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Determining volitional activation (VA) can provide insights on the cause of muscle weakness in orthopedic and neurological populations. Two electrical stimulation techniques are traditionally used to quantify VA: interpolation (IT) and superimposition (CAR). IT allows for a more accurate VA estimation, however it requires individuals to be stimulated twice, compared to once for CAR, and thus increases stimulation associated discomfort. To date, there is no agreement on what is the best practical technique for calculating quadriceps VA. This paper aims to address this problem by determining what reference force (i.e., using either peak force or force at the time of stimulation) and type of stimulation (train of pulses (burst), doublet, and twitch) is the best technique to use. Our findings showed that the IT with the force at the time of stimulation as a reference should be used to determine VA and that when a burst was used, the VA ratio computations were more accurate. Additionally, using a twitch with a 2ms pulse duration produced reliable VA calculations and may be an acceptable alternative for pain-sensitive subjects. Accurate assessment of VA deficits can help clinicians design rehabilitation programs that are based on subject-specific strength impairments and are more effective., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Dynamic resistance training increases the force and speed of muscle contraction, but little is known about modifications to the contractile properties of the main physiological types of motor units (MUs) that contribute to these muscle adaptations. Although the contractile profile of MU muscle fibers is tightly coupled to myosin heavy chain (MyHC) protein expression, it is not well understood if MyHC transition is a prerequisite for modifications to the contractile characteristics of MUs. In this study, we examined MU contractile properties, the mRNA expression of MyHC, parvalbumin, and sarcoendoplasmic reticulum Ca 2+ pump isoforms, as well as the MyHC protein content after 5 wk of volitional progressive weight-lifting training in the medial gastrocnemius muscle in rats. The training had no effect on MyHC profiling or Ca 2+ -handling protein gene expression. Maximum force increased in slow (by 49%) and fast (by 21%) MUs. Within fast MUs, the maximum force increased in most fatigue-resistant and intermediate but not most fatigable MUs. Twitch contraction time was shortened in slow and fast fatigue-resistant MUs. Twitch half-relaxation was shortened in fast most fatigue-resistant and intermediate MUs. The force-frequency curve shifted rightward in fast fatigue-resistant MUs. Fast fatigable MUs fatigued less within the initial 15 s while fast fatigue-resistant units increased the ability to potentiate the force within the first minute of the standard fatigue test. In conclusion, at the early stage of resistance training, modifications to the contractile characteristics of MUs appear in the absence of MyHC transition and the upregulation of Ca 2+ -handling genes., (Copyright © 2016 the American Physiological Society.)

This article reviews the use of transcranial magnetic stimulation (TMS) over the motor cortex to make estimates of the level of voluntary drive to muscles. The method, described in 2003 (Todd et al. J Physiol 551: 661-671, 2003), uses a TMS pulse to produce descending corticospinal volleys that synaptically activate motoneurons, resulting in a muscle twitch. Linear regression of the superimposed twitch amplitude and voluntary force (or torque) can generate an "estimated" resting twitch for muscles involved in a task. This procedure has most commonly been applied to elbow flexors but also to knee extensors and other muscle groups. Data from 44 papers using the method were tabulated. We identify and discuss five major technical challenges, and the frequency with which they are addressed. The technical challenges include inadvertent activation of the cortical representation of antagonist muscles, the role of antagonist torques at the studied joint, uncertainty about the effectiveness of the TMS pulse in activating the motoneuron pool, the linearity of the voluntary force (or torque) and superimposed twitch relationship, and variability in the TMS-evoked EMG and force/torque responses. The ideal situation in which the descending corticospinal volleys recruit all of the agonist motoneurons and none of the antagonist motoneurons is unlikely to ever occur, and hence results must be carefully examined to assess the authenticity of the voluntary activation estimates in the context of the experimental design. A partial compromise lies in the choice of stimulus intensity. We also identify aspects of the procedure that require further investigation., (Copyright © 2016 the American Physiological Society.)

This study examined the contribution of muscle fiber orientation at different knee angles to regional activation identified with high-density surface electromyography (HDsEMG). Monopolar HDsEMG signals were collected using a grid of 13 × 5 electrodes placed over the vastus medialis (VM). Intramuscular electrical stimulation was used to selectively activate two regions within VM. The distribution of EMG responses to stimulation was obtained by calculating the amplitude of the compound action potential for each channel; the position of the peak amplitude was tracked across knee angles to describe shifts of the active muscle regions under the electrodes. In a separate experiment, regional activation was investigated in 10 knee flexion-extension movements against a fixed resistance. Intramuscular stimulation of different VM regions resulted in clear differences in amplitude distribution along the columns of the electrode grid (P < 0.001); changes in knee angle resulted in consistent shifts along the rows (P < 0.01) and negligible shifts along the columns of the electrode grid. Regional VM activation was identified in dynamic movement, with distal shifts of the EMG distribution in the eccentric phase of the movement (P < 0.05) and at more flexed knee angles (P < 0.05). HDsEMG was used to describe regional activation across the VM that was not attributable to anatomic factors. Changes in muscle fiber orientation associated with knee joint angle mainly influence the amplitude distribution along the fiber direction. Future studies are needed to understand possible functional roles for regional activation within the VM in dynamic tasks., (Copyright © 2016 the American Physiological Society.)

Clinically available myoelectric control does not enable simultaneous proportional control of prosthetic degrees of freedom. Multiple studies have proposed systems that provide simultaneous control, though few have investigated whether subjects voluntarily use simultaneous control or how they implement it. Additionally, few studies have explicitly evaluated the effect of providing proportional velocity control. The objective of this study was to evaluate factors influencing when and how subjects use simultaneous myoelectric control, including the ability to proportionally control the velocity and the required task precision. Five able-bodied subjects used simultaneous myoelectric control systems with and without proportional velocity control in a virtual Fitts' Law task. Though subjects used simultaneous control to a substantial degree when proportional velocity control was present, they used very little simultaneous control when using constant-velocity control. Furthermore, use of simultaneous control varied significantly with target distance and width, reflecting a strategy of using simultaneous control for gross cursor positioning and sequential control for fine corrective movements. These results provide insight into how users take advantage of simultaneous control and highlight the need for real-time evaluation of simultaneous control algorithms, as the potential benefit of providing simultaneous control may be affected by other characteristics of the myoelectric control system.

Background: Recently, high-frequency repetitive transcranial magnetic stimulation (rTMS) is reported to evaluating the corticospinal pathway and improving both cortical excitability and motor function significantly in subjects. According to some previous reports, the maximum voluntary muscle contraction (MVC) of target muscle can reinforce the influence by rTMS. The aim of this study was to confirm 5 Hz rTMS with MVC in healthy individuals is an effective method to facilitate motor neuron excitability and the efficiency can last at least 30 min post stimulation., Objective: To compare the motor evoked potentials (MEPs) elicited by 5Hz rTMS and 5Hz rTMS combined with MVC., Methods: In this randomized, controlled, assessor-blinded, crossover trial, 40 healthy right-handed subjects were divided into group A (n=20) and group B (n=20). All subjects received rTMS over the primary motor cortex area (M1) in the left hemisphere. The parameters of rTMS were 5 Hz, 90.of the resting motor threshold (RMT), for a total of 500 pulses in 100 trains (1-sec inter-stimulus and 8- sec inter-interval). Method 1: All subjects received rTMS over the hand area of left M1. Method 2: All subjects received rTMS at the same stimulated point, combined with maximum voluntary hand griping in each 10 trains. Test 1: group A underwent method 1, while group B underwent method 2. Test 2: A week later, group B underwent method 1, while group A underwent method 2. In each test, the MEP amplitude and latency was measured before (P-rTMS), 5 min after (Post 1) and 30 min after (Post 2) the rTMS intervention., Results: MEP amplitude increased significantly from baseline at 5 minutes post intervention under both treatment regimes. However for both sequences, it decreased towards baseline under the rTMS intervention at 30 minutes post intervention but remained relatively high when rTMS was combined with MVC. MEP latency decreased significantly from baseline at 5 minutes post intervention under both treatment regimes. For both sequences, it then increased again towards baseline under both treatment regimes at 30 minutes post intervention. Although there was a trend for a less pronounced increase under the combined treatment, this effect was not significant., Conclusion: Both 5 Hz rTMS and 5 Hz rTMS combined with MVC facilitate motor cortical excitability, but the enhancement in rTMS with MVC is more pronounced and maintained longer than simple rTMS.

