Consistency of motor-unit identification during force-varying static contractions

Due to inter-operator variability, two operators were used to assess the consistency of motor unit (MU) identification during ramp contractions, by the comparison of semi-automatic decompositions of the same recordings. Static shoulder abduction was performed against a force transducer in a position with the upper arms vertical and elbows flexed to 90°. The subjects followed an 8-s force trajectory: 30% maximum voluntary contraction (MVC, 2 s), a reduction in force from 30% to 0% MVC (2 s), 0% MVC (1 s), an increase in force from 0 to 30% MVC (2 s), and 30% MVC (1 s). Muscle activity was recorded from the supraspinatus muscle with a quadripolar needle. From six recordings of 8 s duration, a total of 2527 MU firings were identified by both operators, and 93% of these were identified identically into 31 MUs. Both operators identified 8 of these MUs as continuously firing, 5 as only being active either before or after the 1 s at 0% MVC, and 18 as being de-recruited during force decreases and recruited during force increases. Both operators agreed that 16 of these 18 MUs were de-recruited at a higher force level than that at which they were recruited, which may be due to the electromechanical delay. The coefficient of variation for double determination of the results obtained by operators A and B was 8.5% for the number of MU firings, 4.5% for the MU mean firing rate, and 8.4% for the MU action potential (MUAP) amplitude. Therefore, the operator interactive decomposition method was considered to be valid for studying recruitment and de-recruitment as well as firing rate and MUAP amplitude during static, force-varying ramp contractions. Accepted: 31 May 2000     

The influence of experimental muscle pain on motor unit activity during low-level contraction

In the present study we compared motor unit (MU) activity in a painful extensor carpi ulnaris (ECU) muscle to that of a pain-free control. According to the pain adaptation model the activity of the painful ECU muscle may be inhibited and its antagonist activity increased during wrist extension performed as a pre-defined low-force ramp. The pre-defined low force may then be maintained by increased activity in the pain-free synergist muscles such as the extensor carpi radialis (ECR) muscle. Nine females (31–47 years old) participated in the study. Maximal voluntary contraction (MVC) of the wrist extensors was performed. A catheter was inserted into the ECU muscle to allow the injection of hypertonic saline to evoke muscle pain, and a concentric needle was inserted for the recording of MU activity. Surface electromyograms were recorded from a synergist and an antagonist (ECR and flexor carpi radialis) to the painful ECU muscle. A force ramp of isometric wrist extensions up to 10% MVC, with a force increase of 1% MVC · s⁻¹, were performed followed by 60 s of sustained contraction at 10% MVC. The number of MUs recruited was almost identical for baseline and with pain, and no effect of experimental muscle pain was found on the properties of the MUs (amplitude, area) or their firing characteristics (mean firing rate, firing variability) during low-force ramp contraction. During the sustained 10% MVC, no effect of pain was found for concentric or surface EMG of the forearm muscles. At low force levels no pain-induced modulations were found in MU activity, when the mechanical condition was similar to that of a control situation. Accepted: 30 May 2000     

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     

The fatigability of two agonistic muscles in human isometric voluntary submaximal contraction: an EMG study

Summary The recruitment and firing rate of biceps brachii (BB) and brachioradialis (BR) motor units (MUs) were studied in the course of fatiguing isometric contractions at 20%–30% of maximal voluntary contraction (MVC). MU recruitment generally occurred throughout the maintained contraction and was similar for BB and BR muscles. Newly recruited MUs started to discharge in the form of bursts, the duration of which increased until a continuous rhythmical firing was achieved. Within each burst, the first interval between two consecutive discharges was usually the shortest. MU threshold was lowered just after the limit time of the maintained contraction. The MU's firing rate either increased or remained stable as a function of the elapsed time. It is concluded that (1) in fatiguing isometric contractions at 20%–30% MVC contractile failure is mainly compensated for by MU recruitment and a lowered MU threshold and (2) differences between in surface changes in the electromyogram of BB and BR muscles cannot easily be explained by related differences in MU firing rate and recruitment.     

