Pairs Of Motor Units Research Articles

A robust myoelectric pattern recognition framework based on individual motor unit activities against electrode array shifts

Background and objectiveElectrode shift is always one of the critical factors to compromise the performance of myoelectric pattern recognition (MPR) based on surface electromyogram (SEMG). However, current studies focused on the global features of SEMG signals to mitigate this issue but it is just an oversimplified description of the human movements without incorporating microscopic neural drive information. The objective of this work is to develop a novel method for calibrating the electrode array shifts toward achieving robust MPR, leveraging individual motor unit (MU) activities obtained through advanced SEMG decomposition. MethodsAll of the MUs from decomposition of SEMG data recorded at the original electrode array position were first initialized to train a neural network for pattern recognition. A part of decomposed MUs could be tracked and paired with MUs obtained at the original position based on spatial distribution of their MUAP waveforms, so as to determine the shift vector (describing both the orientation and distance of the shift) implicated consistently by these multiple MU pairs. Given the known shift vector, the features of the after-shift decomposed MUs were corrected accordingly and then fed into the network to finalize the MPR task. The performance of the proposed method was evaluated with data recorded by a 16 × 8 electrode array placed over the finger extensor muscles of 8 subjects performing 10 finger movement patterns. ResultsThe proposed method achieved a shift detection accuracy of 100 % and a pattern recognition accuracy approximating to 100 %, significantly outperforming the conventional methods with lower shift detection accuracies and lower pattern recognition accuracies (p < 0.05). ConclusionsOur method demonstrated the feasibility of using decomposed MUAP waveforms’ spatial distributions to calibrate electrode shift. This study provides a new tool to enhance the robustness of myoelectric control systems via microscopic neural drive information at an individual MU level.

Persistent inward currents in tibialis anterior motoneurons can be reliably estimated within the same session

The response of spinal motoneurons to synaptic input greatly depends on the activation of persistent inward currents (PICs), the contribution of which can be estimated through the paired motor unit technique. Yet, the intra-session test–retest reliability of this measurement remains to be fully established. Twenty males performed isometric triangular dorsiflexion contractions to 20 and 50 % of maximal torque at baseline and after a 15-min resting period. High-density electromyographic signals (HD-EMG) of the tibialis anterior were recorded with a 64-electrode matrix. HD-EMG signals were decomposed, and motor units tracked across time points to estimate the contribution of PICs to motoneuron firing through quantification of motor unit recruitment-derecruitment hysteresis (ΔF). A good intraclass correlation coefficient (ICC = 0.75 [0.63, 0.83]) and a large repeated measures correlation coefficient (rrm = 0.65 [0.49, 0.77]; p < 0.001) were found between ΔF values obtained at both time points for 20 % MVC ramps. For 50 % MVC ramps, a good ICC (0.77 [0.65, 0.85]) and a very large repeated measures correlation coefficient (rrm = 0.73 [0.63, 0.80]; p < 0.001) were observed. Our data suggest that ΔF scores can be reliably investigated in tibialis anterior motor units during both low- and moderate-intensity contractions within a single experimental session.

Caffeine does not influence persistent inward current contribution to motoneuron firing.

The purpose of this study was to investigate whether caffeine consumption would change persistent inward current (PIC) contribution to motoneuron firing at increased contraction intensities and after repetitive sustained maximal contractions. Before and after the consumption of 6 mg·kg-1 of caffeine or placebo, 16 individuals performed isometric triangular-shaped ramp dorsiflexion contractions (to 20% and 40% of peak torque), followed by four maximal contractions sustained until torque production dropped to 60% of maximum, and consecutive 20% triangular-shaped contractions. Tibialis anterior motor unit firing frequencies were analyzed from high-density surface electromyograms. PIC contribution to motor unit firing was estimated by calculating the delta frequency (ΔF) using the paired motor unit technique. Motoneuron peak firing frequencies at 20% and 40% contractions and total torque-time integral during the repetitive sustained maximal contractions were also assessed. ΔF increased 0.69 peaks per second (pps) (95% CI = -0.98, -0.405; d = -0.87) from 20% to 40% contraction intensities and reduced 0.85 pps (95% CI = 0.66, 1.05; d = 0.99) after the repetitive sustained maximal contractions, regardless of caffeine consumption. Participants produced 337 Nm·s (95% CI = 49.9, 624; d = 0.63) more torque integral during the repetitive sustained maximal contractions after caffeine consumption. A strong repeated-measures correlation (r = 0.61; 95% CI = 0.49, 0.69) was observed between reductions of ΔF and peak firing frequencies after the repetitive sustained maximal contractions. PIC contribution to motoneuron firing increases from 20% to 40% contraction intensities, with no effect of caffeine (on rested tibialis anterior). Repetitive sustained maximal contractions reduced PIC contribution to motoneuron firing, regardless of caffeine or placebo consumption, evidencing that changes in intrinsic motoneuron properties contributed to performance loss. Caffeine-attenuated reduction of torque production capacity was unlikely mediated by PICs.NEW & NOTEWORTHY Persistent inward current (PIC) contribution to motoneuron firing increases with contraction intensities and is reduced after repetitive sustained maximal contractions, regardless of caffeine consumption. Reductions of PIC contribution to motoneuron firing and peak firing frequencies were largely associated, evidencing a novel mechanism underpinning decrements in maximal torque production capacity following repetitive sustained maximal contractions. Caffeine consumption attenuated neuromuscular performance reductions-allowing higher time-torque integral production during repetitive sustained maximal contractions. This was unlikely mediated by PIC.

