Graded onset of parasternal intercostal inspiratory activity detected with surface electromyography in healthy young females and males.

Intramuscular recordings of single motor unit activity from parasternal intercostal muscles show a rostrocaudal gradient in timing and amplitude of inspiratory activity. This study determined the feasibility of surface electromyographic activity (EMG) to measure graded parasternal intercostal activity in young females and males during quiet breathing and breathing with inspiratory resistive loads. Surface EMGs were recorded from the 1st-to-5th parasternal intercostal muscles during 10 min of breathing. EMGs were processed to remove 50 Hz and electrocardiogram artifacts and integrated. Amplitude and onset time of inspiratory activity were measured from waveform averages triggered at the onset of inspiratory flow. Onset times were measured independently by two assessors, blinded to interspace and EMG scale, with excellent agreement (ICC3,k = 0.86). The onset of inspiratory activity in the 1st-to-3rd interspaces was at or within ∼400 ms of the start of inspiratory airflow, but activity in the caudal (4th and 5th) spaces was delayed by up to ∼1,000 ms (P < 0.001). There was no main effect of sex on onset time (P = 0.07), but an interaction with interspace (P < 0.001) revealed that inspiratory activity in the caudal interspaces was delayed by 15% of inspiratory time in female participants compared with 30% of inspiratory time in male participants. Inspiratory loads did not affect EMG onset time (P = 0.31). Thus, surface EMG is feasible to assess the onset time of inspiratory activity as a marker of inspiratory neural drive and pattern of activation across spaces, in both females and males.NEW & NOTEWORTHY We demonstrated that surface EMG is a valid method to measure graded inspiratory EMG in the parasternal intercostal muscles in healthy young male and female participants during quiet breathing and loaded breathing. Across the 1st-to-5th interspaces, there was more homogenous activation in women and more graded activity in men across parasternal intercostal muscles during breathing. By recording surface EMG from both male and female participants, we have revealed sex differences in inspiratory activity across intercostal muscles.

Effect of respiratory muscle training on load sensations in people with chronic tetraplegia: a secondary analysis of a randomised controlled trial.

STUDY DESIGN: Secondary analysis of a randomised controlled trial. OBJECTIVES: Our primary study showed that increasing inspiratory muscle strength with training in people with chronic (>1 year) tetraplegia corresponded with reduced sensations of breathlessness when inspiration was loaded. This study investigated whether respiratory muscle training also affected the respiratory sensations for load detection and magnitude perception. SETTING: Independent research institute in Sydney, Australia. METHODS: Thirty-two adults with chronic tetraplegia participated in a 6-week, supervised training protocol. The active group trained the inspiratory muscles through progressive threshold loading. The sham group performed the same protocol with a fixed threshold load (3.6 cmH2O). Primary measures were load detection threshold and perceived magnitudes of six suprathreshold loads reported using the modified Borg scale. RESULTS: Maximal inspiratory pressure (PImax) increased by 32% (95% CI, 18-45) in the active group with no change in the sham group (p =  0.51). The training intervention did not affect detection thresholds in the active (p =  0.24) or sham (p =  0.77) group, with similar overall decreases in Borg rating of 0.83 (95% CI, 0.49-1.17) in active and 0.72 (95% CI, 0.32-1.12) in sham group. Increased inspiratory muscle strength reduced slope magnitude between Borg rating and peak inspiratory pressure (p =  0.003), but not when pressure was divided by PImax to reflect contraction intensity (p =  0.92). CONCLUSIONS: Training reduces the sensitivity of load sensations for a given change in pressure but not for a given change in contraction intensity.

The detection and sensory perception of inspiratory resistive loads in people with chronic tetraplegia.

