Time but not force is transferred between ipsilateral upper and lower limbs

The authors compared the force and time endpoint accuracy of goal-directed ipsilateral upper and lower limb isometric contractions and determined the components of motor performance that can be transferred from 1 limb to the other after practice. Ten young adults (27.4 +/- 4.4 years) performed 100 trials that involved their matching peak force to a force-time target with ankle dorsiflexor and elbow flexor muscles. The peak force error and variability was greater for ankle dorsiflexor contractions than for elbow flexor contractions, whereas the timing error and variability did not significantly vary with limb. There was transfer of timing, but not force, of motor output between upper and lower limbs. The timing error of the elbow flexor contractions decreased by 23% when those contractions were preceded by ankle dorsiflexor contractions, and the timing error of the ankle dorsiflexors decreased by 24% when those contractions were preceded by elbow flexor contractions. These finding therefore suggest that timing of an aiming isometric contraction may be organized at a common part of the brain for the upper and lower limbs.     

History dependence of skeletal muscle force production: implications for movement control

In this study, we review existing evidence on the history dependence of skeletal muscle force production. Specifically, we investigate the steady-state forces following shortening or stretching of an activated skeletal muscle preparation and compare these forces to the corresponding steady-state forces obtained for purely isometric contractions at identical lengths. Force depression following shortening and force enhancement following stretch can reach values of almost 50% of the corresponding isometric reference force, and thus might affect movement control. We also show novel results on history-dependent effects for voluntary contractions in human skeletal muscles, thereby emphasizing that voluntary force production is affected by the contractile history of the target muscles. These results lead to the conclusion that history-dependent force production should be considered in models of movement control and voluntary force production.     

Selective activation of lower leg muscles during maximum voluntary isometric contractions

The pronators and supinators play a key role in the medio-lateral stability of the ankle joint complex (i.e. talo-crural and subtalar joints). We hypothesized that each shank muscle has a specific activation pattern determined by its anatomical course around the axes of the subtalar and talo-crural joints. A secondary objective was to examine the effect of foot posture on these activation patterns. Forty-nine young adults (25 normal-arched feet, 24 flat-arched feet) performed maximum voluntary isometric contractions against manual resistance in four movement directions: plantarflexion (PF), dorsiflexion (DF), pronation (PRO) and supination (SUP). Electromyographic activity was recorded from tibialis posterior (TP) and peroneus longus (PL) with intramuscular electrodes, and gastrocnemius medialis (GM) and tibialis anterior (TA) with surface electrodes. When compared to their agonist function, all muscles were co-activated at significantly lower levels in their synergistic function (GM: 23% during SUP, TA: 72% during SUP; TP: 42% during PF, PL: 52% during PF) (p < 0.001). A significant interaction between foot posture and contraction type was evident for TA. During isometric contractions, the electromyographic activity of the shank muscles is geared to their biomechanical advantage according to their position relative to the subtalar and talo-crural joint axes.     

Effect of knee extensors muscles fatigue on bilateral force accuracy,variability, and coordination

The aim of this study was to test the effect of fatigue of the knee extensors muscles on bilateral force control accuracy, variability, and coordination in the presence and absence of visual feedback. Twenty-two young physically active subjects (18 males, 4 females) were divided into two groups and performed 210 submaximal sustained bilateral isometric contractions of knee extensors muscles with and without visual feedback. One group performed a symmetrical task—both legs were set at identical positions (60° knee flexion)—while the other group performed an asymmetrical task (60° and 30° knee flexion). We used the framework of the uncontrolled manifold hypothesis to quantify two variance components: one of them did not change total force (V_UCM), while the other did (V_ORT). Performance of bilateral isometric contractions reduced voluntary and electrically induced force without changes in bilateral force control variability and accuracy. Bilateral force production stability and accuracy were higher in both tasks with visual feedback. Synergistic (anti-phase) structure of force control between the lower limbs occurred and the values of synergy index were higher only during the performance of the asymmetrical task with visual feedback. In addition, greater bilateral force control accuracy was observed during the performance of the asymmetrical task (with and without visual feedback), despite no differences in within-trial variability of both tasks.     