Effects of jaw clenching and mental stress on persistent inward currents estimated by two different methods.

Spinal motoneuron firing depends greatly on persistent inward currents (PICs), which in turn are facilitated by the neuromodulators serotonin and noradrenaline. The aim of this study was to determine whether jaw clenching (JC) and mental stress (MS), which may increase neuromodulator release, facilitate PICs in human motoneurons. The paired motor unit (MU) technique was used to estimate PIC contribution to motoneuron firing. Surface electromyograms were collected using a 32-channel matrix on gastrocnemius medialis (GM) during voluntary, ramp, plantar flexor contractions. MU discharges were identified, and delta frequency (ΔF), a measure of recruitment-derecruitment hysteresis, was calculated. Additionally, another technique was used (VibStim) that evokes involuntary contractions that persist after cessation of combined Achilles tendon vibration and triceps surae neuromuscular electrical stimulation. VibStim measures of plantar flexor torque and soleus activity may reflect PIC activation. ΔF was not significantly altered by JC (p = .679, n = 18, 9 females) or MS (p = .147, n = 14, 5 females). However, all VibStim variables quantifying involuntary torque and muscle activity during and after vibration cessation were significantly increased in JC (p < .011, n = 20, 10 females) and some, but not all, increased in MS (p = .017-.05, n = 19, 10 females). JC and MS significantly increased the magnitude of involuntary contractions (VibStim) but had no effect on GM ΔF during voluntary contractions. Effects of increased neuromodulator release on PIC contribution to motoneuron firing might differ between synergists or be context dependent. Based on these data, the background level of voluntary contraction and, hence, both neuromodulation and ionotropic inputs could influence neuromodulatory PIC enhancement.

Longitudinal changes in intrinsic motoneuron excitability in amyotrophic lateral sclerosis are dependent on disease progression.

Increased amplitude of persistent inward currents (PICs) is observed in pre-symptomatic genetically modified SOD1 mice models of amyotrophic lateral sclerosis (ALS). However, at the symptomatic stage this reverses and there is a large reduction in PIC amplitude. It remains unclear whether these changes in PICs can be observed in humans, with cross-sectional studies in humans reporting contradictory findings. In people with ALS, we estimated the PIC contribution to self-sustained firing of motoneurons, using the paired-motor unit analysis to calculate the Δfrequency (ΔF), to compare the weaker and stronger muscles during the course of disease. We hypothesised that, with disease progression, ΔFs would relatively increase in the stronger muscles; and decline in the weaker muscles. Forty-three individuals with ALS were assessed in two occasions on average 17weeks apart. Tibialis anterior high-density electromyograms were recorded during dorsiflexion (40% of maximal capacity) ramped contractions, followed by clinical tests. ∆F increased from 3.14 (2.57, 3.71)peaks per second (pps) to 3.55 (2.94, 4.17)pps on the stronger muscles (0.41 (0.041, 0.781)pps, standardised difference (d)=0.287 (0.023, 0.552), P=0.030). ∆F reduced from 3.38 (95% CI 2.92, 3.84)pps to 2.88 (2.40, 3.36)pps on the weaker muscles (-0.50 (-0.80, -0.21)pps, d=0.353 (0.138, 0.567), P=0.001). The ALSFRS-R score reduced 3.9 (2.3, 5.5) points. These data indicate that the contribution of PICs to motoneuron self-sustained firing increases over time in early stages of the disease when there is little weakness before decreasing as the disease progresses and muscle weakness exacerbates, in alignment with the findings from studies using SOD1 mice. KEY POINTS: Research on mouse model of amyotrophic lateral sclerosis (ALS) suggests that the amplitude of persistent inward currents (PICs) is increased in early stages before decreasing as the disease progresses. Cross-sectional studies in humans have reported contradictory findings with both higher and lower PIC contributions to motoneuron self-sustained firing. In this longitudinal (∼17weeks) study we tracked changes in PIC contribution to motoneuron self-sustained firing, using the ΔF calculation (i.e. onset-offset hysteresis of motor unit pairs), in tibialis anterior muscles with normal strength and with clinical signs of weakness in people with ALS. ΔFs decreased over time in muscles with clinical signs of weakness. The PIC contribution to motoneuron self-sustained firing increases before the onset of muscle weakness, and subsequently decreases when muscle weakness progresses.

