Unfused Tetani Research Articles

Estimating the neural spike train from an unfused tetanic signal of low-threshold motor units using convolutive blind source separation

BackgroundIndividual motor units have been imaged using ultrafast ultrasound based on separating ultrasound images into motor unit twitches (unfused tetanus) evoked by the motoneuronal spike train. Currently, the spike train is estimated from the unfused tetanic signal using a Haar wavelet method (HWM). Although this ultrasound technique has great potential to provide comprehensive access to the neural drive to muscles for a large population of motor units simultaneously, the method has a limited identification rate of the active motor units. The estimation of spikes partly explains the limitation. Since the HWM may be sensitive to noise and unfused tetanic signals often are noisy, we must consider alternative methods with at least similar performance and robust against noise, among other factors.ResultsThis study aimed to estimate spike trains from simulated and experimental unfused tetani using a convolutive blind source separation (CBSS) algorithm and compare it against HWM. We evaluated the parameters of CBSS using simulations and compared the performance of CBSS against the HWM using simulated and experimental unfused tetanic signals from voluntary contractions of humans and evoked contraction of rats. We found that CBSS had a higher performance than HWM with respect to the simulated firings than HWM (97.5 ± 2.7 vs 96.9 ± 3.3, p < 0.001). In addition, we found that the estimated spike trains from CBSS and HWM highly agreed with the experimental spike trains (98.0% and 96.4%).ConclusionsThis result implies that CBSS can be used to estimate the spike train of an unfused tetanic signal and can be used directly within the current ultrasound-based motor unit identification pipeline. Extending this approach to decomposing ultrasound images into spike trains directly is promising. However, it remains to be investigated in future studies where spatial information is inevitable as a discriminating factor.

Estimation of contractile parameters of successive twitches in unfused tetanic contractions of single motor units – A proof-of-concept study using ultrafast ultrasound imaging in vivo

During a voluntary contraction, motor units (MUs) fire a train of action potentials, causing summation of the twitch forces, resulting in fused or unfused tetanus. Twitches have been important in studying whole-muscle contractile properties and differentiation between MU types. However, there are still knowledge gaps concerning the voluntary force generation mechanisms. Current methods rely on the spike-triggered averaging technique, which cannot track changes in successive twitches’ properties in response to individual neural firings. This study proposes a method that estimates successive twitches contractile parameters of single MUs during low force voluntary isometric contractions in human biceps brachii. We used a previously developed ultrafast ultrasound imaging method to estimate unfused tetanic activity signals of single MUs. A twitch decomposition model was used to decompose unfused tetanic activity signals into individual twitches. This study found that the contractile parameters varied within and across MUs. There was an association between the inter-spike interval and the contraction time (r = 0.49,p < 0.001) and the half-relaxation time (r = 0.58,p < 0.001), respectively. The method shows the proof-of-concept to study MU contractile properties of individual twitches in vivo, which can provide further insights into the force generation mechanisms of voluntary contractions and response to individual neural discharges.

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.

A model of the rat medial gastrocnemius muscle based on inputs to motoneurons and on an algorithm for prediction of the motor unit force.

