscular activation of the applied muscles and these effects are not dependent on the elongation of the tape.     

A Linked Muscular Activation Model for Movement Generation and Control

A new model for movement control is presented which incorporates characteristics of impulse-variability and mass-spring models. Movements in the model were controlled with phasic torque impulses in agonist and antagonist muscles and a tonic agonist torque.

Characteristics of the phasic agonist and antagonist torque profiles were based on observed properties of movement-related EMGs and muscle isometric torques. Variability of the phasic impulses depended on impulse magnitude as in impulse-variability models. The model therefore predicted a speed-accuracy tradeoff for limb movement. The time of onset and magnitude of the antagonist torque depended on the magnitude of the preceding agonist torque as indicated in studies of movement-related EMGs. This led to the new concept of linkage between the agonist and antagonist muscle forces which was shown to be important for reducing variability of fast movements. Progressive development of linkage during practice could explain the previous findings of decreased movement variability with practice coupled with increased variability of movement-related EMGs.

It was concluded that an inherently variable motor system deals with the variability associated with generation of large muscle forces by linking the forces produced by opposing muscles. In this way, variability in net joint torques and in movements can be decreased without the need for the nervous system to closely regulate the individual torques.     

Transversus abdominis is part of a global not local muscle synergy during arm movement

The trunk muscle transversus abdominis (TrA) is thought to be controlled independently of the global trunk muscles. Methodological issues in the 1990s research such as unilateral electromyography and a limited range of arm movements justify a re-examination of this theory. The hypothesis tested is that TrA bilateral co-contraction is a typical muscle synergy during arm movement. The activity of 6 pairs of trunk and lower limb muscles was recorded using bilateral electromyography during anticipatory postural adjustments (APAs) associated with the arm movements. The integrated APA electromyographical signals were analyzed for muscle synergy using Principle Component Analysis. TrA does not typically bilaterally co-contract during arm movements (1 out of 6 participants did). APA muscle activity of all muscles during asymmetrical arm movements typically reflected a direction specific diagonal pattern incorporating a twisting motion to transfer energy from the ground up. This finding is not consistent with the hypothesis that TrA plays a unique role providing bilateral, feedforward, multidirectional stiffening of the spine. This has significant implications to the theories underlying the role of TrA in back pain and in the training of isolated bilateral co-contraction of TrA in the prophylaxis of back pain.     

An impulse-timing theory for reciprocal control of muscular activity in rapid, discrete movements

Much remains to be learned about how agonist and antagonist muscles are controlled during the production of rapid, voluntary movements. In an effort to summarize a wide body of existing knowledge and stimulate future research on this subject, an impulse-timing theory is presented which attempts to predict the activity of reciprocal muscles based on certain characteristics of a movement. The basic tenet of the theory is that variables of movement time, movement distance and inertial load have fairly predictable effects on the underlying muscular activity of the agonist and antagonist muscles during the production of rapid and discrete, voluntary movements. The theory is derived from the kinematic work of Schmidt, Zelaznik, Hawkins, Frank and Quinn (1979) and supporting evidence from studies which have used electromyographic (EMG) recordings of agonist and antagonist muscles during rapid movements. Issues related to synergistic muscle control, central and peripheral control of reciprocal muscle activity, muscle control, and neurological disorder and the relationship between impulse-timing and mass-spring control are discussed in the final section.     

Functional roles of the proprioceptive system in the control of goal-directed movement.

This article explored functional roles of the proprioceptive system during the control of goal-directed movements. Proprioceptive information contributes to the control of movement through both reflex and central connections. Spinal and transcortical reflex loops establish a servomechanism which provides automatic corrections of unexpected changes in muscle length and allows compensation for undesirable irregularities in the mechanical properties of muscles by modulating limb stiffness at the subconscious level. Central connections provide the control system with information about peripheral states which is used in voluntary components of movement control. Before the initiation of movement, proprioceptive information about initial limb orientation becomes a basis for the programming of motor commands. During a movement, proprioceptive input about velocities and angular displacements of a limb is used to regulate movement by triggering planned sequences of muscle activation and modulating motor commands. After movement, feedback produced by responses is compared with previously stored information, verifying the quality of the movement. Considering potential roles of the reflex and central connections, the proprioceptive system seems to constitute an important aspect of motor control mechanisms, providing the control system with efficiency and flexibility in the regulation of goal-directed movements.     