Two motor neuron synergies, invariant across ankle joint angles, activate the triceps surae during plantarflexion.

Recent studies have suggested that the nervous system generates movements by controlling groups of motor neurons (synergies) that do not always align with muscle anatomy. In this study, we determined whether these synergies are robust across tasks with different mechanical constraints. We identified motor neuron synergies using principal component analysis (PCA) and cross-correlations between smoothed discharge rates of motor neurons. In part 1, we used simulations to validate these methods. The results suggested that PCA can accurately identify the number of common inputs and their distribution across active motor neurons. Moreover, the results confirmed that cross-correlation can separate pairs of motor neurons that receive common inputs from those that do not receive common inputs. In part 2, 16 individuals performed plantarflexion at three ankle angles while we recorded EMG signals from the gastrocnemius lateralis (GL) and medialis (GM) and the soleus (SOL) with grids of surface electrodes. The PCA revealed two motor neuron synergies. These motor neuron synergies were relatively stable, with no significant differences in the distribution of motor neuron weights across ankle angles (P=0.62). When the cross-correlation was calculated for pairs of motor units tracked across ankle angles, we observed that only 13.0% of pairs of motor units from GL and GM exhibited significant correlations of their smoothed discharge rates across angles, confirming the low level of common inputs between these muscles. Overall, these results highlight the modularity of movement control at the motor neuron level, suggesting a sensible reduction of computational resources for movement control. KEY POINTS: The CNS might generate movements by activating groups of motor neurons (synergies) with common inputs. We show here that two main sources of common inputs drive the motor neurons innervating the triceps surae muscles during isometric ankle plantarflexions. We report that the distribution of these common inputs is globally invariant despite changing the mechanical constraints of the tasks, i.e. the ankle angle. These results suggest the functional relevance of the modular organization of the CNS to control movements.

Can local vibration alter the contribution of persistent inward currents to human motoneuron firing?

The response of spinal motoneurons to synaptic input greatly depends on the activation of persistent inward currents (PICs), which in turn are enhanced by the neuromodulators serotonin and noradrenaline. Local vibration (LV) induces excitatory Ia input onto motoneurons and may alter neuromodulatory inputs. Therefore, we investigated whether LV influences the contribution of PICs to motoneuron firing. This was assessed in voluntary contractions with concurrent, ongoing LV, as well as after a bout of prolonged LV. High-density surface electromyograms (HD-EMG) of the tibialis anterior were recorded with a 64-electrode matrix. Twenty males performed isometric, triangular, dorsiflexion contractions to 20% and 50% of maximal torque at baseline, during LV of the tibialis anterior muscle, and after 30-min of LV. HD-EMG signals were decomposed, and motor units tracked across time points to estimate PICs through a paired motor unit analysis, which quantifies motor unit recruitment-derecruitment hysteresis (ΔF). During ongoing LV, ΔF was lower for both 20% and 50% ramps. Although significant changes in ΔF were not observed after prolonged LV, a differential effect across the motoneuron pool was observed. This study demonstrates that PICs can be non-pharmacologically modulated by LV. Given that LV leads to reflexive motor unit activation, it is postulated that lower PIC contribution to motoneuron firing during ongoing LV results from decreased neuromodulatory inputs associated with lower descending corticospinal drive. A differential effect in motoneurons of different recruitment thresholds after prolonged LV is provocative, challenging the interpretation of previous observations and motivating future investigations. KEY POINTS: Neuromodulatory inputs from the brainstem influence motoneuron intrinsic excitability through activation of persistent inward currents (PICs). PICs make motoneurons more responsive to excitatory input. We demonstrate that vibration applied on the muscle modulates the contribution of PICs to motoneuron firing, as observed through analysis of the firing of single motor units. The effects of PICs on motoneuron firing were lower when vibration was concurrently applied during voluntary ramp contractions, likely due to lower levels of neuromodulation. Additionally, prolonged exposure to vibration led to differential effects of lower- vs. higher-threshold motor units on PICs, with lower-threshold motor units tending to present an increased and higher-threshold motor units a decreased contribution of PICs to motoneuron firing. These results demonstrate that muscle vibration has the potential to influence the effects of neuromodulation on motoneuron firing. The potential of using vibration as a non-pharmacological neuromodulatory intervention should be further investigated.

Intrinsic motor neuron excitability is increased after resistance training in older adults.