This study investigated sensations of breathing following tetraplegia. Fifteen people with chronic tetraplegia and fifteen healthy able-bodied controls matched for age, sex, height, and weight participated. Sensations of breathing were quantified by determining the threshold for detecting an added resistance during inspiration. In a separate task, the perceived magnitudes of six suprathreshold resistive loads were determined with a modified Borg scale. The detection threshold of 0.34 cmH2O/L/s [standard deviation (SD) 0.14] in the tetraplegia group was higher than the 0.23 cmH2O/L/s (SD 0.10) threshold for able-bodied controls (P = 0.004). Both participant groups perceived larger loads to be more effortful, with the Borg effort rating increasing linearly with the peak inspiratory pressure generated at each load. The relationship between Borg effort rating and peak inspiratory pressure was steeper in participants with tetraplegia than in able-bodied controls (P = 0.001), but there was no difference when pressure was divided by maximal inspiratory pressure (P = 0.95). Despite a higher detection threshold, the findings suggest that the perceived magnitude of a suprathreshold inspiratory load is not impaired in chronic tetraplegia and that load magnitude perception is related to the maximal, and not absolute, inspiratory muscle force.NEW & NOTEWORTHY Sensations of breathing are thought to be impaired following chronic tetraplegia. The detection threshold for an added resistive load during inspiration was higher in people with tetraplegia than in healthy able-bodied participants. However, for inspiratory loads above the detection threshold, the perceived magnitude of a resistive load as a function of the peak inspiratory pressure was greater in tetraplegia. Load magnitude perception was comparable between participant groups when peak pressure was divided by maximal inspiratory pressure.

Tongue acceleration in humans evoked with intramuscular electrical stimulation of genioglossus.

Genioglossus was stimulated intramuscularly to determine the effect of regional activation of the muscle on tongue movement in eight healthy adults. Stimulation at motor threshold was delivered with a needle electrode inserted to different depths in the anterior and posterior regions of genioglossus. The current amplitude that induced muscle contraction was ∼80% higher for anterior than posterior sites. Evoked tongue movements were determined from stimulus-triggered averages (150 pulses) of the outputs from an accelerometer fixed to the posterosuperior surface of the tongue. The median amplitude [95% confidence intervals] for the resultant acceleration was 0.0 m/s2 [0.0, 0.2] for anterior and 0.6 m/s2 [0.1, 2.8] for posterior sites. There was a positive relationship between acceleration amplitude and stimulation depth in the posterior of genioglossus (p < 0.001), but acceleration amplitude did not vary with stimulation depth in the anterior region (p = 0.83). This heterogeneity in acceleration responses between muscle regions may contribute to differences in collapsibility of the upper airway.

Movement of the ribs in supine humans for small and large changes in lung volume.

An object-tracking algorithm was used on computed tomography (CT) images of the thorax from six healthy participants and nine participants with chronic obstructive pulmonary disease (COPD) to describe the movement of the ribs between the static lung volumes of functional residual capacity (FRC) and total lung capacity (TLC). The continuous motion of the ribs during tidal breathing was also described using four-dimensional CT datasets from seven participants with thoracic esophageal malignancies. Rib motion was defined relative to a local joint coordinate system where rotations about the axes that predominantly affected the anteroposterior and transverse diameters of the rib cage were referred to as pump-handle and bucket-handle movements, respectively. Between TLC and FRC, pump-handle movements were 1.8 times larger in healthy participants than in participants with COPD, in line with their 1.6 times larger inspiratory capacities. However, when rib motion was normalized to the change in lung volume, pump-handle movements were similar for healthy participants and participants with COPD. We found no differences in bucket-handle movements between participant groups before and after normalization. Pump-handle movement was the dominant rib motion between FRC and TLC, on average four times greater than bucket-handle movement in healthy participants. For expiratory tidal volume, pump-handle movements were 20% smaller than bucket-handle movements. When normalized to tidal volume and compared with inspiratory capacity, pump-handle movements were smaller and bucket-handle movements were larger during tidal breathing. The findings suggest that the pump-handle and bucket-handle components of rib motion vary for small and large changes in lung volume.NEW & NOTEWORTHY Rib movements over inspiratory capacity are comparable for healthy participants and participants with chronic obstructive pulmonary disease when normalized to the change in lung volume. The kinematics of the ribs during tidal breathing were described from four-dimensional computed tomography images. For large changes in lung volume with inspiratory capacity, pump-handle movements of the ribs are four times greater than bucket-handle movements, whereas at tidal volume, pump-handle movements are 20% smaller than bucket-handle movements.

Supraspinal fatigue in human inspiratory muscles with repeated sustained maximal efforts.

