The initial increases in force production with resistance training are thought to be primarily underpinned by neural adaptations. This notion is firmly supported by evidence displaying motor unit ...adaptations following resistance training; however, the precise locus of neural adaptation remains elusive. The purpose of this review is to clarify and critically discuss the literature concerning the site(s) of putative neural adaptations to short-term resistance training. The proliferation of studies employing non-invasive stimulation techniques to investigate evoked responses have yielded variable results, but generally support the notion that resistance training alters intracortical inhibition. Nevertheless, methodological inconsistencies and the limitations of techniques, e.g. limited relation to behavioural outcomes and the inability to measure volitional muscle activity, preclude firm conclusions. Much of the literature has focused on the corticospinal tract; however, preliminary research in non-human primates suggests reticulospinal tract is a potential substrate for neural adaptations to resistance training, though human data is lacking due to methodological constraints. Recent advances in technology have provided substantial evidence of adaptations within a large motor unit population following resistance training. However, their activity represents the transformation of afferent and efferent inputs, making it challenging to establish the source of adaptation. Whilst much has been learned about the nature of neural adaptations to resistance training, the puzzle remains to be solved. Additional analyses of motoneuron firing during different training regimes or coupling with other methodologies (e.g., electroencephalography) may facilitate the estimation of the site(s) of neural adaptations to resistance training in the future.
The ability to produce rapid forces requires quick motor unit recruitment, high motor unit discharge rates, and fast motor unit force twitches. The relative importance of these parameters for maximum ...rate of force development (RFD), however, is poorly understood. In this study, we systematically investigated these relationships using a computational model of motor unit pool activity and force. Across simulations, neural and muscular properties were systematically varied in experimentally observed ranges. Motor units were recruited over an interval starting from contraction onset (range: 22-233 ms). Upon recruitment, discharge rates declined from an initial rate (range: 89-212 pulses per second), with varying likelihood of doublet (interspike interval of 3 ms; range: 0-50%). Finally, muscular adaptations were modeled by changing average twitch contraction time (range: 42-78 ms). Spectral analysis showed that the effective neural drive to the simulated muscle had smaller bandwidths than the average motor unit twitch, indicating that the bandwidth of the motor output, and thus the capacity for explosive force, was limited mainly by neural properties. The simulated RFD increased by 1,050 ± 281% maximal voluntary contraction force per second from the longest to the shortest recruitment interval. This effect was more than fourfold higher than the effect of increasing the initial discharge rate, more than fivefold higher than the effect of increasing the chance of doublets, and more than sixfold higher than the effect of decreasing twitch contraction times. The simulated results suggest that the physiological variation of the rate by which motor units are recruited during ballistic contractions is the main determinant for the variability in RFD across individuals.
An important limitation of human performance is the ability to generate explosive movements by means of rapid development of muscle force. The physiological determinants of this ability, however, are poorly understood. In this study, we show using extensive simulations that the rate by which motor units are recruited is the main limiting factor for maximum rate of force development.
Abstract
This work focuses on the hydrothermal synthesis of Na-P
1
zeolite by using a kaolinite rock coming from Romana (Sassari, Italy). The kaolin is calcined at a temperature of 650 °C and then ...mixed with calculated quantities of NaOH. The synthesis runs are carried out at ambient pressure and at variable temperatures of 65 and 100 °C. For the first time compared to the past, the Na-P1 zeolite is synthesized without the use of additives and through a protocol that reduces both temperatures and synthesis times. The synthesis products are analysed by X-ray diffraction, high temperature X-ray diffraction, infrared spectroscopy, scanning electron microscopy and inductively coupled plasma optical emission spectrometry. The cell parameters are calculated using the Rietveld method. Density and specific surface area are also calculated. The absence of amorphous phases and impurities in synthetic powders is verified through quantitative phase analysis using the combined Rietveld and reference intensity ratio methods. The results make the experimental protocol very promising for an industrial transfer.