This study investigated the effects of high-intensity resistance training on estimates of the motor neuron persistent inward current (PIC) in older adults. Seventeen participants (68.5 ± 2.8 yr) completed a 2-wk nonexercise control period followed by 6 wk of resistance training. Surface electromyographic signals were collected with two 32-channel electrodes placed over soleus to investigate motor unit discharge rates. Paired motor unit analysis was used to calculate delta frequency (ΔF) as an estimate of PIC amplitudes during 1) triangular-shaped contractions to 20% of maximum torque capacity and 2) trapezoidal- and triangular-shaped contractions to 20% and 40% of maximum torque capacity, respectively, to understand their ability to modulate PICs as contraction intensity increases. Maximal strength and functional capacity tests were also assessed. For the 20% triangular-shaped contractions, ΔF [0.58-0.87 peaks per second (pps); P ≤ 0.015] and peak discharge rates (0.78-0.99 pps; P ≤ 0.005) increased after training, indicating increased PIC amplitude. PIC modulation also improved after training. During the control period, mean ΔF differences between 20% trapezoidal-shaped and 40% triangular-shaped contractions were 0.09-0.18 pps (P = 0.448 and 0.109, respectively), which increased to 0.44 pps (P < 0.001) after training. Also, changes in ΔF showed moderate to very large correlations (r = 0.39-0.82) with changes in peak discharge rates and broad measures of motor function. Our findings indicate that increased motor neuron excitability is a potential mechanism underpinning training-induced improvements in motor neuron discharge rate, strength, and motor function in older adults. This increased excitability is likely mediated by enhanced PIC amplitudes, which are larger at higher contraction intensities.NEW & NOTEWORTHY Resistance training elicited important alterations in soleus intrinsic motor neuronal excitability, likely mediated by enhanced persistent inward current (PIC) amplitude, in older adults. Estimates of PICs increased after the training period, accompanied by an enhanced ability to increase PIC amplitudes at higher contraction intensities. Our data also suggest that changes in PIC contribution to self-sustained discharging may contribute to increases in motor neuron discharge rates, maximal strength, and functional capacity in older adults after resistance training.

Common synaptic input between motor units from the lateral and medial posterior soleus compartments does not differ from that within each compartment.

The human soleus muscle is anatomically divided into four separate anatomical compartments. The functional role of this compartmentalization remains unclear. Here, we tested the hypothesis that the common synaptic input to motor units between the medial and lateral posterior compartments is less than within each compartment. Fourteen male participants performed three different heel-raise tasks that were considered to place a different mechanical demand on the medial and lateral soleus compartments. High-density electromyography (EMG) signals from the medial and lateral soleus compartments and the medial gastrocnemius of the right leg were decomposed into individual motor unit spike trains. The coherence between cumulative spike trains of the motor units was estimated. The coherence analysis was also repeated for motor units that were matched across all three tasks. Furthermore, we calculated the ratio of significant correlations between the spike trains of pairs of motor units. We observed that the coherence between motor units of the two soleus compartments was similar as the coherence between motor units within each compartment, regardless of the task. The correlation analysis performed on pairs of motor units confirmed these results. We conclude that the level of common synaptic input between the motor units innervating the medial and lateral posterior soleus compartment is not different than the common synaptic input between motor units innervating each of these compartments, which contrasts with findings from previous studies on finger muscles. This suggests that there is no independent neural control for the individual posterior soleus compartments.NEW & NOTEWORTHY The human soleus muscle is anatomically subdivided into four compartments. The functional role for this compartmentalization remains unknown. Here, we showed that, contrary to previous findings in finger muscles, the common synaptic input between motor units innervating the medial and lateral posterior soleus compartment was similar as that between motor units within the individual compartments. We suggest that the contradictory findings with other compartmentalized muscles may be explained by differences in muscle-tendon anatomy and function.

The effects of α-lactalbumin supplementation and handgrip contraction on soleus motoneuron excitability

We tested two strategies that hypothetically increase serotonin availability (α-lactalbumin consumption and a remote submaximal handgrip contraction) on estimates of persistent inward currents (PICs) amplitude of soleus muscle in healthy participants. With a randomised, double-blind, and cross-over design, 13 healthy participants performed triangular-shaped ramp contractions with their plantar flexors (20% of maximal torque), followed by a 30-s handgrip sustained contraction (40% of maximal force) and consecutive repeated triangular-shaped contractions. This was performed before and after the consumption of either 40g of α-lactalbumin, an isonitrogenous beverage (Zein) or an isocaloric beverage (Corn-starch). Soleus motor units discharge rates were analysed from high-density surface electromyography signals. PICs were estimated by calculating the delta frequency (ΔF) of motor unit train spikes using the paired motor unit technique. ΔF (0.19 pps; p = 0.001; d = 0.30) and peak discharge rate (0.20 pps; p < 0.001; d = 0.37) increased after the handgrip contraction, irrespective of the consumed supplement. No effects of α-lactalbumin were observed. Our results indicate that 40g of α-lactalbumin was unable to modify intrinsic motoneuron excitability. However, performing a submaximal handgrip contraction before the plantar flexion triangular contraction was capable of increasing ΔF and discharge rates on soleus motor units. These findings highlight the diffused effects of serotonergic input, its effects on motoneuron discharge behaviour, and suggest a cross-effector effect within human motoneurons.

The control and training of single motor units in isometric tasks are constrained by a common input signal.

