Rate-coding of spinal motoneurons with high-frequency magnetic stimulation of human motor cortex

Rate-coding in spinal motoneurons was studied using high-frequency magnetic stimulation of the human motor cortex. The subject made a weak contraction to cause rhythmic (i.e., tonic) discharge of a single motor unit in flexor (or extensor) carpi radialis or tibialis anterior, while the motor cortical representation of that muscle was stimulated with brief trains of pulses from a Pyramid stimulator (4 Magstim units connected by 3 BiStim modules). An "m@n" stimulus train consisted of m number of pulses (1-4), with an interpulse interval (IPI) of n ms (1-6). Peristimulus time histograms were constructed for each stimulus condition of a given motor unit, and related to the average rectified surface electromyography (EMG) from that muscle. Surface EMG responses showed markedly more facilitation than single-pulse stimulation, with increasing numbers of pulses in the train; responses also tended to increase in magnitude for the longer IPI values (4 and 6 ms) tested. Motor-unit response probability increased in a manner comparable to that of surface EMG. In particular, motoneurons frequently responded twice to a given stimulus train. In addition to recruitment of new motor units, the increased surface EMG responses were, in part, a direct consequence of short-term rate-coding within the tonically discharging motoneuron. Our results suggest that human corticomotoneurons are capable of reliably following high-frequency magnetic stimulation rates, and that this activity pattern is carried over to the spinal motoneuron, enabling it to discharge at extremely high rates for brief periods of time, a pattern known to be optimal for force generation at the onset of a muscle contraction.     

Gravitational force modulates muscle activity during mechanical oscillation of the tibia in humans

Mechanical oscillation (vibration) is an osteogenic stimulus for bone in animal models and may hold promise as an anti-osteoporosis measure in humans with spinal cord injury (SCI). However, the level of reflex induced muscle contractions associated with various loads (g force) during limb segment oscillation is uncertain. The purpose of this study was to determine whether certain gravitational loads (g forces) at a fixed oscillation frequency (30 Hz) increases muscle reflex activity in individuals with and without SCI. Nine healthy subjects and two individuals with SCI sat with their hip and knee joints at 90° and the foot secured on an oscillation platform. Vertical mechanical oscillations were introduced at 0.3, 0.6, 1.2, 3 and 5g force for 20 s at 30 Hz. Non-SCI subjects received the oscillation with and without a 5% MVC background contraction. Peak soleus and tibialis anterior (TA) EMG were normalized to M-max. Soleus and TA EMG were <2.5% of M-max in both SCI and non-SCI subjects. The greatest EMG occurred at the highest acceleration (5g). Low magnitude mechanical oscillation, shown to enhance bone anabolism in animal models, did not elicit high levels of reflex muscle activity in individuals with and without SCI. These findings support the g force modulated background muscle activity during fixed frequency vibration. The magnitude of muscle activity was low and likely does not influence the load during fixed frequency oscillation of the tibia.     

Dynamic dorsoventral stiffness assessment of the ovine lumbar spine

Posteroanterior spinal stiffness assessments are common in the evaluating patients with low back pain. The purpose of this study was to determine the effects of mechanical excitation frequency on dynamic lumbar spine stiffness. A computer-controlled voice coil actuator equipped with a load cell and LVDT was used to deliver an oscillatory dorsoventral (DV) mechanical force to the L3 spinous process of 15 adolescent Merino sheep. DV forces (48 N peak, approximately 10% body weight) were randomly applied at periodic excitation frequencies of 2.0, 6.0, 11.7 and a 0.5-19.7 Hz sweep. Force and displacement were recorded over a 13-22 s time interval. The in vivo DV stiffness of the ovine spine was frequency dependent and varied 3.7-fold over the 0.5-19.7 Hz mechanical excitation frequency range. Minimum and maximum DV stiffness (force/displacement) were 3.86+/-0.38 and 14.1+/-9.95 N/mm at 4.0 and 19.7 Hz, respectively. Stiffness values based on the swept-sine measurements were not significantly different from corresponding periodic oscillations (2.0 and 6.0 Hz). The mean coefficient of variation in the swept-sine DV dynamic stiffness assessment method was 15%, which was similar to the periodic oscillation method (10-16%). The results indicate that changes in mechanical excitation frequency and animal body mass modulate DV spinal stiffness.     

