Determining patterns of motor recruitment during locomotion

2002 ◽  
Vol 205 (3) ◽  
pp. 359-369 ◽  
Author(s):  
James M. Wakeling ◽  
Motoshi Kaya ◽  
Genevieve K. Temple ◽  
Ian A. Johnston ◽  
Walter Herzog

SUMMARY Motor units are the functional units of muscle contraction in vertebrates. Each motor unit comprises muscle fibres of a particular fibre type and can be considered as fast or slow depending on its fibre-type composition. Motor units are typically recruited in a set order, from slow to fast, in response to the force requirements from the muscle. The anatomical separation of fast and slow muscle in fish permits direct recordings from these two fibre types. The frequency spectra from different slow and fast myotomal muscles were measured in the rainbow trout Oncorhynchus mykiss. These two muscle fibre types generated distinct low and high myoelectric frequency bands. The cat paw-shake is an activity that recruits mainly fast muscle. This study showed that the myoelectric signal from the medial gastrocnemius of the cat was concentrated in a high frequency band during paw-shake behaviour. During slow walking, the slow motor units of the medial gastrocnemius are also recruited, and this appeared as increased muscle activity within a low frequency band. Therefore, high and low frequency bands could be distinguished in the myoelectric signals from the cat medial gastrocnemius and probably corresponded, respectively, to fast and slow motor unit recruitment. Myoelectric signals are resolved into time/frequency space using wavelets to demonstrate how patterns of motor unit recruitment can be determined for a range of locomotor activities.

1996 ◽  
Vol 75 (1) ◽  
pp. 26-37 ◽  
Author(s):  
K. E. Tansey ◽  
B. R. Botterman

1. The recruitment order of 64 pairs of motor units, comprising 21 type-identified units, was studied during centrally evoked muscle contractions of the cat medial gastrocnemius (MG) muscle in an unanesthetized, high decerebrate preparation. Motor units were functionally isolated within the MG nerve by intra-axonal (or intramyelin) penetration with conventional glass microelectrodes. 2. Graded stimulation of the mesencephalic locomotor region (MLR) was used to evoke smoothly graded contractions, which under favorable conditions was estimated to reach 40% of maximum tetanic tension of the MG muscle. With this method of activation, 100% of slow twitch (type S) units, 95% of fast twitch, fatigue-resistant (type FR) units, 86% of fast twitch, fatigue-intermediate (type FI) units, and 49% of fast twitch, fatigable (type FF) units studied were recruited. 3. Motoneuron size as estimated by axonal conduction velocity (CV) was correlated with muscle-unit size as estimated by maximum tetanic tension (Po). Although the correlation between these properties was significant among type S and FR units, no significant correlation was found for these properties among type FI and FF units. 4. Motor-unit recruitment was ordered by physiological type (S > F, 100% of pairs; S > FR > FI > FF, 93% of pairs). Although none of the motor-unit properties studied predicted recruitment order perfectly, motor-unit recruitment was found to proceed by increasing Po (89% of pairs), decreasing contraction time (79% of pairs), decreasing fatigue index (80% of pairs), and increasing CV (76% of pairs). These percentages were significantly different from random (i.e., 50%). Statistically, all four motor-unit properties were equivalent in predicting recruitment order. These results are similar to those reported by other investigators for motor-unit recruitment order evoked from other supraspinal centers, as well as from peripheral sites. 5. When, however, motor-unit recruitment within pairs of motor units containing two fast-twitch (type F) units was examined, Po was a significantly better predictor of recruitment order than CV (85% vs. 52% of pairs). One explanation for this observation is that the correlation between Po and CV is high among type S, type FR units, and possibly among the lower-tension type FF units, but not among the remaining higher-tension type FF units. 6. The reproducibility of recruitment order in multiple contractions was investigated in 16 motor-unit pairs. Recruitment order was found to be variable in only three motor-unit pairs, all of which contained units of similar physiological type and recruitment threshold. 7. Analysis of recruitment order by pair-wise testing confirms the general conclusion reached in human studies that the muscle force level at recruitment for a motor unit is highly correlated with its strength. As an additional confirmation, the whole-muscle force level at recruitment for 41 units was measured in a series of contractions in which the rate of rise of muscle tension was limited to rates < 1,000 g/s. For these contractions, a significant correlation was found between muscle tension at recruitment and motor-unit Po.