Recent developments in neural interfaces enable the real-time and non-invasive tracking of motor neuron spiking activity. Such novel interfaces could provide a promising basis for human motor augmentation by extracting potentially high-dimensional control signals directly from the human nervous system. However, it is unclear how flexibly humans can control the activity of individual motor neurons to effectively increase the number of degrees of freedom available to coordinate multiple effectors simultaneously. Here, we provided human subjects (N = 7) with real-time feedback on the discharge patterns of pairs of motor units (MUs) innervating a single muscle (tibialis anterior) and encouraged them to independently control the MUs by tracking targets in a 2D space. Subjects learned control strategies to achieve the target-tracking task for various combinations of MUs. These strategies rarely corresponded to a volitional control of independent input signals to individual MUs during the onset of neural activity. Conversely, MU activation was consistent with a common input to the MU pair, while individual activation of the MUs in the pair was predominantly achieved by alterations in de-recruitment order that could be explained by history-dependent changes in motor neuron excitability. These results suggest that flexible MU recruitment based on independent synaptic inputs to single MUs is unlikely, although de-recruitment might reflect varying inputs or modulations in the neuron's intrinsic excitability.

Effects of reciprocal inhibition and whole-body relaxation on persistent inward currents estimated by two different methods.

Persistent inward currents (PICs) are crucial for initiation, acceleration, and maintenance of motoneuron firing. As PICs are highly sensitive to synaptic inhibition and facilitated by serotonin and noradrenaline, we hypothesised that both reciprocal inhibition (RI) induced by antagonist nerve stimulation and whole‐body relaxation (WBR) would reduce PICs in humans. To test this, we estimated PICs using the well‐established paired motor unit (MU) technique. High‐density surface electromyograms were recorded from gastrocnemius medialis during voluntary, isometric 20‐s ramp, plantarflexor contractions and decomposed into MU discharges to calculate delta frequency (ΔF). Moreover, another technique (VibStim), which evokes involuntary contractions proposed to result from PIC activation, was used. Plantarflexion torque and soleus activity were recorded during 33‐s Achilles tendon vibration and simultaneous 20‐Hz bouts of neuromuscular electrical stimulation (NMES) of triceps surae. ΔF was decreased by RI (n = 15, 5 females) and WBR (n = 15, 7 females). In VibStim, torque during vibration at the end of NMES and sustained post‐vibration torque were reduced by WBR (n = 19, 10 females), while other variables remained unchanged. All VibStim variables remained unaltered in RI (n = 20, 10 females). Analysis of multiple human MUs in this study demonstrates the ability of local, focused inhibition to attenuate the effects of PICs on motoneuron output during voluntary motor control. Moreover, it shows the potential to reduce PICs through non‐pharmacological, neuromodulatory interventions such as WBR. The absence of a consistent effect in VibStim might be explained by a floor effect resulting from low‐magnitude involuntary torque combined with the negative effects of the interventions.Key points Spinal motoneurons transmit signals to skeletal muscles to regulate their contraction. Motoneuron firing partly depends on their intrinsic properties such as the strength of persistent (long‐lasting) inward currents (PICs) that make motoneurons more responsive to excitatory input.In this study, we demonstrate that both reciprocal inhibition onto motoneurons and whole‐body relaxation reduce the contribution of PICs to human motoneuron firing. This was observed through analysis of the firing of single motor units during voluntary contractions.However, an alternative technique that involves tendon vibration and neuromuscular electrical stimulation to evoke involuntary contractions showed less effect. Thus, it remains unclear whether this alternative technique can be used to estimate PICs under all physiological conditions.These results improve our understanding of the mechanisms of PIC depression in human motoneurons. Potentially, non‐pharmacological interventions such as electrical stimulation or relaxation could attenuate unwanted PIC‐induced muscle contractions in conditions characterised by motoneuron hyperexcitability.

Estimates of persistent inward currents in tibialis anterior motor units during standing ramped contraction tasks in humans.

Persistent inward currents (PICs) play an essential role in setting motor neuron gain and shaping motor unit firing patterns. Estimates of PICs in humans can be made using the paired motor unit analysis technique, which quantifies the difference in discharge rate of a lower threshold motor unit at the recruitment onset and offset of a higher threshold motor unit (ΔF). Because PICs are highly dependent on the level of neuromodulatory drive, ΔF represents an estimate of level of neuromodulation at the level of the spinal cord. Most of the estimates of ΔF are performed under constrained, isometric, seated conditions. In the present study, we used high-density surface EMG arrays to discriminate motor unit firing patterns during isometric seated conditions with torque or EMG visual feedback and during unconstrained standing anterior-to-posterior movements with root mean square EMG visual feedback. We were able to apply the paired motor unit analysis technique to the decomposed motor units in each of the three conditions. We hypothesized that ΔF would be higher during unconstrained standing anterior-to-posterior movements compared with the seated conditions, reflecting an increase in the synaptic input to motoneurons drive while standing. In agreement with previous work, we found that there was no evidence of a difference in ΔF between the seated and standing postures, although slight differences in the initial and peak discharge rates were observed. Taken together, our results suggest that both the standing and seated postures are likely not sufficiently different, both being "upright" postures, to result in large changes in neuromodulatory drive.NEW & NOTEWORTHY In the present study, we show that the discharge rate of a lower threshold motor unit at the recruitment onset and offset of a higher threshold motor unit (ΔF) is similar between standing and seated conditions in human tibialis anterior motor units, suggesting that at least for these two upright postures neuromodulatory drive is similar. We also highlight a proposed technological development in using high-density EMG arrays for real-time muscle activity feedback to accomplish standing ramped contraction tasks and demonstrate the validity of the paired motor unit analysis technique during these conditions.

