Fast Motor Units Research Articles

Influence of 21-day antiortostatic hypokinesia on the functional state of the musculoskeletal system of human

The influence of antiorthostatic hypokinesia with an inclination angle relative to the horizon of -6 degrees for 21 days was used as a ground model of the physiological effects of weightlessness. 10 practically healthy male volunteers (30.7 ± 5.4 years, 78.0 ± 8.5 kg, 179.7 ± 5.3 cm) took part in the experimental study, who successfully passed the medical expert commission of the SSC RF-IMBP RAS, familiarized with the study program and signed a voluntary informed consent to participate in the study. The state of the musculoskeletal system was assessed according to the results of speed-strength testing on an isokinetic dynamometer before hypokinesia and on the 3rd day after its completion. Staying in conditions of 21-day anti-orthostatic hypokinesia with a body inclination angle of –6° relative to the horizon, as a model of the physiological effects of weightlessness, leads to changes in the functional state of the musculoskeletal lower limbs which manifests itself during speed-strength testing after hypokinesia by a decrease in maximum voluntary force (MPF) of the knee joint extensor muscles from 9 to 15% compared with the base level. The decrease in MPF did not depend on a change in the force gradient, which reflects the ability to exert greater force in the shortest possible time. This indicated that the decrease in the MPF level after hypokinesia was mainly due to a change in the activity of slow motor units. At the same time, we assume that exposure to hypokinesia did not cause a significant change in the activity of fast motor units. This is confirmed by the results of the analysis of the electromyographic activity of the extensor muscles of the knee joint during testing on an isokinetic dynamometer. Also, after hypokinesia, the possibility of using muscle potential significantly decreased - the physiological cost of work increased with a decrease in strength indicators.

Age-associated impairment in swallow function is concomitant with nucleus ambiguus motor neuron death in Fischer 344 rats

The nucleus ambiguus contains motor neurons (MNs) innervating the muscles of the larynx, pharynx and oesophagus. Many of the motor units these muscles are comprised of are fast fatigueable units — highly vulnerable to neurodegeneration. Loss of the MNs or damage to the nerves innervating the muscles of the larynx, pharynx and oesophagus may impair swallow, vocalization and/or respiratory functions. Of particular importance, human patients with deficits in swallow are at risk of sequelae such as aspiration pneumonia and malnutrition. In aged male Fischer 344-Brown Norway crosses, age-associated loss of nucleus ambiguus MNs contributes to vocalisation deficits. Here, we use female and male Fischer 344 (F344) at young (6-months old) and old (24-months old) to evaluate swallow function and nucleus ambiguus MN survival. Swallow function was evaluated under anesthesia by delivering an 0.3 mL bolus of water to the base of the tongue and videoing the number of subsequent swallows. A pressure catheter was inserted into the oesophagus to evaluate the incidence and duration of wallow-associated apneas and the magnitude of the schluckatmung in 30 s following bolus delivery. Following functional evaluations, rats were euthanized, perfused with paraformaldehyde and brainstem tissue cryopreserved, sectioned and stained with Nissl to stereologically evaluate the number of nucleus ambiguus MNs. In old age, we observed an ~30% reduction in the number of swallows per delivery of the water bolus and an increased incidence and duration of swallow apnea. Schluckatmung was not affected by age. The number of nucleus ambiguus MNs was reduced by ~20% in old F344 rats, with MN size unaffected. We propose that swallow evaluations are a robust readout of oropharyngeal behaviour, with deficits concomitant with MN death. These findings are similar to age-associated weakness in other skeletal muscle being related to MN death. It is likely that the mechanisms underpinning fast fatigue motor unit MN death in brainstem and spinal pools are similar. This research was funded by HL166204. This is the full abstract presented at the American Physiology Summit 2024 meeting and is only available in HTML format. There are no additional versions or additional content available for this abstract. Physiology was not involved in the peer review process.

Functional electrical stimulation therapy for upper extremity rehabilitation following spinal cord injury: a pilot study.