A linked muscular activation model for movement generation and control

A new model for movement control is presented which incorporates characteristics of impulse-variability and mass-spring models. Movements in the model were controlled with phasic torque impulses in agonist and antagonist muscles and a tonic agonist torque. Characteristics of the phasic agonist and antagonist torque profiles were based on observed properties of movement-related EMGs and muscle isometric torques. Variability of the phasic impulses depended on impulse magnitude as in impulse-variability models. The model therefore predicted a speed-accuracy tradeoff for limb movement. The time of onset and magnitude of the antagonist torque depended on the magnitude of the preceding agonist torque as indicated in studies of movement-related EMGs. This led to the new concept of linkage between the agonist and antagonist muscle forces which was shown to be important for reducing variability of fast movements. Progressive development of linkage during practice could explain the previous findings of decreased movement variability with practice coupled with increased variability of movement-related EMGs. It was concluded that an inherently variable motor system deals with the variability associated with generation of large muscle forces by linking the forces produced by opposing muscles. In this way, variability in net joint torques and in movements can be decreased without the need for the nervous system to closely regulate the individual torques.     

An Impulse-Timing Theory for Reciprocal Control of Muscular Activity in Rapid,Discrete Movements

Much remains to be learned about how agonist and antagonist muscles are controlled during the production of rapid, voluntary movements. In an effort to summarize a wide body of existing knowledge and stimulate future research on this subject, an impulse-timing theory is presented which attempts to predict the activity of reciprocal muscles based on certain characteristics of a movement. The basic tenet of the theory is that variables of movement time, movement distance, and inertial load have fairly predictable effects on the underlying muscular activity of the agonist and antagonist muscles during the production of rapid and discrete, voluntary movements. The theory is derived from the kinematic work of Schmidt, Zelaznik, Hawkins, Frank, and Quinn (1979) and supporting evidence from studies which have used electromyographic (EMG) recordings of agonist and antagonist muscles during rapid movements. Issues related to synergistic muscle control, central and peripheral control of reciprocal muscle activity, muscle control, and neurological disorder and the relationship between impulse-timing and mass-spring control are discussed in the final section.     

Task-Dependent Compensatory Responses to Perturbations Applied During Rhythmic Movements in Humans

Modulation of the responses to perturbation applied during different phases of three rhythmic movements in humans—running, cycling, and hopping—was studied. The perturbation was an electrical stimulus. The results showed gating and modulation of the responses in both ipsi-and contralateral limb muscles. The responses during running and cycling were only excitatory in nature, while during hopping an inhibitory response was observed. These responses were not correlated with the normal activity during the movement. The latency of the response in general was not altered for different stimulation phases. The alterations in the step cycle demonstrated overt behavioral changes due to the responses. There were differences between the responses observed during these movements and walking. In running, the major adaptation to perturbations appears to be in the contralateral side as seen in the changes in the step cycle. During cycling (except for one phase) and hopping, the same set of muscles was activated in response to perturbation. This represents a simplifying strategy in response organization. The dependency of the response on the task characteristics, postural stability requirement, and external constraints imposed on the subject is discussed. These studies provide insights into task-dependent strategies adopted by the nervous system to meet unexpected perturbation during rhythmic movement in humans.     

Vision,proprioception and corollary discharge in a movement recall test

Movement recall was investigated in relation to the sensory processes involved in a triangle drawing task.Forty subjects in two groups, one with and one without visual feedback, performed a recall task involving movements of their index finger. All subjects attended different experimental sessions in which (1) all proprioceptive feedback was eliminated by the ischaemic block technique, (2) muscle spindle feedback was distorted by vibration of the muscles and tendons involved in the movement, and (3) proprioceptive feedback was normal.Within each session subjects were required firstly to recall triangular movements made for them passively by the experimenter, and secondly, to recall movements they had made actively. Results indicated comparable accuracy in recall of active movements in all conditions, and a decrement in passive recall dependent on the availability of the alternative sources of feedback. The results indicated a process of integrated contribution of all inputs to the perception of movement; redundancy in information when all channels are available; and a role of corollary discharge in recall of movements.     

Movement Related EMGs Become More Variable During Learning of Fast Accurate Movements

Human subjects performed simple flexion and extension movements about the elbow in a visual step-tracking paradigm. Movements were self-terminated. Subjects were instructed to increase movement velocity while maintaining end-point accuracy during practice. The effects of practice on the pattern and variability of EMG activity of the biceps and triceps muscles were studied. Initial movements were performed using reciprocal phasic activation of agonist and antagonist muscles as indicated by surface EMGs. With practice, increases in movement speed were associated with larger agonist and antagonist bursts and an earlier onset of the antagonist burst. Decreased duration of the premovement antagonist silence was also observed during practice.