The muscle force is the sum of forces of multiple motor units (MUs), which have different contractile properties. During movements, MUs develop unfused tetani, which result from summation of twitch-shape responses to individual stimuli, which are variable in amplitude and duration. The aim of the study was to develop a realistic muscle model that would integrate previously developed models of MU contractions and an algorithm for the prediction of tetanic forces. The proposed model of rat medial gastrocnemius muscle is based on physiological data: excitability and firing frequencies of motoneurons, contractile properties, and the number and proportion of MUs in the muscle. The MU twitches were modeled by a six-parameter analytical function. The excitability of motoneurons was modeled according to a distribution of their rheobase currents measured experimentally. Processes of muscle force regulation were modeled according to a common drive hypothesis. The excitation signal to motoneurons was modeled by two form types: triangular and trapezoid. The discharge frequencies of MUs, calculated individually for each MU, corresponded to those recorded for rhythmic firing of motoneurons. The force of the muscle was calculated as the sum of all recruited MUs. Participation of the three types of MUs in the developed muscle force was presented at different levels of the excitation signal to motoneurons. The model appears highly realistic and open for input data from various skeletal muscles with different compositions of MU types. The results were compared with three other models with different distribution of the input parameters. NEW & NOTEWORTHY The proposed mathematical model of rat medial gastrocnemius muscle is highly realistic because it is based strictly on experimentally determined motor unit contractile parameters and motoneuron properties. It contains the actual number and proportion of motor units and takes into consideration their different contributions to the whole muscle force, depending on the level of the excitation signal. The model is open for input data from other muscles, and additional physiological parameters can also be included.

The sag response in human muscle contraction.

We examined how muscle length and time between stimuli (inter-pulse interval, IPI) influence declines in force (sag) seen during unfused tetani in the human adductor pollicis muscle. A series of 16-pulse contractions were evoked with IPIs between 1 × and 5 × the twitch time to peak tension (TPT) at large (long muscle length) and small (short muscle length) thumb adduction angles. Unfused tetani were mathematically deconstructed into a series of overlapping twitch contractions to examine why sag exhibits length- and IPI-dependencies. Across all IPIs tested, sag was 62% greater at short than long muscle length, and sag increased as IPI was increased at both muscle lengths. Force attributable to the second stimulus increased as IPI was decreased. Twitch force declined from maximal values across all IPI tested, with the greatest reductions seen at short muscle length and long IPI. At IPI below 2 × TPT, the twitch with highest force occurred earlier than the peak force of the corresponding unfused tetani. Contraction-induced declines in twitch duration (TPT + half relaxation time) were only observed at IPI longer than 1.75 × TPT, and were unaffected by muscle length. Sag is an intrinsic feature of healthy human adductor pollicis muscle. The length-dependence of sag is related to greater diminution of twitch force at short relative to long muscle length. The dependence of sag on IPI is related to IPI-dependent changes in twitch duration and twitch force, and the timing of peak twitch force relative to the peak force of the associated unfused tetanus.

Can inorganic phosphate explain sag during unfused tetanic contractions of skeletal muscle?

We test the hypothesis that cytosolic inorganic phosphate (Pi) can account for the contraction‐induced reductions in twitch duration which impair summation and cause force to decline (sag) during unfused tetanic contractions of fast‐twitch muscle. A five‐state model of crossbridge cycling was used to simulate twitch and unfused tetanic contractions. As Pi concentration ([Pi]) was increased from 0 to 30 mmol·L−1, twitch duration decreased, with progressive reductions in sensitivity to Pi as [Pi] was increased. When unfused tetani were simulated with rising [Pi], sag was most pronounced when initial [Pi] was low, and when the magnitude of [Pi] increase was large. Fast‐twitch extensor digitorum longus (EDL) muscles (sag‐prone, typically low basal [Pi]) and slow‐twitch soleus muscles (sag‐resistant, typically high basal [Pi]) were isolated from 14 female C57BL/6 mice. Muscles were sequentially incubated in solutions containing either glucose or pyruvate to create typical and low Pi environments, respectively. Twitch duration was greater (P < 0.05) in pyruvate than glucose in both muscles. Stimuli applied at intervals approximately three times the time to peak twitch tension resulted in sag of 35.0 ± 3.7% in glucose and 50.5 ± 1.4% in pyruvate in the EDL (pyruvate > glucose; P < 0.05), and 3.9 ± 0.3% in glucose and 37.8 ± 2.7% in pyruvate in the soleus (pyruvate > glucose; P < 0.05). The influence of Pi on crossbridge cycling provides a tenable mechanism for sag. Moreover, the low basal [Pi] in fast‐twitch relative to slow‐twitch muscle has promise as an explanation for the fiber‐type dependency of sag.

