Motor units within the normal rat medial gastrocnemius

The contractile properties, fatiguability and axonal conduction velocity were determined for 118 motor units in the medial gastrocnemius of eighteen rats. Fast-twitch and slow-twitch units could be categorized on the basis of the 'sag' test. For the purposes of statistical comparison the fast-twitch units were classified on the basis of their fatiguability as fast-fatiguable (FF), fast intermediate (FI) and fast fatigue-resistant (FR). As such, FR units tended to have a longer isometric twitch time course than other fast-twitch units. On the basis of peak tetanic force FF units were largest (mean 341 +/- 120 mN) followed by FI (145 +/- 85 mN), FR (87.3 +/- 38) and slow units (40.4 +/- 11.0 mN). There were no differences in motor axonal conduction velocity. Although units were categorized, it is clear that for all characteristics investigated the fast-twitch units exist as a continuum. Some of the largest FF units (peak force 350-500 mN) failed to maintain force with high-frequency stimulation (300 Hz) and this was associated with a failure of the EMG signal.     

Recruitment stability in masseter motor units during isometric voluntary contractions

Recruitment of single motor units (SMUs) of the masseter muscle was studied using macro representation (MacroRep) as the indicator of motor unit size. When subjects followed a slow isometric force ramp, units were usually recruited in order of MacroRep size. However, pooling the data from repeated ramps in the same subject resulted in a weak relationship between MacroRep size and force recruitment threshold, probably due to marked variations in the relative contributions of the jaw muscles, and varying levels of cocontraction, in the development of total bite force in each ramp. The force recruitment thresholds of individual SMUs showed marked variability, but recruitment threshold stability was improved when expressed as a percentage of maximum surface electromyographic (SEMG) activity in the ipsilateral masseter. Therefore the SEMG recruitment threshold was concluded to be a more stable and accurate indicator of the SMU's position in the recruitment hierarchy in a given muscle. It was concluded that SMUs in masseter are recruited according to the size principle, and that when investigating recruitment in jaw muscles, SEMG recruitment threshold should be used in preference to force recruitment threshold.     

Mechanical properties and behaviour of motor units in the tibialis anterior during voluntary contractions

The present work was carried out to analyse the properties and behaviour of Tibialis anterior motor units (MUs) during voluntary contractions in humans. A total of 528 single MU mechanical properties was recorded in 10 subjects by means of the spike-triggered averaging (STA) technique. MU recruitment thresholds and discharge frequencies were recorded during linearly increasing maximal voluntary contraction (MVC). The results indicate a mean (+/- SD) MU torque of 25.5 +/- 21.5 mN.m. and a mean time-to-peak of 45.6 +/- 13.6 ms. A comparison of the average MU twitch torque with that of the muscle allowed an estimate of about 300 MUs in the Tibialis anterior. A positive linear relationship was recorded between the MU twitch torque and the recruitment threshold. The mean minimal and maximal discharge frequencies of MUs were 8.4 +/- 3.0 Hz and 33.2 +/- 14.7 Hz, respectively. The results of the present work indicate that MU behaviour during voluntary contractions is different in the tibialis anterior and in the adductor pollicis.     

Hypertrophy of rat plantaris muscle fibers after voluntary running with increasing loads

There have been no systematic comparisons of skeletal muscle adaptations in response to voluntary wheel running under controlled loading conditions. To accomplish this, a voluntary running wheel for rats and mice was developed in which a known load can be controlled and monitored electronically. Five-week-old male Sprague-Dawley rats (10 rats/group) were assigned randomly to either a 1) sedentary control group (Control); 2) voluntary exercised with no load (Run-No-Load) group; or 3) voluntary exercised with additional load (Run-Load) group for 8 wk. The load for the Run-Load group was progressively increased to reach approximately 60% of body weight during the last week of training. The proportions of fast glycolytic (FG), fast oxidative glycolytic (FOG), or slow oxidative (SO) fibers in the plantaris were similar in all groups. The absolute and relative plantaris weights were greater in the Run-Load group compared with the Control and Run-No-Load groups. The mean fiber cross-sectional areas of FG, FOG, and SO fibers were 20, 25, and 15% greater in the Run-Load than in Control rats. In addition, these fiber types were 16, 21, and 12% larger in Run-Load than in Run-No-Load rats. The muscle weights and mean cross-sectional areas of each fiber type were highly correlated with the average running distances and total work performed in the Run-Load, but not the Run-No-Load, group. The slope of the relationship between fiber size and running distance and total work performed was significant for each fiber type but was higher for FG and FOG fibers compared with SO fibers. These data show that the load on a rat running voluntarily can determine the magnitude of a hypertrophic response and the population of motor units that are recruited to perform at a given loading condition.     