Influence of single-level lumbar degenerative disc disease on the behavior of the adjacent segments—A finite element model study

The current study investigated mechanical predictors for the development of adjacent disc degeneration. A 3-D finite element model of a lumbar spine was modified to simulate two grades of degeneration at the L4–L5 disc. Degeneration was modeled by changes in geometry and material properties. All models were subjected to follower preloads of 800 N and moment loads in the three principal directions of motion using a hybrid protocol. Degeneration caused changes in the loading and motion patterns of the segments above and below the degenerated disc. At the level (L3–L4) above the degenerated disc, the motion increased due to moderate degeneration by 21% under lateral bending, 26% under axial rotation and 28% under flexion/extension. At the level (L5-S1) below the degenerated disc, motion increased only during lateral bending by 20% due to moderate degeneration. Both the L3–L4 and L5-S1 segment showed a monotonic increase in both the maximum von Mises stress and shear stress in the annulus as degeneration progressed for all loading directions, expect extension at L3–L4. The most significant increase in stress was observed at the L5-S1 level during axial rotation with nearly a ten-fold increase in the maximum shear stress and 103% increase in the maximum von Mises stress. The L5-S1 segment also showed a progressive increase in facet contact force for all loading directions with degeneration. Nucleus pressure did not increase significantly for any loading direction at either the caudal or cephalic adjacent segment. Results suggest that single-level degeneration can increase the risk for injury at the adjacent levels.     

Predictive equations to estimate spinal loads in symmetric lifting tasks

Response surface methodology is used to establish robust and user-friendly predictive equations that relate responses of a complex detailed trunk finite element biomechanical model to its input variables during sagittal symmetric static lifting activities. Four input variables (thorax flexion angle, lumbar/pelvis ratio, load magnitude, and load position) and four model responses (L4–L5 and L5–S1 disc compression and anterior–posterior shear forces) are considered. Full factorial design of experiments accounting for all combinations of input levels is employed. Quadratic predictive equations for the spinal loads at the L4–S1 disc mid-heights are obtained by regression analysis with adequate goodness-of-fit (R²>98%, p<0.05, and low root-mean-squared-error values compared with the range of predicted spine loads). Results indicate that intradiscal pressure values at the L4–L5 disc estimated based on the predictive equations are in close agreement with available in vivo data measured under similar loadings and postures. Combinations of input (posture and loading) variable levels that yield spine loads beyond the tolerance compression limit of 3400 N are identified using contour plots. Ergonomists and bioengineers, faced with the dilemma of using either complex but more accurate models on one hand or less accurate but simple models on the other hand, have thereby easy-to-use predictive equations that quantifies spinal loads and risk of injury under different occupational tasks of interest.     

Effects of mechanical assistance on muscle activity and motor performance during isometric elbow flexion

Mechanical assistance on joint movement is generally beneficial; however, its effects on cooperative performance and muscle activity needs to be further explored. This study examined how motor performance and muscle activity are altered when mechanical assistance is provided during isometric force control of ramp-down and hold phases. Thirteen right-handed participants (age: 24.7 ± 1.8 years) performed trajectory tracking tasks. Participants were asked to maintain the reference magnitude of 47 N (REF) during isometric elbow flexion. The force was released to a step-down magnitude of either 75% REF or 50% REF and maintained, with and without mechanical assistance. The ramp-down durations of force release were set to 0.5, 2.5, or 5.0 s. Throughout the experiment, we measured the following: (1) the force output using load cells to compute force variability and overshoot ratio; (2) peak perturbation on the elbow movement using an accelerometer; (3) the surface electromyography (sEMG) from biceps brachii and triceps brachii muscles; and (4) EMG oscillation from the biceps brachii muscle in the bandwidth of 15–45 Hz. Our results indicated that mechanical assistance, which involved greater peak perturbation, demonstrated lower force variability than non-assistance (p < 0.01), while EMG oscillation in the biceps brachii muscle from 15 to 45 Hz was increased (p < 0.05). These findings imply that if assistive force is provided during isometric force control, the central nervous system actively tries to stabilize motor performance by controlling specific motor unit activity in the agonist muscle.     

Respiratory syncytial virus infection in anesthetized weanling rather than adult rats prolongs the apneic responses to right atrial injection of capsaicin.