A General Mathematical Algorithm for Predicting the Course of Unfused Tetanic Contractions of Motor Units in Rat Muscle.

An unfused tetanus of a motor unit (MU) evoked by a train of pulses at variable interpulse intervals is the sum of non-equal twitch-like responses to these stimuli. A tool for a precise prediction of these successive contractions for MUs of different physiological types with different contractile properties is crucial for modeling the whole muscle behavior during various types of activity. The aim of this paper is to develop such a general mathematical algorithm for the MUs of the medial gastrocnemius muscle of rats. For this purpose, tetanic curves recorded for 30 MUs (10 slow, 10 fast fatigue-resistant and 10 fast fatigable) were mathematically decomposed into twitch-like contractions. Each contraction was modeled by the previously proposed 6-parameter analytical function, and the analysis of these six parameters allowed us to develop a prediction algorithm based on the following input data: parameters of the initial twitch, the maximum force of a MU and the series of pulses. Linear relationship was found between the normalized amplitudes of the successive contractions and the remainder between the actual force levels at which the contraction started and the maximum tetanic force. The normalization was made according to the amplitude of the first decomposed twitch. However, the respective approximation lines had different specific angles with respect to the ordinate. These angles had different and non-overlapping ranges for slow and fast MUs. A sensitivity analysis concerning this slope was performed and the dependence between the angles and the maximal fused tetanic force normalized to the amplitude of the first contraction was approximated by a power function. The normalized MU contraction and half-relaxation times were approximated by linear functions depending on the normalized actual force levels at which each contraction starts. The normalization was made according to the contraction time of the first contraction. The actual force levels were calculated initially from the recorded tetanic curves and subsequently from the modeled curves obtained from the summation of all models of the preceding contractions (the so called “full prediction”). The preciseness of the prediction was verified by two coefficients estimating the error between the modeled and the experimentally recorded curves. The proposed approach was tested for 30 MUs from the database and for three additional MUs, not included in the initial set. It was concluded that this general algorithm can be successfully used for modeling of a unfused tetanus course of a single MU of fast and slow type.

Prolonged activity evokes potentiation and the "sag" phenomenon in slow motor units of rat soleus.

Slow motor units (MUs) have no sag in their unfused tetani. This study in anesthetized rats shows that the sag can be observed in slow soleus MUs after prolonged activity. Twitches and unfused tetanic contractions were recorded from male (n=35) and female (n=39) MUs before and after the four minutes of the fatigue test (trains of 13 pulses at 40 Hz repeated every second). After this activity twitch contractions potentiated and a shift in the steep part of the force-frequency curve towards lower frequencies was observed in both sexes. Initially no sag was visible in unfused tetani, but after the fatigue test the phenomenon was observed in 77% of male, while in 13% of female MUs, the result consistent with the previously reported higher content of IIa myosin and faster contraction of MUs in male soleus. The decomposition of tetani with sag into trains of twitch-shape responses to consecutive stimuli revealed higher forces of initial decomposed twitches than later. The revealed alterations the force development due to long-lasting activation of slow MUs were sex-related and more pronounced in male soleus.

Inappropriate interpretation of surface EMG signals and muscle fiber characteristics impedes understanding of the control of neuromuscular function

as the final common pathway from the nervous system to muscle, the motor unit transmits an activation signal generated by the nervous system to engage the contractile proteins and produce the muscle forces needed for reflex responses, automatic behaviors, and voluntary actions ([11][1]). The net

Phenol increases intracellular [Ca2+] during twitch contractions in intactXenopusskeletal myofibers

