Motor unit discharge rate modulation during isometric contractions to failure is intensity- and modality-dependent.

The physiological mechanisms determining the progressive decline in the maximal muscle torque production capacity during isometric contractions to task failure are known to depend on task demands. Task-specificity of the associated adjustments in motor unit discharge rate (MUDR), however, remains unclear. This study examined MUDR adjustments during different submaximal isometric knee extension tasks to failure. Participants performed a sustained and an intermittent task at 20% and 50% of maximal voluntary torque (MVT), respectively (Experiment 1). High-density surface EMG signals were recorded from vastus lateralis (VL) and medialis (VM) and decomposed into individual MU discharge timings, with the identified MUs tracked from recruitment to task failure. MUDR was quantified and normalised to intervals of 10% of contraction time (CT). MUDR of both muscles exhibited distinct modulation patterns in each task. During the 20% MVT sustained task, MUDR decreased until ∼50% CT, after which it gradually returned to baseline. Conversely, during the 50% MVT intermittent task, MUDR remained stable until ∼40-50% CT, after which it started to continually increase until task failure. To explore the effect of contraction intensity on the observed patterns, VL and VM MUDR was quantified during sustained contractions at 30% and 50% MVT (Experiment 2). During the 30% MVT sustained task, MUDR remained stable until ∼80-90% CT in both muscles, after which it continually increased until task failure. During the 50% MVT sustained task the increase in MUDR occurred earlier, after ∼70-80% CT. Our results suggest that adjustments in MUDR during submaximal isometric contractions to failure are contraction modality- and intensity-dependent. KEY POINTS: During prolonged muscle contractions a constant motor output can be maintained by recruitment of additional motor units and adjustments in their discharge rate. Whilst contraction-induced decrements in neuromuscular function are known to depend on task demands, task-specificity of motor unit discharge behaviour adjustments is still unclear. In this study, we tracked and compared discharge activity of several concurrently active motor units in the vastii muscles during different submaximal isometric knee extension tasks to failure, including intermittent vs. sustained contraction modalities performed in the same intensity domain (Experiment 1), and two sustained contractions performed at different intensities (Experiment 2). During each task, motor units modulated their discharge rate in a distinct, biphasic manner, with the modulation pattern depending on contraction intensity and modality. These results provide insight into motoneuronal adjustments during contraction tasks posing different demands on the neuromuscular system.

The neural control of movement: a century of in vivo motor unit recordings is the legacy of Adrian and Bronk.

In two papers dated 1928 to 1929 in The Journal of Physiology, Edgar Adrian and Detlev Bronk described recordings from motor nerve and muscle fibres. The recordings from motor nerve fibres required progressive dissection of the nerve until a few fibres remained, from which isolated single fibre activity could be detected. The muscle fibre recordings were performed in humans during voluntary contractions with an intramuscular electrode - the concentric needle electrode - that they describe for the first time in the second paper. They recognised that muscle fibres would respond to each impulse sent by the innervating motor neurone and that therefore muscle fibre recordings provided information on the times of activation of the motor nerve fibres which were as accurate as a direct record from the nerve. These observations and the description of the concentric needle electrode opened the era of motor unit recordings in humans, which have continued for almost a century and have provided a comprehensive view of the neural control of movement at the motor unit level. Despite important advances in technology, many of the principles of motor unit behaviour that would be investigated in the subsequent decades were canvassed in the two papers by Adrian and Bronk. For example, they described the concomitant motor neurones' recruitment and rate coding for force modulation, synchronisation of motor unit discharges, and the dependence of discharge rate on motor unit recruitment threshold. Here, we summarise their observations and discuss the impact of their work. We highlight the advent of the concentric needle, and its subsequent influence on motor control research.

Antagonism of 5-HT2 receptors attenuates self-sustained firing of human motor units.