Adaptations in motor unit discharge activity with force control training in young and older human adults

Six young (aged 18–22 years) and six older (aged 66–76 years) healthy humans participated in a visually guided isometric force modulation training program designed to improve accurate control of force during ankle dorsiflexion. Isometric force and the discharge activity of motor units (MU) supplying the tibialis anterior muscle were sampled concurrently at the beginning of the study, following 2 weeks of force modulation training and again after a 4 week retention period which followed immediately. The initial maximal voluntary force (MVC) and MU discharge rates were similar between young and older adults at 10–60% MVC while MU discharge rates during maximal effort were significantly reduced in older adults. Following the 2 weeks of force modulation training, both young and older adults demonstrated significant improvements in force accuracy (44% young, 48% older) and significantly reduced MU discharge rates at 30%, 40%, and 60% MVC. Young adults also demonstrated increased MVC force (11%), while older adults demonstrated significantly increased (30%) maximal MU discharge rate. Thus, following 2 weeks of force modulation training, young and older individuals demonstrated similar MU discharge rates at all force levels. The MU discharge rate adaptations were retained after the 4 week retention period. In young adults, improved force accuracy and increased MVC force were accompanied by significantly reduced MU recruitment thresholds. In the older subjects, improved force accuracy was accompanied by an increase in the difference between the recruitment-derecruitment force threshold and significantly reduced antagonist co-contraction. Age-related alterations in force regulation and MU discharge activity cannot be explained solely on the basis of contractile changes in senescent muscle. Rather, reliance on compensatory neuromuscular changes including antagonist muscle co-contraction is suggested. Accepted: 27 June 2000     

The effect of prolonged isometric contractions on muscle fluid balance

Ultrasound scanning was performed at three sites above the fossa supraspinata on nine healthy subjects and five patients with myofascial shoulder pain. This method produced a well-defined depiction of the soft tissue layers above the fossa supraspinata and reproducible muscle thickness measurements. In the healthy subjects the average distance from the skin surface to the trapezius muscle was 7.7 mm and the average thickness of the trapezius muscle was 5.3 mm, and the average thickness of supraspinatus muscle was 20.0 mm. The supraspinatus muscle was thinner at the medial measuring site than at the other two sites. In contrast, a tendency towards a larger distance was seen from the skin to trapezius muscle at the medial measuring site than at the other two sites. No statistical differences were found between the two groups of subjects either at rest or during brief shoulder abductions. All the subjects performed a 30° unilateral isometric shoulder abduction test to exhaustion. The median endurance time was 33 min for the healthy subjects and only 5 min for the patients. The ratings of perceived exertion (RPE) were in line with this, since the increment in RPE with time was larger for the patients than for the healthy group. The reduced shoulder abduction endurance time in the patient group may have been related to impaired muscle function and/or pain development. During the 33-min shoulder abduction in the healthy subjects, the thickness of supraspinatus muscle increased by 14%, indicating muscle swelling, whereas the thickness of trapezius muscle remained constant. The fluid imbalance in the supraspinatus muscle compartment may well play a role in the development of muscle fatigue and the disorders found in industry resulting from prolonged work with arms elevated.     

The surface mechanomyogram as a tool to describe the influence of fatigue on biceps brachii motor unit activation strategy. Historical basis and novel evidence

The surface mechanomyogram (MMG) (detectable at the muscle surface as MMG by accelerometers, piezoelectric contact sensors or other transducers) is the summation of the activity of single motor units (MUs). Each MU contribution is related to the pressure waves generated by the active muscle fibres. The first part of this article will review briefly the results obtained by our group studying the possible role of motor unit recruitment and firing rate in determining the characteristics of the MMG during stimulated and voluntary contractions. The second part of this article will study the MMG and EMG during a short isometric force ramp from 0 to 90% of the maximal voluntary contraction (MVC) in fresh and fatigued biceps brachii. The aim is to verify whether changes in motor unit activation strategy in voluntarily fatigued muscle could be specifically reflected in the time and frequency domain parameters of the MMG. MMG-RMS vs. %MVC: at fatigue the MMG-RMS did not present the well known increment, when effort level increases, followed by a clear decrement at near-maximal contraction levels. MMG-MF vs. %MVC: compared to fresh muscle the fatigued biceps brachii showed an MF trend significantly shifted towards lower values and the steeper MF increment, from 65 to 85% MVC, was not present. The alteration in the MMG and EMG parameters vs. %MVC relationships at fatigue seems to be related to the impossibility of recruiting fast, but more fatigable MUs, and to the lowering of the global MUs firing during the short isometric force ramp investigated.     