A fundamental approach for resolving motor deficits in patients suffering from various neurological diseases is to improve the impaired cortical function through the modulation of plasticity. In order to advance clinical practice in this regard, it is necessary to better understand the interactions that occur between functional neuromuscular activity and the resulting cortical plasticity. This study tested whether the voluntary contraction of an antagonist muscle modulates the plasticity-like effect of continuous theta burst stimulation (cTBS) recorded from the agonist. The effects of various opposing torques produced by the antagonist were also measured. As a result, the suppressing effect of cTBS was enhanced by mild antagonist contraction, whereas effortful antagonist contraction suspended the plasticity caused by cTBS. In contrast, the antagonist contractions right after cTBS did not significantly influence the effect of cTBS. The results indicate that the antagonist activity alters the effect of cTBS, especially in protocols with synchronous magnetic stimulation and antagonist contraction. Such modulation on cTBS may be through a reciprocal mechanism within the motor cortex, although the spinal regulation of the motoneuronal pool cannot be fully excluded. The present findings are beneficial for elucidating the mechanism of neuromuscular control and for resolving related neurological disorders., (© 2014 Federation of European Neuroscience Societies and John Wiley & Sons Ltd.)

Objective: Among other neuroimaging techniques, functional magnetic resonance imaging (fMRI) can be useful for studying the development of motor fatigue. The aim of this study was to identify differences in cortical neuronal activation in nine subjects on three motor tasks: right-hand movement with minimum, maximum, and post-fatigue maximum finger flexion., Materials and Methods: fMRI activation maps for each subject and during each condition were obtained by estimating the optimal model of the hemodynamic response function (HRF) out of four standard HRF models and an individual-based HRF model (ibHRF)., Results: ibHRF was selected as the optimal model in six out of nine subjects for minimum movement, in five out of nine for maximum movement, and in eight out of nine for post-fatigue maximum movement. As compared to maximum movement, a large reduction in the total number of active voxels (primary sensorimotor area, supplementary motor area and cerebellum) was observed in post-fatigue maximum movement., Conclusion: This is the first approach to the evaluation of long-lasting contraction effort in healthy subjects by means of the fMRI paradigm with the use of an individual-based hemodynamic response. The results may be relevant for defining a baseline in future studies on central fatigue in patients with neuropathological disorders.

Study Design: A 2 (group) × 2 (abdominal contraction) × 2 (reach activity) crossover mixed design with repeated measures for contraction and activity examined the effects of a loaded (4.6 kg) forward-reach activity and abdominal drawing-in maneuver (ADIM) on transversus abdominis (TrA) contraction in subjects with nonspecific low back pain (NSLBP) history versus controls., Objective: We measured TrA contraction during a loaded forward-reach activity while using the ADIM and examined if a NSLBP history affects TrA activity., Summary of Background Data: The ADIM supports trunk stability during function. Clinicians incorporate ADIM during patients' functional tasks. Pain-free individuals can sustain ADIM during function, such as forward-reach. However, this has not been tested in those with a NSLBP history., Methods: Eighteen normal subjects and 18 subjects with a history of NSLBP participated. A blinded investigator recorded M-mode ultrasound imaging measurements of TrA thickness (mm) during 4 conditions as follows: (1) quiet standing without ADIM; (2) quiet standing with ADIM; (3) loaded forward-reach without ADIM; and (4) loaded forward-reach with ADIM., Results: A mixed analysis of variance demonstrated a significant main effect for group (F [1, 34] = 5.404, P = 0.026; ), where TrA thickness was greater for NSLBP history (7.41 + 2.34 mm) versus controls (5.9 + 2.46 mm). A significant main effect was observed for abdominal contraction (F [1, 34] = 49.57, P < 0.0001; ), where TrA thickness was greater during ADIM (7.47 + 2.7 mm) versus without ADIM (5.84, 1.92 m). A significant main effect was observed for forward-reach activity (F [1, 34] = 12.79, P = 0.001; ), where TrA thickness was greater during a loaded forward-reach (7.04 + 2.6 mm) versus quiet standing (6.2 + 2.4 mm). There were no significant interactions., Conclusion: Individuals can use a volitional pre-emptive ADIM for trunk protection during loaded forward-reach, potentially reducing injury risk. A NSLBP history increases TrA activation during ADIM, suggesting an enhanced protective role., Level of Evidence: 2.

Purpose: To investigate whether the H reflex-M wave recruitment curves obtained during maximal voluntary contraction (MVC) differ from rest and to determine the stimulation intensities allowing to record stable reflex responses., Methods: Full recruitment curves (precision, 2 mA) were obtained from the soleus muscle in 14 volunteers at rest and during plantar flexion MVCs., Results: Maximal M-wave reached significantly larger amplitude during MVC (+2.2 [0.4; 3.9] mV) for a higher stimulation intensity (+7.9 [-0.4; 16] mA). Similarly, maximal H-reflex reached significantly larger amplitude during MVC than at rest (+3.2 [0.9; 5.5] mV) for a much higher stimulation intensity (+17.7 [9.7; 25.7] mA). V-wave amplitude plateaued only when M-wave during MVC plateaued, that is, at higher intensity than M-wave at rest. V-wave was correlated to the maximal H-reflex during MVC (r = 0.79, P < 0.05)., Conclusion: Electrically evoked potentials showed a specific recruitment curve during MVC with higher maximal values attained for higher stimulation intensities. Thus, recording reflex responses during MVC based on intensities determined at rest or as a percentage of M-wave may yield inaccurate results. V-wave presented a plateau for stimulation intensity of 1.5 times the onset of the resting M-wave plateau. Evoked potentials obtained during actual contractions should be normalized to M-waves obtained during contractions of the same force level.

The purpose of the present study was to examine the influence of activation capabilities on the electromyography (EMGRMS) and mechanomyography amplitude (MMGRMS)-force relationships of the vastus lateralis (VL) and rectus femoris (RF). Thirteen men (mean±SD; age=22±3 year) performed nine submaximal contractions (10-90% maximal voluntary contraction [MVC]) with the interpolated twitch technique performed during a separate contraction at 90% MVC to calculate percent voluntary activation (%VA). Nine participants with >90% VA were categorized into the high-activated group with the remaining categorized into the moderate-activated group. Slopes (b terms) were calculated from the log-transformed EMGRMS and MMGRMS-force relationships. The b terms (collapsed across the VL and RF) for the EMGRMS-force relationships were greater for the high- (1.29±0.31) than the moderate-activated (1.10±0.20) group. In contrast, there were no differences in the b terms for the MMGRMS-force relationships between the high- and moderate-activated groups. For the EMGRMS and MMGRMS-force relationships, the b terms were greater for the RF (1.38±0.30, 0.81±0.20) than the VL (1.08±0.19, 0.60±0.13) collapsed across groups. The b terms from the EMGRMS-force relationships, but not the MMGRMS-force relationships, reflected differences in %VA., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

There is some evidence that the Biering-Sorensen endurance test can discriminate low back pain sufferers from healthy individuals and can predict future back pain. This test relies on the subject's ability to voluntarily drive the back muscles. This neural drive, termed voluntary activation (VA) can be measured using the twitch interpolation technique. The aim of the current study was to investigate the relationship between back muscle endurance and VA. Twenty-one healthy volunteers (10 males) participated. Bilateral electromyographic recordings were obtained from erector spinae and rectus abdominis. Back extensor torque was recorded using a dynamometer. The protocol consisted of measurement of VA (using magnetic stimulation of the brain and assessment of the sizes of the evoked twitches) and measurement of endurance. There was a linear correlation (r(2)=1, P<0.01) between voluntary torque and VA. The mean (SEM) endurance time was 174.9 (12.8)s. There was no correlation between endurance and VA at either 100% MVC (r(2)=0.01, P=0.72) or at 50% MVC (r(2)=0.11, P=0.16). These findings indicate that the endurance of the back muscles, as assessed using this widely utilised test does not appear to be related to a subject's ability to drive their back muscles voluntarily either maximally or submaximally., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