5-HT2 receptors on motoneurones play a critical role in facilitating persistent inward currents (PICs). Although facilitation of PICs can enhance self-sustained firing after periods of excitation, the relationship between 5-HT2 receptor activity and self-sustained firing in human motor units (MUs) has not been resolved. MU activity was assessed from the tibialis anterior of 10 healthy adults (24.9 ± 2.8 years) during two contraction protocols. Both protocols featured steady-state isometric contractions with constant descending drive to the motoneurone pool. However, one protocol also included an additional phase of superimposed descending drive. Adding and then removing descending drive in the middle of steady-state contractions altered MU firing behaviour across the motor pool, where newly recruited units in the superimposed phase were unable to switch off (P = 0.0002), and units recruited prior to additional descending drive reduced their discharge rates (P < 0.0001, difference in estimated marginal means (∆) = 2.24 pulses/s). The 5-HT2 receptor antagonist, cyproheptadine, was then administered to determine whether changes in MU firing were mediated by serotonergic mechanisms. 5-HT2 receptor antagonism caused reductions in MU discharge rate (P < 0.001, ∆ = 1.65 pulses/s), recruitment threshold (P = 0.00112, ∆ = 1.09% maximal voluntary contraction) and self-sustained firing duration (P < 0.0001, ∆ = 1.77s) after the additional descending drive was removed in the middle of the steady-state contraction. These findings indicate that serotonergic neuromodulation plays a key role in facilitating discharge and self-sustained firing of human motoneurones, where adaptive changes in MU recruitment must occur to meet the demands of the contraction. KEY POINTS: Animal and cellular preparations indicate that somato-dendritic 5-HT2 receptors regulate the intrinsic excitability of motoneurones. 5-HT2 receptor antagonism reduces estimates of persistent inward currents in motoneurones, which contribute to self-sustained firing when synaptic inputs are reduced or removed. This human study employed a contraction task that slowly increased (and then removed) the additional descending drive in the middle of a steady-state contraction where marked self-sustained firing occurred when the descending drive was removed. 5-HT2 receptor antagonism caused widespread reductions in motor unit (MU) discharge rates during contractions, which was accompanied by reduced recruitment threshold and attenuation of self-sustained firing duration after the removal of the additional descending drive to motoneurones. These findings support the role that serotonergic neuromodulation is a key facilitator of MU discharge and self-sustained firing of human motoneurones, where adaptative changes in MU recruitment must occur to meet the demands of the contraction.

Onion skin is not a universal firing pattern for spinal motoneurons: simulation study.

Muscle force is modulated by sequential recruitment and firing rates of motor units (MUs). However, discrepancies exist in the literature regarding the relationship between MU firing rates and their recruitment, presenting two contrasting firing-recruitment schemes. The first firing scheme, known as "onion skin," exhibits low-threshold MUs firing faster than high-threshold MUs, forming separate layers akin to an onion. This contradicts the other firing scheme, known as "reverse onion skin" or "afterhyperpolarization (AHP)," with low-threshold MUs firing slower than high-threshold MUs. To study this apparent dichotomy, we used a high-fidelity computational model that prioritizes physiological fidelity and heterogeneity, allowing versatility in the recruitment of different motoneuron types. Our simulations indicate that these two schemes are not mutually exclusive but rather coexist. The likelihood of observing each scheme depends on factors such as the motoneuron pool activation level, synaptic input activation rates, and MU type. The onion skin scheme does not universally govern the encoding rates of MUs but tends to emerge in unsaturated motoneurons (cells firing < their fusion frequency that generates peak force), whereas the AHP scheme prevails in saturated MUs (cells firing at their fusion frequency), which is highly probable for slow (S)-type MUs. When unsaturated, fast fatigable (FF)-type MUs always show the onion skin scheme, whereas S-type MUs do not show either one. Fast fatigue-resistant (FR)-type MUs are generally similar but show weaker onion skin behaviors than FF-type MUs. Our results offer an explanation for the longstanding dichotomy regarding MU firing patterns, shedding light on the factors influencing the firing-recruitment schemes.NEW & NOTEWORTHY The literature reports two contrasting schemes, namely the onion skin and the afterhyperpolarization (AHP) regarding the relationship between motor units (MUs) firing rates and recruitment order. Previous studies have examined these schemes phenomenologically, imposing one scheme on the firing-recruitment relationship. Here, we used a high-fidelity computational model that prioritizes biological fidelity and heterogeneity to investigate motoneuron firing schemes without bias toward either scheme. Our objective findings offer an explanation for the longstanding dichotomy on MU firing patterns.

Characterising sex-related differences in lower- and higher-threshold motor unit behaviour through high-density surface electromyography.