Differences in Human Motoneuron Excitability Between Functionally Diverse Muscles

Introduction: Spinal motoneurons (MN) transmit neural commands from the brain to the muscles they innervate and, as a result, produce functional movement. However, MNs are not simply passive conduits of these command and, instead, actively shape motor output through alterations in intrinsic excitability. We hypothesize that the excitability of MNs is not fixed across the body; instead, MNs are functionally tuned to the tasks they control. Here, we investigate this mapping of MN excitability across motor pools.&#x0D; Methods: High-density surface electromyography of the tibialis anterior (TA) and first dorsal interosseous (FDI) was recorded from four neurologically intact participants while they performed low-level, isometric contractions. The data were decomposed into underlying motor unit action potentials and paired motor unit analyses were subsequently performed on these spike trains to quantify MN excitability (ΔF).&#x0D; Results: 1,638 motor unit spike trains were extracted across all contractions. Mann-Whitney U test revealed that all subjects (4/4) had significantly higher maximal discharge rates in FDI, 19.13 [17.62 – 20.59] pps, when compared to the TA, 13.08 [11.51 – 15.46] pps. All subjects (4/4) had a higher ΔF in the TA (4.22 [2.89 – 5.61] pps) than the FDI (3.62 [1.23 – 5.94] pps), with 3/4 reaching statistical significance.&#x0D; Conclusions: Our findings suggest that the discharge rate and intrinsic excitability of human MNs differs across TA and FDI motor pools during similar isometric tasks. These results support the notion that motor pools are functionally tuned to their environmental demands.

An ePIC Assessment of Quadriceps Motor Unit Firing Patterns During Isometric Knee Extension: A High Density Surface Electromyography Study

Motor neurons (MNs) are the ‘final common path’ through which all motor commands must converge to generate motor output. MN firing patterns are readily accessible in humans due to the one‐to‐one spike ratio between MNs and the muscle fibers that they innervate. MNs are, therefore, the only neurons in the body in which the firing patterns can be directly associated with functional behaviors. Observed firing patterns vary systematically across the muscles of the human body, reflecting the interplay between the synaptic organization of motor commands, structure of the musculoskeletal system, and diversity of motor tasks. The goal of our lab is to develop a “human motor output map”, using newly developed high‐density surface electromyography (EMG) array electrodes that discriminate motor unit firing patterns based on convolutive blind source separation. Two main advantages of these arrays are: 1) the relatively high motor unit yield, and 2) the fact that motor units can be recorded with minimally invasive methods. We estimate the level of neuromodulatory drive (estimated persistent inward current, i.e. ePIC) to MNs using the well‐established paired motor unit analysis technique, which compares the onset and offset of a high‐threshold motor unit with respect to a low‐threshold unit and quantifies discharge rate hysteresis (ΔF). Preliminary studies in our lab suggest a strong proximal to distal gradient along the arm for motor unit firing patterns. Proximal muscles (i.e. Anterior Deltoid, Triceps Brachii, and Biceps Brachii) generate motor unit firing patterns consistent with high neuromodulatory drive (i.e. ΔF = ~4–6 spikes/s), whereas distal muscles (i.e. finger flexors or extensors, and intrinsic hand muscles) show lower neuromodulatory drive (i.e. ΔF = ~2–3 spikes/s). In the legs, the ankle dorsi‐ and plantarflexors generate motor unit firing patterns with moderate neuromodulatory drive (i.e. ΔF = ~3–5 spikes/s). However, it is unknown whether the proximal to distal gradient found in the arm also occurs in the leg. In the present study, we used high‐density surface EMG arrays to decompose motor unit firing patterns of three of the four quadriceps muscles: vastus lateralis, rectus femoris, and vastus medialis oblique. Decomposition of the EMG yielded ~7 motor units per trial, which is fewer than the decomposed motor unit yield of the lower leg, though still considerably higher than more traditional in‐dwelling EMG electrodes used for motor unit discrimination. Our data indicate that activation of the quadriceps generates motor unit firing patterns consistent with moderate neuromodulatory drive (i.e. ΔF = ~3.5 spikes/s), similar to the more distal muscles. These findings suggest that the proximal to distal gradient in motor unit firing patterns observed in the upper limbs is not analogous in the lower limbs of humans, likely reflecting the differences in functional organization between the arms and legs in bipeds.