Pilot study. To examine if functional electrical stimulation therapy (FEST) improves neuromuscular factors underlying upper limb function in individuals with SCI. A tertiary spinal cord rehabilitation center specialized in spinal cord injury care in Canada. We examined 29 muscles from 4 individuals living with chronic, cervical, and incomplete SCI. The analysis was focused on the changes in muscle activation, as well as on how the treatment could change the ability to control a given muscle or on how multiple muscles would be coordinated during volitional efforts. There was evidence of gains in muscle strength, activation, and median frequency after the FEST. Gains in muscle activation indicated the activation of a greater number of motor units and gains in muscle median frequency the involvement of higher threshold, faster motor units. In some individuals, these changes were smaller but accompanied by increased control over muscle contraction, evident in a greater ability to sustain a volitional contraction, reduce the co-contraction of antagonist muscles, and provide cortical drive. FEST increases muscle strength and activation. Enhanced control of muscle contraction, reduced co-contraction of antagonist muscles, and a greater presence of cortical drive were some of the findings supporting the effects of FEST at the sensory-motor integration level.

Electromyostimulation in sport

Electromyostimulation (EMS) is a widely used therapeutic tool of low-frequency electrotherapy in several areas of rehabilitation, with gradual use increasingly in sports as well. The main goal of our article is to approach the use of EMS in sports based on available studies. EMS can be called as a training method used to obtain muscle activation through externally applied electric currents to the muscles to achieve a functional increase in sports performance by training fast motor units. Even if, based on the available study database, there are no clear conclusions regarding the effectiveness and application of EMS in the sense of sports, we can say that EMS represents one of the means by which we can influence muscle functionality, not only in terms of rehabilitation, but also as a possible way of increasing sports performance.

Early deficits in GABA inhibition parallels an increase in L-type Ca2+ currents in the jaw motor neurons of SOD1G93A mouse model for ALS

Amyotrophic Lateral Sclerosis (ALS) involves protracted pre-symptomatic periods of abnormal motor neuron (MN) excitability occurring in parallel with central and peripheral synaptic perturbations. Focusing on inhibitory control of MNs, we first compared longitudinal changes in pre-synaptic terminal proteins for GABA and glycine neurotransmitters around the soma of retrogradely identified trigeminal jaw closer (JC) MNs and ChAT-labeled midbrain extraocular (EO) MNs in the SOD1G93A mouse model for ALS. Fluorescence immunocytochemistry and confocal imaging were used to quantify GAD67 and GlyT2 synaptic bouton density (SBD) around MN soma at pre-symptomatic ages ∼P12 (postnatal), ∼P50 (adult) and near disease end-stage (∼P135) in SOD1G93A mice and age-matched wild-type (WT) controls. We noted reduced GAD67 innervation in the SOD1G93A trigeminal jaw closer MNs around P12, relative to age-matched WT and no significant difference around P50 and P135. In contrast, both GAD67 and GlyT2 innervation were elevated in the SOD1G93A EO MNs at the pre-symptomatic time points. Considering trigeminal MNs are vulnerable in ALS while EO MNs are spared, we suggest that upregulation of inhibition in the latter might be compensatory. Notable contrast also existed in the innate co-expression patterns of GAD67 and GlyT2 with higher mutual information (co-dependency) in EO MNs compared to JC in both SOD1G93A and WT mice, especially at adult stages (P50 and P135). Around P12 when GAD67 terminals expression was low in the mutant, we further tested for persistent GABA inhibition in those MNs using in vitro patch-clamp electrophysiology. Our results show that SOD1G93A JC MNs have reduced persistent GABA inhibition, relative to WT. Pharmacological blocking of an underlying tonically active GABA conductance using the GABA-α5 subunit inverse agonist, L-655-708, disinhibited WT JC MNs and lowered their recruitment threshold, suggesting its role in the control of intrinsic MN excitability. Quantitative RT-PCR in laser dissected JC MNs further supported a reduction in GABA-α5 subunit mRNA expression in the mutant. In light of our previous report that JC MNs forming putative fast motor units have lower input threshold in the SOD1G93A mice, we suggest that our present result on reduced GABA-α5 tonic inhibition provides for a mechanism contributing to such imbalance. In parallel with reduced GABA inhibition, we noted an increase in voltage-gated L-type Ca2+ currents in the mutant JC MNs around P12. Together these results support that, early modifications in intrinsic properties of vulnerable MNs could be an adaptive response to counter synaptic deficits.