Decreases in variability of movements during practice were not accompanied by equivalent decreases in variability of the associated EMGs. Surprisingly, both agonist and antagonist EMGs were more variable in faster, practiced movements. The combined agonist-antagonist EMG variability depended on both movement speed and trajectory variability. Lower variability in movements in the presence of greater variability in the related EMGs occurred because of linked variations in agonist and antagonist muscle activities. Variations in the first agonist burst were often compensated for by associated variations in the antagonist and late agonist bursts. These linked variations maintained the limb trajectory relatively constant in spite of large variations in the first agonist burst. Modifications to impulse-variability models are therefore needed to explain compensations for variability in accelerative impulses (produced by the first agonist burst) by linked variations in impulses for deceleration (produced by the antagonist and late agonist bursts).     

Movement related EMGs become more variable during learning of fast accurate movements

Human subjects performed simple flexion and extension movements about the elbow in a visual step-tracking paradigm. Movements were self-terminated. Subjects were instructed to increase movement velocity while maintaining end-point accuracy during practice. The effects of practice on the pattern and variability of EMG activity of the biceps and triceps muscles were studied. Initial movements were performed using reciprocal phasic activation of agonist and antagonist muscles as indicated by surface EMGs. With practice, increases in movement speed were associated with larger agonist and antagonist bursts and an earlier onset of the antagonist burst. Decreased duration of the premovement antagonist silence was also observed during practice. Decreases in variability of movements during practice were not accompanied by equivalent decreases in variability of the associated EMGs. Surprisingly, both agonist and antagonist EMGs were more variable in faster, practiced movements. The combined agonist-antagonist EMG variability depended on both movement speed and trajectory variability. Lower variability in movements in the presence of greater variability in the related EMGs occurred because of linked variations in agonist and antagonist muscle activities. Variations in the first agonist burst were often compensated for by associated variations in the antagonist and late agonist bursts. These linked variations maintained the limb trajectory relatively constant in spite of large variations in the first agonist burst. Modifications to impulse-variability models are therefore needed to explain compensations for variability in accelerative impulses (produced by the first agonist burst) by linked variations in impulses for deceleration (produced by the antagonist and late agonist bursts).     

Time-dependence between upper arm muscles activity during rapid movements: Observation of the proportional effects predicted by the kinematic theory

Rapid human movements can be assimilated to the output of a neuromuscular system with an impulse response modeled by a Delta-Lognormal equation. In such a model, the main assumption concerns the cumulative time delays of the response as it propagates toward the effector following a command. To verify the validity of this assumption, delays between bursts in electromyographic (EMG) signals of agonist and antagonist muscles activated during a rapid hand movement were investigated. Delays were measured between the surface EMG signals of six muscles of the upper limb during single rapid handwriting strokes. From EMG envelopes, regressions were obtained between the timing of the burst of activity produced by each monitored muscle. High correlation coefficients were obtained supporting the proportionality of the cumulative time delays, the basic hypothesis of the Delta-Lognormal model. A paradigm governing the sequence of muscle activities in a rapid movement could, in the long run, be useful for applications dealing with the analysis and synthesis of human movements.     

Detection and execution of movements

Summary Detection thresholds for movements imposed on the relaxed joints in upper limbs, when expressed in terms of angular or linear displacement, differ from joint to joint. However, when they are expressed in terms of proportional changes in the lengths of fascicles of the muscles serving the joints, they are found to be similar. When the execution of finely graded voluntary movements is analysed, performances of similar accuracy occur when the movements at different joints require alterations of the lengths of active muscle fascicles by similar proportions. These findings suggest that muscle length is a variable of importance to the CNS in both the detection and execution of movements. For faster contractions, another category of movement must be considered. This is the triggered response, which can be voluntarily pre-formulated and stored in the brain, to be released subsequently by some sensory input. Such triggered responses can be demonstrated in experiments in which subjects respond to masked stimuli — low-intensity sensory stimuli which, while readily detected when presented alone, are not detected when followed very soon afterwards by a high-intensity stimulus. Subjects are able to react with simple and more complex movements to low-intensity stimuli whether these are detected (delivered alone) or undetected.     

Changes in the relationship between movement velocity and movement distance in primary focal hand dystonia

The authors examined the relationship between movement velocity and distance and the associated muscle activation patterns in 18 individuals with focal hand dystonia (FHD) compared with a control group of 18 individuals with no known neuromuscular condition. Participants performed targeted voluntary wrist and elbow flexion movements as fast as possible across 5 movement distances. Individuals with FHD were slower than controls across all distances, and this difference was accentuated for longer movements. Muscle activation patterns were triphasic in the majority of individuals with FHD, and muscle activation scaled with distance in a similar manner to controls. Cocontraction did not explain movement slowing in individuals with dystonia, but there was a trend toward underactivation of the 1st agonist burst in the dystonic group. The authors concluded that slowness is a consistent feature of voluntary movement in FHD and is present even in the absence of dystonic posturing. Underactivation of the 1st agonist burst appears to be the most likely reason to explain slowing.     