Estimates of Motor Neuron Excitability During Postural Sways in Lower Leg Muscles

Motor system investigations are often performed under isometric conditions, but real‐world motor behaviors are seldom isometric. During tasks where isometric force or torque measurements are not feasible (i.e. during unconstrainedconditions), EMG is often used as a surrogate of the level of muscle activity. However, the linear relationship between EMG and force generation breaks down during non‐isometric contractions. It is not clear how this affects investigators’ ability to discriminate motor units using state‐of‐the‐art high‐density surface EMG arrays. Additionally, the paired motor unit analysis technique, which generates an estimate of the intrinsic excitability of a ‘test’ motor unit by determining discharge rate hysteresis (ΔF), has not been evaluated in unconstrained conditions. Consequently, we asked if 1) visual feedback provided by the subject’s own EMG activity allow for subjects to adequately perform triangular ramp contractions in constrained conditions and 2) if the EMG visual feedback is provided in unconstrained conditions, are ΔF values in subjects similar to those in the constrained condition? We hypothesized that during voluntary isometric ramp contractions, ΔF will be similar regardless of the feedback source. We further hypothesized that standing postural adjustments using EMG feedback in unconstrained conditions will produce similar ΔF values to those calculated during the isometric ramps. We expected that although the mean values across conditions will be similar, the inter‐subject coefficient of variation of ΔF will be greater in the less constrained condition. Accordingly, surface EMG using high‐density array electrodes was collected from five subjects under three conditions: 1) seated in a dynamometer in the isometric mode with torque feedback, 2) in the isometric dynamometer with filtered EMG feedback, and 3) during postural adjustments while standing freely with EMG feedback. In agreement with our hypothesis, we found that ΔF values across all three conditions were similar. In the constrained conditions with either torque or EMG feedback ΔF was 3.10 ± 0.70 imp/s and 3.40 ± 0.75 imp/s, respectively; during the unconstrained case with EMG feedback, ΔF was 3.08 ± 0.73 imp/s (all n = 15, mean ± SE). Contrary to our expectations, the coefficient of variation was not greater in the unconstrained standing condition. These results demonstrate that real‐time EMG feedback is sufficiently similar to torque feedback during both isometric and near‐isometric, unconstrained contractions, suggesting that detailed investigations of motor unit activity may be expanded to more functional movements and used relatively easily in more clinical settings.

Impact of parameter selection on estimates of motoneuron excitability using paired motor unit analysis

Objective. Noninvasive estimation of motoneuron excitability in human motoneurons is achieved through a paired motor unit analysis (ΔF) that quantifies hysteresis in the instantaneous firing rates at motor unit recruitment and de-recruitment. The ΔF technique provides insight into the magnitude of neuromodulatory synaptic input and persistent inward currents (PICs). While the ΔF technique is commonly used for estimating motoneuron excitability during voluntary contractions, computational parameters used for the technique vary across studies. A systematic investigation into the relationship between these parameters and ΔF values is necessary. Approach. We assessed the sensitivity of the ΔF technique with several criteria commonly used in selecting motor unit pairs for analysis and methods used for smoothing the instantaneous motor unit firing rates. Using high-density surface EMG and convolutive blind source separation, we obtained a large number of motor unit pairs (5409) from the triceps brachii of ten healthy individuals during triangular isometric contractions. Main results. We found an exponential plateau relationship between ΔF and the recruitment time difference between the motor unit pairs and an exponential decay relationship between ΔF and the de-recruitment time difference between the motor unit pairs, with the plateaus occurring at approximately 1 s and 1.5 s, respectively. Reduction or removal of the minimum threshold for rate–rate correlation of the two units did not affect ΔF values or variance. Removing motor unit pairs in which the firing rate of the control unit was saturated had no significant effect on ΔF. Smoothing the filter selection had no substantial effect on ΔF values and ΔF variance; however, filter selection affected the minimum recruitment and de-recruitment time differences. Significance. Our results offer recommendations for standardized parameters for the ΔF approach and facilitate the interpretation of findings from studies that implement the ΔF analysis but use different computational parameters.

Passive muscle stretching reduces estimates of persistent inward current strength in soleus motor units.

Prolonged (≥60 s) passive muscle stretching acutely reduces maximal force production at least partly through a suppression of efferent neural drive. The origin of this neural suppression has not been determined; however, some evidence suggests that reductions in the amplitude of persistent inward currents (PICs) in the motoneurons may be important. The aim of the present study was to determine whether acute passive (static) muscle stretching affects PIC strength in gastrocnemius medialis (GM) and soleus (SOL) motor units. We calculated the difference in instantaneous discharge rates at recruitment and de-recruitment (ΔF) for pairs of motor units in GM and SOL during triangular isometric plantar flexor contractions (20% maximum) both before and immediately after a 5min control period and immediately after five 1min passive plantar flexor stretches. After stretching, there was a significant reduction in SOL ΔF (-25.6%; 95% confidence interval, CI=-45.1% to -9.1%, P=0.002) but not GM ΔF These data suggest passive muscle stretching can reduce the intrinsic excitability, via PICs, of SOL motor units. These findings (1) suggest that PIC strength might be reduced after passive stretching, (2) are consistent with previously established post-stretch decreases in SOL but not GM EMG amplitude during contraction, and (3) indicate that reductions in PIC strength could underpin the stretch-induced force loss.