Stretch of activated muscles leads to enhanced forces compared to isometric contractions at the same muscle length and the same level of activation. This so-called residual force enhancement (RFE) is thought to be a property of all muscles and preparations. However, observations concerning the existence, amount and duration of RFE are inconsistent, especially for voluntary activated large human muscles. Therefore, physiological relevance for daily activity is still questionable and the purpose of this study was to examine whether RFE is present in voluntary sub-maximal activated quadriceps femoris (QF). Seated in a rotational dynamometer with EMG attached to superficial parts of QF, 30 subjects performed isometric and isometric-eccentric-isometric contractions (20° stretch, ω=60°s(-1)) at 30% and 60% of maximum voluntary activation (MVA) and contraction (MVC). To account for the complexity of the multi-headed QF, a compensation model based on physiological cross-sectional area and individual EMG-torque relations was used to interpret EMG data. For both levels of intensity and both feedback control strategies, ANOVA identified significant RFE (at the same level of activation) and reduced activation (at the same level of torque). Against expectations, RFE was independent of the level of activation., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

We studied postural adjustments associated with a quick voluntary postural sway under two conditions, self-paced and simple reaction-time. Standing subjects were required to produce quick discrete shifts of the center of pressure (COP) forward. About 400-500ms prior to the instructed COP shift, there were deviations of the COP in the opposite direction (backwards) accompanied by changes in the activation levels of several postural muscles. Under the reaction-time conditions, the timing of those early postural adjustments did not change (repeated measures MANOVA: p>0.05) while its magnitude increased significantly (confirmed by repeated measures MANOVA: p<0.05). These observations are opposite to those reported for anticipatory postural adjustments under simple reaction time conditions (a significant change in the timing without major changes in the magnitude). We conclude that there are two types of feed-forward postural adjustments. Early postural adjustments prepare the body for the planned action and/or expected perturbation. Some of these preparatory actions may be mechanically necessary. Later, anticipatory postural adjustments generate net forces and moments of force acting against those associated with the expected perturbation. Both types of adjustments fit well the referent configuration hypothesis, which offers a unified view on movement-posture control., (Published by Elsevier Ltd.)

To assess the surface electromyographic spectral characteristics of masticatory and neck muscles during the performance of maximum voluntary clench (MVC) tasks, 29 healthy young adults (15 men, 14 women, mean age 22years) were examined. Electromyography of masseter, temporalis and upper trapezius muscles was performed during 5-s MVCs either on cotton rolls or in intercuspal position. Using a fast Fourier transform, the median power frequency (MPF) was obtained for the first and last seconds of clench, and compared between sexes, muscles, sides, tests and time intervals using ANOVAs. On average, the MPFs did not differ between sexes or sides (p>0.05), but significant effects of muscle (MPF temporalis larger than masseter, larger than trapezius muscles), test (larger MPFs when clenching in intercuspal position than when clenching on cotton rolls) and time (larger MPFs in the first than in the fifth second of clench) were found. In conclusion, a set of data to characterize the sEMG spectral characteristics of jaw and neck muscles in young adult subjects performing MVC tasks currently in use within the dental field was obtained. Reference values may assist in the assessment of patients with alterations in the cranio-cervical-mandibular system., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Agonist-mediated reciprocal inhibition (RI) in distal skeletal muscles is an important neurophysiological phenomenon leading to improved movement coordination and efficiency. It has been shown to be reduced in aged and clinical populations, so the development of interventions augmenting RI is an important research goal. We examined the efficacy of using chronic passive muscle stretching to augment RI. The influence of 3 wk of plantarflexor stretching (4 × 30 s, two times/day) on RI of soleus and gastrocnemius initiated by tonic, voluntary dorsiflexion contractions [20% of maximum voluntary contraction (MVC)] was examined in 11 healthy men who performed stretch training and in nine nontraining controls. Hoffmann's reflexes (H-reflexes) were elicited by tibial nerve stimulation during both weak isometric (2% MVC) plantarflexions and dorsiflexion contractions at 20% MVC. Changes were examined at three joint angles, normalized to each subject's range of motion (ROM; plantarflexed = 10 ± 0°, neutral = -3.3 ± 2.9°, dorsiflexed = -16.5 ± 5.6°). No changes were detected in controls. A 20% increase in ROM in the stretch subjects was associated with a significant decrease in maximum H-reflex (H(max)): maximum evoked potential (M(max)), measured during 2% plantarflexion at the plantarflexed and neutral angles in soleus and at the plantarflexed angle in gastrocnemius (P < 0.05-0.01). By contrast, decreases in H(max):M(max) during 20% dorsiflexion contract were also seen at each angle in soleus and at the dorsiflexed angle in gastrocnemius. However, a greater decrease in H(max):M(max) measured during voluntary dorsiflexion rather than during plantarflexion, which indicates a specific change in RI, was detected only at the dorsiflexed angle (-30.7 ± 9.4% and -35.8 ± 6.8% for soleus and gastrocnemius, respectively). These results demonstrate the efficacy of soleus-gastrocnemius stretch training in increasing agonist-mediated RI from tibialis anterior onto soleus-gastrocnemius in young, healthy individuals at dorsiflexed, but not plantarflexed, joint angles.

We examined elbow muscle activities and movement kinematics to determine how subjects combine elementary control actions in performing movements with one and two trajectory segments. In reaching, subjects made a rapid elbow flexion to a visual target before stabilizing the limb with either a low or a higher level of elbow flexor/extensor coactivity (CoA), which was cued by target diameter. Cursor diameter provided real-time biofeedback of actual muscle CoA. In reversing, the limb was to reverse direction within the target and return to the origin with minimal CoA. We previously reported that subjects overshoot the goal when attempting a reversal after first having learned to reach accurately to the same target. Here we test the hypothesis that this hypermetria results because reversals co-opt the initial feedforward control action from the preceding trained reach, thereby failing to account for task-dependent changes in limb impedance induced by differences in flexor/extensor coactivity as the target is acquired (higher in reaching than reversing). Instructed increases in elbow CoA began mid-reach, thus increasing elbow impedance and reducing transient oscillations present in low CoA movments. Flexor EMG alone increased at movement onset. Test reversals incorporated the initial agonist activity of previous reaches but not the increased coactivity at the target, thus leading to overshoot. Moreover, we observed elevated coactivity in reversals upon returning to the origin even though coactivity in reaching was centered at the goal target. These findings refute the idea that the brain necessarily invokes distinct unitary control actions for reaches and reversals made to the same target. Instead, reaches and reversals share a common control action that initiates trajectories toward their target and another later control action that terminates movement and stabilizes the limb about its final resting posture, which differs in the two tasks.

Introduction: The effects of postactivation potentiation (PAP) on evoked contractions are well understood, but less is known about the effect of PAP on voluntary submaximal contractions. Using a measure of neuromuscular efficiency (NME) [NME = (mV EMG / Nm torque)] we explored the effects of PAP in the triceps brachii at two muscle lengths., Methods: Evoked twitch and NME were compared at short (40° elbow flexion) and long (120°) muscle lengths. At each length, 12 subjects performed a contraction of 25% maximum voluntary contraction (MVC) torque before and after a potentiating MVC., Results: Twitch torque potentiated more at short length (216.9 ± 169.3%) than at long length (77.3 ± 32.6%), but PAP moderately improved NME only at short length (12.2 ± 8.7%)., Conclusions: The greater capacity for PAP at the short length is reflected by greater NME. Compared with evoked responses, the relatively small change in NME suggests a different and more modest role of PAP during voluntary submaximal contractions., (Copyright © 2011 Wiley Periodicals, Inc.)

