Collateral nerve sprouting and twitch forces of single motor units in conditions with partial denervation in man

Single motor units (MUs) were studied in the first dorsal interosseus muscle of 7 patients with slight partial denervation and in 10 controls. MU action potentials were recorded using the macro-EMG technique and their size was taken to assess collateral nerve sprouting. Simultaneously, the muscle force was monitored to determine the voluntary recruitment thresholds of the MUs and spike-triggered-averaging was applied to measure their twitch forces. At comparable force recruitment thresholds, both macro-EMG potentials and twitch forces were increased in the patients. We conclude, that collateral nerve sprouting increases MU force and can compensate for MU loss.     

Human motor unit activity during concentric and eccentric movements

Single motor units (MUs) activity was investigated in human m. biceps brachii during movements against an elastic load. A total of sixty-five MUs were studied by means of subcutaneously placed fine-wire branched electrodes. Subjects were asked to perform active shortening and lengthening of the muscle with approximately constant velocities at two different speeds--slow and fast. Both recruitment (RT) and decruitment (DT) thresholds of MU were found to be lower in movement with higher velocity. The recruitment order of MUs was approximately one and the same during concentric movements with a different but constant velocity. The firing onset of MUs is organized so that the peak of the first twitch contraction occurs at approximately the same force level irrespective of how fast the movement is. In contrast, during the eccentric movements the peak of the last twitch contraction of MU occurs at different torque levels depending on the velocity. The decruitment of the MUs during eccentric movement was in a reverse order to their recruitment during concentric movements. Generally, at one and the same velocity the RT of a given MU was lower than DT. Nevertheless, the peaks of the first and the last twitch contractions during concentric and eccentric movements with one and the same velocity occurred at approximately one and the same torque level.     

Physiological properties of the motor units of the wrist extensor muscles in man

Summary The physiological properties of 355 motor units (MUs) recorded in the extensor carpi radialis muscles were studied in 34 healthy human subjects during isometric contractions. MU selective twitches were educed from the whole muscle force using the spike-triggered averaging method. The twitch contraction times and twitch forces were measured. From these data it was attempted to estimate the distribution of fast and slow MUs in the muscles studied. MU recruitment thresholds were systematically measured during stereotyped slow ramp contractions (force increase=0.25 N·s^-1). Degrees of correlation between contraction times, twitch forces and recruitment thresholds were pair analysed by computing simple regression curves and correlation coefficients. The degrees of correlation were compared between 245 MUs recorded in 34 subjects and 66 MUs recorded in a single subject. Analysis of the instantaneous discharge frequency of 132 MUs showed the existence of a remarkable degree of correlation (correlation coefficient, r=-0.75) between the frequency rise times (discharge onset to maximal frequency) and the MU twitch contraction times; i.e., the frequency rise times increase when the twitch contraction times decrease. The possibility that muscle contraction may be differentially modulated on the basis of this discharge property of the MUs is discussed. The results are compared to previous data and the limitations of the spike-triggered averaging method applied to long muscles in man are extensively discussed.     

Motor unit recruitment order during voluntary and electrically induced contractions in the tibialis anterior

 The recruitment order of motor units (MU) was compared during voluntary and electrically induced contractions. With the use of spike-triggered averaging, a total of 302 MUs with recruitment thresholds ranging from 1% to 88% of maximal voluntary contraction were recorded in the human tibialis anterior muscle in five subjects. The mean (±SD) MU force was 98.3±93.3 mN (mean torque 16.8±15.9 mNm) and the mean contraction time (CT) 46.2±12.7 ms. The correlation coefficients (r) between MU twitch force and CT versus the recruitment threshold in voluntary contractions were +0.68 and –0.38 (P<0.001), respectively. In voluntary contractions, MUs were recruited in order of increasing size except for only 6% of the cases; whereas, during transcutaneous electrical stimulation (ES) at the muscle motor point, MU pairs showed a reversal of recruitment order in 28% and 35% of the observations, respectively, when the pulse durations were 1.0 ms or 0.1 ms. This recruitment reversal during ES was not related to the magnitude of the difference in voluntary recruitment thresholds between MUs. It is concluded that if the reversal of MU recruitment observed during ES is biophysically controlled by differences in their nerve axon input impedance, in percutaneous stimulation at the motor point, other factors such as the size and the morphological organisation of the axonal branches can also influence the order of activation. Received: 24 May 1996 / Accepted: 30 September 1996     