Single-motor-unit discharge characteristics in lumbar multifidus muscle of acute low back pain patients

Acute low back pain (ALBP) causes rapid deterioration of paraspinal muscle function. The underlying neurophysiology is poorly understood. We therefore carried out this observational study in patients with ALBP to characterize motor unit (MU) activity in deep lumbar multifidus (LM) muscle and compare with our previous findings from pain-free subjects. Nine subjects (1 woman; age 26-59 yr) with ALBP duration of 1-21 days were recruited from outpatient clinics. Fine wire electromyography (EMG) electrodes were implanted bilaterally at the painful spinal level under computer tomography guidance. EMG was recorded during spontaneous sitting and standing, and during voluntary force production. Linear mixed models were utilized to test or control for the effects of a number of predefined variables. Compared with sitting, standing increased total duration of EMG activity, median MU discharge rate, interspike interval variability, and common drive measured as common drive coefficients (CDC) derived from concurrently active MU pairs. Median discharge rate in 73 MUs was 5.5 and 6.6 pulses per second (pps) during spontaneous sitting and standing, and 7.2 pps during voluntary force production. Interspike interval variability was lower during voluntary tasks than during spontaneous force production. Common drive was less pronounced in bilateral vs. unilateral unit pairs, also in spontaneous standing. This difference was not seen in our previous pain-free subjects, suggesting altered bilateral control of the spine in ALBP. The distribution of CDC values was not a homogeneous continuum but could be seen as two partially overlapping populations of CDC distributions.NEW & NOTEWORTHY We implanted fine-wire electrodes in the deepest part of axial postural muscles in patients with acute low back pain and characterized their motor unit activity. We found less pronounced common drive to the two sides of the spine compared with pain-free subjects, suggesting a different postural control strategy in patients with acute low back pain. An unexpected finding was that common drive coefficient values appeared to consist of two partially overlapping populations of normal distributions.

Assessment of human motoneuron excitability during functional motor tasks

Detailed investigations of the human motor system are often performed under isometric conditions, although few real‐world activities incorporate isometric contractions. Because of their one‐to‐one relationship with muscle fibers and high safety factor, spinal motoneurons are the only cells in the central nervous system from which firing patterns can be readily accessed in humans. Recent advances in motor unit action potential (MUAP) decomposition techniques using high‐density surface array electrodes allow large numbers of MUAPs to be discriminated relatively non‐invasively. The activity of multiple motor units can allow for the quantification of motoneuron excitability. For example, the paired motor unit analysis technique compares the onset and offset of a high‐threshold motor unit with respect to a low‐threshold unit and quantifies discharge rate hysteresis (ΔF), which is proportional to the intrinsic excitability of the spinal motoneuron. During movements where joint torque measurements are not feasible (i.e. unconstrained by the dynamometer) electromyography (EMG) is often used as a surrogate for the level of muscle activity. However, whether activity level feedback provided by EMG measurements or torque during voluntary contractions elicits similar ΔF values is unknown. We hypothesized that during voluntary isometric ramp contractions, ΔF is similar whether feedback is provided with a torque signal or conditioned EMG. We further hypothesized that standing postural adjustments using EMG feedback in unconstrained conditions will produce similar ΔF values to those calculated during the isometric ramps. We expected that although the mean values across conditions will be similar, the inter‐subject coefficient of variation of ΔF will be greater in the less constrained condition. Accordingly, surface EMG using high‐density array electrodes was collected from five subjects under three conditions: 1) seated in a dynamometer in the isometric mode with activity level feedback provided by the torque (ISO‐TOR), 2) in the isometric dynamometer with feedback provided by EMG (ISO‐EMG), and 3) during postural adjustments while standing freely with feedback provided by EMG (POS‐EMG). In agreement with our hypothesis, we found that ΔF values across all three conditions were similar. In the ISO‐TOR and ISO‐EMG conditions, ΔF was 2.70 ± 0.70 imp/s and 3.40 ± 0.75 imp/s, respectively; during POS‐EMG it was 3.08 ± 0.73 imp/s (all n = 5, mean ± SE). Contrary to our expectations, the coefficient of variation was not greater in the unconstrained standing condition. These results demonstrate that real‐time EMG feedback is sufficiently similar to torque feedback during both isometric and near‐isometric, unconstrained contractions, suggesting that detailed investigations of motor unit activity may be expanded to more functional movements.Support or Funding InformationFunded by R01 NS098509‐01 and T32 HD007418‐27This abstract is from the Experimental Biology 2019 Meeting. There is no full text article associated with this abstract published in The FASEB Journal.