2021 ◽  
Vol 18 (174) ◽  
pp. 20200765
Author(s):  
Adrian K. M. Lai ◽  
Taylor J. M. Dick ◽  
Andrew A. Biewener ◽  
James M. Wakeling

The nervous system is faced with numerous strategies for recruiting a large number of motor units within and among muscle synergists to produce and control body movement. This is challenging, considering multiple combinations of motor unit recruitment may result in the same movement. Yet vertebrates are capable of performing a wide range of movement tasks with different mechanical demands. In this study, we used an experimental human cycling paradigm and musculoskeletal simulations to test the theory that a strategy of prioritizing the minimization of the metabolic cost of muscle contraction, which improves mechanical efficiency, governs the recruitment of motor units within a muscle and the coordination among synergist muscles within the limb. Our results support our hypothesis, for which measured muscle activity and model-predicted muscle forces in soleus—the slower but stronger ankle plantarflexor—is favoured over the weaker but faster medial gastrocnemius (MG) to produce plantarflexor force to meet increased load demands. However, for faster-contracting speeds induced by faster-pedalling cadence, the faster MG is favoured. Similar recruitment patterns were observed for the slow and fast fibres within each muscle. By contrast, a commonly used modelling strategy that minimizes muscle excitations failed to predict force sharing and known physiological recruitment strategies, such as orderly motor unit recruitment. Our findings illustrate that this common strategy for recruiting motor units within muscles and coordination between muscles can explain the control of the plantarflexor muscles across a range of mechanical demands.


1996 ◽  
Vol 76 (3) ◽  
pp. 1503-1516 ◽  
Author(s):  
C. J. de Luca ◽  
P. J. Foley ◽  
Z. Erim

1. The purpose of this study was 1) to characterize the decrease observed in mean firing rates of motor units in the first 8-15 s of isometric constant-force contractions and 2) to investigate possible mechanisms that could account for the ability to maintain force output in the presence of decreasing motor unit firing rates. 2. The decrease in mean firing rates was characterized by investigating myoelectric signals detected with a specialized quadrifilar needle electrode from the first dorsal interosseus (FDI) and the tibialis anterior (TA) muscles of 19 healthy subjects during a total of 85 constant-force isometric contractions at 30, 50, or 80% of maximal effort. The firing times of motor units were obtained from the myoelectric signals with the use of computer algorithms to decompose the signal into the constituent motor unit action potentials. Time-varying mean firing rates and recruitment thresholds were also calculated. 3. Motor units detected from the TA muscle were found to have a continual decrease in their mean firing rates in 36 of 44 trials performed during isometric ankle dorsiflexion at force values ranging from 30 to 80% of maximal effort and a duration of 8-15 s. Likewise, motor units detected in the FDI muscle displayed a decrease in firing rate in 32 of 41 trials performed during constant-force isometric index finger abduction for contractions ranging from 30 to 80% of maximal effort. In 14 contractions (16% of total), firing rates were essentially constant, whereas in 3 contractions (4%), firing rates appeared to increase. 4. Motor units with the higher recruitment thresholds and lower firing rates tended to display the greater decreases in firing rate over the constant-force interval, whereas motor units with lower recruitment thresholds and higher firing rates had lesser rates of decrease. Furthermore, increasing contraction levels tended to intensify the decrease in the motor unit firing rates. 5. Three possible mechanisms were considered as factors responsible for the maintaining of force output while motor units decreased their firing rates: motor unit recruitment, agonist/antagonist interaction, and twitch potentiation. Of these, motor unit recruitment was discarded first because none was observed during the 8-15 s duration of any of the 85 contractions. Furthermore, contractions outside the physiological range of motor unit recruitment (at 80% of maximal effort) revealed the same decreasing trend in firing rates, ruling out recruitment as the means of sustaining force output. 6. The role of agonist or antagonist muscle interaction was investigated with the use of the muscles controlling the wrist joint. Myoelectric signals were recorded with quadrifilar needle electrodes from the wrist extensor muscles while myoelectric activity in the wrist flexor muscles was concurrently monitored with surface electrodes during constant-force isometric wrist extension at 50% of maximal effort. Firing rates of the motor units in the wrist extensor muscles simultaneously decreased while the flexor muscles were determined to be inactive. 7. All the findings of this study regarding the behavior of the firing rates could be well explained by the reported characteristics of twitch potentiation that have been previously documented in animals and humans. 8. The results of this study, combined with the results of other investigators, provide the following scenario to explain how a constant-force isometric contraction is sustained. As the contraction progresses, the twitch force of the muscle fibers undergoes a potentiation followed by a decrease. Simultaneously, the "late adaptation" property of the motoneuron decreases the firing rate of the motor unit. Findings of this study suggest that voluntary reduction in firing rates also cannot be ruled out as a means to augment the adaptation in motoneurons. (ABSTRACT TRUNCATED)