Modeling variability of force during isometric contractions of the quadriceps femoris

The authors modeled variability of force during continuous isometric contractions of the quadriceps femoris. Twenty adults (aged 25 +/- 6 years old) performed isometric leg extensions. Target forces were 11 percentages of maximum voluntary contraction (%MVC), ranging from 2 to 95%, and 5 absolute levels, from 25 to 225 N. The authors used standard deviation of absolute force, coefficient of variation, and signal-to-noise ratio as measures of variability. The results suggested a nonlinear relationship between variability and level of force, which could best be expressed as %MVC and not as absolute force. Variability for continuous isometric contractions was described best by a sigmoidal logistic function. The sigmoidal pattern of variability as a function of %MVC is consistent with physiological mechanisms.     

Muscle coordination and force variability during static and dynamic tracking tasks

This study examined muscular activity patterns of extensor and flexor muscles and variability of forces during static and dynamic tracking tasks using compensatory and pursuit display. Fourteen volunteers performed isometric actions in two conditions: (i) a static tracking task consisting of flexion/pronation, ulnar deviation, extension/supination and radial deviation of the wrist at 20% maximum voluntary contraction (MVC), and (ii) a dynamic tracking task aiming at following a moving target at 20% MVC in the four directions of contraction. Surface electromyography (SEMG) from extensor carpi ulnaris, extensor carpi radialis, flexor carpi ulnaris and flexor digitorum superficialis muscles and exerted forces in the transverse and sagittal plane were recorded. Normalized root mean square and mutual information (index of functional connectivity within muscles) of SEMGs and the standard deviation and sample entropy of force signals were extracted. Larger SEMG amplitudes were found for the dynamic task (p < .05), while normalized mutual information between muscle pairs was larger for the static task (p < .05). Larger size of variability (standard deviation of force) concomitant with smaller sample entropy was observed for the dynamic task compared with the static task (p < .01 for both). These findings underline a rescaling of the muscles’ respective contribution influencing force variability relying on feedback and feed-forward control strategies in relation to display modes during static and dynamic tracking tasks.     

The interaction of respiration and visual feedback on the control of force and neural activation of the agonist muscle

The purpose of this study was to compare force variability and the neural activation of the agonist muscle during constant isometric contractions at different force levels when the amplitude of respiration and visual feedback were varied. Twenty young adults (20–32 years, 10 men and 10 women) were instructed to accurately match a target force at 15% and 50% of their maximal voluntary contraction (MVC) with abduction of the index finger while controlling their respiration at different amplitudes (85%, 100% and 125% normal) in the presence and absence of visual feedback. Each trial lasted 22 s and visual feedback was removed from 8–12 and 16–20 s. Each subject performed three trials with each respiratory condition at each force level. Force variability was quantified as the standard deviation of the detrended force data. The neural activation of the first dorsal interosseus (FDI) was measured with bipolar surface electrodes placed distal to the innervation zone. Relative to normal respiration, force variability increased significantly only during high-amplitude respiration (∼63%). The increase in force variability from normal- to high-amplitude respiration was strongly associated with amplified force oscillations from 0 to 3 Hz (R² ranged from .68 to .84, p < .001). Furthermore, the increase in force variability was exacerbated in the presence of visual feedback at 50% MVC (vision vs. no-vision: .97 vs. .87 N) and was strongly associated with amplified force oscillations from 0 to 1 Hz (R² = .82) and weakly associated with greater power from 12 to 30 Hz (R² = .24) in the EMG of the agonist muscle. Our findings demonstrate that high-amplitude respiration and visual feedback of force interact and amplify force variability in young adults during moderate levels of effort.     

The stability of precision grip force in older adults

The exceedingly large grip forces that many older adults employ when lifting objects with a precision pinch grip (Cole, 1991) may compensate for a reduced capability to produce a stable isometric force. That is, their grip force may fluctuate enough from moment to moment to yield grip forces that approach the force at which the object would slip from grasp. We examined the within-trial variability of isometric force in old (68-85 years, n = 13) and young (n = 11) human subjects (a) when they were asked to produce a constant pinch force at three target levels (0.49, 2.25, and 10.5 N) with external support of the arm, hand, and force transducer and (b) when they were asked to grasp, lift, and hold a small test object with a precision grip. Pinch force produced in the first task was equally stable across the two subject groups during analysis intervals that lasted 4 s. The elderly subjects produced grip forces when lifting objects that averaged twice as much as those produced by the young subjects. The force variability during the static (hold) phase of the lift for the old subjects was comparable with that used by the young subjects, after adjusting for the difference in grip force. The failure to observe less stable grip force in older adults contradicts a similar recent study. Differences in task (isometric grip force versus isometric abduction torque of a single digit) may account for this conflict, however. Thumb and finger forces for grip are produced through coactivation of many muscles and thus promote smooth force output through temporal summation of twitches. We conclude that peripheral reorganization of muscle in older adults does not yield increased instability of precision grip force and therefore does not contribute directly to increased grip forces in this population. However, force instability may affect other grip configurations (e.g., lateral pinch) or manipulation involving digit abduction or adduction forces.     