Mitochondrial adaptations to inactivity in diaphragm muscle fibers.

Type I and IIa diaphragm muscle (DIAm) fibers comprise slow and fast fatigue-resistant motor units that are recruited to accomplish breathing and thus have a high duty cycle. In contrast, type IIx/IIb fibers comprise more fatigable fast motor units that are infrequently recruited for airway protective and straining behaviors. We hypothesize that mitochondrial structure and function in type I and IIa DIAm fibers adapt in response to inactivity imposed by spinal cord hemisection at C2 (C2SH). At 14 days after C2SH, the effect of inactivity on mitochondrial structure and function was assessed in DIAm fibers. Mitochondria in DIAm fibers were labeled using MitoTracker Green (Thermo Fisher Scientific), imaged in three-dimensions (3-D) by fluorescence confocal microscopy, and images were analyzed for mitochondrial volume density (MVD) and complexity. DIAm homogenate from either side was assessed for PGC1α, Parkin, MFN2, and DRP1 using Western blot. In alternate serial sections of the same DIAm fibers, the maximum velocity of the succinate dehydrogenase reaction (SDHmax) was determined using a quantitative histochemical technique. In all groups and both sides of the DIAm, type I and IIa DIAm fibers exhibited higher MVD, with more filamentous mitochondria and had higher SDHmax normalized to both fiber volume and mitochondrial volume compared with type IIx/IIb Diam fibers. In the inactive right side of the DIAm, mitochondria became fragmented and MVD decreased in all fiber types compared with the intact side and sham controls, consistent with the observed reduction in PGC1α and increased Parkin and DRP1 expression. In the inactive side of the DIAm, the reduction in SDHmax was found only for type I and IIa fibers. These results show that there are intrinsic fiber-type-dependent differences in the structure and function of mitochondria in DIAm fibers. Following C2SH-induced inactivity, mitochondrial structure (MVD and fragmentation) and function (SDHmax) were altered, indicating that inactivity influences all DIAm fiber types, but inactivity disproportionately affected SDHmax in the more intrinsically active type I and IIa fibers.NEW & NOTEWORTHY Two weeks of diaphragm (DIAm) inactivity imposed by C2SH caused reduced mitochondrial volume density, mitochondrial fragmentation, and a concomitant reduction of SDHmax in type I and IIa DIAm fibers on the lesioned side. Type I and IIa DIAm fibers were far more sensitive to inactivation than type IIx/IIb fibers, which exhibited little pathology. Our results indicate that mitochondria in DIAm fibers are plastic in response to varying levels of activity.

Mitochondria Adapt to Diaphragm Muscle Inactivity Imposed by Cervical Spinal Cord Injury