Phenol is a neurolytic agent used for management of spasticity in patients with either motoneuron lesions or stroke. In addition, compounds that enhance muscle contractility (i.e., polyphenols, etc.) may affect muscle function through the phenol group. However, the effects of phenol on muscle function are unknown, and it was, therefore, the purpose of the present investigation to examine the effects of phenol on tension development and Ca(2+) release in intact skeletal muscle fibers. Dissected intact muscle fibers from Xenopus laevis were electrically stimulated, and cytosolic Ca(2+) concentration ([Ca(2+)](c)) and tension development were recorded. During single twitches and unfused tetani, phenol significantly increased [Ca(2+)](c) and tension without affecting myofilament Ca(2+) sensitivity. To investigate the phenol effects on Ca(2+) channel/ryanodine receptors, single fibers were treated with different concentrations of caffeine in the presence and absence of phenol. Low concentrations of phenol significantly increased the caffeine sensitivity (P < 0.01) and reduced the caffeine concentrations necessary to produce nonstimulated contraction (contracture). However, at high phenol concentrations, caffeine did not increase tension or Ca(2+) release. These results suggest that phenol affects the ability of caffeine to release Ca(2+) through an effect on the ryanodine receptors, or on the sarcoplasmic reticulum Ca(2+) pump. During tetanic contractions inducing fatigue, phenol application decreased the time to fatigue. In summary, phenol increases intracellular [Ca(2+)] during twitch contractions in muscle fibers without altering myofilament Ca(2+) sensitivity and may be used as a new agent to study skeletal muscle Ca(2+) handling.

Summation of motor unit forces in rat medial gastrocnemius muscle

The summation of contractile forces of motor units (MUs) was analyzed by comparing the recorded force during parallel stimulation of two and four individual MUs or four groups of MUs to the algebraic sum of their individual forces. Contractions of functionally-isolated single MUs of the medial gastrocnemius muscle were evoked by electrical stimulation of thin filaments of the split L5 or L4 ventral roots of spinal nerves. Additionally, contractions of large groups of MUs were evoked by stimuli delivered to four parts of the divided L5 ventral root. Single twitches, 40 Hz unfused tetani, and 150 Hz fused maximum tetani were recorded. In these experimental situations the summation was more effective for unfused tetani than for twitches or maximum tetani. The results obtained for pairs of MUs were highly variable (more- or less-than-linear summation), but coactivation of more units led to progressively weaker effects of summation, which were usually less-than-linear in comparison to the algebraic sums of the individual forces. The variability of the results highlights the importance of the structure of the muscle and the architecture of its MUs. Moreover, the simultaneous activity of fast and slow MUs was considerably more effective than that of two fast units.

Experimentally verified mathematical approach for the prediction of force developed by motor units at variable frequency stimulation patterns

During normal daily activity, muscle motor units (MUs) develop unfused tetanic contractions evoked by trains of motoneuronal firings at variable interpulse intervals (IPIs). The mechanical responses of a MU to successive impulses are not identical. The aim of this study was to develop a mathematical approach for the prediction of each response within the tetanus as well as the tetanic force itself. Experimental unfused tetani of fast and slow rat MUs, evoked by trains of stimuli at variable IPIs, were decomposed into series of twitch-shaped responses to successive stimuli using a previously described algorithm. The relationships between the parameters of the modeled twitches and the tetanic force level at which the next response begins were examined and regression equations were derived. Using these equations, profiles of force for the same and different stimulation patterns were mathematically predicted by summating modeled twitches. For comparison, force predictions were made by the summation of twitches equal to the first one. The recorded and the predicted tetanic forces were compared. The results revealed that it is possible to predict tetanic force with high accuracy by using regression equations. The force predicted in this way was much closer to the experimental record than the force obtained by the summation of equal twitches, especially for slow MUs. These findings are likely to have an impact on the development of realistic muscle models composed of MUs, and will assist our understanding of the significance of the neuronal code in motor control and the role of biophysical processes during MU contractions.