Emerging questions in neuromuscular physiology revolve around whether males and females share similar neural control in diverse tasks across a broad range of intensities. In order to explore these features, high-density electromyography was used to record the myoelectrical activity of biceps brachii during trapezoidal isometric contractions at 35% and 70% of maximal voluntary force (MVF) on 11 male and 13 female participants. Identified motor units were then classified as lower-threshold (recruited at ≤30%MVF) and higher-threshold (recruited at >30%MVF). The discharge rate, interspike interval variability, recruitment and derecruitment thresholds, and estimates of neural drive to motor neurons were assessed. Female lower-threshold motor units showed higher neural drive (P < 0.001), accompanied by higher discharge rate at recruitment (P = 0.006), plateau (P = 0.001) and derecruitment (P = 0.001). On the other hand, male higher-threshold motor units showed greater neural drive (P = 0.04), accompanied by higher discharge rate at recruitment (P = 0.005), plateau (P = 0.04) and derecruitment (P = 0.01). Motor unit discharge rate normalised by the recruitment threshold was significantly higher in female lower-threshold motor units (P < 0.001), while no differences were observed in higher-threshold motor units. Recruitment and derecruitment thresholds are higher in males across all intensities (P < 0.01). However, males and females have similar activation and deactivation strategies, as evidenced by similar recruitment-to-derecruitment ratios (P > 0.05). This study encompasses a broad intensity range to analyse motor unit sex-related differences, highlighting higher neural drive and discharge rates in female lower-threshold motor units, elevated recruitment and derecruitment thresholds in males, and convergences in activation and deactivation strategies. HIGHLIGHTS: What is the central question of the study? Do male and female motor units behave similarly in low- and high-intensity contractions? What is the main finding and its importance? Female motor units show higher discharge rates in low-intensity tasks and lower discharge rates in high-intensity tasks, with no differences in recruitment behaviour. A broader inter-spike interval variability was also observed in females. These findings underline that there are sex-specific differences concern the firing strategies based on task intensity.

Changes in motor unit behaviour across repeated bouts of eccentric exercise.

Unaccustomed eccentric exercise (EE) is protective against muscle damage following a subsequent bout of similar exercise. One hypothesis suggests the existence of an alteration in motor unit (MU) behaviour during the second bout, which might contribute to the adaptive response. Accordingly, the present study investigated MU changes during repeated bouts of EE. During two bouts of exercise where maximal lengthening dorsiflexion (10 repetitions × 10 sets) was performed 3 weeks apart, maximal voluntary isometric torque (MVIC) and MU behaviour (quantified using high-density electromyography; HDsEMG) were measured at baseline, during (after set 5), and post-EE. The HDsEMG signals were decomposed into individual MU discharge timings, and a subset were tracked across each time point. MVIC was reduced similarly in both bouts post-EE (Δ27 vs. 23%, P = 0.144), with a comparable amount of total work performed (∼1,300 J; P = 0.905). In total, 1,754 MUs were identified and the decline in MVIC was accompanied by a stepwise increase in discharge rate (∼13%; P < 0.001). A decrease in relative recruitment was found immediately after EE in Bout 1 versus baseline (∼16%; P < 0.01), along with reductions in derecruitment thresholds immediately after EE in Bout 2. The coefficient of variation of inter-spike intervals was lower in Bout 2 (∼15%; P < 0.001). Our data provide new information regarding a change in MU behaviour during the performance of a repeated bout of EE. Importantly, such changes in MU behaviour might contribute, at least in part, to the repeated bout phenomenon.

Motor unit number estimation via MScanFit MUNE in spinal muscular atrophy.

INTRODUCTION/AIMS: MScanFit MUNE (MScanFit) is a novel tool to derive motor unit number estimates (MUNEs) from compound muscle action potential (CMAP) scans. Few studies have explored its utility in 5q spinal muscular atrophy (SMA5q) patients, assessing only the abductor pollicis brevis (APB) muscle. We aimed to assess different distal muscles in pediatric and adult SMA5q patients, further evaluating clinical-electrophysiological correlations. METHODS: We analyzed MScanFit parameters reflecting the extent of denervation (MUNE; N50) and parameters of collateral reinnervation in APB, abductor digiti minimi (ADM), and tibialis anterior (TA) muscles. SMA patients were clinically evaluated using standardized motor function clinical scales, including the Hammersmith Functional Motor Scale - Expanded and the Revised Upper Limb Module. RESULTS: A total of 23 SMA5q (9 SMA type 2 and 14 SMA type 3) and 12 age-matched healthy controls (HCs) were enrolled. SMA patients showed lower MUNE and N50 values and higher parameters of collateral sprouting in all muscles compared to HC (p < .001). SMA type 2 patients demonstrated lower MUNE and higher collateral reinnervation values in APB and TA compared to SMA type 3 (p < .05). Walker patients showed higher values of MUNE and N50, and lower parameters of reinnervation in all muscles compared to sitters (p < .05). MScanFit parameters showed strong correlations (Rho-values ranging from .72 to .83) with clinical measurements. MUNE values were abnormal in muscles that were not clinically affected. DISCUSSION: MScanFit parameters showed promise as an outcome measure. Further studies, particularly longitudinal ones, are needed to evaluate MScanFit in measuring response to treatments.