Apnea is a common complication in infants infected by respiratory syncytial virus (RSV). A recent study has shown that intranasal inoculation of RSV in conscious weanling rats strengthens the apneic responses to right atrial injection of capsaicin (CAP), leading to 66% mortality. The objectives of the present study were to determine 1) whether RSV infection changes baseline minute ventilation (Ve) and arterial blood gases in anesthetized rats; 2) what the effects of RSV infection are on the respiratory responses to CAP; and 3) whether the RSV-strengthened apneic responses are age dependent. Our experiments were conducted in anesthetized and spontaneously breathing rats divided into four groups of weanling and adult rats that received either intranasal inoculation of RSV or virus-free medium. Two days after RSV infection (0.7 ml/kg), animal blood gases, baseline Ve, and Ve responses to right atrial injection of three doses of CAP (4, 16, and 64 microg/kg) were measured and compared among the four groups. Our results showed that RSV infection increased respiratory frequency (approximately 25%, P<0.05) in weanling but not adult rats, with little effect on arterial blood gases. RSV infection amplified the apneic responses to CAP in weanling but not adult rats, characterized by increases in the initial (40%) and the longest apneic duration (650%), the number of apneic episodes (139%), and the total duration of apneas (60%). These amplifications led to 50% mortality (P<0.05). We conclude that RSV infection increases respiratory frequency and strengthens the apneic responses to CAP only in anesthetized weanling but not adult rats.     

Studies on the excitability of the central program generator in the spinal cord of the terrapin Pseudemys scripta elegans

We present observations on the multicyclic scratch reflex in spinal terrapins as produced by electrical stimuli applied to the shell at the specific regions at which a mechanical stimulus produces the reflex. EMGs and hip and knee movements are recorded. The responses to the electrical stimuli are similar to the responses to mechanical stimuli. There is a three phase EMG pattern (Stein and Grossman, 1980), to which the movement pattern is related (Bakker and Crowe, 1982). A response may consist of a series of up to 25 movement cycles with a total time course of up to about 30 sec. The initial cycles of a response are relatively fast (less than 1 sec), but the cycles at the expiration of the response may have a duration of 2-3 sec. A single electrical stimulus pulse is often insufficient to trigger a series response. Instead, a weak EMG burst of a few tenths of a second duration, together with a slight movement, is often seen. However, a second pulse can set the cycle series in motion even after an interval of 40 sec between the pulses. A further booster stimulus pulse given while a reflex response is taking place can increase the speed of the movement. If the booster pulse is given just after cessation of reflex activity it can restart the activity, but this second cycle series is often shorter than the first one. The results indicate that the excitability of the central program generator is not constant. Long duration changes in the excitability are produced within the spinal cord.     

Search for critical loading condition of the spine–A meta analysis of a nonlinear viscoelastic finite element model

The relative vulnerability of spinal motion segments to different loading combinations remains unknown. The meta-analysis described here using the results of a validated L2–L3 nonlinear viscoelastic finite element model was designed to investigate the critical loading and its effect on the internal mechanics of the human lumbar spine. A Box-Behnken experimental design was used to design the magnitude of seven independent variables associated with loads, rotations and velocity of motion. Subsequently, an optimization method was used to find the primary and secondary variables that influence spine mechanical output related to facet forces, disc pressure, ligament forces, annulus matrix compressive/shear stresses and anulus fibers strain. The mechanical responses with respect to the two most-relevant variables were then regressed linearly using the response surface quadratic model. Axial force and sagittal rotation were identified as the most-relevant variables for mechanical responses. The procedure developed can be used to find the critical loading for finite element models with multi input variables. The derived meta-models can be used to predict the risk associated with various loading parameters and in setting safer load limits.     

The effect of degenerative morphological changes of the intervertebral disc on the lumbar spine biomechanics: a poroelastic finite element investigation

Intervertebral disc degeneration involves changes in the spinal anatomical structures. The mechanical relevance of the following changes was investigated: disc height, endplate sclerosis, disc water content, permeability and depressurisation. A poroelastic nonlinear finite element model of the L4–L5 human spine segments was employed. Loads represented a daily cycle (500 N compression combined with flexion–extension motion for 16 h followed by 200 N compression for 8 h). In non-degenerative conditions, the model predicted a diurnal axial displacement of 1.32 mm and a peak intradiscal pressure of 0.47 MPa. Axial displacement, facet force and range of motion in flexion–extension are decreased by decreasing disc height. By decreasing the initial water content, axial displacement, facet force and fluid loss were all reduced. Endplate sclerosis did not have a significant influence on the calculated results. Depressurisation determined an increase of the disc effective stress, possibly inducing failure. Degenerative instability was not calculated in any simulations.     