Composition of newly forming motor units in prenatal rat intercostal muscle

We have examined the composition of rat intercostal motor units during the period of late gestation, when most muscle fibres are formed, in order to see the pattern of the contacts initially made between single motoneurons and myotubes. At this early stage, the muscle contains two types of myotubes, primary and secondary myotubes, and a major aim was to see whether individual motoneurons preferentially made contact with a particular myotube type. The technique used to define myotubes contacted by a single motoneuron was anterograde labelling of the neuron, followed by electron microscopic detection of labelled terminals and their postsynaptic targets. We find that prenatal motor units are inhomogeneous with respect to their primary/secondary myotube composition. Most individual motoneurons show many permutations of contact with primary myotubes, secondary myotubes, and undifferentiated cells, including single nerve terminals which contact both primary and secondary myotubes. Our results are interpreted in terms of changes to the composition of both the muscle and of the motor units during the final 5 days of gestation. We demonstrate that motoneurons necessarily make their initial contacts on primary myotubes, but that these are surprisingly sparse. As secondary myotubes appear and become innervated, motor units are at first all similar and all heterogeneous. However, primary myotubes are represented more often in motor units than in the muscle as a whole. This probably reflects the relative densities of polyinnervation of primary vs. secondary myotubes. By embryonic day 20, motor units have become divergent in composition, with some dominated by primary myotubes and others by secondaries. We propose that motoneurons initially establish contacts at random on either myotube type, but then begin to express preference for one type or the other and reorganise their periphery. Refining of motor unit composition towards homogeneity in the postnatal period probably involves other elements, such as mutability of muscle fibre and/or motoneuron characteristics as a function of usage and muscle position, perhaps influenced by sensory feedback mechanisms.     

The orderly recruitment of motor units of the masseter and temporal muscles during voluntary isometric contraction in man

1. The contractile properties of the motor units of the masseter and temporal muscles of human subjects were studied during voluntary isometric contractions, using a method previously employed to examine units of a small hand muscle. 2. Over the range of forces studied (0-6 kg), the units of both muscles were recruited in an orderly fashion, with a nearly linear relationship between the voluntary force at which units were recruited and their measured twitch tensions. 3. The range of contraction times (25-90 msec) was similar to that observed for the hand muscle. In some subjects it seemed that small units, recruited at low forces, exhibited shorter contraction times.     

A longitudinal study of the pathophysiological changes in single human thenar motor units in amyotrophic lateral sclerosis

The sequence of pathophysiological changes in amyotrophic lateral sclerosis (ALS) at the single motor unit (MU) level is not well understood. Using a recently described technique, a comprehensive range of physiological properties in two thenar MUs in ALS were intensively studied. In the first MU, despite a marked decline in the ability of the subject to voluntarily recruit the MU, the physiological properties of this MU remained remarkably stable over a 2-year period. In contrast, the physiological properties of the other MU declined rapidly over 5 months despite the fact that this MU could be recruited with ease throughout the study period. These differences between the progressively dysfunctional changes in these two MUs illustrates the value of such longitudinal studies of specific MUs in improving our understanding of the evolution of changes in single motoneurons in ALS. The broader application of longitudinally tracking the pathophysiological changes of the surviving MUs may prove to be a sensitive measure of disease progression and in evaluating the effectiveness of treatments.     

The reorganization of motor units in motor neuron disease

We studied 78 patients with motor neuron disease (MND) using concentric needle electromyography. Analysis on weak and maximal effort was performed using our own, fully automated, computer method, EMG-LAB. In addition to the conventional parameters of single motor unit action potentials (MUAPs) and interference pattern, new criteria were applied: the range of the acting motor units and the functional recruitment order. A total of 375 muscles of MND patients and 120 control muscles were investigated. The electromyographic data were analyzed separately in five groups of muscles, classified A, B, C, D, and E according to their clinical condition. Those results allowed us to discern six neurophysiological stages (N(0,1,2,3,4,5)) from the early to the most advanced phase. It has been confirmed that reinnervation in MND is adequate to compensate for the loss of over 50% of motor neurons but it is only a transitory phase in the morbid course. At stages N(O-5), the electrophysiological data reflect structural and functional integrity of the functioning motor units. Evaluation of not only single MUAPs but also of the full range of acting motor units and their recruitment order allowed a deeper look into the underlying pathophysiological mechanisms.     

Medial gastrocnemius is more activated than lateral gastrocnemius in sural nerve induced reflexes during human gait

In humans the sural nerve was stimulated at one of 16 phases of the step cycle. In MG (medial gastrocnemius) the amplitude of the P2 responses (latency 80-93 ms) was on average 1.3 times larger than the corresponding background activity while this was 0.9 for LG (lateral gastrocnemius; predominantly suppressive responses). It is speculated that such differences contribute to an exorotation moment during gait.     