Motoneuron-driven computational muscle modelling with motor unit resolution and subject-specific musculoskeletal anatomy.

The computational simulation of human voluntary muscle contraction is possible with EMG-driven Hill-type models of whole muscles. Despite impactful applications in numerous fields, the neuromechanical information and the physiological accuracy such models provide remain limited because of multiscale simplifications that limit comprehensive description of muscle internal dynamics during contraction. We addressed this limitation by developing a novel motoneuron-driven neuromuscular model, that describes the force-generating dynamics of a population of individual motor units, each of which was described with a Hill-type actuator and controlled by a dedicated experimentally derived motoneuronal control. In forward simulation of human voluntary muscle contraction, the model transforms a vector of motoneuron spike trains decoded from high-density EMG signals into a vector of motor unit forces that sum into the predicted whole muscle force. The motoneuronal control provides comprehensive and separate descriptions of the dynamics of motor unit recruitment and discharge and decodes the subject's intention. The neuromuscular model is subject-specific, muscle-specific, includes an advanced and physiological description of motor unit activation dynamics, and is validated against an experimental muscle force. Accurate force predictions were obtained when the vector of experimental neural controls was representative of the discharge activity of the complete motor unit pool. This was achieved with large and dense grids of EMG electrodes during medium-force contractions or with computational methods that physiologically estimate the discharge activity of the motor units that were not identified experimentally. This neuromuscular model advances the state-of-the-art of neuromuscular modelling, bringing together the fields of motor control and musculoskeletal modelling, and finding applications in neuromuscular control and human-machine interfacing research.

Neural Drive and Motor Unit Characteristics of the Serratus Anterior in Individuals With Scapular Dyskinesis.

OBJECTIVE: Scapular dyskinesis is one of the causes of shoulder disorders and involves muscle weakness in the serratus anterior. This study investigated whether motor unit (MU) recruitment and firing property, which are important for muscle exertion, have altered in serratus anterior of the individuals with scapular dyskinesis. METHODS: Asymptomatic adults with (SD) and without (control) scapular dyskinesis were analyzed. Surface electromyography (sEMG) waveforms were collected at submaximal voluntary contraction of the serratus anterior. The sEMG waveform was decomposed into MU action potential amplitude (MUAPAMP), mean firing rate (MFR), and recruitment threshold. MUs were divided into low, moderate, and high thresholds, and MU recruitment and firing properties of the groups were compared. RESULTS: High-threshold MUAPAMP was significantly smaller in the SD group than in the control group. The control group also exhibited recruitment properties that reflected the size principle, however, the SD group did not. Furthermore, the SD group had a lower MFR than the control group. CONCLUSIONS: Individuals with scapular dyskinesis exhibit altered MU recruitment properties and lower firing rates of the serratus anterior; this may be detrimental to muscle performance. Thus, it may be necessary to improve the neural drive of the serratus anterior when correcting scapular dyskinesis.

Different discrete motor-unit activation patterns in the flexor carpi radialis in boys and men.