Effect of elbow joint angle on force–EMG relationships in human elbow flexor and extensor muscles

The purpose of this study was to examine the effect of joint angle on the relationship between force and electromyogram (EMG) amplitude and median frequency, in the biceps, brachioradialis and triceps muscles. Surface EMG were measured at eight elbow angles, during isometric flexion and extension at force levels from 10% to 100% of maximum voluntary contraction (MVC). Joint angle had a significant effect on MVC force, but not on MVC EMG amplitude in all of the muscles examined. The median frequency of the biceps and triceps EMG decreased with increasing muscle length, possibly due to relative changes in electrode position or a decrease in muscle fibre diameter. The relationship between EMG amplitude and force, normalised with respect to its maximum force at each angle, did not vary with joint angle in the biceps or brachioradialis muscles over all angles, or in the triceps between 45° and 120° of flexion. These results suggest that the neural excitation level to each muscle is determined by the required percentage of available force rather than the absolute force required. It is, therefore, recommended that when using surface EMG to estimate muscle excitation, force should be normalised with respect to its maximum value at each angle.     

Modulation of the visceromotor reflex by a lumbosacral ventral root avulsion injury and repair in rats

Increased abdominal muscle wall activity may be part of a visceromotor reflex (VMR) response to noxious stimulation of the bladder. However, information is sparse regarding the effects of cauda equina injuries on the VMR in experimental models. We studied the effects of a unilateral L6-S1 ventral root avulsion (VRA) injury and acute ventral root reimplantation (VRI) into the spinal cord on micturition reflexes and electromyographic activity of the abdominal wall in rats. Cystometrogram (CMG) and electromyography (EMG) of the abdominal external oblique muscle (EOM) were performed. All rats demonstrated EMG activity of the EOM associated with reflex bladder contractions. At 1 wk after VRA and VRI, the duration of the EOM EMG activity associated with reflex voiding was significantly prolonged compared with age-matched sham rats. However, at 3 wk postoperatively, the duration of the EOM responses remained increased in the VRA series but had normalized in the VRI group. The EOM EMG duration was normalized for both VRA and VRI groups at 8-12 wk postoperatively. CMG recordings show increased contraction duration at 1 and 3 wk postoperatively for the VRA series, whereas the contraction duration was only increased at 1 wk postoperatively for the VRI series. Our studies suggest that a unilateral lumbosacral VRA injury results in a prolonged VMR to bladder filling using a physiological saline solution. An acute root replantation decreased the VMR induced by VRA injury and provides earlier sensory recovery.     

Tension- and afferent input-associated responses of neuromuscular system of rats to hindlimb unloading and/or tenotomy

Responses of electromyogram (EMG) in soleus muscle and both afferent and efferent neurograms at the fifth lumbar (L(5)) segmental level of spinal cord were investigated during acute and chronic unloading induced by hindlimb suspension and/or tenotomy in adult rats. The soleus EMG and afferent neurogram decreased 88 and 37%, respectively, relative to those at quadrupedal posture on the floor after acute hindlimb suspension that causes passive shortening of soleus due to ankle plantarflexion. However, the afferent neurogram (P < 0.05) and soleus EMG (P > 0.05) recorded on the floor increased after tenotomy of synergists. Furthermore, the afferent input was inhibited when the soleus EMG disappeared after tenotomy of soleus. The afferent neurogram and EMG of the soleus showed correlated responses to a variety of treatments, suggesting that the afferent neurogram recorded at the L(5) segmental level reflects the neural input associated with the activity level of the soleus predominantly. The level of efferent neurogram decreased after acute hindlimb suspension but was not influenced significantly by tenotomy of synergists and/or soleus itself. The EMG and afferent neurograms remained low up to the 4th day but recovered to the preexperimental levels within 14 days, due to reorganization of sarcomere number and length, as well as the shortening of muscle fiber length and recovery of tension development. It is suggested that the levels of EMG and afferent neurogram associated with antigravity muscle are closely related to the tension development of the muscle.     

The differential effects of fatigue on reflex response timing and amplitude in males and females.