Motor unit discharge and force tremor in skill- and strength-trained individuals

We examined motor unit (MU) discharge properties (mean interspike interval, ISI, discharge variability, short-term synchronization, common drive) and force tremor in the first dorsal interosseous (FDI) muscle of five musicians (skill-trained), five weight-lifters (strength-trained) and six untrained subjects during low-force isometric abduction of the index finger. Mean MU ISI was slightly shorter in skill-trained subjects than in untrained subjects. Discharge variability of FDI MUs did not differ significantly between groups. The mean strength of MU synchronization (expressed as the frequency of extra synchronous discharges above chance) was different in skill-trained (0.22±0.02 s^–1, 162 MU pairs), untrained (0.32±0.02 s^–1, 199 MU pairs) and strength-trained subjects (0.44±0.03 s^–1, 183 MU pairs). FDI MU synchrony was weak and of equivalent strength in both hands of skill-trained subjects and the dominant (skilled) hand of untrained subjects. The stronger FDI MU synchrony in the non-dominant hand of untrained subjects was equivalent to that found in both hands of strength-trained subjects. The extent of common modulation of firing rates (common drive) was assessed for a subset of MU pairs and was weaker in skill-trained subjects (0.30±0.04, n=14) than untrained (0.43±0.3, n=14) and strength-trained (0.48±0.03, n=21) subjects. Force tremor was quantified for each hand in the same subjects during isometric index finger abduction at target forces of 0.5 N and 3.5 N. Tremor rms amplitude and peak power in the force frequency spectrum were significantly lower in skill-trained subjects than strength-trained subjects with the 3.5-N target force. The peak tremor frequency was similar in the three groups. The relatively more independent discharge of pairs of FDI MUs in skill-trained subjects was not responsible for the reduced tremor amplitudes in these subjects. Correlations between the overall extent of MU synchrony and common drive in FDI muscles and tremor measures obtained during the same experimental session were all non-significant. Differences in the central descending command signals are the most likely explanation for the more independent discharge of FDI MUs in skill-trained hands, while neural or peripheral muscular factors may be responsible for the weaker tremor. Received: 15 May 1997 / Accepted: 15 August 1997     

Intramuscular laser-Doppler flowmetry in the supraspinatus muscle during isometric contractions

To study the regulation of microvascular blood flow in a compartment muscle, laser-Doppler measurements of muscle microcirculation were recorded in the supraspinatus muscle in eight volunteers during and following submaximal isometric muscle contractions. The subjects performed isometric shoulder abductions at five contraction levels from 5% to 50% maximal voluntary contraction for I min each and a sustained 30° shoulder abduction for 20 min. The subjects' perceived exertion increased from no perceived exertion to near maximal exertion during the 20-min period with 30° shoulder abduction. Microcirculation increased during all 1-min contractions. Following the contractions at 20%, 30% and 50% MVC post-exercise reactive hyperaemia was seen for a period of at least 1 min. The reactive hyperaemia increased in magnitude in response to increasing contraction level. The results showed the same time-history of the blood flow at microvascular level as previously seen in larger peripheral vessels in response to muscle contractions. During the 20-min contraction microcirculation increased in line with the findings during the brief contractions. However, in contrast to the brief contractions no postexercise reactive hyperaemia occurred following the prolonged contraction. Lack of postexercise reactive hyperaemia following the prolonged shoulder abduction would suggest insufficient regulation of the vascular resistance. Alternatively, lack of hyperaemia could be taken as an indication of sufficient microcirculation during the preceding contraction. From previous studies on intramuscular pressure and metabolism the latter alternative would seem unlikely.     

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.     