Mechanomyograms recorded during evoked contractions of single motor units in the rat medial gastrocnemius muscle

Acoustic phenomena accompanying contractions of single motor units (MUs) have previously received little attention. Therefore, in the present study, the mechanomyographic (MMG) signals during evoked contractions of single MUs have been recorded from the medial gastrocnemius muscle of the rat. A piezoelectric transducer immersed in a paraffin-oil pool was used for the measurement of these signals. Muscle fibre action potentials, tension and MMG were recorded in parallel during twitch (the weakest) and fused tetanic (the strongest) MU contractions. It was observed that the onset of the MMG signals was coincident with the beginning of the increase in tension for both the twitch and tetanus. Weaker MMG signals than those accompanying the beginning of the first phase of the fused tetanus were seen during the beginning of the relaxation after tetanic contraction. During contraction and relaxation, MMG signals were characterised by the reverse-direction of the first extreme phase, positive and negative, respectively. No MMG signals were observed when the tension was constant during the fused tetanus. The amplitude of MMG signals was correlated with both the tension increase and the velocity of tension increase during both the twitch and the fused tetanus. The strongest MUs (fast fatiguable) generated MMG signals of the highest amplitude. MMG signals were not detected for some of the weakest slow MUs (with tension increases of < or = 2 mN). These results indicate a strong correlation between the MMG and the change of tension. Therefore, we believe that MMG signals are generated by muscle deformation that occurs during the contraction of MU muscle fibres. We conclude that the number of active muscle fibres, their topography, and their localisation in relation to the muscle surface (which is variable for different types of MUs) influence these MMG phenomena.     

Mechanomyograms recorded during evoked contractions of single motor units in the rat medial gastrocnemius muscle

Acoustic phenomena accompanying contractions of single motor units (MUs) have previously received little attention. Therefore, in the present study, the mechanomyographic (MMG) signals during evoked contractions of single MUs have been recorded from the medial gastrocnemius muscle of the rat. A piezoelectric transducer immersed in a paraffin-oil pool was used for the measurement of these signals. Muscle fibre action potentials, tension and MMG were recorded in parallel during twitch (the weakest) and fused tetanic (the strongest) MU contractions. It was observed that the onset of the MMG signals was coincident with the beginning of the increase in tension for both the twitch and tetanus. Weaker MMG signals than those accompanying the beginning of the first phase of the fused tetanus were seen during the beginning of the relaxation after tetanic contraction. During contraction and relaxation, MMG signals were characterised by the reverse-direction of the first extreme phase, positive and negative, respectively. No MMG signals were observed when the tension was constant during the fused tetanus. The amplitude of MMG signals was correlated with both the tension increase and the velocity of tension increase during both the twitch and the fused tetanus. The strongest MUs (fast fatiguable) generated MMG signals of the highest amplitude. MMG signals were not detected for some of the weakest slow MUs (with tension increases of ≤2 mN). These results indicate a strong correlation between the MMG and the change of tension. Therefore, we believe that MMG signals are generated by muscle deformation that occurs during the contraction of MU muscle fibres. We conclude that the number of active muscle fibres, their topography, and their localisation in relation to the muscle surface (which is variable for different types of MUs) influence these MMG phenomena. Accepted: 29 May 2000     