BACKGROUND: Lower activation of higher threshold (type-II) motor units (MUs) has been suggested in children compared with adults. We examined child-adult differences in discrete MU activation of the flexor carpi radialis (FCR). METHODS: Fifteen boys (10.2 ± 1.4 years), and 17 men (25.0 ± 2.7 years) completed 2 laboratory sessions. Following a habituation session, maximal voluntary isometric wrist flexion torque (MVIC) was determined before completing trapezoidal isometric contractions at 70%MVIC. Surface electromyography was captured by Delsys Trigno Galileo sensors and decomposed into individual MU action potential trains. Recruitment threshold (RT), and MU firing rates (MUFR) were calculated. RESULTS: MVIC was significantly greater in men (10.19 ± 1.92 Nm) than in boys (4.33 ± 1.47 Nm) (p < 0.05), but not statistically different after accounting for differences in body size. Mean MUFR was not different between boys (17.41 ± 7.83 pps) and men (17.47 ± 7.64 pps). However, the MUFR-RT slope was significantly (p < 0.05) steeper (more negative) in boys, reflecting a progressively greater decrease in MUFR with increasing RT. Additionally, boys recruited more of their MUs early in the ramped contraction. CONCLUSION: Compared with men, boys tended to recruit their MUs earlier and at a lower percentage of MVIC. This difference in MU recruitment may explain the greater decrease in MUFR with increasing RT in boys compared with men. Overall, these findings suggest an age-related difference in the neural strategy used to develop moderate-high torque in wrist flexors, where boys recruit more of their MUs earlier in the force gradation process, possibly resulting in a narrower recruitment range.

Sex differences in laterality of motor unit firing behavior of the first dorsal interosseous muscle in strength-matched healthy young males and females.

PURPOSE: The purpose of this study was to compare laterality in motor unit firing behavior between females and males. METHODS: Twenty-seven subjects (14 females) were recruited for this study. The participants performed ramp up and hold isometric index finger abduction at 10, 30, and 60% of their maximum voluntary contraction (MVC). High-density surface electromyography (HD-sEMG) signals were recorded in the first dorsal interosseous (FDI) muscle and decomposed into individual motor unit (MU) firing behavior using a convolution blind source separation method. RESULTS: In total, 769 MUs were detected (females, n = 318 and males, n = 451). Females had a significantly higher discharge rate than males at each relative torque level (10%: male dominant hand, 13.4 ± 2.7 pps vs. female dominant hand, 16.3 ± 3.4 pps; 30%: male dominant hand, 16.1 ± 3.9 pps vs. female dominant hand, 20.0 ± 5.0 pps; and 60%: male dominant hand, 19.3 ± 3.8 vs. female dominant hand, 25.3 ± 4.8 pps; p < 0.0001). The recruitment threshold was also significantly higher in females than in males at 30 and 60% MVC. Furthermore, males exhibited asymmetrical discharge rates at 30 and 60% MVC and recruitment thresholds at 30 and 60% MVC, whereas no asymmetry was observed in females. CONCLUSION: In the FDI muscle, compared to males, females exhibited different neuromuscular strategies with higher discharge rates and recruitment thresholds and no asymmetrical MU firing behavior. Notably, the findings that sex differences in neuromuscular activity also occur in healthy individuals provide important information for understanding the pathogenesis of various diseases.

Motor unit firing rates increase in prepubescent youth following linear periodization resistance exercise training.

PURPOSE: The purpose was to examine the effects of 8-weeks (3 days/week) of linear periodization resistance exercise training (RET) on neuromuscular function in prepubescent youth. METHODS: Twenty-five healthy prepubescent youth (11 males, 14 females, age = 9.1 ± 0.8 years) completed the RET (n = 17) or served as controls (CON, n = 8). Isometric maximal voluntary contractions (MVCs) and trapezoidal submaximal contractions at 35 and 60% MVC of the right leg extensors were performed with surface electromyography (EMG) recorded from the leg extensors [vastus lateralis (VL), rectus femoris, and vastus medialis] and flexors (biceps femoris and semitendinosus). EMG amplitude of the leg extensors and flexors were calculated during the MVCs. Motor unit (MU) action potential trains were decomposed from the surface EMG of the VL for the 35 and 60% MVCs. MU firing rates and action potential amplitudes were regressed against recruitment threshold with the y-intercepts and slopes calculated for each contraction. Total leg extensor muscle cross-sectional area (CSA) was collected using ultrasound images. ANOVA models were used to examine potential differences. RESULTS: Isometric strength increased post-RET (P = 0.006) with no changes in leg extensor and flexor EMG amplitude. Furthermore, there were no changes in total CSA or the MU action potential amplitude vs. recruitment threshold relationships. However, there were increases in the firing rates of the higher-threshold MUs post-RET as indicated with greater y-intercepts (P = 0.003) from the 60% MVC and less negative slope (P = 0.004) of the firing rates vs. recruitment threshold relationships at 35% MVC. CONCLUSIONS: MU adaptations contribute to strength increases following RET in prepubescent youth.