The influence of experimentally induced pain on shoulder muscle activity

Muscle function is altered in painful shoulder conditions. However, the influence of shoulder pain on muscle coordination of the shoulder has not been fully clarified. The aim of the present study was to examine the effect of experimentally induced shoulder pain on shoulder muscle function. Eleven healthy men (range 22–27 years), with no history of shoulder or cervical problems, were included in the study. Pain was induced by 5% hypertonic saline injections into the supraspinatus muscle or subacromially. Seated in a shoulder machine, subjects performed standardized concentric abduction (0°–105°) at a speed of approximately 120°/s, controlled by a metronome. During abduction, electromyographic (EMG) activity was recorded by intramuscular wire electrodes inserted in two deeply located shoulder muscles and by surface-electrodes over six superficially located shoulder muscles. EMG was recorded before pain, during pain and after pain had subsided and pain intensity was continuously scored on a visual analog scale (VAS). During abduction, experimentally induced pain in the supraspinatus muscle caused a significant decrease in activity of the anterior deltoid, upper trapezius and the infraspinatus and an increase in activity of lower trapezius and latissimus dorsi muscles. Following subacromial injection a significantly increased muscle activity was seen in the lower trapezius, the serratus anterior and the latissimus dorsi muscles. In conclusion, this study shows that acute pain both subacromially and in the supraspinatus muscle modulates coordination of the shoulder muscles during voluntary movements. During painful conditions, an increased activity was detected in the antagonist (latissimus), which support the idea that localized pain affects muscle activation in a way that protects the painful structure. Further, the changes in muscle activity following subacromial pain induction tend to expand the subacromial space and thereby decrease the load on the painful structures.     

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     

Motor impairment in the human hand following eccentric exercise

Motor impairment was induced by having subjects perform two sets of 50 maximal contractions, using the first dorsal interosseus (FDI) muscle to abduct the index finger, while the muscle was being stretched. Tests were conducted prior to the exercise (pre-exercise) and 24 h following the exercise (post-exercise). There were declines of 19% in maximal abduction torque and 15% in maximal flexion torque at the metacarpaphalangeal joint, during isometric contraction post-exercise compared to pre-exercise. The ability to stabilize the metacarpophalangeal joint about the abduction/adduction axis was reduced by 14% post-exercise, and the variability in tracking an isometric torque target increased by 30%. There was a decrement of 7%–10% in the median frequency of the power density spectrum of FDI electromyogram (EMG) throughout a 60 s maintained abduction at 50% maximal voluntary contraction. The mean rectified EMG, on the other hand, increased by 100%–175% for torque levels below 40% of maximal voluntary contraction, post-exercise. The results were consistent with preferential injury of type II muscle fibres in FDI. Although non-exercised synergist muscles appeared to be inhibited during maximal voluntary flexion, there was evidence that they compensated for injured FDI muscle fibres during maintained contraction at sub-maximal flexion torque. Accepted: 19 September 2000     

Comparison of F-waves of motor unit action potentials activated during voluntary contraction

The system for classifying F-waves was developed to study the properties of F-wave and to compare single motor unit (MU) F-waves with motor unit action potentials (MUAPs) activated during voluntary contraction. The F-waves evoked by submaximal stimulation as well as the EMG signals during voluntary contraction at 6 levels of 10-100% of maximum voluntary contraction (MVC) were measured in the tibialis anterior muscles of 3 healthy volunteers. Nine channel F-wave waveforms in a selected electrode array were classified using a template-matching method. After the detection procedure of MUAPs in voluntary EMG signals, the MUAPs were also classified by the same method. Most of the F-waves (88.4%) were composed of a single MUAP. The numbers of MU classified from single MU F-waves in 3 subjects were 12, 12 and 15, and the numbers of MU classified from the voluntary EMG signals at 6 contraction levels were 20, 27 and 24, respectively. A total of 26 single MU F-waves were identified with the MUs extracted from the data during voluntary contractions. The results suggest that the F-waves are composed of a population of the MUs, which are recruited at a wide range of contraction levels. The classification procedures of F-waves and voluntary EMG signals have made it possible to recognize the same MU in both signals and to analysis the firing thresholds of F-waves.     

Effect of experimental muscle pain on motor unit firing rate and conduction velocity