We examined the effects of fatigue on patellar tendon reflex responses in males and females. A spring-loaded reflex hammer elicited a standardized tendon tap with the knee positioned in an isokinetic dynamometer and flexed to 85 degrees. We recorded vastus lateralis activity (SEMG) and knee extension force production at the distal tibia (force transducer). Reflex trials were performed before and after (immediate, 2, 4, and 6 min) an isokinetic fatigue protocol to 50% MVC (90 degrees /s). For each event, pre-motor time (PMT), electromechanical delay (EMD), and total motor time (TMT) were obtained, as well as EMG amplitude (EMG(amp)), time to peak EMG (EMG(tpk)), peak force amplitude (F(amp)), time to peak force (F(tpk)), EMG:force ratio (E:F), and rate of force production (F(rate)=N/ms). TMT increased significantly in females following fatigue, while males showed no change. The increased TMT was due to an increased EMD with fatigue, while PMT was unaffected. EMG(amp) and F(amp) were somewhat diminished in females yet significantly augmented in males following fatigue, likely accounting for the differential changes in EMD noted. Results suggest males and females may respond differently to isokinetic fatigue, with males having a greater capacity to compensate for contraction force failure when responding to mechanical perturbations.     

Electromyography of trunk muscles in isometric graded axial rotation.

This study was conducted to determine the pattern, magnitude, and phasic inter-relationship of the trunk muscles in maximal isometric and graded isometric axial rotational contractions and compare them with those previously observed from the same subjects in the same experimental session in dynamic conditions. In 50 normal young healthy subjects (27 male and 23 female), after a suitable skin preparation, bipolar silver-silver chloride recessed pregelled surface electrodes were placed on external oblique, internal oblique, rectus abdominis, pectoralis major, latissimus dorsi, erector spinae at T(10) and L(3) levels bilaterally with 2 cm interelectrode distance. EMG signals from grounded subjects were suitably preamplified and amplified by a fully isolated system. These subjects were stabilized in an upright-seated posture in the Axial Rotation Tester (AROT), which was placed in isometric mode for force and rotation output from the AROT. The 14 channels of EMG, the force and the rotation were sampled at 1 kHz. The subjects initially registered their isometric maximal voluntary contraction (MVC) on both sides which was used for reference and then performed their 25%, 50% and 75% of MVC bilaterally in an isometric mode in a random order. The EMG magnitude, the slope of the rise of the EMG, and the phasic interrelationship of muscles were analyzed. The results showed that female sample generated only 65% of torque of their male counterparts. There were no significant differences between the male and the female samples in the EMG variables. Exertions to the left and to the right were not significantly different from each other for the measured variables. However, the magnitude contribution of the muscles and the slope of rise of EMG were significantly different in two directions (p<0.001). The phasic interrelationship of the external obliques, the latissimus dorsi and the erector spinae were different from other muscles (p<0.01). With the increasing grades of contraction the latissimus dorsi and the external obliques increased their magnitude significantly whereas that of the erectores spinae underwent a decrease in proportionate terms (but not in absolute magnitude) suggesting their role as stabilizers but not as rotators.     

Responses of thoracic spinal neurons to activation and desensitization of cardiac TRPV1-containing afferents in rats

The purpose of this study was to examine how upper thoracic spinal neurons responded to activation and desensitization of cardiac transient receptor potential vanilloid-1 (TRPV1)-containing afferent fibers. Extracellular potentials of single T3 spinal neurons were recorded in pentobarbital-anesthetized, paralyzed, and ventilated male rats. To activate cardiac nociceptive receptors, a catheter was placed in the pericardial sac to administer various chemicals: bradykinin (BK; 10 microg/ml, 0.2 ml), capsaicin (CAP, 10 microg/ml, 0.2 ml), or a mixture of algesic chemicals (AC; 0.2 ml) containing adenosine 10(-3) M, BK, serotonin, histamine, and PGE(2), 10(-5) M for each. Spinal neurons that responded to intrapericardial BK and/or CAP were used in this study. Results showed that 81% (35/43) of the neurons had excitatory responses to both intrapericardial BK and CAP, and the remainder responded to either BK or CAP. Intrapericardial resiniferatoxin (RTX) (0.2 microg/ml, 0.2 ml, 1 min), which desensitizes TRPV1-containing nerve endings, abolished excitatory responses to both BK (n = 8) and CAP (n = 7), and to AC (n = 5) but not to somatic stimuli. Intrapericardial capsazepine (1 mg/ml, 0.2 ml, 3 min), a specific antagonist of TRPV1, sharply attenuated excitatory responses to CAP in 5/5 neurons, but responses to BK in 5/5 neurons was maintained. Additionally, intrapericardial capsazepine had no significant effect on excitatory responses to AC in 3/3 neurons. These data indicated that intrapericardial BK-initiated spinal neuronal responses were linked to cardiac TRPV1-containing afferent fibers, but were not dependent on TRPV1. Intraspinal signaling for cardiac nociception was mediated through CAP-sensitive afferent fibers innervating the heart.     