Visual exposure enhances stimulus encoding and persistence in primary cortex.

The brain adapts to the sensory environment. For example, simple sensory exposure can modify the response properties of early sensory neurons. How these changes affect the overall encoding and maintenance of stimulus information across neuronal populations remains unclear. We perform parallel recordings in the primary visual cortex of anesthetized cats and find that brief, repetitive exposure to structured visual stimuli enhances stimulus encoding by decreasing the selectivity and increasing the range of the neuronal responses that persist after stimulus presentation. Low-dimensional projection methods and simple classifiers demonstrate that visual exposure increases the segregation of persistent neuronal population responses into stimulus-specific clusters. These observed refinements preserve the representational details required for stimulus reconstruction and are detectable in postexposure spontaneous activity. Assuming response facilitation and recurrent network interactions as the core mechanisms underlying stimulus persistence, we show that the exposure-driven segregation of stimulus responses can arise through strictly local plasticity mechanisms, also in the absence of firing rate changes. Our findings provide evidence for the existence of an automatic, unguided optimization process that enhances the encoding power of neuronal populations in early visual cortex, thus potentially benefiting simple readouts at higher stages of visual processing.

Analysis of the suprahyoid muscles during tongue elevation: High-density surface electromyography as a novel tool for swallowing-related muscle assessment.

BACKGROUND: High-density surface electromyography (HD-sEMG) has enabled non-invasive analysis of motor unit (MU) activity and recruitment, but its application to swallowing-related muscles is limited. OBJECTIVE: We aimed to investigate the utility of HD-sEMG for quantitatively evaluating the MU recruitment characteristics of the suprahyoid muscles during tongue elevation. METHODS: We measured the sEMG activity of the suprahyoid muscles of healthy participants during tongue elevation using HD-sEMG. Maximum voluntary contraction (MVC) was measured, followed by data collection during sustained and ramp-up tasks to capture suprahyoid muscle activity. Changes in the temporal/spatial MU recruitment patterns within individual suprahyoid muscles were analysed. RESULTS: This study enrolled 16 healthy young adults (mean age: 27.8 ± 5.3 years; eight males and eight females). Increasing muscle force corresponded to a decrease in modified entropy and correlation coefficient and an increase in the coefficient of variation. No significant differences were observed between male and female participants. CONCLUSION: The results of this study, consistent with those observed in other muscles, such as the vastus lateralis muscle, suggest that HD-sEMG is a valuable and reliable tool for quantitatively evaluating MU recruitment in the suprahyoid muscles. This measurement technique holds promise for novel assessments of swallowing function.

Blockade of 5-HT2 receptors suppresses motor unit firing and estimates of persistent inward currents during voluntary muscle contraction in humans.

Serotonergic neuromodulation contributes to enhanced voluntary muscle activation. However, it is not known how the likely motoneurone receptor candidate (5-HT2 ) influences the firing rate and activation threshold of motor units (MUs) in humans. The purpose of this study was to determine whether 5-HT2 receptor activity contributes to human MU behaviour during voluntary ramped contractions of differing intensity. High-density surface EMG (HDsEMG) of the tibialis anterior was assessed during ramped isometric dorsiflexions at 10, 30, 50 and 70% of maximal voluntary contraction (MVC). MU characteristics were successfully extracted from HDsEMG of 11 young adults (four female) pre- and post-ingestion of 8 mg cyproheptadine or a placebo. Antagonism of 5-HT2 receptors caused a reduction in MU discharge rate during steady-state muscle activation that was independent of the level of contraction intensity [P < 0.001; estimated mean difference (∆) = 1.06 pulses/s], in addition to an increase in MU derecruitment threshold (P < 0.013, ∆ = 1.23% MVC), without a change in force during MVC (P = 0.652). A reduction in estimates of persistent inward current amplitude was observed at 10% MVC (P < 0.001, ∆ = 0.99 Hz) and 30% MVC (P = 0.003, ∆ = 0.75 Hz) that aligned with 5-HT changes in MU firing behaviour attributable to 5-HT2 antagonism. Overall, these findings indicate that 5-HT2 receptor activity has a role in regulating the discharge rate in populations of spinal motoneurones when performing voluntary contractions. This study provides evidence of a direct link between MU discharge properties, persistent inward current activity and 5-HT2 receptor activity in humans. KEY POINTS: Activation of 5-HT receptors on the soma and dendrites of motoneurones regulates their excitability. Previous work using chlorpromazine and cyproheptadine has demonstrated that the 5-HT2 receptor regulates motoneurone activity in humans with chronic spinal cord injury and non-injured control subjects. It is not known how the 5-HT2 receptor directly influences motor unit (MU) discharge and MU recruitment in larger populations of human motoneurones during voluntary contractions of differing intensity. Despite the absence of change in force during maximal voluntary dorsiflexions, 5-HT2 receptor antagonism caused a reduction in MU discharge rate during submaximal steady-state muscle contraction, in addition to an increase in MU derecruitment threshold, irrespective of the submaximal contraction intensity. Reductions in estimates of persistent inward currents after 5-HT2 receptor antagonism support the viewpoint that the 5-HT2 receptor plays a crucial role in regulating motor activity, whereby a persistent inward current-based mechanism is involved in regulating the excitability of human motoneurones.