2014 ◽  
Vol 112 (7) ◽  
pp. 1685-1691 ◽  
Author(s):  
Christopher J. Dakin ◽  
Brian H. Dalton ◽  
Billy L. Luu ◽  
Jean-Sébastien Blouin

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


2021 ◽  
Vol 14 (3) ◽  
pp. 112
Author(s):  
Kai Shi

We attempted to comprehensively decode the connectedness among the abbreviation of five emerging market countries (BRICS) stock markets between 1 August 2002 and 31 December 2019 not only in time domain but also in frequency domain. A continuously varying spillover index based on forecasting error variance decomposition within a generalized abbreviation of vector-autoregression (VAR) framework was computed. With the help of spectral representation, heterogeneous frequency responses to shocks were separated into frequency-specific spillovers in five different frequency bands to reveal differentiated linkages among BRICS markets. Rolling sample analyses were introduced to allow for multiple changes during the sample period. It is found that return spillovers dominated by the high frequency band (within 1 week) part declined with the drop of frequencies, while volatility spillovers dominated by the low frequency band (above 1 quarter) part grew with the decline in frequencies; the dynamics of spillovers were influenced by crucial systematic risk events, and some similarities implied in the spillover dynamics in different frequency bands were found. From the perspective of identifying systematic risk sources, China’s stock market and Russia’s stock market, respectively, played an influential role for return spillover and volatility spillover across BRICS markets.


2021 ◽  
Vol 18 ◽  
Author(s):  
Luoyu Wang ◽  
Qi Feng ◽  
Mei Wang ◽  
Tingting Zhu ◽  
Enyan Yu ◽  
...  

Background: As a potential brain imaging biomarker, amplitude of low frequency fluc-tuation (ALFF) has been used as a feature to distinguish patients with Alzheimer’s disease (AD) and amnestic mild cognitive impairment (aMCI) from normal controls (NC). However, it remains unclear whether the frequency-dependent pattern of ALFF alterations can effectively distinguish the different phases of the disease. Methods: In the present study, 52 AD and 50 aMCI patients were enrolled together with 43 NC in total. The ALFF values were calculated in the following three frequency bands: classical (0.01-0.08 Hz), slow-4 (0.027-0.073 Hz) and slow-5 (0.01-0.027 Hz) for the three different groups. Subsequently, the local functional abnormalities were employed as features to examine the effect of classification among AD, aMCI and NC using a support vector machine (SVM). Results: We found that the among-group differences of ALFF in the different frequency bands were mainly located in the left hippocampus (HP), right HP, bilateral posterior cingulate cortex (PCC) and bilateral precuneus (PCu), left angular gyrus (AG) and left medial prefrontal cortex (mPFC). When the local functional abnormalities were employed as features, we identified that the ALFF in the slow-5 frequency band showed the highest accuracy to distinguish among the three groups. Conclusion: These findings may deepen our understanding of the pathogenesis of AD and suggest that slow-5 frequency band may be helpful to explore the pathogenesis and distinguish the phases of this disease.