Normalized surface EMG requires known muscle activity from a well defined reference contraction, often a maximum voluntary isometric exertion (MVIE). One factor that may impact the activity obtained during a MVIE is the use of a bilateral or unilateral version of the test. The objective of this work was to investigate how performing MVIE tests unilaterally or bilaterally affects maximum voluntary electrical activity (MVE) for selected shoulder muscles. Further, we examined if within-participant reproducibility could be improved with the use of bilateral or unilateral MVIE tests. Sixteen participants completed five repetitions of a palm press, empty can, and external rotation MVIE tests both bilaterally and unilaterally. Muscle activity was recorded from seven muscles acting at the glenohumeral joint during the performance of each test. Repeated measures ANOVAs were used to determine within-participant differences on muscle specific MVEs and hand force measures between the two versions of the test. Gender was compared as a between participants factor. Within-participant reproducibility in muscle activity between bilateral and unilateral MVIEs was compared using coefficients of variation. Performing a bilateral empty can MVIE test elicited significantly higher MVEs for the upper trapezius (14%) and supraspinatus muscles (15%) than a unilateral test in this study population. MVEs during the palm press and external rotation were not affected by performing the tests bilaterally or unilaterally. Reliability was similar between unilateral and bilateral exertions across MVIE tests. Hence, we recommend that the empty can MVIE test be performed bilaterally, while either bilateral or unilateral configurations are equivalent during the palm press and external rotation MVIE tests., (Copyright © 2010 Elsevier Ltd. All rights reserved.)

Force enhancement following muscle stretching and force depression following muscle shortening are well-accepted properties of skeletal muscle contraction. However, the factors contributing to force enhancement/depression remain a matter of debate. In addition to factors on the fiber or sarcomere level, fiber length and angle of pennation affect the force during voluntary isometric contractions in whole muscles. Therefore, we hypothesized that differences in fiber lengths and angles of pennation between force-enhanced/depressed and reference states may contribute to force enhancement/depression during voluntary contractions. The purpose of this study was to test this hypothesis. Twelve subjects participated in this study, and force enhancement/depression was measured in human tibialis anterior. Fiber lengths and angles of pennation were quantified using ultrasound imaging. Neither fiber lengths nor angles of pennation were found to differ between the isometric reference contractions and any of the force-enhanced or force-depressed conditions. Therefore, we rejected our hypothesis and concluded that differences in fiber lengths or angles of pennation do not contribute to the observed force enhancement/depression in human tibialis anterior, and speculate that this result is likely true for other muscles too.

The present study aimed to investigate neurophysiologic mechanisms mediating the newly discovered phenomenon of respiratory-motor interactions and to explore its potential clinical application for motor recovery. First, young and healthy subjects were instructed to breathe normally (NORM); to exhale (OUT) or inhale (IN) as fast as possible in a self-paced manner; or to voluntarily hold breath (HOLD). In experiment 1 (n = 14), transcranial magnetic stimulation (TMS) was applied during 10% maximal voluntary contraction (MVC) finger flexion force production or at rest. The motor-evoked potentials (MEPs) were recorded from flexor digitorum superficialis (FDS), extensor digitorum communis (EDC), and abductor digiti minimi (ADM) muscles. Similarly, in experiment 2 (n = 11), electrical stimulation (ES) was applied to FDS or EDC during the described four breathing conditions while subjects maintained 10%MVC of finger flexion or extension and at rest. In the exploratory clinical experiments (experiment 3), four patients with chronic neurological disorders (three strokes, one traumatic brain injury) received a 30-min session of breathing-controlled ES to the impaired EDC. In experiment 1, the EDC MEP magnitudes increased significantly during IN and OUT at both 10%MVC and rest; the FDS MEPs were enhanced only at 10%MVC, whereas the ADM MEP increased only during OUT, compared with NORM for both at rest and 10%MVC. No difference was found between NORM and HOLD for all three muscles. In experiment 2, when FDS was stimulated, force response was enhanced during both IN and OUT, but only at 10%MVC. When EDC was stimulated, force response increased at both 10%MVC and rest, only during IN, but not OUT. The averaged response latency was 83 ms for the finger extensors and 79 ms for the finger flexors. After a 30-min intervention of ES to EDC triggered by forced inspiration in experiment 3, we observed a significant reduction in finger flexor spasticity. The spasticity reduction lasted for ≥ 4 wk in all four patients. TMS and ES data, collectively, support the phenomenon that there is an overall respiration-related enhancement on the motor system, with a strong inspiration-finger extension coupling during voluntary breathing. As such, breathing-controlled electrical stimulation (i.e., stimulation to finger extensors delivered during the voluntary inspiratory phase) could be applied for enhancing finger extension strength and finger flexor spasticity reduction in poststroke patients.

Visualization of the geometric deformation and associated displacement patterns of tubular abdominal organs to mechanical stimuli provides a quantitative measure that is useful in modeling their elastic properties. The origin of the stimulus may be the result of direct and voluntary muscle contraction or in response to a triggered reflex activity. Using trans-perineal 2D ultrasound imaging we examined the characteristics of deformation and displacement of these organs in response to voluntary activity, contraction, straining, and fast reflex responses to stimuli such as coughing. The relative time sequence in movement was examined by serially segmenting the outline of these structures and mapping their temporal characteristics.

Currently, contrasting views exist regarding which body and arm postures are most effective for eliciting maximal voluntary exertions in the shoulder muscles. Informed exertion standardization may improve comparisons between subjects and muscle groups for normalized electromyography values. Additionally, identifying exertions that can produce equivalent maximal electrical activity values can reduce experimental setup time and reduce the likelihood of fatigue development. This research study examined twelve posture and force direction defined test exertions to identify those that elicited maximal electrical activity from the deltoid (anterior and middle fibres) and pectoralis major (clavicular and sternal heads). Further, the question of whether a single test exertion could obtain maximal electrical activity from multiple muscle fascicles was explored. Maximal activation was demonstrated for the deltoid during several exertions that incorporated an upward force exertion and the pectoralis major for multiple exertions that included an inward force direction. Finally, two test exertions produced maximal electrical activity from both muscles of interest. This research supports the notion that a range of exertions can elicit maximal electrical activity from a muscle, rather than one specific exertion. This suggests that researchers may be able to leverage a smaller set of test exertions to evaluate multiple muscles simultaneously without loss of data quality, and thereby decrease overall experimental data collection time while maintaining high fidelity data., (Copyright (c) 2009 Elsevier Ltd. All rights reserved.)

Superimposition of electrical stimulation during voluntary contractions is used to produce functional movements in individuals with central nervous system impairment, to evaluate the ability to activate a muscle, to characterize the nature of fatigue, and to improve muscle strength during postsurgical rehabilitation. Currently, the manner in which voluntary contractions and electrically elicited forces summate is not well understood. The objective of the present study is to develop a model that predicts the forces obtained when electrical stimulation is superimposed on a volitional contraction. Quadriceps femoris muscles of 12 able-bodied subjects were tested. Our results showed that the total force produced when electrical stimulation was superimposed during a volitional contraction could be modeled by the equation T=V+S[(MaxForce-V)/MaxForce]N, where T is the total force produced, V is the force in response to volitional contraction alone, S is the force response to the electrical stimulation alone, MaxForce is the maximum force-generating ability of the muscle, and N is a parameter that we posit depends on the differences in the motor unit recruitment order and firing rates between volitional and electrically elicited contractions. In addition, our results showed that the model predicted accurately (intraclass correlation coefficient>or=0.97) the total force in response to a wide range of stimulation intensities and frequencies superimposed on a wide range of volitional contraction levels. Thus the model will be helpful to clinicians and scientists to predict the amount of stimulation needed to produce the targeted force levels in individuals with partial paralysis.

This study aimed to determine whether the time-course of maximal resting H-reflex amplitude (H(max)) recovery after a prolonged moderate-intensity muscle contraction differs according to the optimal stimulation intensity used (predetermined vs. readjusted). Thirteen males performed a sustained isometric plantar flexion at 40% of their maximal voluntary contraction torque output until exhaustion. H(max) of the soleus muscle was recorded before and 2, 6, 10, and 14 min after the end of the contraction, then normalized by the respective maximal M-wave to form the H(max)/M(max) ratio. During recovery, pre- and redetermined optimal stimulation intensities (mini-recruitment curve drawn before each recovery measurement) were applied randomly to measure H(max). When using redetermined stimulation intensities, normalized H-reflex values were systematically greater (+11, +16, +15, and +15% after 2-, 6-, 10-, and 14-min recovery periods, respectively) than those obtained with the predetermined intensity. Keeping the stimulation intensity constant to evoke H(max) after a sustained muscle contraction can underestimate the H-reflex facilitation occurring after exhaustive exercise. It is therefore more appropriate to redefine the optimal stimulation intensity to evoke H(max) (using mini-recruitment curves) when the purpose is to analyze spinal modulation during the recovery phase.