The aim of this human study was to investigate the relationship between experimentally induced muscle pain intensity (i.e., amount of nociceptive activity) and motor unit (MU) firing decrease and MU conduction velocity (CV). In 12 healthy subjects, nociceptive afferents were stimulated in the right tibialis anterior muscle by three intramuscular injections of hypertonic saline (0.2, 0.5, and 0.9 ml) separated by 140 s. The subjects performed six isometric contractions (20 s long) at 10% of the maximal voluntary contraction during the experimental muscle pain. The same set of six contractions was performed without any infusion before the painful condition on the right leg. The procedure was repeated for the left leg with infusion of isotonic (nonpainful) saline. Intramuscular and surface electromyographic (EMG) signals were collected to assess MU firing rate and CV. The firing rate of the active MUs [range: 7.4-14.8 pulses/s (pps)] did not change significantly in the three control conditions (without infusion for the right and left leg and with infusion of isotonic saline in the left leg). There was, on the contrary, a significant decrease (on average, mean +/- SE, 1.03 +/- 0.21 pps) of the firing rates during the painful condition. Moreover, MU firing rates were inversely significantly correlated with the subjective scores of pain intensity. Single MU CV was 3.88 +/- 0.03 m/s (mean +/- SE, over all the MUs) with no statistical difference among any condition, i.e., the injection of hypertonic saline did not alter the muscle fiber membrane properties of the observed MUs. Progressively increased muscle pain intensity causes a gradual decrease of MU firing rates. This decrease is not associated with a change in MU membrane properties, indirectly assessed by CV. This study demonstrates a central inhibitory motor control mechanism with an efficacy correlated to the nociceptive activity.     

Task-related behaviour of motor units in the human temporalis muscle

Muscle activity patterns in some complex human jaw muscles appear to be task sensitive. However, it is presently uncertain how changes in motor task affect motor unit (MU) behaviour in the human temporalis muscle. In this study, activity was recorded from 40 MUs in the anterior region of the muscle. The lowest sustainable firing frequency (LSFF) was reached by slow increases and decreases in firing rate, then firing was maintained at the lowest possible rate without significant pauses. An array of consecutive interspike intervals (ISI) were sampled digitally and used to measure the LSFF for each task associated with the MU. In a controlled paradigm, MU reflex inhibition was measured during the performance of different tasks. Single electrical pulses of non-noxious intensity were delivered to the gingiva near the maxillary canine tooth. During continuous MU firing at a controlled firing frequency of 10 Hz, series of pulses were delivered with increasing delays, after preselected spikes. The MUs fired continuously during the performance of 1–4 postural and tooth-contact tasks. There were significant differences in LSFFs between tasks in those MUs associated with multiple tasks. In the reflex study, all MUs were inhibited, but the magnitude of the inhibition was highly task dependent. Thus, both LSFF and reflex inhibition of temporalis MUs appear to vary with the motor task and are sensitive to the position of the jaw and the direction and location of tooth contact along the tooth row. This behaviour most likely reflects task-related changes in output from orofacial and muscle afferents.     

Upper trapezius muscle conduction velocity during fatigue in subjects with and without work-related muscular disorders: a non-invasive high spatial resolution approach

The development of fatigue at the muscle fibre level can be assessed in terms of a decrease in conduction velocity (CV). The present study aimed to investigate if work-related muscular disorders in occupations characterised by static loads of long duration affect fatigue resistance in the painful muscle. A group of eight secretaries suffering from bilateral chronic muscle pain in the shoulder/neck region was compared to a group of healthy subjects. The upper trapezius muscle was studied under isometric contractions, holding the arm in the horizontal plane up to the endurance point. Changes in CV estimated at the motor unit level were investigated using a non-invasive high spatial resolution electromyographic (EMG) approach. In addition, the number of motor unit potentials per second (PPS), and the root mean square (RMS) of bipolar signals were assessed, and the results reported as the mean and standard error for each value. Subjects with work-related disorders showed less pronounced changes in CV with respect to healthy subjects. No differences between subjects with and without work-related disorders were encountered for the PPS and RMS. The present findings on CV indicate an increased fatigue-related recruitment of MUs in the painful muscle with respect to a healthy muscle. The fact that this recruitment is not reflected in the PPS and RMS estimates might be due to a fatigue-induced decrease in the firing rate and/or the de-recruitment of fatigued MUs. Furthermore, methodological limitations of the adopted method in the estimation of ‘global’ parameters such as the PPS and RMS have to be considered.     