1987 ◽  
Vol 57 (4) ◽  
pp. 1227-1245 ◽  
Author(s):  
R. C. Foehring ◽  
G. W. Sypert ◽  
J. B. Munson

We tested whether the muscle innervated may influence the expression of motoneuron electrical properties. Properties of individual motor units were examined following cross-reinnervation (X-reinnervation) of cat lateral gastrocnemius (LG) and soleus muscles by the medial gastrocnemius (MG) nerve. We examined animals at two postoperative times: 9-10 wk (medX) and 9-11 mo (longX). For comparison, normal LG and soleus motoneuron properties were also studied. Motor units were classified on the basis of their contractile responses as fast contracting fatigable, fast intermediate fast contracting fatigue resistant, and slow types FF, FI, FR, or S, respectively) (9, 21). Motoneuron electrical properties (rheobase, input resistance, axonal conduction velocity, afterhyperpolarization) were measured. After 9-11 mo, MG motoneurons that innervated LG muscle showed recovery of electrical properties similar to self-regenerated MG motoneurons. The relationships between motoneuron electrical properties were largely similar to self-regenerated MG. For MG motoneurons that innervated LG, motoneuron type (65) predicted motor-unit type in 74% of cases. LongX-soleus motoneurons differed from longX-LG motoneurons or self-regenerated MG motoneurons in mean values for motoneuron electrical properties. The differences in overall means reflected the predominance of type S motor units. The relationships between motoneuron electrical properties were also different than in self-regenerated MG motoneurons. In all cases, the alterations were in the direction of properties of type S units, and the relationship between normal soleus motoneurons and their muscle units. Within motor-unit types, the mean values were typical for that type in self-regenerated MG. Motoneuron type (65) was a fairly strong predictor of motor-unit type in longX soleus. MG motoneurons that innervated soleus displayed altered values for axonal conduction velocity, rheobase, and input resistance, which could indicate incomplete recovery from the axotomized state. However, although mean afterhyperpolarization (AHP) half-decay time was unaltered by axotomy (25), this parameter was significantly lengthened in MG motoneurons that innervated soleus muscle. There were, however, individual motoneuron-muscle-unit mismatches, which suggested that longer mean AHP half-decay time may also be due to incomplete recovery of a subpopulation of motoneurons. Those MG motoneurons able to specify soleus muscle-fiber type exhibited motoneuron electrical properties typical of that same motoneuron type in self-regenerated MG.(ABSTRACT TRUNCATED AT 400 WORDS)


1994 ◽  
Vol 76 (6) ◽  
pp. 2663-2671 ◽  
Author(s):  
L. J. Einsiedel ◽  
A. R. Luff

The aim of the study was to determine whether increased motoneuron activity induced by treadmill walking would alter the extent of motoneuron sprouting in the partially denervated rat medial gastrocnemius muscle. An extensive partial denervation was effected by unilateral section of the L5 ventral root, and it is very likely that all units remaining in the medial gastrocnemius were used in treadmill walking. Rats were trained for 1.5 h/day and after 14 days were walking at least 1 km/day. Motor unit characteristics were determined 24 days after the partial denervation and were compared with units from partially denervated control (PDC) animals and with units from normal (control) animals. In PDC rats, force developed by slow, fast fatigue-resistant, and fast intermediate-fatigable motor units increased substantially compared with control animals; that of fast-fatigable units did not increase. In partially denervated exercised animals, force developed by slow and fast-fatigue-resistant units showed no further increase, but fast-intermediate- and fast-fatigable units showed significant increases compared with those in PDC animals. The changes in force were closely paralleled by changes in innervation ratios. We concluded that neuronal activity is an important factor in determining the rate of motoneuron sprouting.