Type I and IIa diaphragm muscle (DIAm) fibers comprise fatigue resistant slow and fast motor units that are recruited to accomplish breathing. In contrast, type IIx/IIb fibers comprise more fatigable fast motor units that are infrequently recruited only during airway protective and straining behaviors. The activity and energetic demand for type I and IIa DIAm fibers is markedly increased compared to type IIx/IIb fibers. Cervical spinal cord hemisection at C2 (C2SH) removes excitatory drive for inspiratory activity of the DIAm and thereby imposing prolonged inactivity of type I and IIa fibers. We hypothesize that mitochondrial structure and function in type I and IIa DIAm fibers adapt in response to imposed inactivity induced by spinal cord hemisection at C2 (C2SH). After C2SH, inactivity was confirmed by the absence of DIAm EMG, and after 14 days, and the effect of inactivity on mitochondrial structure and function was assessed in type identified fibers. Mitochondria in DIAm fibers were labeled using MitoTracker Green and imaged in 3‐D by fluorescence confocal microscopy. Mitochondrial volume density and morphometry were analyzed using ImageJ. In alternate serial sections of the same DIAm fibers, the maximum velocity of the succinate dehydrogenase reaction (SDHmax) was determined using a quantitative histochemical technique. Overall, mitochondrial volume density was higher in type I and IIa DIAm fibers and mitochondria were more filamentous compared to type IIx/IIb fibers. Inactivity reduced mitochondrial volume density in type I and IIa DIAm fibers and induced mitochondrial fragmentation. The SDHmax values normalized for fiber volume and for mitochondrial volume were both higher in type I and IIa DIAm fibers compared to type IIx/IIb fibers. After 14 days of inactivity induced by C2SH, SDHmax (normalized for fiber volume and for mitochondrial volume) decreased in type I and IIa. These results are consistent with mitochondrial structural and functional adaptions to the marked reduction in the energetic demand of type I and IIa DIAm fibers imposed by C2SH.

Contractions dynamic of “fast” and “slow” rat muscle under spinal shock and modulators of contraction

Aim. To study the dynamics of neuromotor regulation of the contractile function of fast and slow muscles in rodents during spinal shock by spinal cord transection at the level Тh11Тh12.
 Methods. The experiments were carried out on laboratory rats weighing 140180 g. The animals were divided into two groups: Control (8 rats) and Spinal shock (6 rats). The lower leg muscles, m. soleus and m. extensor digitorum longus (m. EDL), were dissected by partially isolating without disrupting the connection with the body's circulatory system. The sciatic nerve was stimulated with single electrical impulses (10 V, 0.5 ms). Contractions of both muscles caused by electrical stimulation of the sciatic nerve before and after the injection of the substances into the femoral artery tubocurarine (1 mM) or norepinephrine (10 mM) were recorded in animals of both groups. After spinalization, muscle contractions were re-recorded during electrical stimulation of the sciatic nerve before and 10 minutes after the injection of tubocurarine or noradrenaline into the femoral artery in the same concentrations.
 Results. After spinalization of the animal, the contraction force of the muscle m. EDL fibers increased to 0.430.03 g (p=0.040), but the temporal parameters remained unchanged. M. soleus, on the contrary, showed a decrease in the contraction time to 0.0530.005 s (p=0.045), and no change in the contraction force was observed under these conditions. Intra-arterial administration of norepinephrine in the control group resulted in an increase of m. soleus contractions up to 1.210.17 g (p=0.048), and m. EDL up to 0.570.07 g (p=0.043). The administration of norepinephrine in spinalized animals caused an increase in the contraction of m. soleus up to 1.210.09 g (p=0.047), and m. EDL up to 0.660.05 g (p=0.043). The blocker of postsynaptic cholinergic receptors tubocurarine administration reduced the force of contraction of both muscle types in both control [m. soleus up to 0.390.03 g (p=0.039), m. EDL up to 0.110.02 g (p=0.042)] and spinalized [m. soleus up to 0.340.05 g (p=0.039), m. EDL up to 0.150.04 g (p=0.040)] animals.
 Conclusion. The data obtained demonstrate the presence of significant differences in the mechanisms of control of contractile activity in the fast and slow skeletal muscles of warm-blooded animals; the persistence of the similar effect of the basic modulators on the contraction of both muscles with such a striking reaction to spinalization highlights the contribution of neurotrophic control to the functioning of fast and slow motor units.