Occurrence of widespread motor-unit firing correlations in muscle contractions: their role in the generation of tremor and time-varying voluntary force

The firing behavior of motor units (MUs) of the first dorsal interrosseus muscle of the hand was examined during both constant-force and varying-force (sinusoidal or broadband random variations) isometric contractions in healthy adults. The emphasis was on the analysis of MU synchrony with an efficient and sensitive method. In static contractions, widespread and strong MU firing correlations, with the MUs in phase, were present at the frequency of muscle tremor, when the tremor was regular (narrowband) and large. MU correlations could also exist in contractions where the tremor of a subject was irregular (broadband) overall, but they were generally weak. These correlations were at the frequency of the subject's regular tremor, and the corresponding distinct tremor component was sometimes discernible within the broad tremor-band. In contrast, the MUs did not show any such correlations in the case of purely irregular and small tremor. On the basis of these observations, it is concluded that the rhythms in the force contributions of the last- recruited, large MUs, which fire near their threshold rate, compose the broadband frequency content of physiological muscle tremor in every contraction. Within this band, there is an additional distinct tremor component when MU correlations are present. For widespread and strong MU correlations, this component dominates and constitutes the observed regular tremor. In dynamic contractions, the firing of all MUs was modulated in the frequency band of both the sinusoidal and the complex variations of the force. The MU modulations showed a time-lead over the force variations and were strongly correlated both to these variations and among themselves. Thus widespread and strong correlations of MU firing modulations seem to provide a mechanism for generation of time-varying voluntary force, under general dynamic conditions. Finally, when regular tremor was present in dynamic contractions, widespread and fairly strong MU correlations also existed at the tremor frequency. It is concluded that at least two mechanisms can cause widespread MU synchrony, and they can act in parallel. They involve two types of correlated inputs to the alpha-motoneurons (presumably from the muscle spindles and the cortex), whose effects combine at the level of the membrane potential of the cells.     

Oxygen availability and motor unit activity in humans

Summary Six men were studied to determine the interrelationships among blood supply, motor unit (MU) activity and lactate concentrations during intermittent isometric contractions of the hand grip muscles. The subjects performed repeated contractions at 20% of maximal voluntary contraction (MVC) for 2 s followed by 2-s rest for 4 min with either unhindered blood circulation or arterial occlusion given between the 1st and 2nd min. The simultaneously recorded intramuscular MU spikes and surface electromyogram (EMG) data indicated that mean MU spike amplitude, firing frequency and the parameters of surface EMG power spectra (mean power frequency and root mean square amplitude) remained constant during the experiment with unhindered circulation, providing no electrophysiological signs of muscle fatigue. Significant increases in mean MU spike amplitude and frequency were, however, evident during the contractions with arterial occlusion. Similar patterns of significant changes in the surface EMG spectra parameters and venous lactate concentration were also observed, while the integrated force-time curves remained constant. These data would suggest that the metabolic state of the active muscles may have played an important role in the regulation of MU recruitment and rate coding patterns during exercise.     

Changes in conduction velocity,median frequency,and root mean square-amplitude of the electromyogram during 25% maximal voluntary contraction of the triceps brachii muscle,to limit of endurance

Summary The objective of the present study was to investigate whether isometric contraction of the right triceps brachii muscle, of maximal duration and at 25% of the maximal voluntary contraction (MVC), would reduce mean fibre conduction velocity (CV) for the active motor units (MU). In addition to the cross-correlation of surface electromyograms (EMG) for CV determination, median frequency (f _m) and root-mean-square amplitude (rms-amplitude) were calculated. The initial 5 min of the recovery of the three parameters was also investigated. The MVC were performed before and after the sustained contraction. Seven males — six in their twenties and one aged 43 - participated in the investigation. Mean CV for the unfatigued muscle was 4.5 m·s^–1, SD 0.38. On average, CV decreased less than 10% during the sustained contraction (P<0.05). Thef _m decreased almost linearly (46%) during the endurance time, while three quarters of the 250% increase in rms-amplitude took place during the last 50% of the contraction (P<0.001, both parameters). The MVC was reduced by 39% immediately after exhaustion (P<0.05). During the 1st min of recovery the rms-amplitude decreased by 50%, and the fm increased from 54% to 82% of the initial value (bothP<0.05. No measurable simultaneous CV restitution occurred. A parallel 15% increase inf _m and CV took place during the last 4 min of recovery (bothP<0.001), while the amplitude remained constant. Since mean CV was essentially unchanged during the last 50% of the endurance time where large changes inf _m and rms-amplitude occurred, factors supplementary to CV probably caused the striking changes in fatigue EMG, notably —MU recruitment, synchronization of MU activity, and lowering of MU firing frequencies. Nevertheless, during the last 4 min of recovery the entire increase inf _m could be accounted for by the simultaneous increase in CV.