1997 ◽  
Vol 77 (5) ◽  
pp. 2605-2615 ◽  
Author(s):  
John B. Munson ◽  
Robert C. Foehring ◽  
Lorne M. Mendell ◽  
Tessa Gordon

Munson, John B., Robert C. Foehring, Lorne M. Mendell, and Tessa Gordon. Fast-to-slow conversion following chronic low-frequency activation of medial gastrocnemius muscle in cats. II. Motoneuron properties. J. Neurophysiol. 77: 2605–2615, 1997. Chronic stimulation (for 2–3 mo) of the medial gastrocnemius (MG) muscle nerve by indwelling electrodes renders the normally heterogeneous MG muscle mechanically and histochemically slow (type SO). We tested the hypothesis that motoneurons of MG muscle thus made type SO by chronic stimulation would also convert to slow phenotype. Properties of all single muscle units became homogeneously type SO (slowly contracting, nonfatiguing, nonsagging contraction during tetanic activation). Motoneuron electrical properties were also modified in the direction of type S, fatigue-resistant motor units. Two separate populations were identified (on the basis of afterhyperpolarization, rheobase, and input resistance) that likely correspond to motoneurons that had been fast (type F) or type S before stimulation. Type F motoneurons, although modified by chronic stimulation, were not converted to the type S phenotype, despite apparent complete conversion of their muscle units to the slow oxidative type (type SO). Muscle units of the former type F motor units were faster and/or more powerful than those of the former type S motor units, indicating some intrinsic regulation of motor unit properties. Experiments in which chronic stimulation was applied to the MG nerve cross-regenerated into skin yielded changes in motoneuron properties similar to those above, suggesting that muscle was not essential for the effects observed. Modulation of group Ia excitatory postsynaptic potential (EPSP) amplitude during high-frequency trains, which in normal MG motoneurons can be either positive or negative, was negative in 48 of 49 chronically stimulated motoneurons. Negative modulation is characteristic of EPSPs in motoneurons of most fatigue-resistant motor units. The general hypothesis of a periphery-to-motoneuron retrograde mechanism was supported, although the degree of control exerted by the periphery may vary: natural type SO muscle appears especially competent to modify motoneuron properties. We speculate that activity-dependent regulation of the neurotrophin-(NT) 4/5 in muscle plays an important role in controlling muscle and motoneuron properties.


1991 ◽  
Vol 65 (4) ◽  
pp. 952-967 ◽  
Author(s):  
C. J. Heckman ◽  
M. D. Binder

1. A pool of 100 simulated motor units was constructed in which the steady-state neural and mechanical properties of the units were very closely matched to the available experimental data for the cat medial gastrocnemius motoneuron pool and muscle. The resulting neural network generated quantitative predictions of whole system input-output functions based on the single unit data. The results of the simulations were compared with experimental data on normal motor system behavior in humans and animals. 2. We considered only steady-state, isometric conditions. All motoneurons received equal proportions of the synaptic input, and no feedback loops were operative. Thus the intrinsic properties of the motor unit population alone determined the form of the system input-output function. Expressing the synaptic input in terms of effective synaptic current allowed the simulated motoneuron input-output functions to be specified by well-known firing rate-injected current relations. The motor unit forces were determined from standard motor unit force-frequency relations, and the system output at any input level was assumed to be the linear sum of the forces of the active motor units. 3. The steady-state input-output function of the simulated motoneuron pool had a roughly sigmoidal shape that was quite different from those derived from previous recruitment models, which did not incorporate frequency modulation. Frequency modulation in combination with the skewed distribution of thresholds (low values much more frequent than high) restricted upward curvature to low input levels, whereas frequency modulation alone was responsible for the final gradual approach to the maximum force output. 4. Sensitivity analyses were performed to assess the importance of several assumptions that were required to deal with gaps and uncertainties in the available experimental data. The shape of the input-output function was not critically dependent on any of these assumptions, including those specifying linear summation of inputs and outputs. 5. A key assumption of the model was that systematic variance in motor unit properties was much more important than random variance for determining the input-output function. Addition of random variance via Monte Carlo techniques showed that this assumption was correct. These results suggest that the output of a motoneuron pool should be quite tolerant of random variance in the distribution of synaptic inputs and yet substantially altered by any systematic differences, such as unequal distribution of inputs among different motor unit types.(ABSTRACT TRUNCATED AT 400 WORDS)


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