To investigate the involvement of supraspinal fatigue in the loss of maximal inspiratory pressure (Pimax), we fatigued the inspiratory muscles. Six participants performed 5 sustained maximal isometric inspiratory efforts (15-s contractions, duty cycle ∼75%) which reduced Pimax, as measured from esophageal and mouth pressure, to around half of their initial maximums. Transcranial magnetic stimulation (TMS) delivered over the motor cortex near the beginning and end of each maximal effort evoked superimposed twitch-like increments in the ongoing Pimax, increasing from ∼1.0% of Pimax in the unfatigued contractions to ≥40% of ongoing Pimax for esophageal and mouth pressures. The rate of increase in the superimposed twitch as Pimax decreased with fatigue was not significantly different between the esophageal and mouth pressure measures. The inverse relationship between superimposed twitch pressure and Pimax indicates a progressive decline in the ability of motor cortical output to drive the inspiratory muscles maximally, leading to the development of supraspinal fatigue. TMS also evoked silent periods in the electromyographic recordings of diaphragm, scalenes, and parasternal intercostal. The duration of the silent period increased with fatigue in all three muscles, which suggests greater intracortical inhibition, with the largest change observed in the diaphragm. The peak rate of relaxation in pressure during the silent period slowed as fatigue developed, indicating peripheral contractile changes in the active inspiratory muscles. These changes in the markers of fatigue show that both central and peripheral fatigue contribute to the loss in Pimax when inspiratory muscles are fatigued with repeated sustained maximal efforts.NEW & NOTEWORTHY When the inspiratory muscles are fatigued with repeated sustained maximal efforts, supraspinal fatigue, a component of central fatigue, contributes to the loss in maximal inspiratory pressure. The presence of supraspinal fatigue was confirmed by the increase in amplitude of twitch-like increments in pressure evoked by motor cortical stimulation during maximal efforts, indicating that motor cortical output was not maximal as extra muscle force could be generated to increase inspiratory pressure.

Genioglossus motor unit activity in supine and upright postures in obstructive sleep apnea.

This study investigated whether a change in posture affected the activity of the upper-airway dilator muscle genioglossus in participants with and without obstructive sleep apnea (OSA). During wakefulness, a monopolar needle electrode was used to record single motor unit activity in genioglossus in supine and upright positions to alter the gravitational load that causes narrowing of the upper airway. Activity from 472 motor units was recorded during quiet breathing in 17 males, nine of whom had OSA. The mean number of motor units for each participant was 11.8 (SD 3.4) in the upright and 16.0 (SD 4.2) in the supine posture. For respiratory-modulated motor units, there were no significant differences in discharge frequencies between healthy controls and participants with OSA. Within each breath, genioglossus activity increased through the recruitment of phasic motor units and an increase in firing rate, with an overall increase of ~6 Hz (50%) across both postures and participant groups. However, the supine posture did not lead to compensatory increases in the peak discharge frequencies of inspiratory and expiratory motor units, despite the increase in gravitational load on the upper airway. Posture also had no significant effect on the discharge frequency of motor units that showed no respiratory modulation during quiet breathing. We postulate that, in wakefulness, any increase in genioglossus activity to compensate for the gravitational effects on the upper airway is achieved primarily through the recruitment of additional motor units in both healthy controls and participants with OSA.

Reflex response to airway occlusion in human inspiratory muscles when recruited for breathing and posture.

Briefly occluding the airway during inspiration produces a short-latency reflex inhibition in human inspiratory muscles. This occlusion reflex seems specific to respiratory muscles; however, it is not known whether the reflex inhibition has a uniform effect across a motoneuron pool when a muscle is recruited concurrently for breathing and posture. In this study, participants were seated and breathed through a mouthpiece that occluded inspiratory airflow for 250 ms at a volume threshold of 0.2 liters. The reflex response was measured in the scalene and sternocleidomastoid muscles during 1) a control condition with the head supported in space and the muscles recruited for breathing only, 2) a postural condition with the head unsupported and the neck flexors recruited for both breathing and to maintain head posture, and 3) a large-breath condition with the head supported and the volume threshold raised to between 0.8 and 1.0 liters to increase inspiratory muscle activity. When normalized to its preocclusion mean, the reflex response in the scalene muscles was not significantly different between the large-breath and control conditions, whereas concomitant recruitment of these muscles for posture control reduced the reflex response by half compared with the control condition. A reflex response occurred in sternocleidomastoid when it contracted phasically as an accessory muscle for inspiration during the large-breath condition. These results indicate that the occlusion reflex does not produce a uniform effect across the motoneuron pool and that afferent inputs for this reflex most likely act via intersegmental networks of premotoneurons rather than at a motoneuronal level. NEW & NOTEWORTHY In this study, we investigated the effect of nonrespiratory activity on the reflex response to brief sudden airway occlusions in human inspiratory muscles. We show that the reflex inhibition in the scalene muscles was not uniform across the motoneuron pool when the muscle was recruited concurrently for breathing and postural control. The reflex had a larger effect on respiratory-driven motoneurons than those recruited to maintain head posture.