Correlation between motor unit discharge times, often referred to as motor unit synchronization, is determined by common synaptic input to motor neurons. Although it has been largely speculated that ...synchronization should influence the rate of force development, the association between the degree of motor unit synchronization and rapid force generation has not been determined. In this study, we examined this association with both simulations and experimental motor unit recordings. The analysis of experimental motor unit discharges from the tibialis anterior muscle of 20 healthy individuals during rapid isometric contractions revealed that the average motor unit discharge rate was associated with the rate of force development. Moreover, the extent of motor unit synchronization was entirely determined by the average motor unit discharge rate (
> 0.7,
< 0.0001). The simulation model demonstrated that the relative proportion of common synaptic input received by motor neurons, which determines motor unit synchronization, does not influence the rate of force development (
= 0.03,
> 0.05). Nonetheless, the estimates of correlation between motor unit spike trains were significantly correlated with the rate of force generation (
> 0.8,
< 0.0001). These results indicate that the average motor unit discharge rate, but not the degree of motor unit synchronization, contributes to most of the variance of human contractile speed among individuals. In addition, estimates of correlation between motor unit discharge times depend strongly on the number of identified motor units and therefore are not indicative of the strength of common input.
It is commonly assumed that motor unit synchronization has an impact on the rate of force development of a muscle. Here we present computer simulations and experimental data of human tibialis anterior motor units during rapid contractions that show that motor unit synchronization is not a determinant of the rate of force production. This conclusion clarifies the neural determinants of rapid force generation.
Resistance training is frequently performed with the goal of stimulating muscle hypertrophy. Due to the key roles motor unit recruitment and mechanical tension play to induce muscle growth, when ...programming, the manipulation of the training variables is oriented to provoke the correct stimulus. Although it is known that the nervous system is responsible for the control of motor units and active muscle force, muscle hypertrophy researchers and trainers tend to only focus on the adaptations of the musculotendinous unit and not in the nervous system behaviour. To better guide resistance exercise prescription for muscle hypertrophy and aiming to delve into the mechanisms that maximize this goal, this review provides evidence-based considerations for possible effects of neural behaviour on muscle growth when programming resistance training, and future neurophysiological measurement that should be tested when training to increase muscle mass. Combined information from the neural and muscular structures will allow to understand the exact adaptations of the muscle in response to a given input (neural drive to the muscle). Changes at different levels of the nervous system will affect the control of motor units and mechanical forces during resistance training, thus impacting the potential hypertrophic adaptations. Additionally, this article addresses how neural adaptations and fatigue accumulation that occur when resistance training may influence the hypertrophic response and propose neurophysiological assessments that may improve our understanding of resistance training variables that impact on muscular adaptations.
The surface interference EMG signal provides some information on the neural drive to muscles. However, the association between neural drive to muscle and muscle activation has long been debated with ...controversial indications due to the unavailability of motor unit population data. In this study, we clarify the potential and limitations of interference EMG analysis to infer motor unit recruitment strategies with an experimental investigation of several concurrently active motor units and of the associated features of the surface EMG. For this purpose, we recorded high-density surface EMG signals during linearly increasing force contractions of the tibialis anterior muscle, up to 70% of maximal force. The recruitment threshold (RT), conduction velocity (MUCV), median frequency (MDF
), and amplitude (RMS
) of action potentials of 587 motor units from 13 individuals were assessed and associated with features of the interference EMG. MUCV was positively associated with RT (
= 0.64 ± 0.14), whereas MDF
and RMS
showed a weaker relation with RT (
= 0.11 ± 0.11 and 0.39 ± 0.24, respectively). Moreover, the changes in average conduction velocity estimated from the interference EMG predicted well the changes in MUCV (
= 0.71), with a strong association to ankle dorsiflexion force (
= 0.81 ± 0.12). Conversely, both the average EMG MDF and RMS were poorly associated with motor unit recruitment. These results clarify the limitations of EMG spectral and amplitude analysis in inferring the neural strategies of muscle control and indicate that, conversely, the average conduction velocity could provide relevant information on these strategies.
The surface EMG provides information on the neural drive to muscles. However, the associations between EMG features and neural drive have been long debated due to unavailability of motor unit population data. Here, by using novel highly accurate decomposition of the EMG, we related motor unit population behavior to a wide range of voluntary forces. The results fully clarify the potential and limitation of the surface EMG to provide estimates of the neural drive to muscles.
Musculoskeletal (MSK) pain disorders represent a group of highly prevalent and often disabling conditions. Investigating the structure of motor variability in response to pain may reveal novel motor ...impairment mechanisms that may lead to enhanced management of motor dysfunction associated with MSK pain disorders. This review aims to systematically synthesize the evidence on the influence of MSK pain disorders on muscle synergies.
Nine electronic databases were searched using Medical Subject Headings and keywords describing pain, electromyography and synergies. Relevant characteristics of included studies were extracted and assessed for generalizability and risk of bias. Due to the significant heterogeneity, a qualitative synthesis of the results was performed.