Plastyczność układu nerwowo-mięśniowego - adaptacja do zmienionego poziomu aktywności ruchowej

Plastyczność układu nerwowo-mięśniowego to trwająca całe życie zdolność do reorganizacji struktury i funkcji pod wpływem doświadczenia. Obok plastyczności związanej z rozwojem organizmu, procesami uczenia się lub odnową funkcji po uszkodzeniach, wyróżnia się plastyczność adaptacyjną, która polega na dostosowaniu właściwości neuronów i/lub włókien mięśniowych do zmienionego poziomu aktywności ruchowej. Celem niniejszego artykułu było zebranie aktualnej wiedzy na temat zmian adaptacyjnych jednostek ruchowych w oparciu o wyniki kilku serii eksperymentów wykonanych na modelach zwierzęcych. Podobieństwo struktury i mechanizmów sterowania pracą mięśni u wszystkich ssaków uzasadniają wnioskowanie na temat podobnych procesów u człowieka. W pierwszych rozdziałach prezentujemy pojęcie jednostki ruchowej, składającej się z jednego motoneuronu i grupy włókien mięśniowych unerwianych przez ten motoneuron, przedstawiamy charakterystykę histologiczną, biochemiczną i elektrofizjologiczną trzech typów jednostek (S, FR i FF) o odmiennych cechach skurczu oraz omawiamy ich udział w różnych formach wysiłku fizycznego. W kolejnych rozdziałach prezentujemy wyniki projektów badawczych, przeprowadzonych w celu odpowiedzi na pytania, w jaki sposób odmienne rodzaje treningu fizycznego (prowadzonego w celu wzrostu wytrzymałości lub zwiększenia siły), przeciążenie mięśni, unieruchomienie kończyny lub ograniczenie ruchu wpływają na cechy skurczu włókien mięśniowych oraz właściwości elektrofizjologiczne motoneuronów różnych typów jednostek ruchowych. Omówione obserwacje doświadczalne dowodzą, że plastyczność jednostek ruchowych to procesy zachodzące równolegle w neuronach i włóknach mięśniowych. W zależności od rodzaju treningu i jego intensywności oraz od czasu trwania zwiększonej lub ograniczonej aktywności ruchowej mogą one polegać na transformacji włókien mięśniowych jednego typu w inny, modulacji podstawowych cech skurczu, zmian odporności na zmęczenie, zmian pobudliwości motoneuronów, lub częstotliwości generowanych przez nie impulsów nerwowych. Zwykle jednostki ruchowe szybkie (FF i FR) wykazują większą dynamikę procesów adaptacyjnych niż jednostki wolne (S).

Biomechanical conditioning of the motor unit transitory force decrease following a reduction in stimulation rate

BackgroundThe biomechanical background of the transitory force decrease following a sudden reduction in the stimulation frequency under selected experimental conditions was studied on fast resistant motor units (MUs) of rat medial gastrocnemius in order to better understand the mechanisms of changes in force transmission.MethodsFirstly, MUs were stimulated with three-phase trains of stimuli (low–high–low frequency pattern) to identify patterns when the strongest force decrease (3–36.5%) following the middle high frequency stimulation was observed. Then, in the second part of experiments, the MUs which presented the largest force decrease in the last low-frequency phase were alternatively tested under one of five conditions to analyse the influence of biomechanical factors of the force decrease: (1) determine the influence of muscle stretch on amplitude of the force decrease, (2) determine the numbers of interpulse intervals necessary to evoke the studied phenomenon, (3) study the influence of coactivation of other MUs on the studied force decrease, (4) test the presence of the transitory force decrease at progressive changes in stimulation frequency, (5) and perform mathematical analysis of changes in twitch-shape responses to individual stimuli within a tetanus phase with the studied force decrease.ResultsResults indicated that (1) the force decrease was highest when the muscle passive stretch was optimal for the MU twitch (100 mN); (2) the middle high-frequency burst of stimuli composed of at least several pulses was able to evoke the force decrease; (3) the force decrease was eliminated by a coactivation of 10% or more MUs in the examined muscle; (4) the transitory force decrease occured also at the progressive decrease in stimulation frequency; and (5) a mathematical decomposition of contractions with the transitory force decrease into twitch-shape responses to individual stimuli revealed that the force decrease in question results from the decrease of twitch forces and a shortening in contraction time whereas further force restitution is related to the prolongation of relaxation.ConclusionsHigh sensitivity to biomechanical conditioning indicates that the transitory force decrease is dependent on disturbances in the force transmission predominantly by collagen surrounding active muscle fibres.