The aim of this study was to compare the possible changes in muscle activation level between a first and second bout of damaging eccentric exercise performed at 2 weeks interval (i.e. repeated bout effect). To that purpose, ten physically active males took part in this study. The eccentric exercise consisted of 10 sets of 12 maximal voluntary contractions (MVC) produced by the knee extensors during movements performed at a constant speed of 160 degrees s(-1). Changes in voluntary and electrically evoked torque in concentric and/or isometric conditions were assessed at the following time points: pre-exercise, and 2 min, 1 and 24 h after each eccentric exercise. At the same time points, voluntary activation was quantified by the superimposed electrical stimulation technique. Muscle soreness and plasma CK activity were measured within 48 h after the eccentric exercise. The results showed that the decrease in eccentric peak torque was linear throughout the exercise protocol. At the end of bouts 1 and 2, torque was significantly reduced by 27.7 +/- 9.1 and 23.4 +/- 11.2, respectively, with no difference between bouts (P > 0.05). At 24 h post-exercise, a lower reduction (P < 0.05) in MVC (17.8 +/- 5.4%) and electrically evoked (16.7 +/- 4.6%) isometric torque was observed for bout 2. In contrast, no statistical difference was found in the deficit in voluntary activation between the two bouts. In conclusion, our results indicate that the repeated bout effect of eccentric exercise appears to reduce muscle damage, but does not influence the level of voluntary activation.

The normalization of electromyographic signals to a maximum voluntary reference contraction is common practice within the ergonomics research paradigm. However, there is a lack of support for a common protocol for obtaining maximum repeatable exertions. Specifically, there is minimal evidence to support the use of providing force magnitude feedback during the production of voluntary maximum exertions (MVE) in terms of both signal amplitude and repeatability. Therefore, the purpose of this study was to determine (1) if an MVE force magnitude feedback protocol increased both the force exerted and corresponding muscle activity, (2) if force magnitude feedback improved the within-participant reproducibility of the force or activity observed, and (3) if the surface electromyography (sEMG) signal processing method affected the repeatability of determining peak muscle activity. Seventeen participants completed a series of MVEs; first without feedback of the forces they produced, then with feedback of the forces they were producing, and again without feedback to determine if providing force feedback influenced their ability to produce a maximum force. Hand force and sEMG from the anterior deltoid, middle deltoid, and infraspinatus were measured during each exertion. The results showed that the highest forces and muscle activities were achieved when force feedback was provided. Force magnitude feedback resulted in a 7-22% increase in magnitude (for force and activity) and a decrease of 11-46% in the coefficient of variation specifying an improvement in the within-participant reproducibility. Signal processing techniques also affected within-participant reproducibility, however to a much lesser extent. The peak value from a 500-ms moving window average of the linear enveloped or root mean squared sEMG was the most reproducible technique tested.

Interpolated twitch technique (ITT) is a non-invasive method for assessing the completeness of muscle activation in clinical settings. Voluntary activation level (VA), measured by ITT and estimated by a conventional linear model, was reported to have a non-linear relationship with true voluntary contraction force at higher activation levels. The relationship needs to be further clarified for the correct use by clinicians and researchers. This study was to established a modified voluntary activation (modified VA) and define a valid range by fitting a non-linear logistic growth model. Eight healthy male adults participated in this study. Each subject performed three sets of voluntary isometric ankle plantar flexions at 20, 40, 60, 80 and 100% maximal voluntary contraction (MVC) with real-time feedback on a computer screen. A supramaximal electrical stimulation was applied on tibia nerve at rest and during contractions. The estimated VA was calculated for each contraction. The relationship between the estimated VA and the actual voluntary contraction force was fitted by a logistic growth model. The result showed that according to the upper and lower limit points of the logistic curve, the valid range was between the 95.16% and 10.55% MVC. The modified VA estimated by this logistic growth model demonstrated less error than the conventional model. This study provided a transfer function for the voluntary activation level and defined the valid range which would provide useful information in clinical applications.

Dynamic joint stiffness defines the dynamic relationship between the position of the joint and the torque acting about it. Joint stiffness consists of two components: intrinsic and reflex stiffness. Previous work from our lab has shown that subjects can alter their reflex stiffness voluntarily and independently of intrinsic stiffness. Numerous studies have investigated whether reflex stiffness is altered in a task-dependent fashion; however the results of these studies are inconclusive. We designed an experimental paradigm where subjects were faced with 3 tasks: one task where joint stiffness aided subjects, a second where joint stiffness hindered the subjects and a third where joint stiffness had no effect. We found that subjects did not alter their joint stiffness to perform the different tasks. Rather, they performed the tasks by voluntarily producing the appropriate torque based on visual feedback. Thus, with the paradigm used in this study, reflex stiffness was not modulated in a task-dependent manner.

Background: Middle-aged adults show a higher incidence of falls compared to young adults when performing outdoor physical activities. This study investigated whether or not the patterns and quantitative characteristics of the trunk and lower extremity muscle activations associated with stepping, which represents an important movement for arresting falls, differ between middle-aged adults and young adults., Methods: Nine healthy young adults (age=22[3] years) and nine healthy middle-aged adults (age=52[8] years), performed a single-step, volitional, fast forward stepping movement with each leg. The stepping movement was divided into the step-initiation, single-leg-support, and landing phases based on foot-switch signals. The activation sequence, occurrence rate, onset latency, burst duration, and normalized co-contraction duration of the tibialis anterior, medial gastrocnemius, rectus femoris, biceps femoris, and gluteus medius of the stance and swing legs and that of bilateral erector spinae muscles were analyzed using surface electromyography. We defined the essential muscle activation as exhibiting an occurrence rate of 90% or more in all of these trials., Findings: As compared to young adults, the middle-aged adults demonstrated several additional essential bursts throughout the stepping movement. Middle-aged adults also displayed significantly longer burst durations of the biceps femoris and medial gastrocnemius of the swing leg after landing, as well as longer co-contraction of the rectus femoris and biceps femoris of the stance leg in the single-leg-support phase (P<0.05)., Interpretation: Age-related changes in step-related neuromuscular control exist in healthy middle-aged adults. We propose that training focused on improving or maintaining neuromuscular control associated with volitional leg movements may benefit middle-aged individuals.

Stretch reflexes and motor-evoked potentials (MEPs) of a muscle are facilitated when performing intensive contraction of muscles located in a different segment (remote effect). We investigated to what extent the remote effect on MEPs in the flexor carpi radialis (FCR) in humans is modulated during sustained maximal and submaximal voluntary contractions of the ipsilateral quadriceps (remote muscle). We found that even when the force of maximal voluntary contraction (MVC) of the remote muscle declined during sustained MVC, the magnitude of the remote effect on MEPs remained constant. Maximal electrical stimulation of the remote muscle and transcranial magnetic stimulation of the corresponding motor cortex revealed that the level of voluntary activation gradually decreased during the sustained MVC. The motor response in the FCR following magnetic stimulation at the level of the foramen magnum, which preferentially elicits muscle response as a direct response of the corticospinal tract, was not modified by the remote effect during the sustained MVC. This finding suggested that the excitability of the spinal motoneuron pool remained constant. In contrast to the sustained MVC, during sustained submaximal contraction of the remote muscle, the magnitude of the remote effect on MEPs gradually increased as muscle fatigue developed. These findings suggest that the remote effect on MEPs was dependent on the level of effort driving the remote muscle, but not on the actual level of force output of the remote muscle, and that the origin of the remote effect was supraspinal, putatively upstream of the primary motor cortex.