Sensitivity of lumbar spine response to follower load and flexion moment: finite element study

The follower load (FL) combined with moments is commonly used to approximate flexed/extended posture of the lumbar spine in absence of muscles in biomechanical studies. There is a lack of consensus as to what magnitudes simulate better the physiological conditions. Considering the in-vivo measured values of the intradiscal pressure (IDP), intervertebral rotations (IVRs) and the disc loads, sensitivity of these spinal responses to different FL and flexion moment magnitudes was investigated using a 3D nonlinear finite element (FE) model of ligamentous lumbosacral spine. Optimal magnitudes of FL and moment that minimize deviation of the model predictions from in-vivo data were determined. Results revealed that the spinal parameters i.e. the IVRs, disc moment, and the increase in disc force and moment from neutral to flexed posture were more sensitive to moment magnitude than FL magnitude in case of flexion. The disc force and IDP were more sensitive to the FL magnitude than moment magnitude. The optimal ranges of FL and flexion moment magnitudes were 900–1100 N and 9.9–11.2 Nm, respectively. The FL magnitude had reverse effect on the IDP and disc force. Thus, magnitude for FL or flexion that minimizes the deviation of all the spinal parameters together from the in-vivo data can vary. To obtain reasonable compromise between the IDP and disc force, our findings recommend that FL of low magnitude must be combined with flexion moment of high intensity and vice versa.     

The phototropic response of Triticum aestivum coleoptiles under conditions of low gravity

Wheat (Triticum aestivum L. cv Broom) coleoptiles were stimulated by unilateral blue light pulses, the duration of which varied between 3 s and 30 min, under microgravity conditions on the IML-1 Spacelab mission and in ground controls. The stimuli covered first positive, indifferent and second positive response regions. When phototropic responses were observed under low-gravity conditions, slightly fewer seedlings responded compared with 1 g conditions. The latent period was similar in flight and 1g treatments (10–20 min), except for the response to 300,μmol m^?2 in the indifferent response region, where a positive response in flight plants followed a 130 min latent period, while no response was observed in 1 g plants. First positive responses at Og were slightly enhanced, both in magnitude and in duration, compared to the 1 g controls, but not to the extent predicted by clinostat studies. The response kinetics in the second positive region at 1g showed a plateau at 120 min, in contrast to the single maximum at Og. The fluence-response relationship was similar in both flight and ground controls. Only the responses to 4 and 6μmolm-2 showed significantly greater curvatures at 9g. This contrasts with previous clinostat studies, which reported substantial response enhancement at all fluence levels.     

Assessment of Neuromuscular Function Using Percutaneous Electrical Nerve Stimulation

Percutaneous electrical nerve stimulation is a non-invasive method commonly used to evaluate neuromuscular function from brain to muscle (supra-spinal, spinal and peripheral levels). The present protocol describes how this method can be used to stimulate the posterior tibial nerve that activates plantar flexor muscles. Percutaneous electrical nerve stimulation consists of inducing an electrical stimulus to a motor nerve to evoke a muscular response. Direct (M-wave) and/or indirect (H-reflex) electrophysiological responses can be recorded at rest using surface electromyography. Mechanical (twitch torque) responses can be quantified with a force/torque ergometer. M-wave and twitch torque reflect neuromuscular transmission and excitation-contraction coupling, whereas H-reflex provides an index of spinal excitability. EMG activity and mechanical (superimposed twitch) responses can also be recorded during maximal voluntary contractions to evaluate voluntary activation level. Percutaneous nerve stimulation provides an assessment of neuromuscular function in humans, and is highly beneficial especially for studies evaluating neuromuscular plasticity following acute (fatigue) or chronic (training/detraining) exercise.