Isokinetic Plantar Flexion Endurance. Reliability and Validity of output/excitation Measurements

The reliability and validity of isokinetic measurement of plantar flexion endurance has been studied by a method previously described by us which utilizes simultaneous measurements of mechanical contractional work (CW) and integrated electromyogram (iEMG). The reliability was gauged by test/re-test with a two year's interval; while validity was assessed by myoelectric power spectrum analyses. The output/input balance (CW/iEMG) remained unchanged for the group as well as inter-individually. Changes in myoelectric power spectrum as function of number of contractions were clearly indicative of fatigue. Under the present conditions fatigue of fast twitch motor units may explain the rapid decreases in output and excitation followed by nearly steady state levels of all registered parameters. As is the case for non-fatigued isokinetic plantar flexions, the motor unit recruitment order appears quite stereotyped during plantar flexion fatigue. The findings of significantly lower mean power spectrum during the first part of each contraction than during the second part may support the size-principle described by Henneman.

Estimation of self-sustained activity produced by persistent inward currents using firing rate profiles of multiple motor units in humans.

Persistent inward calcium and sodium currents (IP) activated during motoneuron recruitment help synaptic inputs maintain self-sustained firing until derecruitment. Here, we estimate the contribution of the IP to self-sustained firing in human motoneurons of varying recruitment threshold by measuring the difference in synaptic input needed to maintain minimal firing once the IP is fully activated compared with the larger synaptic input required to initiate firing before full IP activation. Synaptic input to ≈20 dorsiflexor motoneurons simultaneously recorded during ramp contractions was estimated from firing profiles of motor units decomposed from high-density surface electromyography (EMG). To avoid errors introduced when using high-threshold units firing in their nonlinear range, we developed methods where the lowest threshold units firing linearly with force were used to construct a composite (control) unit firing rate profile to estimate synaptic input to higher threshold (test) units. The difference in the composite firing rate (synaptic input) at the time of test unit recruitment and derecruitment (ΔF = Frecruit - Fderecruit) was used to measure IP amplitude that sustained firing. Test units with recruitment thresholds 1-30% of maximum had similar ΔF values, which likely included both slow and fast motor units activated by small and large motoneurons, respectively. This suggests that the portion of the IP that sustains firing is similar across a wide range of motoneuron sizes.NEW & NOTEWORTHY A new method of estimating synaptic drive to multiple, simultaneously recorded motor units provides evidence that the portion of the depolarizing drive from persistent inward currents that contributes to self-sustained firing is similar across motoneurons of different sizes.

Initial force production before sag is enhanced by prior contraction followed by a 3-minute rest period in fast motor units of the rat medial gastrocnemius

Unfused tetanic contractions evoked in fast motor units exhibit extra-efficient force production at the onset of contraction, an effect called “boost”. Boost is diminished in subsequent contractions if there is a short rest period between contractions, but can be re-established with a longer period of rest. We tested the hypothesis that contractile activity and rest could enhance boost-related metrics. Two sets of 3 unfused tetani were evoked 3 min apart in fast fatigable (FF) and fast fatigue-resistant (FR) motor units of the rat medial gastrocnemius. The greatest changes occurred in the first unfused tetanic contractions. Relative to the first contraction in the first set, the first contraction in the second set exhibited higher peak force during boost in a subset of motor units (76% of FF and 48% of FR). Enhanced force during boost was influenced by interaction of slowing of twitch contraction time (up to 20% and 25%, for FF and FR motor units, respectively), half-relaxation time (up to 37% and 49% for FF and FR motor units, respectively), and potentiation of the first twitch (up to 13% and 5% for FF and FR motor units, respectively). Examination of twitches evoked between sets suggested opportunity for greater enhancement of boost with shorter intervening rest periods. The phenomenon of enhanced boost following motor unit activity may interest sports scientists.