Time-related changes of motor unit properties in the rat medial gastrocnemius muscle after the spinal cord injury. II. Effects of a spinal cord hemisection

The contractile properties of motor units (MUs) were investigated in the medial gastrocnemius (MG) muscle in rats after the spinal cord hemisection at a low thoracic level. Hemisected animals were divided into 4 groups: 14, 30, 90 and 180 days after injury. Intact rats formed a control group. The mass of the MG muscle did not change significantly after spinal cord hemisection, hind limb locomotor pattern was almost unchanged starting from two weeks after injury, but contractile properties of MUs were however altered. Contraction time (CT) and half-relaxation time (HRT) of MUs were prolonged in all investigated groups of hemisected rats. The twitch-to-tetanus ratio (Tw/Tet) of fast MUs after the spinal cord hemisection increased. For slow MUs Tw/Tet values did not change in the early stage after the injury, but significantly decreased in rats 90 and 180 days after hemisection. As a result of hemisection the fatigue resistance especially of slow and fast resistant MU types was reduced, as well as fatigue index (Fat I) calculated for the whole examined population of MUs decreased progressively with the time. After spinal cord hemisection a reduced number of fast MUs presented the sag at frequencies 30 and 40 Hz, however more of them revealed sag in 20 Hz tetanus in comparison to control group. Due to considerable changes in twitch contraction time and disappearance of sag effect in unfused tetani of some MUs in hemisected animals, the classification of MUs in all groups of rats was based on the 20 Hz tetanus index (20 Hz Tet I) but not on the standard criteria usually applied for MUs classification. MU type differentiations demonstrated some clear changes in MG muscle composition in hemisected animals consisting of an increase in the proportion of slow MUs (likely due to an increased participation of the studied muscle in tonic antigravity activity) together with an increase in the percentage of fast fatigable MUs.

Time-related changes of motor unit properties in the rat medial gastrocnemius muscle after spinal cord injury. I. Effects of total spinal cord transection

The contractile properties of motor units (MUs) were electrophysiologically investigated in the medial gastrocnemius (MG) muscle in 17 Wistar three-month-old female rats: 14, 30, 90 and 180 days after the total transection of the thoracic spinal cord and compared to those in intact (control) rats. A sag phenomenon, regularly observed in unfused tetani of fast units in intact animals at 40 Hz stimulation, almost completely disappeared in spinal rats. Therefore, the MUs of intact and spinal rats were classified as fast or slow types basing on 20 Hz tetanus index, the value of which was lower or equal 2.0 for fast and higher than 2.0 for slow MUs. The MUs composition of MG muscle changed with time after the spinal cord transection: an increasing proportion of fast fatigable (FF) units starting one month after injury and a disappearance of slow (S) units within the three months were observed. In all MUs investigated the twitch contraction and half-relaxation time were significantly prolonged after injury ( p < 0.01, Mann–Whitney U-test). Moreover, a decrease of the fatigue index for fast resistant (FR) and slow MUs was observed in subsequent groups of spinal rats. No significant changes were found between twitch forces in all MU types of spinal animals ( p > 0.05). However, due to a decrease of the maximal tetanic force, a significant rise of the twitch-to-tetanus ratio of all MUs in spinal rats was detected ( p < 0.01). The considerable reduction of ability to potentiate the force was noticed for fast, especially FF type MUs. In conclusion, the spinal cord transection leads to changes in the proportion of the three MU types in rat MG muscle. The majority of changes in MUs’ contractile properties were developed progressively with time after the spinal cord injury. However, the most intensive alterations of twitch-time parameters were observed in rats one month after the transection.