Adaptations in motor unit properties underlying changes in recruitment, rate coding, and maximum force.

Changes in the discharge characteristics of motor units as well as in the maximum force-producing capacity of the muscle are observed following training, aging, and fatiguability. The ability to measure the adaptations in the neuromuscular properties underlying these changes experimentally, however, is limited. In this study we used a computational model to systematically investigate the effects of various neural and muscular adaptations on motor unit recruitment thresholds, average motor unit discharge rates in submaximal contractions, and maximum force. The primary focus was to identify candidate adaptations that can explain experimentally observed changes in motor unit discharge characteristics after 4 wk of strength training (Del Vecchio A, Casolo A, Negro F, Scorcelletti M, Bazzucchi I, Enoka R, Felici F, Farina D. J Physiol 597: 1873-1887, 2019). The simulation results indicated that multiple combinations of adaptations, likely involving an increase in maximum discharge rate across motor units, may occur after such training. On a more general level, we found that the magnitude of the adaptations scales linearly with the change in recruitment thresholds, discharge rates, and maximum force. In addition, the combination of multiple adaptations can be predicted as the linear sum of their individual effects. Together, this implies that the outcomes of the simulations can be generalized to predict the effect of any combination of neural and muscular adaptations. In this way, the study provides a tool for estimating potential underlying adaptations in neural and muscular properties to explain any change in commonly used measures of rate coding, recruitment, and maximum force.NEW & NOTEWORTHY Our ability to measure adaptations in neuromuscular properties in vivo is limited. Using a computational model, we quantify the effect of multiple neuromuscular adaptations on common measures of motor unit recruitment, rate coding, and force-producing capacity. Scaling and combining adaptations had a near-linear effect on these measures, indicating that the results can explain and predict neuromuscular adaptations in a wide range of conditions, including, but not limited to, strength training.

Context-specific early recruitment of small motor units in the shoulder muscle reflects a reach movement plan.

Understanding how motor plans are transformed into appropriate patterns of muscle activity is a central question in motor control. Although muscle activity during the delay period has not been reported using conventional electromyographic (EMG) approaches, we isolated motor unit activity using a high-density surface EMG signal from the anterior deltoid muscle to test whether heterogeneity in motor units could reveal early preparatory activity. Consistent with our previous work (Rungta SP, Basu D, Sendhilnathan N, Murthy A. J Neurophysiol 126: 451-463, 2021), we observed early selective recruitment of small amplitude size motor units during the delay period for hand movements similar to the observed early recruitment of small-amplitude motor units in neck muscles of nonhuman primates performing delayed saccade tasks. This early activity was spatially specific and increased with time and resembled an accumulation to threshold model that correlated with movement onset time. Such early recruitment of ramping motor units was observed at the single trial level as well. In contrast, no such recruitment of large amplitude size motor units, called nonrampers, was observed during the delay period. Instead, nonrampers became spatially specific and predicted movement onset time after the delay period. Interestingly, spatially specific delay period activity was only observed for hand movements but was absent for isometric force-driven cursor movements. Nonetheless, muscle activity was correlated with the time it took to initiate movements in both task conditions for nonrampers. Overall, our results reveal a novel heterogeneity in the EMG activity that allows the expression of early motor preparation via small amplitude size motor units that are differentially activated during movement initiation.NEW & NOTEWORTHY We studied the spatial and temporal aspects of response preparation in the anterior deltoid muscle using high-density surface EMG. Our results show that early spatially specific ramping activity that predicted reaction times could be accessed from muscle activity but was absent during isometric force-driven cursor movements. Such ramping activity could be quantified using an accumulator framework across trials, as well as within single trials, but was not observed in isometric reach tasks involving cursor movements.