Microendoscopy detects altered muscular contractile dynamics in a mouse model of amyotrophic lateral sclerosis

Amyotrophic lateral sclerosis (ALS) is a fatal disease involving motor neuron degeneration. Effective diagnosis of ALS and quantitative monitoring of its progression are crucial to the success of clinical trials. Second harmonic generation (SHG) microendoscopy is an emerging technology for imaging single motor unit contractions. To assess the potential value of microendoscopy for diagnosing and tracking ALS, we monitored motor unit dynamics in a B6.SOD1G93A mouse model of ALS for several weeks. Prior to overt symptoms, muscle twitch rise and relaxation time constants both increased, consistent with a loss of fast-fatigable motor units. These effects became more pronounced with disease progression, consistent with the death of fast fatigue-resistant motor units and superior survival of slow motor units. From these measurements we constructed a physiological metric that reflects the changing distributions of measured motor unit time constants and effectively diagnoses mice before symptomatic onset and tracks disease state. These results indicate that SHG microendoscopy provides a means for developing a quantitative, physiologic characterization of ALS progression.

Keep calm and hang on: EMG activation in the forelimb musculature of three-toed sloths (Bradypus variegatus).

Sloths exhibit below branch locomotion whereby their limbs are loaded in tension to support the body weight. Suspensory behaviors require both strength and fatigue resistance from the limb flexors; however, skeletal muscle mass of sloths is reduced compared with other arboreal mammals. Although suspensory locomotion demands that muscles are active to counteract the pull of gravity, it is possible that sloths minimize muscle activation and/or selectively recruit slow motor units to maintain support, thus indicating neuromuscular specializations to conserve energy. Electromyography (EMG) was evaluated in a sample of three-toed sloths (Bradypus variegatus; N=6) to test this hypothesis. EMG was recorded at 2000 Hz via fine-wire electrodes implanted into two suites of four muscles in the left forelimb while sloths performed suspensory hanging (SH), suspensory walking (SW) and vertical climbing (VC). All muscles were minimally active for SH. During SW and VC, sloths moved slowly (duty factor: 0.83) and activation patterns were consistent between behaviors; the flexors were activated early and for a large percentage of limb contact, whereas the extensors were activated for shorter burst durations on average and showed biphasic (contact and swing) activity. Muscle activities were maximal for the elbow flexors and lowest for the carpal/digital flexors, and overall activity was significantly greater for SW and VC compared with SH. Wavelet analysis indicated high mean EMG frequencies from the myoelectric intensity spectra coupled with low burst intensities for SH, although the opposite pattern occurred for SW and VC, with the shoulder flexors and elbow flexor, m. brachioradialis, having extremely low mean EMG frequencies that are consistent with recruitment of slow fibers. Collectively, these findings support the hypothesis and suggest that sloths may selectively recruit smaller, fast motor units for suspensory postures but have the ability to offset the cost of force production by recruitment of large, slow motor units during locomotion.

Neural and muscular determinants of maximal rate of force development.

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.NEW & NOTEWORTHY 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.

Factors contributing to sag in unfused tetanic contractions of fast motor units in rat medial gastrocnemius

The sag phenomenon can be observed in fast motor units (MUs) as a transitional decline in force during unfused tetanic contractions; however, its mechanisms are poorly understood. The study aimed to identify in the rat muscle factors that contribute to sag in two types of fast MUs: fast fatigable (FF) and fast resistant to fatigue (FR). First, we performed mathematical decomposition of sagging tetanic contractions of FF and FR MUs into twitch-like responses to consecutive stimuli. This process indicated an increase in the amplitudes of a few initial responses (up to the 2nd–3rd for FF and up to the 2nd–7th for FR MUs), followed by a decrease in the amplitudes of later responses. In comparison to the first twitch, the relative increase in force amplitudes of the several subsequent decomposed responses was smaller, and their contraction and relaxation times were shorter for FF than for FR units, which corresponded to observed differences in their sag profiles. Additionally, after occlusion of the blood circulation, sag disappeared, but it reappeared after restoration of the blood supply. This indicates that the presence of sag depends on the proper circulation in the muscle.

Does a two-element muscle model offer advantages when estimating ankle plantar flexor forces during human cycling?