Control of myotube contraction using electrical pulse stimulation for bio-actuator

The contractility of tissue-engineered muscle on the application of electrical signals is required for the development of bio-actuators and for muscle tissue regeneration. Investigations have already reported on the contraction of myotubes differentiated from myoblasts and the construction of tissue-engineered skeletal muscle using electrical pulses. However, the relationship between myotube contraction and electrical pulses has not been quantitatively evaluated. We quantitatively investigated the effect of electrical pulse frequency on the excitability of myotubes and developed bio-actuators made of tissue-engineered skeletal muscle. C2C12 cells were seeded on a collagen-coated dish and in collagen gel and were cultured in Dulbecco's modified Eagle's medium (DMEM) containing 10% fetal bovine serum and antibiotics. When the cells reached confluence or after 2 days in culture, the medium was shifted to DMEM containing 7% horse serum to allow them to differentiate to C2C12 myotubes. We electrically stimulated the myotubes and tissue-engineered skeletal muscle, and contractions were observed under a microscope. The myotubes contracted synchronously with electrical pulses between 0.5 and 5 Hz and unfused tetanus was generated at 10 Hz. The contractile performance of tissue-engineered skeletal muscle made of collagen gel and C2C12 was similar to that of the myotubes. Both the rheobase and chronaxie of the myotubes were lowest when the electric field was applied parallel to the myotube axis, and the values were 8.33 +/- 2.78 mA and 1.19 +/- 0.38 ms, respectively. The motion of C2C12 myotube contraction depended on the pulse frequency and showed anisotropy in the electric field. These results suggest that a tissue-engineered bio-actuator may be controlled using electrical signals.

Model-generated decomposition of unfused tetani of motor units evoked by random stimulation

Unfused tetani of motor units (MUs) evoked by stimulation at variable interpulse intervals at mean frequencies of 20, 25, 33, 40 and 50 Hz were studied using ten functionally isolated fast-type MUs from the medial gastrocnemius muscle of adult Wistar rats. A previously proposed algorithm and computer program for mathematical decomposition of unfused tetani into a series of twitches, representing responses to individual pulses, were used. Analysis of the parameters of the decomposed twitches showed considerable variability in force of successive contractions. These twitches were extremely variable with up to 2-fold higher forces and longer contraction times than a single twitch evoked by one stimulus. However, when the stimulation frequency was decreased, the decomposed twitches became similar to the single twitch with respect to amplitude and contraction time. It was found that the basic contractile parameters of decomposed twitches could be predicted with high accuracy on the basis of the tetanus force level at which the next contraction begins. This analysis of the parameters of decomposed twitches demonstrated that the contractile responses of the muscle fibers to successive action potentials generated by motoneurons are highly variable and depend on the previous MU state.

Variability of successive contractions subtracted from unfused tetanus of fast and slow motor units

Stimulation of motor units (MUs) with repeated pulses evokes tetanic contractions, which consist of overlapping mechanical responses. The summation of these responses into tetanus is a nonlinear process due to the dynamic changes in the amplitudes and time parameters of the successive components. In order to study these changes, two MUs (one fast and one slow) of rat medial gastrocnemius muscle were stimulated with a progressively increasing number of pulses, from one ( i = 1) to sixteen ( i = 16) at a frequency of 15 Hz for the slow MU and 60 Hz for the fast MU. The individual responses were calculated by subtracting the ( i)th from the ( i + 1)th tetanus recording. The contractions obtained following subtraction were modeled using a novel 6-parameter analytical function. The main conclusions of this study are (1) the newly presented analytical function is able to precisely describe the variable shape of all subtracted experimental contractions; (2) the shapes of successive contractions are variable and the subtracted contractions differ from the individual twitches; (3) as the pulse number increases, the parameters of the subtracted contractions change in a different manner for the slow and fast MUs: for the slow MU, the maximal forces and the time parameters increase considerably up to the 4th response, after which they remain nearly constant or show only a slight increase; for the fast MU, the maximal forces and durations also increase, whereas the remaining time parameters initially increase and then maintain a constant level or decrease, which explains the sag phenomenon visible in the unfused tetanus of fast MUs.