This study documents intra-session and inter-day reproducibility (coefficient of variation [V%]) and single measurement reliability (intra-class correlations [R(I)]; standard error of a single measurement [SEM%] [95% confidence limits]) of indices of neuromuscular performance elicited during peripheral nerve magnetic stimulation. Twelve adults (five men and seven women) completed 3 assessment sessions on 3 days, during which multiple assessments of knee flexor volitional and magnetically-evoked indices of electromechanical delay (EMD(V); EMD(E)), rate of force development (RFD(V); RFD(E)), peak force (PF(V); P(T)F(E)), and compound muscle action potential latency (LAT(E)) and amplitude (AMP(E)) were obtained. Results showed that magnetically-evoked indices of neuromuscular performance offered statistically equivalent levels of measurement reproducibility (V%: 4.3-31.2%) and reliability (R(I): 0.98-0.51) compared to volitional indices (V%: 3.7-25.2%; R(I): 0.98-0.64), which support the efficacy of both approaches to assessment and the indices PF(V), EMD(V), EMD(E) and LAT(E) offer the greatest practical utility for assessing neuromuscular performance.

This study investigated the feasibility of measuring voluntary activation of the trapezius muscle with twitch interpolation. Subjects (n=8) lifted the right shoulder or both shoulders against fixed force transducers. Stimulation of the accessory nerve in the neck was used to evoke maximal twitches in right trapezius. The twitch-like increments in force (superimposed twitches) evoked during different strength voluntary contractions were linearly related to voluntary force (r=-0.82 to -0.99). Hence, voluntary activation could be quantified by twitch interpolation with this stimulus. Comparison of unilateral and bilateral MVCs showed that maximal voluntary force was greater in unilateral than bilateral efforts (92.7+/-2.9% and 82.3+/-5.8% MVC, respectively) but voluntary activation was similar (88.6+/-9.6% and 91.7+/-5.2%). Trapezius is commonly affected in work-related musculoskeletal disorders. Measurement of voluntary activation will be a useful technique to demonstrate whether the reduced maximal voluntary force reported in such disorders is due to muscular or neural factors.

Along with afferent information, centrally generated motor command signals may play a role in joint position sense. Isometric muscle contractions can produce a perception of joint displacement in the same direction as the joint would move if unrestrained. Contradictory findings of perceived joint displacement in the opposite direction have been reported. As this only occurs if muscle spindle discharge in the contracting muscle is initially low, it may reflect increased muscle spindle firing from fusimotor activation, rather than central motor command signals. Methodological differences including the muscle contraction task and use of muscle conditioning could underlie the opposing findings. Hence, we tested perceived joint position during two contraction tasks ('hold force' and 'hold position') at the same joint (wrist) and controlled muscle spindle discharge with thixotropic muscle conditioning. We expected that prior conditioning of the contracting muscle would eliminate any effect of increased fusimotor activation, but not of central motor commands. Muscle conditioning altered perceived wrist position as expected. Further, during muscle contractions, subjects reported wrist positions displaced ~12 degrees in the direction of contraction, despite no change in wrist position. This was similar for 'hold force' and 'hold position' tasks and occurred despite prior conditioning of the agonist muscle. However, conditioning of the antagonist muscle did reduce the effect of voluntary contraction on position sense. The errors in position sense cannot be explained by fusimotor activation. We propose that central signals combine with afferent signals to determine limb position and that multiple sources of information are weighted according to their reliability.

Fluctuations in muscle force during steady voluntary contractions result from the summation of twitch forces produced by asynchronous activation of multiple motor units. We hypothesized that oscillatory lateral muscle displacement, measured with a non-contact high-resolution laser displacement sensor, is correlated with force fluctuations during steady, voluntary contractions with a human muscle. Eight healthy young adults (20-33 yrs) performed steady isometric contractions with the first dorsal interosseus muscle. Contraction intensity ranged from 2.5% to 60% of the maximal voluntary contraction force. Oscillatory lateral displacement of the muscle surface was measured with a high-resolution laser displacement sensor (0.5 microm resolution), concurrently with abduction force of the index finger. In the time-domain analysis, there was a significant positive peak in the cross-correlation function between lateral muscle displacement and force fluctuations. In addition, the amplitude increased linearly with contraction intensity in both signals. In the frequency-domain analysis, frequency content was similar in both signals, and there was significant coherence between signals for the major frequency range of the signals (<5 Hz). In conclusion, laser-detected lateral displacement of a hand muscle is correlated with force fluctuations across a wide range of contraction intensity during steady voluntary contractions in humans.

The purpose of this study was to examine the effects of postactivation potentiation (PAP) on the torque and rate of torque development for contractions evoked by short trains of stimuli at different frequencies, in young and elderly adults. Individual mechanical contributions to each electrical stimulus within trains were also analysed. Single pulse, and two- (PT2) and three-pulse trains (PT3) delivered at 20, 50, 80 and 100 Hz were evoked before and during a 10 min period after a 6-s conditioning MVC. The results show that PAP of the torque for PT2 decreased with the increase of the stimulation frequency for young (from 184.5 to 140.4% of control values) and elderly (from 140.5 to 109.6%). Regardless of the stimulation frequency, the peak of PAP was greater in young than in elderly and occurred immediately after the conditioning MVC but was delayed for the 100 Hz condition in elderly adults. For PT3, the results were similar although the extent of PAP was less. The PAP of the mechanical contributions within the trains also decreased with the augmentation of the stimulation frequency. For most of the frequencies above 20 Hz, the peak of PAP for each mechanical contribution was delayed by 1 min after the conditioning MVC. These results indicate an age- and frequency-related PAP saturation of the successive mechanical contributions within a train of stimuli that decrease with time. The functional implication of the findings is that PAP effect is lower and delayed at high compared with low activation rate.

The contribution of central fatigue during and after low- and high-force isometric contractions sustained until failure with age is not established. We compared the time to failure and changes in voluntary activation measured using motor point stimulation of 15 young and 15 old adults for an isometric contraction sustained with the elbow flexor muscles at 20% and 80% of maximal voluntary contraction (MVC) force. Young adults had a briefer time to task failure than old adults for the 20% MVC fatiguing contraction, but a similar duration for the 80% task. Voluntary activation was reduced at the end of the 20% MVC task, but by greater magnitudes for old than young adults. The reduction in MVC torque after the low-force task was associated with the reduction in voluntary activation. After the 80% task, voluntary activation declined to similar levels for the young and old adults. Electromyographic activity levels (% MVC) of the biceps brachii and brachioradialis muscles during the fatiguing contraction were greater for the old than young for the 20% MVC task, but similar with age for the 80% MVC task. Our findings indicate that intensity and duration of contraction can be manipulated in young and old adults to induce varying magnitudes of fatigue within the central nervous system. Aging increases: (1) fatigue within the central nervous system immediately after a low-force fatiguing contraction, and (2) the potential for large neural adaptations during neuromuscular rehabilitation in old adults.

We used the framework of the uncontrolled manifold hypothesis to explore the effects of practice on the composition of muscle groups (M-modes) and multi-M-mode synergies stabilizing the location of the center of pressure (COP). In particular, we tested a hypothesis that practice could lead to a transition from co-contraction muscle activation patterns to reciprocal patterns. We also tested a hypothesis that new sets of M-modes would form stronger synergies stabilizing COP location. Subjects practiced load release tasks for five days while standing on a board with a narrow support surface (unstable board). Their M-modes and indices of multi-M-mode synergies were computed during standing without instability and during standing on an unstable board before practice, in the middle of practice, and at the end of practice. During standing without instability, subjects showed two consistent M-modes uniting dorsal and ventral muscles of the body respectively (reciprocal modes). While standing on an unstable board, prior to practice, subjects commonly showed M-modes uniting muscle pairs with opposing actions at major leg joints-co-contraction modes. Such sets of M-modes failed to stabilize the COP location in the anterior-posterior direction. Practice led to better task performance reflected in fewer incidences of lost balance. This was accompanied by a drop in the occurrence of co-contraction M-modes and the emergence of multi-mode synergies stabilizing the COP location. We conclude that the central nervous system uses flexible sets of elemental variables (modes) to ensure stable trajectories of important performance variables (such as COP location). Practice can lead to adjustments in both the composition of M-modes and M-mode co-variation patterns resulting in stronger synergies stabilizing COP location.