Influence of respiratory mechanics and drive on genioglossus movement under ultrasound imaging.

METHODS: Twenty healthy subjects (10 males, age 28±5 years [mean ± SD]) lay supine, awake, with the head in a neutral position. Ventilation was monitored with inductance bands. Real-time B-mode ultrasound movies were analysed. We measured genioglossus motion (i) during spontaneous breathing, voluntary targeted breathing (normal tidal volume Vt), and voluntary hyperpnoea (at 1.5Vt and 2 Vt); (ii) during inspiratory flow resistive loading; (iii) with changes in end-expiratory lung volume (EELV). RESULTS: Average peak inspiratory displacement of the infero-posterior region of genioglossus was 0.89±0.56 mm; 1.02±0.88 mm; 1.27±0.70 mm respectively for voluntary Vt, and during voluntary hyperpnoea at 1.5Vt and 2Vt. A change in genioglossus motion was observed with increased Vt. During increasing inspiratory resistive loading, the genioglossus displaced less anteriorly (p = 0.005) but more inferiorly (p = 0.027). When lung volume was altered, no significant changes in genioglossus movement were observed (p = 0.115). CONCLUSION: In healthy subjects, we observed non-uniform heterogeneous inspiratory motion within the inferoposterior part of genioglossus during spontaneous quiet breathing with mean peak displacement between 0.5-2 mm, with more displacement in the posterior region than the anterior. This regional heterogeneity disappeared during voluntary targeted breathing. This may be due to different neural drive to genioglossus during voluntary breathing. During inspiratory resistive loading, the observed genioglossus motion may serve to maintain upper airway patency by balancing intraluminal negative pressure with positive pressure generated by upper airway dilatory muscles. In contrast, changes in EELV were not accompanied by major changes in genioglossus motion.

Motor unit territories in human genioglossus estimated with multichannel intramuscular electrodes.

The discharge patterns of genioglossus motor units during breathing have been well-characterized in previous studies, but their localization and territories are not known. In this study, we used two newly developed intramuscular multichannel electrodes to estimate the territories of genioglossus motor units in the anterior and posterior regions of the muscle. Seven healthy men participated. Each electrode contained fifteen bipolar channels, separated by 1 mm, and was inserted percutaneously below the chin, perpendicular to the skin, to a depth of 36 mm. Single motor unit activity was recorded with subjects awake, supine, and breathing quietly through a nasal mask for 180 s. Motor unit territories were estimated from the spike-triggered averages of the electromyographic signal from each channel. A total of 30 motor units were identified: 22 expiratory tonic, 1 expiratory phasic, 2 tonic, 3 inspiratory tonic, and 2 inspiratory phasic. Motor units appeared to be clustered based on unit type, with peak activities for expiratory units predominantly located in the anterior and superficial fibers of genioglossus and inspiratory units in the posterior region. Of these motor unit types, expiratory tonic units had the largest estimated territory, a mean 11.3 mm (SD 1.9). Estimated territories of inspiratory motor units ranged from 3 to 6 mm. In accordance with the distribution of motor unit types, the estimated territory of genioglossus motor units varied along the sagittal plane, decreasing from anterior to posterior. Our findings suggest that genioglossus motor units have large territories relative to the cross-sectional size of the muscle. NEW & NOTEWORTHY In this study, we used a new multichannel intramuscular electrode to address a fundamental property of human genioglossus motor units. We describe the territory of genioglossus motor units in the anterior and posterior regions of the muscle and show a decrease in territory size from anterior to posterior and that expiratory-related motor units have larger estimated territories than inspiratory-related motor units.

Transformation of Vestibular Signals for the Control of Standing in Humans.