The search resulted in a total of 1312 hits, of which seven articles were deemed eligible. There was unclear consistency that pain reduced the number of muscle synergies. There were low consistencies of evidence that the synergy vector (W weights) and activation coefficient (C weights) differed in painful compared to asymptomatic conditions. There was a high consistency that muscle synergies were dissimilar between painful and asymptomatic conditions.
MSK pain alters the structure of variability in muscle control, although its specific nature remains unclear. Greater consistency in muscle synergy analysis may be achieved with appropriate selection of muscles assessed and ensuring consistent achievement of motor task outcomes. Synergy analysis is a promising method to reveal novel understandings of altered motor control, which may facilitate the assessment and treatment of MSK pain disorders.
Because of the biophysical relation between muscle fibre diameter and the propagation velocity of action potentials along the muscle fibres, motor unit conduction velocity could be a non‐invasive ...index of muscle fibre size in humans. However, the relation between motor unit conduction velocity and fibre size has been only assessed indirectly in animal models and in human patients with invasive intramuscular EMG recordings, or it has been mathematically derived from computer simulations. By combining advanced non‐invasive techniques to record motor unit activity in vivo, i.e. high‐density surface EMG, with the gold standard technique for muscle tissue sampling, i.e. muscle biopsy, here we investigated the relation between the conduction velocity of populations of motor units identified from the biceps brachii muscle, and muscle fibre diameter. We demonstrate the possibility of predicting muscle fibre diameter (R2 = 0.66) and cross‐sectional area (R2 = 0.65) from conduction velocity estimates with low systematic bias (∼2% and ∼4% respectively) and a relatively low margin of individual error (∼8% and ∼16%, respectively). The proposed neuromuscular interface opens new perspectives in the use of high‐density EMG as a non‐invasive tool to estimate muscle fibre size without the need of surgical biopsy sampling. The non‐invasive nature of high‐density surface EMG for the assessment of muscle fibre size may be useful in studies monitoring child development, ageing, space and exercise physiology, although the applicability and validity of the proposed methodology need to be more directly assessed in these specific populations by future studies.
Key points
Because of the biophysical relation between muscle fibre size and the propagation velocity of action potentials along the sarcolemma, motor unit conduction velocity could represent a potential non‐invasive candidate for estimating muscle fibre size in vivo.
This relation has been previously assessed in animal models and humans with invasive techniques, or it has been mathematically derived from simulations.
By combining high‐density surface EMG with muscle biopsy, here we explored the relation between the conduction velocity of populations of motor units and muscle fibre size in healthy individuals.
Our results confirmed that motor unit conduction velocity can be considered as a novel biomarker of fibre size, which can be adopted to predict muscle fibre diameter and cross‐sectional area with low systematic bias and margin of individual error.
The proposed neuromuscular interface opens new perspectives in the use of high‐density EMG as a non‐invasive tool to estimate muscle fibre size without the need of surgical biopsy sampling.
figure legend In this study, we investigated the relation between the conduction velocity of populations of motor units identified from biceps brachii muscle and muscle fibre size. We adopted high‐density surface EMG to decode the activity of voluntarily activated motor units and estimated their conduction velocity. Similarly, we adopted muscle biopsy to measure muscle fibre size. We revealed the possibility of accurately transforming motor unit conduction velocity values into estimated measures of muscle fibre size, which in turn showed a good degree of association with the muscle fibre size measured directly by muscle biopsy. Furthermore, we demonstrated that the proposed neuromuscular interface allows the prediction of the mean measured fibre diameter and cross‐sectional area from an EMG‐derived parameter with a relatively low bias and error, thus opening new perspectives in the use of high‐density EMG as a non‐invasive tool to estimate muscle fibre size without the need of surgical biopsy sampling.
Key points
We propose and validate a method for accurately identifying the activity of populations of motor neurons during contractions at maximal rate of force development in humans.
The behaviour ...of the motor neuron pool during rapid voluntary contractions in humans is presented.
We show with this approach that the motor neuron recruitment speed and maximal motor unit discharge rate largely explains the individual ability in generating rapid force contractions.
The results also indicate that the synaptic inputs received by the motor neurons before force is generated dictate human potential to generate force rapidly.
This is the first characterization of the discharge behaviour of a representative sample of human motor neurons during rapid contractions.