Regional recruitment and differential behavior of motor units during postural control in older adults.

Aging is associated with neuromuscular system changes that may have implications for the recruitment and firing behaviors of motor units (MUs). In previous studies, we observed that young adults recruit subpopulations of triceps surae MUs during tasks that involved leaning in five directions: common units that were active during different leaning directions and unique units that were active in only one leaning direction. Furthermore, the MU subpopulation firing behaviors [average firing rate (AFR), coefficient of variation (CoVISI), and intermittent firing] modulated with leaning direction. The purpose of this study was to examine whether older adults exhibited this regional recruitment of MUs and firing behaviors. Seventeen older adults (aged 74.8 ± 5.3 yr) stood on a force platform and maintained their center of pressure leaning in five directions. High-density surface electromyography recordings from the triceps surae were decomposed into single MU action potentials. A MU tracking analysis identified groups of MUs as being common or unique across the leaning directions. Although leaning in different directions did not affect the AFR and CoVISI of common units (P > 0.05), the unique units responded to the leaning directions by increasing AFR and CoVISI, albeit modestly (F = 18.51, P < 0.001). The unique units increased their intermittency with forward leaning (F = 9.22, P = 0.003). The mediolateral barycenter positions of MU activity in both subpopulations were found in similar locations for all leaning directions (P > 0.05). These neuromuscular changes may contribute to the reduced balance performance seen in older adults.NEW & NOTEWORTHY In this study, we observed differences in motor unit recruitment and firing behaviors of distinct subpopulations of motor units in the older adult triceps surae muscle from those observed in the young adult. Our results suggest that the older adult central nervous system may partially lose the ability to regionally recruit and differentially control motor units. This finding may be an underlying cause of balance difficulties in older adults during directionally challenging leaning tasks.

Assessment of motor unit firing by high-density surface electromyography detects motor neuronal hyperexcitable state in amyotrophic lateral sclerosis.

INTRODUCTION/AIMS: In amyotrophic lateral sclerosis (ALS), the impact of motor neuron dysfunction on the motor unit (MU) firing pattern remains to be elucidated. The aim of this study was to clarify the characteristics of the MU firing rate and its association with clinical factors in ALS patients using high-density surface electromyography (HDSEMG) and MU decomposition analysis. METHODS: Nineteen ALS patients and 20 controls prospectively underwent HDSEMG recording of the vastus lateralis muscle during ramp-up (30% of maximum voluntary contraction) and sustained (10% of maximal voluntary contraction for 60 seconds) contractions on performing isometric knee extension. After decomposition analysis, instantaneous firing rates (IFRs) of individually identified MUs were calculated. Comparison of IFRs and clinical variables between ALS patients and controls and analysis of the correlation between individual mean IFR and clinical variables in ALS patients were performed. RESULTS: The number of identified MUs was lower in ALS patients than in controls (P = .017). Mean IFRs of MUs (i.e., mean MU firing rates) were higher in ALS patients than in controls at some force levels on ramp-up contraction (P < .05) and at 50 to 60 seconds during sustained contraction (9.1 [ALS] vs 8.3 [controls] pulses per second; P = .036). There was no correlation between the clinical parameters and mean IFR of each patient. DISCUSSION: ALS patients had a higher MU firing rate during muscle contraction at a low force level. Noninvasive assessment of the MU firing rate by HDSEMG can detect a motor neuronal hyperexcitable state in ALS patients.

Neuromechanics of the Rate of Force Development.

The rate at which an individual can develop force during rapid voluntary contractions can be influenced by both the neural drive to a muscle and its intrinsic musculotendinous properties. We hypothesize that the maximal rate of force development across human individuals is mainly attributable to the rate of motor unit recruitment.