During standing balance, vestibular signals encode head movement and are transformed into coordinates that are relevant to maintaining upright posture of the whole body. This transformation must account for head-on-body orientation as well as the muscle actions generating the postural response. Here, we investigate whether this transformation is dependent upon a muscle's ability to stabilize the body along the direction of a vestibular disturbance. Subjects were braced on top of a robotic balance system that simulated the mechanics of standing while being exposed to an electrical vestibular stimulus that evoked a craniocentric vestibular error of head roll. The balance system was limited to move in a single plane while the vestibular error direction was manipulated by having subjects rotate their head in yaw. Vestibular-evoked muscle responses were greatest when the vestibular error was aligned with the balance direction and decreased to zero as the two directions became orthogonal. This demonstrates that muscles respond only to the component of the error that is aligned with the balance direction and thus relevant to the balance task, not to the cumulative afferent activity, as expected for vestibulospinal reflex loops. When we reversed the relationship between balancing motor commands and associated vestibular sensory feedback, the direction of vestibular-evoked ankle compensatory responses was also reversed. This implies that the nervous system quickly reassociates new relationships between vestibular sensory signals and motor commands related to maintaining balance. These results indicate that vestibular-evoked muscle activity is a highly flexible balance response organized to compensate for vestibular disturbances. SIGNIFICANCE STATEMENT: The postural corrections critical to standing balance and navigation rely on transformation of sensory information into reference frames that are relevant for the required motor actions. Here, we demonstrate that the nervous system transforms vestibular sensory signals of head motion according to a muscle's ability to stabilize the body along the direction of a vestibular-evoked disturbance. By manipulating the direction of the imposed vestibular signal relative to a muscle's action, we show that the vestibular contribution to muscle activity is a highly flexible and organized balance response. This study provides insight into the neural integration and central processing associated with transformed vestibulomotor relationships that are essential to standing upright.

Feedforward consequences of isometric contractions: effort and ventilation.

The onset of voluntary muscle contractions causes rapid increases in ventilation and is accompanied by a sensation of effort. Both the ventilatory response and perception of effort are proportional to contraction intensity, but these behaviors have been generalized from contractions of a single muscle group. Our aim was to determine how these relationships are affected by simultaneous contractions of multiple muscle groups. We examined the ventilatory response and perceived effort of contraction during separate and simultaneous isometric contractions of the contralateral elbow flexors and of an ipsilateral elbow flexor and knee extensor. Subjects made 10-sec contractions at 25, 50, and 100% of maximum during normocapnia and hypercapnia. For simultaneous contractions, both muscle groups were activated at the same intensities. Ventilation was measured continuously and subjects rated the effort required to produce each contraction. As expected, ventilation and perceived effort increased proportionally with contraction intensity during individual contractions. However, during simultaneous contractions, neither ventilation nor effort reflected the combined muscle output. Rather, the ventilatory response was similar to when contractions were performed separately, and effort ratings showed a small but significant increase for simultaneous contractions. Hypercapnia at rest doubled baseline ventilation, but did not affect the difference in perceived effort between separate and simultaneous contractions. The ventilatory response and the sense of effort at the onset of muscle activity are not related to the total output of the motor pathways, or the working muscles, but arise from cortical regions upstream from the motor cortex.

Vestibular contribution to balance control in the medial gastrocnemius and soleus.

The soleus (Sol) and medial gastrocnemius (mGas) muscles have different patterns of activity during standing balance and may have distinct functional roles. Using surface electromyography we previously observed larger responses to galvanic vestibular stimulation (GVS) in the mGas compared with the Sol muscle. However, it is unclear whether this difference is an artifact that reflects limitations associated with surface electromyography recordings or whether a compensatory balance response to a vestibular error signal activates the mGas to a greater extent than the Sol. In the present study, we compared the effect of GVS on the discharge behavior of 9 Sol and 21 mGas motor units from freely standing subjects. In both Sol and mGas motor units, vestibular stimulation induced biphasic responses in measures of discharge timing [11 ± 5.0 (mGas) and 5.6 ± 3.8 (Sol) counts relative to the sham (mean ± SD)], and frequency [0.86 ± 0.6 Hz (mGas), 0.34 ± 0.2 Hz (Sol) change relative to the sham]. Peak-to-trough response amplitudes were significantly larger in the mGas (62% in the probability-based measure and 160% in the frequency-based measure) compared with the Sol (multiple P < 0.05). Our results provide direct evidence that vestibular signals have a larger influence on the discharge activity of motor units in the mGas compared with the Sol. More tentatively, these results indicate the mGas plays a greater role in vestibular-driven balance corrections during standing balance.

More conditioning stimuli enhance synaptic plasticity in the human spinal cord.