Tetanic potentiation in motor units of rat medial gastrocnemius

Tetanic potentiation is a phenomenon, which expresses the ability of a motor unit (MU) to increase its force output in tetanic contractions above that predicted with an assumption of algebraic summation of single twitch responses. To quantify tetanic potentiation, a coefficient TPC (tetanic potentiation coefficient) was defined as a ratio of the areas below tetanic force recording corresponding to the single stimulus contribution and that of the single twitch. Single MUs (27 Slow, 71 Fast, Fatigue Resistant, and 47 Fast, Fatigable) were isolated from the rat medial gastrocnemius muscle (MG) by ventral root splitting. TPC value was rate-dependent, with the maximum TPCmax at a certain optimal rate. The largest values of the TPCmax were obtained in the weakest and most fatigue resistant (S and weak FR) MUs. The different manifestation of staircase effect, post-tetanic and tetanic potentiation in individual MU types indicates that these phenomena may be independent of each other. We suggest that these phenomena as well as the fatigue resistance should be reexamined with protocols adjusted to the MU optimal frequency.     

Short-term synchronization of motor units in human extensor digitorum communis muscle: relation to contractile properties and voluntary control

Synchronous activity was studied in relation to the contractile properties of pairs of motor units (MUs) recorded with independent microelectrodes in the right extensor digitorum communis muscle (EDC) of human subjects during isometric finger extension. MU contrac tile properties were characterized in terms of the rise time and amplitude of twitch tensions extracted by spike-trig gered averages of the extension force. Synchronization of MU discharges appeared in the form of narrow central peaks in the cross-correlograms of 35 of 50 pairs of MUs, suggesting the contribution of common last-order neu rons. Synchronization peaks tended to be briefer and higher among pairs of MUs with slower and smaller twitches than among pairs of MUs with faster and larger twitches. The higher peaks of slow-contracting MUs sug gest a greater effectiveness of the common synaptic in puts. The broader peaks of fast-contracting MUs might reflect an additional synchronization of the inputs to fast MUs at high force levels. The areas of the cross-correlo gram peaks were similar for both groups and suggest that under our conditions, about three motoneurons would discharge synchronously for a given motoneuron spike. To test whether the amount of MU synchronization could be altered voluntarily, four subjects attempted to increase or decrease synchrony, using as feedback clicks triggered by coincident firings of the recorded MUs. In nine of 15 conditioning sessions, the magnitudes of the synchronization peaks showed significant changes in the intended direction. These results imply that supraspinal centers can control the relative amount of inputs that contribute to the synchronization of motoneuron dis charges during voluntary contraction of EDC.     

Relation between the motor units recruitment threshold and their potentials propagation velocity at isometric activity.

The relationship between the propagation velocity of the excitation along the muscle fibers of the motor units (MUs) and their threshold of recruitment at different level of isometric voluntary contraction was investigated. The threshold of recruitment was measured by the value of the muscle force, expressed in percents from the maximal voluntary contraction (MVC) at which the first impulse of the MU appeared. A wire subcutaneous branched electrode was used to select the potentials from a single MU. The selected in this way MU impulses were used as a trigger to average two electromyographic (EMG) signals picked up by means of two monopolar surface electrodes with small leading-off areas mounted on a common plate at a distance of 10 mm from one another. The propagation velocity of the extraterritorial potentials of the MUs increased non-linearly with the increase of the recruitment threshold. The relationship was fitted as V = square root of a+b.theta, where v is the propagation velocity, theta is the threshold of recruitment and a and b are constants. The consideration of the velocity of propagation as a "size principle parameter" was discussed and the limitations of the latter are pointed out.     

EMG activation patterns during force production in precision grip. III. Synchronisation of single motor units