Changes in contractile properties of motor units of the rat medial gastrocnemius muscle after spinal cord transection

The effects of complete transection of the spinal cord at the level of Th9/10 on contractile properties of the motor units (MUs) in the rat medial gastrocnemius (MG) muscle were investigated. Our results indicate that 1 month after injury the contraction time (time-to-peak) and half-relaxation time were prolonged and the maximal tetanic force in most of the MUs in the MG muscle of spinal rats was reduced. The resistance to fatigue also decreased in most of the MUs in the MG of spinal animals. Moreover, the post-tetanic potentiation of twitches in MUs diminished after spinal cord transection. Criteria for the division of MUs into three types, namely slow (S), fast fatigue resistant (FR) and fast fatigable (FF), applied in intact animals, could not be directly used in spinal animals owing to changes in contractile properties of MUs. The 'sag' phenomenon observed in unfused tetani of fast units in intact animals essentially disappeared in spinal rats and it was only detected in few units, at low frequencies of stimulation only. Therefore, the MUs in spinal rats were classified as fast or slow on the basis of an adjusted borderline of 20 ms, instead of 18 ms as in intact animals, owing to a slightly longer contraction time of those fast motor units with the 'sag'. We conclude that all basic contractile properties of rat motor units in the medial gastrocnemius muscle are significantly changed 1 month after complete spinal cord transection, with the majority of motor units being more fatigable and slower than those of intact rats.

Changes in the time course of potentiated and fatigued contractions of fast motor units in rat muscle.

The contraction and relaxation times of the twitches and the last contractions within 32 unfused tetani of FF and 27 unfused tetani of FR motor units in the rat medial gastrocnemius muscle were studied during prolonged activity. The pattern of the MU stimulation included single pulses (to evoke twitches) and series of three trains of stimuli at 40, 50 and 60 Hz (to evoke unfused tetani), repeated 30 times. The analysis concerned changes of force and time parameters at the beginning of activity, during the potentiation and then during the fatigue. It was found that changes of force during the potentiation and the fatigue were mainly accompanied by changes in the course of relaxation. The significant prolongation of the half-relaxation time during the potentiation of either twitches or unfused tetani was revealed in both types of fast MU. The twitch contraction time did not change markedly, whereas significantly shortened in the last contractions of unfused tetani during the potentiation. These changes of time parameters correlated to the increase of the fusion degree. During the fatigue, the time parameters shortened, however, changes of the half-relaxation times were remarkably higher. The shortening of relaxation was responsible for the decrease of the fusion degree. Changes of the fusion index exceeding 0.75 during the potentiation or decreasing below this value during the fatigue, were accompanied by respective appearance or disappearance of the biphasic relaxation.

Modeling of summation of individual twitches into unfused tetanus for various types of rat motor units

Repeated stimulation of motor units (MUs) causes an increase of the force output that cannot be explained by linear summation of equal twitches evoked by the same stimulation pattern. To explain this phenomenon, an algorithm for reconstructing the individual twitches, that summate into an unfused tetanus is described in the paper. The algorithm is based on an analytical function for the twitch course modeling. The input parameters of this twitch model are lead time, contraction and half-relaxation times and maximal force. The measured individual twitches and unfused tetani at 10, 20, 30 and 40 Hz stimulation frequency of three rat motor units (slow, fast resistant to fatigue and fast fatigable) are processed. It is concluded that: (1) the analytical function describes precisely the course of individual twitches; (2) the summation of equal twitches does not follow the results from the experimentally measured unfused tetani, the differences depend on the type of the MU and are bigger for higher values of stimulation frequency and fusion index; (3) the reconstruction of individual twitches from experimental tetanic records can be successful if the tetanus is feebly fused (fusion index up to 0.7); (4) both the maximal forces and time parameters of individual twitches subtracted from unfused tetani change and influence the course of each tetanus. A discrepancy with respect to the relaxation phase was observed between experimental results and model prediction for tetani with fusion index exceeding 0.7. This phase was predicted longer than the experimental one for better fused tetani. Therefore, a separate series of physiological experiments and then, more complex model are necessary for explanation of this distinction.