Development of unimanual versus bimanual task performance in an isometric task

Sixty-three children between 5 and 12 years of age and 15 adults performed a unimanual and a bimanual isometric force task. The performance of the preferred hand in the unimanual task was compared to the performance of the preferred hand in the bimanual task. It was hypothesized that in the bimanual task the absolute error will be higher, there will be more irregularity and the participants will need more time due to the additional effort from the central nervous system, especially with respect to the communication between the hemispheres. Furthermore, in younger children bimanual force variability was expected to be higher due to developmental aspects concerning callosal maturation and attention. It was found that with respect to force generation the preferred hand was not affected by bilateral isometric force generation, but with respect to force regulation it was. The coefficient of variation (CV) of the force was 34% larger in the bimanual task as compared to the unimanual task. For the time to target force, the increase was 28%. With repetition of the trials the CV decreased in the bimanual task, but only in the youngest age group. During development there was no change in absolute error, yet there was a major reduction in force variability in the bimanual task. It is suggested that improvement in interhemispheric communication and in the ability to focus attention plays a role in the decrease in variability with age.     

Effects of electromyostimulation on knee extensors and flexors strength and steadiness in older adults

It is known that electromyostimulation (EMS) alone or superimposed over voluntary contraction (EV) can effectively improve muscle strength. However, the effect of this type of training on the ability to control force production at submaximal levels is unknown. The authors examined the effects of EV training on steadiness in force production of knee extensors and flexors in older adults. Forty participants, including 20 men and 20 women, 60-77 years of age, were randomly allocated into a control group (CG) and an electromyostimulation superimposed over voluntary contraction (EVG) group. The EVG performed 30 bilateral isometric knee extension and flexion contractions per session, 3 training sessions per week, for 6 weeks. The variations in force production, expressed in absolute (standard deviation [SD]) and relative (coefficient of variation [CV]) terms, were assessed in isometric contractions at 5%, 15% and 25% maximal voluntary contraction (MVC) levels. Results indicated that MVC increased in knee extension and flexion in EVG (p < .05) after the training; steadiness CV also improved at 15% MVC in knee flexion (p < .05) but no significant changes were found in knee extension and steadiness SD. The training-induced changes in MVC were not correlated to steadiness CV that might indicate different mechanisms underlying these adaptations.     

Practice effects on decreasing and increasing force-control during periodic isometric movements of the index finger

The present study examined whether improvement in control while decreasing force to achieve a lower force target would be facilitated by comparison of performance while increasing force to achieve a higher force target. Participants practiced control of isometric force and timing during a unimanual force production task cycling between 5 and 10% of maximum voluntary contraction with a target interval of 500 msec. Although errors and variability of both peak and valley forces and interval decreased during early practice, the valley force was still more inaccurate and variable than the peak force in the final practice. Variabilities of both forces did not decrease when the valley force was synchronized with an audible metronome pulse but did decrease when the peak force was synchronized with it.     

Force matching errors following eccentric exercise

During eccentric exercise contracting muscles are forcibly lengthened, to act as a brake to control motion of the body. A consequence of eccentric exercise is damage to muscle fibres. It has been reported that following the damage there is disturbance to proprioception, in particular, the senses of force and limb position. Force sense was tested in an isometric force-matching task using the elbow flexor muscles of both arms before and after the muscles in one arm had performed 50 eccentric contractions at a strength of 30% of a maximum voluntary contraction (MVC). The exercise led to an immediate reduction of about 40%, in the force generated during an MVC followed by a slow recovery over the next four days, and to the development of delayed onset muscle soreness (DOMS) lasting about the same time. After the exercise, even though participants believed they were making an accurate match, they made large matching errors, in a direction where the exercised arm developed less force than the unexercised arm. This was true whichever arm was used to generate the reference forces, which were in a range of 5-30% of the reference arm's MVC, with visual feedback of the reference arm's force levels provided to the participant. The errors were correlated with the fall in MVC following the exercise, suggesting that participants were not matching force, but the subjective effort needed to generate the force: the same effort producing less force in a muscle weakened by eccentric exercise. The errors were, however, larger than predicted from the measured reduction in MVC, suggesting that factors other than effort might also be contributing. One factor may be DOMS. To test this idea, force matches were done in the presence of pain, induced in unexercised muscles by injection of hypertonic (5%) saline or by the application of noxious heat to the skin over the muscle. Both procedures led to errors in the same direction as those seen after eccentric exercise.     