Motor unit (MU) synchronisation during isometric force production in the precision grip was analysed in five subjects performing a visually guided steptracking motor task with three different force levels. With this aim multi-unit electromyographic (EMG) activity of 14 intrinsic and extrinsic finger muscles from 15 experimental sessions was decomposed into the potentials of single MUs. The behaviour of 62 intrinsic and 30 extrinsic MUs in the motor task was quantified. Most MUs displayed a positive correlation between firing rate and grip force. Compared to MUs in extrinsic muscles, intrinsic MUs had steeper regression lines with negative intercepts indicating higher force sensitivity and higher recruitment thresholds. A cross-correlation analysis was performed for 69 intra- and 166 intermuscular MU pairs while steady grip force was exerted at the three force levels. Synchronisation, for at least one force level, was found in 78% of the intra- and 45% of the intermuscular pairs. The occurrence of synchronisation was not stable over the force range tested. Factors influencing the fluctuations in occurrence and strength of synchronisation were investigated. Force increase was not paralleled by increased synchronisation; in contrast, in most MU pairs, especially intermuscular pairs, synchronisation occurred preferentially at the lower force levels. The recruitment threshold appeared to play a determining role in synchronisation: the more similar the thresholds of two MUs, the greater the probability of them being synchronised at this force level. Synchronised MUs fired on average at a lower frequency than non-synchronised ones. Finally, synchronisation at the multi-unit EMG level does not indicate that all underlying MUs are synchronised, nor does the absence of temporal coupling at the multi-unit level indicate that none of the MUs is synchronised.     

Nonlinear tension summation of different combinations of motor units in the anesthetized cat peroneus longus muscle

The purpose of this study was to examine the linearity of summation of the forces produced by the stimulation of different combinations of type identified motor units (MUs) in the cat peroneus longus muscle (PL) under isometric conditions. The muscle was fixed at its twitch optimal length, and the tension produced by the single MU was recorded during 24- and 72-Hz stimulation. The summation analysis was first carried out for MUs belonging to the same functional group, and then different combinations of fast fatigable (FF) MUs were added to the nonfatigable slow (S) and fatigue resistant (FR) group. The tension resulting from the combined stimulation of increasing numbers of MUs (measured tension) was evaluated and compared with the linearly predicted value, calculated by adding algebraically the tension produced by the individual MUs assembled in the combination (calculated tension). Tension summation displayed deviations from linearity. S and FR MUs mainly showed marked more than linear summation; FF MUs yielded either more or less than linear summation; and, when the FF units were recruited after the S and FR MUs, less than linear summation always occurred. The magnitude of the nonlinear summation appeared stimulus frequency dependent for the fatigable FF and FI group. The relationship between measured tension and calculated tension for each MU combination was examined, and linear regression lines were fitted to each set of data. The high correlation coefficients and the different slope values for the different MU-type combinations suggested that the nonlinear summation was MU-type specific. The mechanisms of nonlinear summations are discussed by considering the consequences of internal shortening and thus the mechanical interactions among MUs and shifts in muscle fiber length to a more or less advantageous portion of single MU length-tension curves.     

Motor Unit Properties in the Soleus Muscle after Its Distal Tendon Transfer to the Plantaris Muscle Tendon in the Rat

The aim of this study was to evaluate how a modification in the mechanical conditions under which a muscle is used could induce changes in the characteristics and the spinal drive of its motor units (MU). The distal tendon of the soleus muscle of Wistar rats was transferred to the distal stump of the plantaris muscle tendon. The EMG activity of the soleus was chronically recorded for 8 weeks, every other day, during a 1-min treadmill walk. After spinal ventral root splitting, individual MU contractile properties were measured in control soleus (102 MUs) or in transposed soleus muscles after 4 weeks (41 MUs) or 8 weeks (28 MUs). Muscle/body weight ratio did not vary after transposition, nor did MU tetanic forces. A decrease in MU twitch contraction times and in their half relaxation times was observed at weeks 4 and 8. MU tension-frequency curves varied significantly after tendon transfer, becoming closer to the curves of the fast MUs of the control group. During locomotion, we observed no change in the amplitude of rectified-filtered electromyographic activity, but a significant decrease in mean burst duration and an increase in the median frequency of the power density spectrum. Tendon transposition of the soleus muscle brought about adaptations in MU contractile properties and soleus spinal control.     