OBJECTIVE: To examine whether more paired corticospinal-motoneuronal stimulation (PCMS) is more effective at inducing spinal level plasticity. METHODS: To produce facilitation, corticospinal volleys evoked by motor cortical transcranial magnetic stimulation (TMS) were timed to arrive at corticospinal-motoneuronal synapses prior to antidromic potentials evoked in motoneurones by electrical brachial plexus stimulation. Paired stimuli were delivered repeatedly. 50-pair conditioning (50-PCMS) was compared to 100 pairs in single block (100-PCMSsingle) and spaced (2 blocks of 50, 15-min break; 100-PCMSspaced) patterns, and to 50 single, unpaired TMS (50-TMS). Biceps responses to cervicomedullary stimulation (cervicomedullary motor evoked potentials, CMEPs) and TMS (motor evoked potentials, MEPs) were measured before and for 1h after conditioning (recorded each 5 min). RESULTS: After 100-PCMS, average CMEP areas were increased by 46 ± 55% (mean ± SD; n=10; 100-PCMSsingle) and 71 ± 99% (100-PCMSspaced). 50-PCMS produced a non-significant 6 ± 40% increase. After 100-PCMSsingle and 100-PCMSspaced, CMEPs were larger than those after 50-TMS from 0 to 60 min (p<0.05). 100-PCMSsingle and 100-PCMSspaced produced more reliable changes than 50-PCMS. Overall, MEPs were larger at 35-60 min; however there were no differences between conditioning protocols. CONCLUSIONS: More PCMS produces more reliable enhancement of corticospinal transmission. SIGNIFICANCE: This technique has therapeutic potential to improve muscle control in patients with reduced descending drive.

TMS-evoked silent periods in scalene and parasternal intercostal muscles during voluntary breathing.

Transcranial magnetic stimulation (TMS) during voluntary muscle contraction causes a period of reduced electromyographic (EMG) activity (EMG). This is attributed to cortical inhibition and is known as the 'silent period'. Silent periods were compared in inspiratory muscles following TMS during voluntary inspiratory efforts during normocapnia, hypercapnia, and hypocapnia. TMS was delivered during isometric and dynamic contractions of scalenes and parasternal intercostals at 25% maximum inspiratory pressure. Changing end-tidal CO2 did not affect the duration of the silent period nor suppression of EMG activity during the silent period. In scalenes, silent periods were shorter for dynamic compared to isometric contractions (p<0.05); but contraction type did not alter the degree of suppression of EMG during the silent period. In parasternal intercostal, no significant differences in silent period parameters occurred for the different contraction types. The lack of effect of end-tidal CO2 suggests that descending drive from the medullary respiratory centres does not independently activate the inspiratory muscles during voluntary inspiratory efforts.

Neurogenic changes in the upper airway of obstructive sleep apnoea.

Obstructive sleep apnoea (OSA) is linked to local neural injury that evokes airway muscle remodelling. The upper airway muscles of patients with OSA are exposed to intermittent hypoxia as well as vibration induced by snoring. A range of electrophysiological and other studies have established altered motor and sensory function of the airway in OSA. The extent to which these changes impair upper airway muscle function and their relationship to the progression of OSA remains undefined. This review will collate the evidence for upper airway remodelling in OSA, particularly the electromyographic changes in upper airway muscles of patients with OSA.

Effects of tongue position and lung volume on voluntary maximal tongue protrusion force in humans.

Maximal voluntary protrusion force of the human tongue has not been examined in positions beyond the incisors or at different lung volumes. Tongue force was recorded with the tongue tip at eight positions relative to the incisors (12 and 4mm protrusion, neutral and 4, 12, 16, 24 and 32mm retraction) at functional residual capacity (FRC), total lung capacity (TLC) and residual volume (RV) in 15 healthy subjects. Maximal force occurred between 12mm and 32mm retraction (median 16mm). Maximum force at FRC was reproducible at the optimal tongue position across sessions (P=0.68). Across all positions at FRC the average force was highest at 24mm retraction (28.3±5.3N, mean±95% CI) and lowest at 12mm protrusion (49.1±4.6% maximum; P<0.05). Across all tongue positions, maximal force was on average 9.3% lower at FRC than TLC and RV (range: 4.5-12.7% maximum, P<0.05). Retracted positions produce higher-force protrusions with a small effect of lung volume.

Rectification is required to extract oscillatory envelope modulation from surface electromyographic signals.