During rapid contractions, motor neurons are recruited in a short burst and begin to discharge at high frequencies (up to >200 Hz). In the present study, we investigated the behaviour of relatively large populations of motor neurons during rapid (explosive) contractions in humans, applying a new approach to accurately identify motor neuron activity simultaneous to measuring the rate of force development. The activity of spinal motor neurons was assessed by high‐density electromyographic decomposition from the tibialis anterior muscle of 20 men during isometric explosive contractions. The speed of motor neuron recruitment and the instantaneous motor unit discharge rate were analysed as a function of the impulse (the time–force integral) and the maximal rate of force development. The peak of motor unit discharge rate occurred before force generation and discharge rates decreased thereafter. The maximal motor unit discharge rate was associated with the explosive force variables, at the whole population level (r2 = 0.71 ± 0.12; P < 0.001). Moreover, the peak motor unit discharge and maximal rate of force variables were correlated with an estimate of the supraspinal drive, which was measured as the speed of motor unit recruitment before the generation of afferent feedback (P < 0.05). We show for the first time the full association between the effective neural drive to the muscle and human maximal rate of force development. The results obtained in the present study indicate that the variability in the maximal contractile explosive force of the human tibialis anterior muscle is determined by the neural activation preceding force generation.
Key points
We propose and validate a method for accurately identifying the activity of populations of motor neurons during contractions at maximal rate of force development in humans.
The behaviour of the motor neuron pool during rapid voluntary contractions in humans is presented.
We show with this approach that the motor neuron recruitment speed and maximal motor unit discharge rate largely explains the individual ability in generating rapid force contractions.
The results also indicate that the synaptic inputs received by the motor neurons before force is generated dictate human potential to generate force rapidly.
This is the first characterization of the discharge behaviour of a representative sample of human motor neurons during rapid contractions.
Key points
Previous studies have indicated that several weeks of strength training is sufficient to elicit significant adaptations in the neural drive sent to the muscles.
There are few data, ...however, on the changes elicited by strength training in the recruitment and rate coding of motor units during voluntary contractions. We show for the first time that the discharge characteristics of motor units in the tibialis anterior muscle tracked across the intervention are changed by 4 weeks of strength training with isometric voluntary contractions.
The specific adaptations included significant increases in motor unit discharge rate, decreases in the recruitment‐threshold force of motor units and a similar input–output gain of the motor neurons.
The findings suggest that the adaptations in motor unit function may be attributable to changes in synaptic input to the motor neuron pool or to adaptations in intrinsic motor neuron properties.
The strength of a muscle typically begins to increase after only a few sessions of strength training. This increase is usually attributed to changes in the neural drive to muscle as a result of adaptations at the cortical or spinal level. We investigated the change in the discharge characteristics of large populations of longitudinally tracked motor units in tibialis anterior before and after 4 weeks of strength training the ankle‐dorsiflexor muscles with isometric contractions. The adaptations exhibited by 14 individuals were compared with 14 control subjects. High‐density electromyogram grids with 128 electrodes recorded the myoelectric activity during isometric ramp contractions to the target forces of 35%, 50% and 70% of maximal voluntary force. The motor unit recruitment and derecruitment thresholds, discharge rate, interspike intervals and estimates of synaptic inputs to motor neurons were assessed. The normalized recruitment‐threshold forces of the motor units were decreased after strength training (P < 0.05). Moreover, discharge rate increased by 3.3 ± 2.5 pps (average across subjects and motor units) during the plateau phase of the submaximal isometric contractions (P < 0.001). Discharge rates at recruitment and derecruitment were not modified by training (P < 0.05). The association between force and motor unit discharge rate during the ramp‐phase of the contractions was also not altered by training (P < 0.05). These results demonstrate for the first time that the increase in muscle force after 4 weeks of strength training is the result of an increase in motor neuron output from the spinal cord to the muscle.
Key points
Previous studies have indicated that several weeks of strength training is sufficient to elicit significant adaptations in the neural drive sent to the muscles.
There are few data, however, on the changes elicited by strength training in the recruitment and rate coding of motor units during voluntary contractions. We show for the first time that the discharge characteristics of motor units in the tibialis anterior muscle tracked across the intervention are changed by 4 weeks of strength training with isometric voluntary contractions.
The specific adaptations included significant increases in motor unit discharge rate, decreases in the recruitment‐threshold force of motor units and a similar input–output gain of the motor neurons.
The findings suggest that the adaptations in motor unit function may be attributable to changes in synaptic input to the motor neuron pool or to adaptations in intrinsic motor neuron properties.