Variability of the twitch parameters of the rat medial gastrocnemius motor units--experimental and modeling study

In the present study a previously proposed model of a twitch based on an analytical function with four-parameters (lead, contraction and half-relaxation times and maximum force of the twitch) was validated on 115 motor units (MUs), divided into slow (S), fast-fatigue resistant (FR) and fast fatigable (FF) types. The original records were collected from electrophysiological experiments performed on MUs from the medial gastrocnemius muscle of five rats. Besides the easy calculation of the twitch parameters and their variability, the usefulness of the model was confirmed by eliminating artifacts and noise in the original twitch records, as well as by calculations of the velocity of force increase and decrease, the area under force records, and by normalization of all twitches with respect to the maximal force and contraction time. It was concluded that: (1) the four-parameter twitch model describes precisely the individual contractions of various MUs; (2) all physiological twitch parameters are distributed continuously and located within overlapping intervals for different MU types; this distribution is not linear, but exponential; (3) S MUs can be distinguished from fast ones on the basis of some twitch parameters (contraction and half-relaxation times, velocity of contraction), but the same cannot be applied for FF and FR MUs; (4) the analysis of the normalized twitches reveals the differences in shapes for different types of MUs, which shows that twitches of different MUs cannot be obtained from one standard pattern scaled in time and force. These results may have functional implications for studying effectiveness of twitch summation during tetanic contractions and the work performed by various types of MUs.     

Motor unit recruitment and rate coding in response to fatiguing shoulder abductions and subsequent recovery

The purpose of the present study was to investigate motor unit (MU) recruitment and firing rate, and the MU action potential (MUAP) characteristics of the human supraspinatus muscle during prolonged static contraction and subsequent recovery. Eight female subjects sustained a 30° shoulder abduction, requiring 11–12% of maximal voluntary contraction (MVC), for 30 min. At 10 and 30 min into the recovery period, the shoulder abduction was repeated for 1 min. The rating of perceived exertion for the shoulder region increased to “close to exhaustion” during the prolonged contraction, and the surface electromyography (EMG) recorded from the deltoid and trapezius muscles showed signs of local muscle fatigue. From the supraspinatus muscle, a total of 23,830 MU firings from 265 MUs were identified using needle electrodes. Of the identified MUs, 95% were continuously active during the 8-s recordings, indicating a low degree of MU rotation. The mean (range) MU firing rate was 11.2 (5.7–14.5) Hz, indicating the relative force contribution of individual MUs to be larger than the overall mean shoulder muscle load. The average MU firing rate remained stable throughout the prolonged abduction, although firing rate variability increased in response to fatigue. The average concentric MUAP amplitude increased by 38% from the beginning (0–6 min) to the end (24–29 min) of the contraction period, indicating recruitment of larger MUs in response to fatigue. In contrast, after 10 min of recovery the average MU amplitude was smaller than seen initially in the prolonged contraction, but not different after 30 min, while the MU firing rate was higher during both tests. In conclusion, MU recruitment plays a significant role during fatigue, whereas rate coding has a major priority during recovery. Furthermore, a low degree of MU rotation in combination with a high relative load at the MU level may imply a risk of overloading certain MUs during prolonged contractions. Accepted: 6 June 2000     

Linear and nonlinear effects in the interactions of motor units and muscle spindle afferents

Summary Successive motor unit (MU) twitches often do not sum linearly. Also, muscle spindle (MS) afferents may react nonlinearly to MU contractions occurring at short intervals. Little data is presently available on the interactions between two (or more) MUs regarding their effects on tension output and MS responses. We have studied these effects in cat Mm. gastrocnemius medialis (MG), soleus and semitendinosus. In adult anaesthetized cats, MUs of the muscle under study were electrically stimulated via their ventral root axons with random sequences of brief pulses having mean rates between 6 and 12 pulses per second. Isometric tension fluctuations were recorded from the muscle under study, and discharge patterns of MS afferents (Ia and group II) were recorded from dorsal root filaments. A crosscorrelation analysis was performed to display linear and nonlinear effects evoked by selected time constellations of MU activations. 1) 18 (67%) of 27 MG MUs showed marked potentiation of the second of two twitches in response to pairs of stimuli separated by 5 to about 25 ms. The remainder of these and 16 of the soleus MUs did not exhibit conspicuous nonlinearities. — 2) MS responses to such pairs of MU activations usually showed a prolonged spindle pause. — 3) About 28% of 36pairs of MG MUs produced twitch tension less than expected for linear summation if activated nearly simultaneously. — 4) If two MUs both produced a spindle pause and possibly a relaxation discharge in an MS afferent, the near-synchronous activation of the units produced respective discharge variations that were less than expected for linear summation. If one MU produced an early discharge, contraction of another MU would often prevent it. — These results are discussed in regard to mechanisms of tremor suppression.     