Rectification of surface electromyographic (EMG) recordings prior to their correlation with other signals is a widely used form of preprocessing. Recently this practice has come into question, elevating the subject of EMG rectification to a topic of much debate. Proponents for rectifying suggest it accentuates the EMG spike timing information, whereas opponents indicate it is unnecessary and its nonlinear distortion of data is potentially destructive. Here we examine the necessity of rectification on the extraction of muscle responses, but for the first time using a known oscillatory input to the muscle in the form of electrical vestibular stimulation. Participants were exposed to sinusoidal vestibular stimuli while surface and intramuscular EMG were recorded from the left medial gastrocnemius. We compared the unrectified and rectified surface EMG to single motor units to determine which method best identified stimulus-EMG coherence and phase at the single-motor unit level. Surface EMG modulation at the stimulus frequency was obvious in the unrectified surface EMG. However, this modulation was not identified by the fast Fourier transform, and therefore stimulus coherence with the unrectified EMG signal failed to capture this covariance. Both the rectified surface EMG and single motor units displayed significant coherence over the entire stimulus bandwidth (1-20 Hz). Furthermore, the stimulus-phase relationship for the rectified EMG and motor units shared a moderate correlation (r = 0.56). These data indicate that rectification of surface EMG is a necessary step to extract EMG envelope modulation due to motor unit entrainment to a known stimulus.

Absence of lateral gastrocnemius activity and differential motor unit behavior in soleus and medial gastrocnemius during standing balance.

In a standing position, the vertical projection of the center of mass passes in front of the ankle, which requires active plantar-flexor torque from the triceps surae to maintain balance. We recorded motor unit (MU) activity in the medial (MG) and lateral (LG) gastrocnemius muscle and the soleus (SOL) in standing balance and voluntary isometric contractions to understand the effect of functional requirements and descending drive from different neural sources on motoneuron behavior. Single MU activity was recorded in seven subjects with wire electrodes in the triceps surae. Two 3-min standing balance trials and several ramp-and-hold contractions were performed. Lateral gastrocnemius MU activity was rarely observed in standing. The lowest thresholds for LG MUs in ramp contractions were 20-35 times higher than SOL and MG MUs (P < 0.001). Compared with MUs from the SOL, MG MUs were intermittently active (P < 0.001), had higher recruitment thresholds (P = 0.022), and greater firing rate variability (P < 0.001); this difference in firing rate variability was present in standing balance and isometric contractions. In SOL and MG MUs, both recruitment of new MUs (R(2) = 0.59-0.79, P < 0.01) and MU firing rates (R(2) = 0.05-0.40, P < 0.05) were associated with anterior-posterior and medio-lateral torque in standing. Our results suggest that the two heads of the gastrocnemius may operate in different ankle ranges with the larger MG being of primary importance when standing, likely due to its fascicle orientation. These differences in MU discharge behavior were independent of the type of descending neural drive, which points to a muscle-specific optimization of triceps surae motoneurons.

Blood pressure and the contractility of a human leg muscle.

These studies investigate the relationships between perfusion pressure, force output and pressor responses for the contracting human tibialis anterior muscle. Eight healthy adults were studied. Changing the height of tibialis anterior relative to the heart was used to control local perfusion pressure. Electrically stimulated tetanic force output was highly sensitive to physiological variations in perfusion pressure showing a proportionate change in force output of 6.5% per 10 mmHg. This perfusion-dependent change in contractility begins within seconds and is reversible with a 53 s time constant, demonstrating a steady-state equilibrium between contractility and perfusion pressure. These stimulated contractions did not produce significant cardiovascular responses, indicating that the muscle pressor response does not play a major role in cardiovascular regulation at these workloads. Voluntary contractions at forces that would require constant motor drive if perfusion pressure had remained constant generated a central pressor response when perfusion pressure was lowered. This is consistent with a larger cortical drive being required to compensate for the lost contractility with lower perfusion pressure. The relationship between contractility and perfusion for this large postural muscle was not different from that of a small hand muscle (adductor pollicis) and it responded similarly to passive peripheral and active central changes in arterial pressure, but extended over a wider operating range of pressures. If we consider that, in a goal-oriented motor task, muscle contractility determines central motor output and the central pressor response, these results indicate that muscle would fatigue twice as fast without a pressor response. From its extent, timing and reversibility we propose a testable hypothesis that this change in contractility arises through contraction- and perfusion-dependent changes in interstitial K(+) concentration.