Effects of Electromyostimulation on Knee Extensors and Flexors Strength and Steadiness in Older Adults

It is known that electromyostimulation (EMS) alone or superimposed over voluntary contraction (EV) can effectively improve muscle strength. However, the effect of this type of training on the ability to control force production at submaximal levels is unknown. The authors examined the effects of EV training on steadiness in force production of knee extensors and flexors in older adults. Forty participants, including 20 men and 20 women, 60–77 years of age, were randomly allocated into a control group (CG) and an electromyostimulation superimposed over voluntary contraction (EVG) group. The EVG performed 30 bilateral isometric knee extension and flexion contractions per session, 3 training sessions per week, for 6 weeks. The variations in force production, expressed in absolute (standard deviation [SD]) and relative (coefficient of variation [CV]) terms, were assessed in isometric contractions at 5%, 15% and 25% maximal voluntary contraction (MVC) levels. Results indicated that MVC increased in knee extension and flexion in EVG (p < .05) after the training; steadiness CV also improved at 15% MVC in knee flexion (p < .05) but no significant changes were found in knee extension and steadiness SD. The training-induced changes in MVC were not correlated to steadiness CV that might indicate different mechanisms underlying these adaptations.     

Online feedback and the regulation of degrees of freedom in motor control

We tested the hypothesis that the degree to which online feedback is used to control movement influences the regulation of degrees of freedom in a task. Ten participants performed an isometric force production task with their two index fingers with the goal of matching the total force to a target waveform. The role of online feedback was manipulated by changing three factors - the tracking mode, the profile of the target waveform, and the visual gain. The results showed that the coupling between the finger forces was lower in conditions where participants used online feedback to correct their movements compared to conditions where more feedforward strategies were used. The availability of online feedback is dependent on the nature of the task and this contributes to task-dependent changes in the regulation of the degrees of freedom.     

Force and timing variability in rhythmic unimanual tapping

In 3 experiments the interdependencies between timing and force production in unimanual paced and self-paced rhythmic tapping tasks were examined as participants (N = 6 in each experiment) tapped (a) to 1 of 3 target periods (333 ms, 500 ms, and 1,000 ms), while they simultaneously produced a constant peak force (PF) over a 50-s trial; (b) to produce 1 of 3 target forces (5, 10, and 15 N) at their preferred frequency, while keeping their rhythm as invariant as possible; and (c) to all combinations of target force and period. The results showed that (a) magnitudes of force and period were largely independent; (b) variability in timing increased proportionally with tapping period, and the variability in force increased with peak force; (c) force variability decreased at faster tapping rates; and (d) timing variability decreased with increasing force levels. (e) Analysis of tap-to-tap variability revealed adjustments over sequences of taps and an acceleration in the tapping rate in unpaced conditions. The interdependencies of force and time are discussed with respect to the challenges they provide for an oscillator-based account.     

Visuomotor and audiomotor processing in continuous force production of oral and manual effectors

The authors examined force control in oral and manual effectors as a function of sensory feedback (i.e., visual and auditory). Participants produced constant isometric force via index finger flexion and lower lip elevation to 2 force levels (10% and 20% maximal voluntary contraction) and received either online visual or online auditory feedback. Mean, standard deviation, and coefficient of variation of force output were used to quantify the magnitude of force variability. Power spectral measures and approximate entropy of force output were calculated to quantify the structure of force variability. Overall, it was found that the oral effector conditions were more variable (e.g., coefficient of variation) than the manual effector conditions regardless of sensory feedback. No effector differences were found for the structure of force variability with visual or auditory feedback. Oral and manual force control appears to involve different control mechanisms regulating continuous force production in the presence of visual or auditory feedback.     