Postnatal loss of synaptic terminals in the normal mouse soleus muscle

Using conventional physiological techniques for measuring unitary contractions and end-plate potentials (epps), the number, size and segmental properties of motor units (MUs) in the soleus muscle of the mouse during postnatal development have been examined. The number of MUs remains constant after birth, and there is no evidence of segmental preferences in the innervation pattern, before, during or after the postnatal elimination of redundant terminals. In neonates, MU size estimates based on twitch contractions are 30% smaller than tetanic estimates. Intracellular recording of epps shows that this is caused by facilitation of epps on repetitive stimulation. The frequency of occurrence of epps in the muscle from a few, isolated motor axons shows that the average motoneuron contacts 36% of the fibres in the muscle neonatally. A substantial fraction of the contacts is subthreshold for twitch activation of their fibre. The MU size remains constant up to day 5. During the next 10 days, the MU size is reduced to the mature value of 5% of the fibres in the muscle. It is concluded that the neonatal loss of synaptic terminals in this muscle takes place without concomitant loss of entire motor neurons, and that it is independent of possible segmental preferences in the innervation of the muscle.     

Relations among motor unit types,generated forces and muscle length in single motor units of anaesthetized cat peroneus longus muscle

The active length-tension curves of identified single motor units (MUs) belonging to peroneus longus muscle (PL) of anaesthetized adult cats were obtained by eliciting isometric single twitches and tetani. The recorded responses were evaluated by measuring the peak tension amplitude and the tension-time area at muscle lengths extending throughout the physiological length range of the muscle (mean 5.5 mm, standard deviation ±0.8). The muscle lengths at which each tested MU developed its maximal twitch (L _tw) and tetanic (L _te) tensions were determined and compared with the muscle length (L _o) at which the stimulation of all the -axons, innervating PL and contained in L7 ventral root, developed their maximal twitch tension. The mean of single MU L _tw values was at L _o+1.08±1.1 mm. Slow MUs showed the longest values of L _tw(L _o+1.6±1.0 mm). Single MUs stimulated at tetanic frequencies presented their L _te at values shorter than L _o (L _o–2.8±1.7 mm). Slow MUs had the shortest L _te (L _o–3.4±1.5 mm). For all the units L _te was shorter than L _tw. L _tw and L _te were, respectively, negatively and positively correlated with the developed tension. Optimal length values also appeared to be related to the MU types. The possibility is discussed that the muscle and tendon compliances and the high non-linearities to the applied forces are the main factors which can determine the differences among L _o, L _tw and L _te values. The relationships between MU type and optimal length values are suggested to be, at least partly, an epiphenomenon due to the different contraction strengths of the various MU types. However, the heterogeneous distribution of the MU types is brought into account to explain the dependence of L _tw and L _te values on MU type.     