Following a suprathreshold transcranial magnetic stimulation (TMS) to the primary motor cortex (M1) during voluntary muscle contraction, a motor evoked potential (MEP) occurs in the target muscle followed by a silent period (SP) in the electromyographic (EMG) activities. The present study investigated how short interval intracortical inhibition (SICI) and intracortical facilitation (ICF) change during the SP. The time course of MEP and motor threshold during the SP were examined in the right first dorsal interosseous muscle. Using a triple-pulse protocol, SICI and ICF were tested at different times during the SP. The effects of different intensities of the conditioning stimulation (CS) for SICI and ICF were also investigated during the SP and at rest. During the SP, MEP was inhibited and motor threshold was increased, whereas MEP latency and background EMG level were same as those at rest. SICI decreased during the SP over a wide range of CS intensities. ICF increased at higher CS intensity. We conclude that SICI is suppressed and ICF is facilitated during the SP and the effects are separate from the interruption of voluntary drive.

In cases of partial deficiency of muscle activation capacity, force augmentation can be achieved by hybrid activation, i.e., by combining electrical stimulation (ES) with volitional activation. In this activation modality the shares of the volitional and induced torques within the overall hybrid torque are unknown. The purpose of this study was to suggest a computational approach to parcel out the volitional and stimulation induced components of joint torque generated during combined voluntary and electrical activation of the Tibialis Anterior muscle (TA). For this purpose, isometric contraction of the TA was studied on 5 healthy subjects, using an activation protocol involving ES alone, volitional activation alone and hybrid activation. Ankle torque and TA EMG were measured. A computational algorithm was developed to dissociate the volitional from the overall torque, based on EMG filtering and on pre-measured calibration curves of volitional torque versus EMG. The results indicated that for a certain hybrid torque there is a linear decaying relationship between the induced torque and the volitional torque shares. Moreover, based on a defined enhancement ratio, the results indicate that within the range of stimulation intensities, there exist regions of increased facilitation, in which the stimulation efficiency is higher under combined compared to isolated conditions.

Previous research on human balance recovery suggests that, prior to an externally triggered postural perturbation, healthy subjects can pre-select their postural response based on the environmental context, but it is unclear whether this pre-selection includes the selection of a stepping leg when performing compensatory steps. We sought to determine how pre-selecting a stepping limb affects the compensatory steps and stability of young, healthy subjects when responding to postural perturbations. Nine healthy subjects (24-37 years of age) stepped in response to backward translations of a platform under their feet when, prior to the perturbations, the subjects either knew whether they were to step with their left or right leg to a visual target (the Predictable condition) or did not know whether to step with their left or right leg until one of two targets appeared at perturbation onset (the Unpredictable condition). The Unpredictable condition also included randomly inserted trials of toes-up rotations and backward translations without targets (catch trials). The results showed that, in the Predictable condition, the subjects consistently exhibited one anticipatory postural adjustment (APA; a lateral weight shift toward the stance limb) before stepping accurately to the target with the correct leg. In the Unpredictable condition, the subjects either (1) exhibited multiple APAs, late step onsets, and forward center-of-mass (CoM) displacements that were farther beyond their base of support, or (2) exhibited an early step with only one APA and kept their CoM closer to the base of support, but also stepped more often with the incorrect leg. Thus, when the subjects had to select a stepping leg at perturbation onset, they either became more unstable and used multiple APAs to delay stepping in order to provide enough time to select the correct stepping leg, or they stepped earlier to remain stable but often stepped with the incorrect leg. In addition, responses to catch trials in the Unpredictable condition included distorted step placements that resembled steps to anticipated targets, despite allowing the subjects to step with a leg of their choice and to a location of their choice. Lastly, the subjects' voluntary stepping latencies to visual targets presented without perturbations were twice as long as their stepping latencies to the backward platform translations. Therefore, healthy subjects appear to pre-select their stepping limb, even when the perturbation characteristics are unpredictable, because relying on visual input provided at perturbation onset requires a delayed response that leads to greater instability.

The effect of age on mechanomyography (MMG) has not been examined for electrically evoked contractions. Similar to torque, we expected that postactivation potentiation of the MMG would differ between young and old subjects. Additionally, under voluntary conditions, we compared normalized MMG and electromyographic (EMG) signals in relation to torque, and expected that MMG, unlike EMG, would be affected by age. In 10 young and 10 old men, electrical stimulation was delivered before and after a 10-s maximal voluntary contraction (MVC) to assess potentiation of contractile (twitch torque; Pt), electrical (M-wave amplitude), and mechanical (MMG amplitude) properties of the dorsiflexors. Subsequently, subjects performed voluntary contractions at 20%, 40%, 60%, 80%, and 100% MVC for calculation of normalized MMG-torque and EMG-torque relationships. Following the MVC, Pt and evoked MMG were larger than at rest in both groups, but M-wave amplitude was unchanged. Twitch potentiation was greater in young than old, whereas evoked MMG was unaffected by age. Under voluntary conditions, values for MMG and EMG were similar between groups, except for greater MMG at MVC in young men. The shape of MMG and EMG relationships to torque was similar only in young men. Using the aging model, our results indicate that potentiation of the mechanical components (MMG) differ from those of twitch torque. Furthermore, the comparison of normalized voluntary MMG with age provides additional support for the concept of age-related motor unit remodeling.

A characteristic of skilled movement is the ability of the CNS to predict the consequences of motor commands. When we lift an object there is an anticipatory increase in grip force that prevents a grasped object from slipping. When an object is pulled from our grasp by an external force, a reflexive modulation in grip force prevents slippage. Here we examine how external perturbations to a grasped object influence anticipatory grip force during object manipulation using a bimanual task, with each hand holding a computer-controlled object. Subjects were instructed to maintain the position of the object held in the right hand. Loading was applied to this restrained object: either self-generated by the action of their left hand or externally generated by a motor. The magnitude of the grip force response to self-generated loading increased after the object was loaded, and the latency of this response remained predictive of load force. This implies that external and self-generated loading increase the anticipatory grip force response. Unlinked trials, where the subject's moved their left hand but no loading was generated on the right-hand object were used to assess the presence of purely predictive control of grip force. External loading soon after self-generated loading maintained an existing predictive response once the linkage between the subject's action and object loading had been removed. However, external loading had no influence as the existing prediction decays. Therefore, the predictive grip force response during object manipulation can be significantly modified by object loading from an external source.

There are conflicting observations of the effects of fatigue on the sensitivity of large diameter Ia afferents. Our goal was to characterize any fatigue-related changes in the spinal reflex pathways during fatigue. Manipulation of the Ia afferent response by vibration and tendon tap, in which the motor neuron pool is modulated by both short- and long-loop activation from muscle spindles, were elicited before and after a fatigue task. The fatigue task consisted of intermittent submaximal and maximal voluntary contractions (MVCs). Percent voluntary activation fell from 98.75% MVC to 80.92% MVC following the fatigue task as measured by the twitch interpolation technique. Voluntary contractions of the same force profile as the force produced by 30 s of vibration were produced by having participants (n = 10) follow the trajectory on a computer monitor, before and after the fatigue task. Recruitment thresholds (RTs) of voluntarily activated units showed no change during fatigue; however, units activated via the reflex pathway were recruited approximately 30% sooner during fatigue (P < 0.05). The ratio of the electrical-to-mechanical response of the tendon tap increased significantly with fatigue. Our findings of decreased RTs in response to vibration and increased EMG activity during the tendon tap following the fatigue task indicate that Ia afferent input to the motoneuron pool was increased. The decrease in MVC force indicates that during this time the descending drive was compromised. These results provide evidence that the gain of the gamma loop is increased during fatigue, indicating possible peripheral neural compensation to the motor neuron pool in order to preserve force output.