Changes in muscle and joint coordination in learning to direct forces

While it has been suggested that bi-articular muscles have a specialized role in directing external reaction forces, it is unclear how humans learn to coordinate mono- and bi-articular muscles to perform force-directing tasks. Participants were asked to direct pedal forces in a specified target direction during one-legged cycling. We expected that with practice, performance improvement would be associated with specific changes in joint torque patterns and mono- and bi-articular muscular coordination. Nine male participants practiced pedaling an ergometer with only their left leg, and were instructed to always direct their applied pedal force perpendicular to the crank arm (target direction) and to maintain a constant pedaling speed. After a single practice session, the mean error between the applied and target pedal force directions decreased significantly. This improved performance was accompanied by a significant decrease in the amount of ankle angular motion and a smaller increase in knee and hip angular motion. This coincided with a re-organization of lower extremity joint torques, with a decrease in ankle plantarflexor torque and an increase in knee and hip flexor torques. Changes were seen in both mono- and bi-articular muscle activity patterns. The mono-articular muscles exhibited greater alterations, and appeared to contribute to both mechanical work and force-directing. With practice, a loosening of the coupling between bi-articular thigh muscle activation and joint torque co-regulation was observed. The results demonstrated that participants were able to learn a complex and dynamic force-directing task by changing the direction of their applied pedal forces through re-organization of joint torque patterns and mono- and bi-articular muscle coordination.     

Modulation of motor variability related to experimental muscle pain during elbow-flexion contractions

Experimental muscle pain typically reorganizes the motor control. The pain effects may decrease when the three-dimensional force components are voluntarily adjusted, but it is not known if this could have negative consequences on other structures of the motor system. The present study assessed the effects of acute pain on the force variability during sustained elbow flexion when controlling task-related (one-dimensional) and all (three-dimensional) contraction force components via visual feedback. Experimental muscle pain was induced by bolus injection of hypertonic saline into m. biceps brachii, and isotonic saline was used as control. Twelve subjects performed sustained elbow flexion at different levels of the maximal voluntary contraction (5–30% MVC) before, during, and after the injections. Three-dimensional force components were measured simultaneously with surface electromyography (EMG) from elbow flexors and auxiliary muscles. Results showed that force variability was increased during pain compared to baseline for contractions using one-dimensional feedback (P < .05), but no significant differences were found for three-dimensional feedback. During painful contractions (1) EMG activity from m. trapezius was increased during contractions using both one-dimensional and three-dimensional feedback (P < .05), and (2) the complexity of EMG from m. triceps brachii and m. deltoid was higher for the three-dimensional feedback (P < .05). In conclusion, the three-dimensional feedback reduced the pain-related functional distortion at the cost of a more complex control of synergistic muscles.     

Short-term memory effects of an auditory biofeedback on isometric force control: is there a differential effect as a function of transition trials?

The aim of the present study was to investigate memory effects, force accuracy, and variability during constant isometric force at different force levels, using auditory biofeedback. Two types of transition trials were used: a biofeedback-no biofeedback transition trial and a no biofeedback-biofeedback transition trial. The auditory biofeedback produced a low- or high-pitched sound when participants produced an isometric force lower or higher than required, respectively. To achieve this goal, 16 participants were asked to produce and maintain two different isometric forces (30 ± 5% and 90 N ± 5%) during 25 s. Constant error and standard deviation of the isometric force were calculated. While accuracy and variability of the isometric force varied according to the transition trial, a drift of the force appeared in the no biofeedback condition. This result suggested that the degradation of information about force output in the no biofeedback condition was provided by a leaky memory buffer which was mainly dependent on the sense of effort. Because this drift remained constant whatever the transition used, this memory buffer seemed to be independent of short-term memory processes.     

Intermittent visual information and the multiple time scales of visual motor control of continuous isometric force production

In an experiment, we examined the effect of intermittency (from 25.6 Hz to 0.2 Hz) of visual information on continuous isometric force production as a function of force level (5%, 10%, 25%, and 50% of maximal voluntary contraction [MVC]). The amount of force variability decreased and the irregularity of force output increased as a function of increased visual intermittency rate. Vision was found to have an influence on the frequency structure of force output up to 12 Hz, and the 25% MVC force level had more high-frequency modulations with higher rates of visual information. The effective use of intermittent visual information is mediated nonlinearly by force level, and there are multiple time scales of visual control (range, approximately 0 - 12 Hz) that are postulated to be a function of both feedback and feedforward control processes.