Traditional Hill-type muscle models, parameterized using high-quality experimental data, are often “too weak” to reproduce the joint torques generated by healthy adults during rapid, high force tasks. This study investigated whether the failure of these models to account for different types of motor units contributes to this apparent weakness; if so, muscle-driven simulations may rely on excessively high muscle excitations to generate a given force. We ran a series of forward simulations that reproduced measured ankle mechanics during cycling at five cadences ranging from 60 to 140 RPM. We generated both “nominal” simulations, in which an abstract ankle model was actuated by a 1-element Hill-type plantar flexor with a single contractile element (CE), and “test” simulations, in which the same model was actuated by a 2-element plantar flexor with two CEs that accounted for the force-generating properties of slower and faster motor units. We varied the total excitation applied to the 2-element plantar flexor between 60 and 105% of the excitation from each nominal simulation, and we varied the amount distributed to each CE between 0 and 100% of the total. Within this test space, we identified the excitation level and distribution, at each cadence, that best reproduced the plantar flexor forces generated in the nominal simulations. Our comparisons revealed that the 2-element model required substantially less total excitation than the 1-element model to generate comparable forces, especially at higher cadences. For instance, at 140 RPM, the required excitation was reduced by 23%. These results suggest that a 2-element model, in which contractile properties are “tuned” to represent slower and faster motor units, can increase the apparent strength and perhaps improve the fidelity of simulations of tasks with varying mechanical demands.

Modifying motor unit territory placement in the Fuglevand model.

The Fuglevand model is often used to address challenging questions in neurophysiology; however, there are elements of the neuromuscular system unaccounted for in the model. For instance, in some muscles, slow and fast motor units (MUs) tend to reside deep and superficially in the muscle, respectively, necessarily altering the development of surface electromyogram (EMG) power during activation. Thus, the objective of this study was to replace the randomized MU territory (MUT) placement algorithm in the Fuglevand model with an optimized method capable of reflecting these observations. To accomplish this, a weighting term was added to a previously developed optimization algorithm to encourage regionalized MUT placement. The weighting term consequently produced significantly different muscle fibre type content in the deep and superficial portions of the muscle. The relation between simulated EMG and muscle force was found to be significantly affected by regionalization. These changes were specifically a function of EMG power, as force was unaffected by regionalization. These findings suggest that parameterizing MUT regionalization will allow the model to produce a larger variety of EMG-force relations, as is observed physiologically, and could potentially simulate the loss of specific MU types as observed in ageing and clinical populations.

Role of histidyl dipeptides in contractile function of fast and slow motor units in rat skeletal muscle.

The physiological role of the muscle histidyl dipeptides carnosine and anserine in contractile function of various types of muscle fibers in vivo is poorly understood. Ten adult male Wistar rats were randomly assigned to two groups: control and supplemented for 10 wk with beta-alanine, the precursor of carnosine (∼640 mg·kg body wt(-1)·day(-1)). Thereafter, contractile properties and fatigability of isolated fast fatigable (FF), fast resistant to fatigue (FR), and slow motor units (MUs) from the medial gastrocnemius were determined in deeply anaesthetized animals. The fatigue resistance was tested with a 40-Hz fatigue protocol followed by a second protocol at 40 Hz in fast and 20 Hz in slow units. In the supplemented rats, histidyl dipeptide concentrations significantly increased (P < 0.05) by 25% in the red portion of the gastrocnemius, and carnosine increased by 94% in the white portion. The twitch force of FF units and maximum tetanic force of FR units were significantly increased (P < 0.05), and the half-relaxation time was prolonged in slow units (P < 0.05). FF units showed less fatigue during the first 10 s, and FR units showed higher forces between 10 and 60 s during the 40-Hz fatigue test. In slow units, forces declined less during the first 60 s of the 20-Hz test. In conclusion, this in vivo experiment demonstrates that an elevation in muscle histidyl dipeptide content elicits beneficial changes in MU contractile characteristics and fatigue resistance. Carnosine and anserine seem to play an important yet divergent role in various MUs.