Humans are known to show anticipatory adjustments in the grip force prior to a self-generated or predictable action or perturbation applied to a hand-held object. We investigated whether humans can also adjust covariation of individual finger forces (multi-finger synergies) prior to self-triggered perturbations. To address this issue, we studied adjustments in multi-digit synergies associated with applied load/torque perturbations while the subjects held a customized handle steadily. The main hypothesis was that the subjects would be able to demonstrate the phenomenon of anticipatory covariation, that is changes in covariation patterns among digit forces and moments of force in anticipation of a perturbation, but only when the perturbation was triggered by the subjects themselves. Based on the principle of superposition (decoupled grasping force and resultant torque control), we also expected to see different adjustments in indices of multi-digit synergies stabilizing the total gripping force and the total moment of force. The task for the subjects (n = 8) was to return the initial handle position as quickly as possible after a perturbation, which consisted of removing one of three loads hanging from the handle. There were six experimental conditions: two types of perturbations (self-triggered and experimenter-triggered) by three positions of the load (left, center, and right). Three-dimensional forces and moments of force recorded from each digit contact were used for the analysis. Indices of covariation among digit forces and among moments of force, previously employed for studying motor synergies, were computed across trials. Positive values of the indices reflected negative covariations of individual digit forces and moments of force (their inter-compensatory changes) to stabilize the total force and moment acting on the handle. In steady-state conditions, subjects showed strong positive indices for both digit forces and digit moments. Under the self-triggered conditions, changes in the indices of digit force and moment covariation were seen about 150 ms prior to the perturbation, while such changes were observed only after the perturbation under the experimenter-triggered conditions. Immediately following a perturbation, the indices of force and moment covariation rapidly changed to negative revealing the lack of inter-compensation among the individual digit forces and moments. Later, both indices showed a recovery to positive values; the recovery was faster in the self-triggered conditions than in the experimenter-triggered ones. During the steady-state phase after the perturbation, the indices of force and moment covariation decreased and increased, respectively, as compared to their values during the steady-state phase prior to the perturbation. We conclude that humans are able to adjust multi-digit synergies involved in prehensile tasks in anticipation of a self-triggered perturbation. These conclusions speak against hypotheses on the organization of multi-element actions based on optimal control principles. Different changes in the indices of force and moment covariation after a perturbation corroborate the principle of superposition. We discuss relations of anticipatory covariation to anticipatory postural adjustments.

During strong voluntary contractions, activity is not restricted to the target muscles. Other muscles, including contralateral muscles, often contract. We used transcranial magnetic stimulation (TMS) to analyse the origin of these unintended contralateral contractions (termed "associated" contractions). Subjects (n = 9) performed maximal voluntary contractions (MVCs) with their right elbow-flexor muscles followed by submaximal contractions with their left elbow flexors. Electromyographic activity (EMG) during the submaximal contractions was matched to the associated EMG in the left biceps brachii during the right MVC. During contractions, TMS was delivered to the motor cortex of the right or left hemisphere and excitatory motor evoked potentials (MEPs) and inhibitory (silent period) responses recorded from left biceps. Changes at a spinal level were investigated using cervicomedullary stimulation to activate corticospinal paths (n = 5). Stimulation of the right hemisphere produced silent periods of comparable duration in associated and voluntary contractions (218 vs 217 ms, respectively), whereas left hemisphere stimulation caused a depression of EMG but no EMG silence in either contraction. Despite matched EMG, MEPs elicited by right hemisphere stimulation were approximately 1.5-2.5 times larger during associated compared to voluntary contractions (P < 0.005). Similar inhibition of the associated and matched voluntary activity during the silent period suggests that associated activity comes from the contralateral hemisphere and that motor areas in this (right) hemisphere are activated concomitantly with the motor areas in the left hemisphere. Comparison of the MEPs and subcortically evoked potentials implies that cortical excitability was greater in associated contractions than in the matched voluntary efforts.

The purpose of this research was to examine the effects of voluntarily manipulating muscle activation and localized muscle fatigue on tibial response parameters, including peak tibial acceleration, time to peak tibial acceleration, and the acceleration slope, measured at the knee during unshod heel impacts. A human pendulum delivered consistent impacts to 15 female and 15 male subjects. The tibialis anterior and lateral gastrocnemius were examined using electromyography, thus allowing voluntary contraction to various activation states (baseline, 15%, 30%, 45%, and 60% of the maximum activation state) and assessing localized muscle fatigue. A skin-mounted uniaxial accelerometer, preloaded medial to the tibial tuberosity, allowed tibial response parameter determination. There were significant decreases in peak acceleration during tibialis anterior fatigue, compared to baseline and all other activation states. In females, increased time to peak acceleration and decreased acceleration slope occurred during fatigue compared to 30% and 45%, and compared to 15% through 60% of the maximum activation state, respectively. Slight peak acceleration and acceleration slope increases, and decreased time to peak acceleration as activation state increased during tibialis anterior testing, were noted. When examining the lateral gastrocnemius, the time to peak acceleration was significantly higher across gender in the middle activation states than at the baseline and fatigue states. The acceleration slope decreased at all activation states above baseline in females, and decreased at 60% of the maximum activation state in males compared to the baseline and fatigue states. Findings agree with localized muscle fatigue literature, suggesting that with fatigue there is decreased impact transmission, which may protect the leg. The relative effects of leg stiffness and ankle angle on tibial response need to be verified.

This study was designed to investigate the modulations of H-reflex and V-wave responses during passive and maximal active dynamic actions. Experiments were performed on 16 healthy males [age: 24 +/- 4 (SD) yr]. Maximal H-reflexes (Hmax) and M-waves (MmaxR) were evoked at the same muscle length during passive isometric, shortening and lengthening actions and during maximal voluntary isometric, concentric, and eccentric plantar-flexion. In all contraction types, supra-maximal stimulus intensity was used to evoke the superimposed maximal M wave (MmaxA) and V wave (V) of the soleus muscle. At rest, the Hmax/MmaxR ratio was significantly reduced during lengthening with respect to isometric and shortening actions (P < 0.05). For each action type, the ratio between H reflex superimposed to the contraction (Hsup) and MmaxA was not different from Hmax/MmaxR ratio. When plantar flexors were maximally voluntary activated, the Hsup/MmaxA ratio was still lower during eccentric contraction as compared with isometric and concentric efforts (0.33 +/- 0.03 vs. 0.47 +/- 0.02 and 0.50 +/- 0.03, P < 0.001), whereas V/MmaxA ratios were similar for all contraction types (isometric 0.26 +/- 0.02; concentric 0.23 +/- 0.03, and eccentric 0.24 +/- 0.02; P > 0.05). The V/MmaxA ratio was significantly lower than Hsup/MmaxA during isometric and concentric MVC (P < 0.001). No difference was observed between V/MmaxA and Hsup/MmaxA ratios during eccentric efforts. The H-reflex modulations, present during lengthening actions, were mainly attributed to presynaptic inhibition of Ia afferents and to homosynaptic postactivation depression. Results on V wave and H reflex suggest that during eccentric MVC, the spinal loop is specifically modulated by the supra-spinal centers and/or neural mechanisms at spinal level.

Glycogenolysis results in the selective catabolism of individual glycogen granules by glycogen phosphorylase. However, once the carbohydrate portion of the granule is metabolized, the fate of glycogenin, the protein primer of granule formation, is not known. To examine this, male subjects (n = 6) exercised to volitional exhaustion (Exh) on a cycle ergometer at 75% maximal O2 uptake. Muscle biopsies were obtained at rest, 30 min, and Exh (99 +/- 10 min). At rest, total glycogen concentration was 497 +/- 41 and declined to 378 +/- 51 mmol glucosyl units/kg dry wt following 30 min of exercise (P < 0.05). There were no significant changes in proglycogen, macroglycogen, glycogenin activity, or mRNA in this period (P > or = 0.05). Exh resulted in decreases in total glycogen, proglycogen, and macroglycogen as well as glycogenin activity (P < 0.05). These decrements were associated with a 1.9 +/- 0.4-fold increase in glycogenin mRNA over resting values (P < 0.05). Glycogenolysis in the initial exercise period (0-30 min) was not adequate to induce changes in glycogenin; however, later in exercise when concentration and granule number decreased further, decrements in glycogenin activity and increases in glycogenin mRNA were demonstrated. Results show that glycogenin becomes inactivated with glycogen catabolism and that this event coincides with an increase in glycogenin gene expression as exercise and glycogenolysis progress.