Non-invasive assessment of single motor unit mechanomyographic response and twitch force by spike-triggered averaging

A method for non-invasive assessment of single motor unit (MU) properties from electromyographic (EMG), mechanomyographic (MMG) and force signals is proposed. The method is based on the detection and classification of single MU action potentials from interference multichannel surface EMG signals and on the spike-triggered average of the MMG (detected by an accelerometer) and force signals. The first dorsal interosseous (FDI) and abductor digiti minimi (ADM) muscles were investigated at contraction levels of 2% and 5% of the maximum voluntary contraction (MVC) force. A third contraction was performed by selective activation of a single MU with surface MU action potential visual feedback provided to the subject. At 5% MVC, the mean (±standard error) single MU MMG peak-to-peak value was 11.0±1.8 mm s⁻² (N=17) and 32.3±6.5 mm s⁻² (N=20) for the FDI and AMD muscles, respectively. The peak of the twitch force was, at the same contraction livel, 7.41±1.34 mN and 14.42±2.92 mN, for the FDI and ADM muscles, respectively. The peak-to-peak value of the MMG was significantly different for the same MU at different contraction levels, indicating a non-linear summation of the single MU contributions. For the FDI muscle, the MMG peak-to-peak value of individual MUs was 21.5±7.8 mm s⁻², when such MUs were activated with visual feedback provided to the subject, whereas, for the same MUs, it was 11.8±3.8 mm s⁻², when the subject maintained a constant force level of 2% MVC. The method proposed allows the non-invasive assessment of single MU membrane and contractile properties during voluntary contractions.     

Localization and contractile properties of intrinsic longitudinal motor units of the rat tongue

Tongue dysfunction is a hallmark of many human clinical disorders, yet we lack even a rudimentary understanding of tongue neural control. Here, the location and contractile properties of intrinsic longitudinal motor units (MUs) of the rat tongue body are described to provide a foundation for developing and testing theories of tongue motor control. One hundred and sixty-five MUs were studied by microelectrode penetration and stimulation of individual motor axons coursing in the terminal portion of the lateral (retrusor) branch of the hypoglossal nerve in the rat. Uniaxial MU force was recorded by a transducer attached to the protruded tongue tip, and MU location was estimated by electromyographic (EMG) electrodes implanted into the anterior, middle, and posterior portions of the tongue body. All MUs produced retrusive force. MU twitch force ranged from 2-129 mg (mean = 35 mg) and tetanic force ranged from 9-394 mg (mean = 95 mg). MUs reached maximal twitch force in 8-33 ms (mean = 15 ms) and were resistant to fatigue; following 2 min of stimulation, MUs (n = 11) produced 78-131% of initial force. EMG data were collected for 105 MUs. For 65 of these MUs, the EMG response was confined to a single electrode location: for 26 MUs to the anterior, 21 MUs to the middle, and 18 MUs to the posterior portion of the tongue. Of the remaining MUs, EMG responses were observed in two (38/40) or all three (2/40) tongue regions. These data provide the first contractile measures of identified intrinsic tongue body MUs and the first evidence that intrinsic longitudinal MUs are restricted to a portion of tongue length. Localization of MU territory suggests a role for intrinsic MU in the regional control of the mammalian tongue observed during feeding and speech.     

Discharge behavior of motor units in knee extensors during the initial stage of constant-force isometric contraction at low force level

Abstract. To elucidate the strategy of the activity of motor units (MUs) to maintain a constant-force isometric contraction, I examined the behavior of MUs in knee extensor muscles [(vastus medialis (VM), vastus lateralis (VL) and rectus femoris (RF)] during a sustained contraction at 5% of maximal voluntary contraction for 5 min. In all cases, the spike interval exhibited an elongating trend, and two discharge patterns were observed, continuous discharge and decruitment. In continuous-discharge MUs, the trend slope was steep immediately after the onset of constant force (steep phase), and then became gentle (gentle phase). Decruitments were observed frequently during each phase, and additional MU recruitment was observed throughout the contraction. The mean value of recruitment threshold force did not differ among the extensors. The mean spike interval at the onset of constant-force isometric contractions was shorter in RF than in VL. However, there were no differences in the duration and extent of the elongating trend, decruitment time and recruitment time among the extensors. The electromyogram of the antagonist biceps femoris muscle revealed no compensatory change for extensor activity. These results indicated that at a low force level, the strategy employed by the central nervous system to maintain constant force appears to involve cooperation among elongating trends in the spike interval, decruitment following elongation, and additional MU recruitment in synergistic muscles. Electronic Publication