The SYBAR system: Integrated recording and display of video, EMG, and force plate data

A new system—called SYBAR—is introduced, that employs digital video for registration of the movements of a patient while simultaneously recording electromyogram signals of relevant muscles and ground reaction forces (for the lower extremities in gait studies). All information is stored in a multimedia record, which can be viewed by the clinician with a simple user interface. This setup allows an integrated and more detailed view of the movement of the patient and related information (i.e., muscle physiology). It is used by clinicians to assess the causes of movement disorders in their patients. This paper describes the SYBAR system and focuses on the employed methods of data synchronization for both the time and the spatial domains. It is concluded that, although SYBAR was developed for clinical gait studies, the technology can be applied in all situations in which the relation between physiological signals and human or animal behavior is studied.     

Stretch reflexes in the human masticatory muscles: a brief review and a new functional role

Stretch reflexes play a vital role in fine-tuning movements and in automatically maintaining posture. This article briefly reviews the operation of the stretch reflex in the human masticatory system. The conventional approach of stretching muscles in an open-loop manner has yielded much valuable information on the operation of this reflex. In particular, it has revealed that stretching the jaw-closing muscles evokes a reflex response with two major components. The short-latency reflex is favoured when stretches are brisk, but slower stretches evoke an additional long-latency component. In the hand muscles, the long-latency response is transcortical: in the masticatory muscles, it is not. In addition to its role in servo-control of muscle length during chewing, the stretch reflex in the jaw-closing muscles maintains the vertical position of the mandible during vigorous head movements such as those that occur during running, jumping, hopping and other vigorous whole-body movements in which the head moves briskly up and down. This is an interesting model system in which to investigate stretch reflexes with natural stimuli under unrestrained, physiological conditions.     

The postural control can be optimized by the first movement initiation condition encountered when submitted to muscle fatigue

We investigated whether and how the movement initiation condition (IC) encountered during the early movements performed following focal muscle fatigue affects the postural control of discrete ballistic movements. For this purpose, subjects performed shoulder flexions in a standing posture at maximal velocity under two movement IC, i.e., in self-paced conditions and submitted to a Stroop-like task in which participants had to trigger fast shoulder flexions at the presentation of incongruent colors. Shoulder flexion kinematics, surface muscle activity of focal and postural muscles as well as center-of-pressure kinematics were recorded. The initial IC and the order in which subjects were submitted to these two conditions were varied within two separate experimental sessions. IC schedule was repeated before and after fatigue protocols involving shoulder flexors. The aim of this fatigue procedure was to affect acceleration-generating capacities of focal muscles. In such conditions, the postural muscle activity preceding and accompanying movement execution is expected to decrease. Following fatigue, when subjects initially moved in self-paced conditions, postural muscle activity decreased and scaled to the lower focal peak acceleration. This postural strategy then transferred to the Stroop-like task. In contrast, when subjects initially moved submitted to the Stroop-like task, postural muscle activity did not decrease and this transferred to self-paced movements. Regarding the center-of-pressure peak velocity, which is indicative of the efficiency of the postural actions generated in stabilizing posture, no difference appeared between the two sessions post-fatigue. This highlights an optimization of the postural actions when subjects first moved in self-paced conditions, smaller postural muscle activation levels resulting in similar postural consequences. In conclusion, the level of neuromuscular activity associated with the postural control is affected and can be optimized by the initial movement IC experienced post-fatigue. Beyond the fundamental contributions arising from these results, we point out potential applications for trainers and sports instructors.     

Task-dependent compensatory responses to perturbations applied during rhythmic movements in humans

Modulation of the responses to perturbation applied during different phases of three rhythmic movements in humans-running, cycling, and hopping-was studied. The perturbation was an electrical stimulus. The results showed gating and modulation of the responses in both ipsi- and contralateral limb muscles. The responses during running and cycling were only excitatory in nature, while during hopping an inhibitory response was observed. These responses were not correlated with the normal activity during the movement. The latency of the response in general was not altered for different stimulation phases. The alterations in the step cycle demonstrated overt behavioral changes due to the responses. There were differences between the responses observed during these movements and walking. In running, the major adaptation to perturbations appears to be in the contralateral side as seen in the changes in the step cycle. During cycling (except for one phase) and hopping, the same set of muscles was activated in response to perturbation. This represents a simplifying strategy in response organization. The dependency of the response on the task characteristics, postural stability requirement, and external constraints imposed on the subject is discussed. These studies provide insights into task-dependent strategies adopted by the nervous system to meet unexpected perturbation during rhythmic movements in humans.