Neuromuscular contacts of expanded motor units in rat soleus muscles are rescued by leupeptin

In the soleus muscle of the rat following section of the L5 ventral ramus (partial denervation) the remaining motor axons increase their territory by sprouting. Nerve sprouts are first seen two to three days after the operation, their number peaks at 10-14 days and subsequently remains at this level. The time course of the initial sprouting in partially denervated muscles is not altered by paralysing the muscles with alpha-bungarotoxin, and the initial extent of the sprouting is, if anything, greater in the paralysed muscles. However, unlike in controls, this level of sprouting is not maintained and neuromuscular contacts are lost when muscles recover from the paralysis. The loss of these contacts can be prevented by treatment of these partially denervated paralysed muscles with leupeptin, an inhibitor of calcium-activated neutral protease. Interestingly, more contacts are rescued when leupeptin is applied 10 days after alpha-bungarotoxin treatment, when sprouting has reached high levels, than at three days, when sprouting has just begun. The neuromuscular connections rescued by leupeptin are functional. Maximum tetanic tension produced by untreated soleus muscles two to five months after partial denervation is 66 +/- 9% of contralateral control muscles, but only 39 +/- 8% when the muscles were paralysed with alpha-bungarotoxin for 12-14 days after partial denervation. However, when partially denervated paralysed muscles were treated with leupeptin three and 10 days after alpha-bungarotoxin treatment their tension output is 74 +/- 3% and 81 +/- 8%, respectively. After partial denervation alone, motor units are twice their normal size. Short-term paralysis with alpha-bungarotoxin prevents this increase in motor unit territory. However, the application of leupeptin to the paralysed muscles rescues neuromuscular contacts, allowing motor unit size to remain expanded, at around 2-2.5-fold. Thus, following recovery from temporary paralysis with alpha-bungarotoxin, there is a sudden withdrawal of neuromuscular contacts and these can be rescued by treatment with leupeptin.     

Rank-ordered regulation of motor units

Myoelectric signals were detected from the tibialis anterior muscle of 5 subjects with a quadrifilar needle electrode while the subjects generated isometric forces that increased linearly with time (10% of maximal voluntary contraction/s) up to maximal voluntary level. Motor unit firing rates were studied as a function of force throughout the full range of muscle force output. The relationship between force and firing rate was found to contain three distinct regions. At recruitment and near maximal force levels, firing rates increased more rapidly with force than in the intermediate region. Furthermore, in the regions with rapid increases, the rate of change of firing rate was correlated to the recruitment threshold, with higher recruitment threshold motor units displaying greater rates of change. In the intermediate region, all motor units had similar rates of change of firing rate. A weak positive correlation was found between initial firing rate and recruitment threshold. Firing rates of motor units at any instant were found to be ordered according to the recruitment order: at any given time in the contraction motor units with lower recruitment thresholds had higher firing rates than units with higher recruitment thresholds. Firing rates of all motor units were observed to converge to the same value at maximal forces. Mechanisms underlying motor unit recruitment and firing rate modulation are discussed in the context of a conceptual model.     

The voluntary control of facial action units in adults.

We investigated adults' voluntary control of 20 facial action units theoretically associated with 6 basic emotions (happiness, fear, anger, surprise, sadness, and disgust). Twenty young adults were shown video excerpts of facial action units and asked to reproduce them as accurately as possible. Facial Action Coding System (FACS; Ekman & Friesen, 1978a) coding of the facial productions showed that young adults succeeded in activating 18 of the 20 target actions units, although they often coactivated other action units. Voluntary control was clearly better for some action units than for others, with a pattern of differences between action units consistent with previous work in children and adolescents. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Loss of motor units in Alzheimer's disease

Little attention has been paid in the literature to the state of the peripheral nervous system (PNS) in Alzheimer's disease (AD). We conducted a comprehensive electrophysiological study in 15 AD patients looking for functional abnormalities within their PNSs. A reduction of the number of functioning motor units (MU) was found in the thenar and soleus muscles of most of these patients without enlargement of the remaining MUs territories, while the motor and sensory conduction velocities of the peripheral nerves were preserved. These results suggest dysfunction of the spinal motoneurones in patients afflicted with this condition.     

Responses of human single motor units to transcranial magnetic stimulation

The authors analyse the results of 51 unicompartmental knee prostheses with 1 to 12 years follow-up (mean follow-up: 5 years). The results were evaluated using the scoring system of the "Knee Group" of the SO.B.C.O.T. (Société Belge de Chirurgie Orthopédique et de Traumatologie). This analysis demonstrates that the quality of the results depends on implant positioning. The authors suggest positioning the tibial implant parallel with the healthy plateau and slightly distal, i.e. to position the tibial implant perpendicular to the epiphyseal axis and not to the mechanical axis, as is systematically done with the usual tibial cutting guides. When this ideal positioning was respected, 77.5% of the patients had a score above 90 points (out of a possible maximum of 100 points) and 12.5% had a score between 75 and 89 points. When this condition was not respected, none of the knees obtained more than 75 points. The difference was statistically significant (p = 0.0001).     

Directional tuning of single motor units

The directional activity of whole muscles has been shown to be broadly and often multimodally tuned, raising the question of how this tuning is subserved at the level of single motor units (SMUs). Previously defined rules of SMU activation would predict that units of the same muscle (or at least of the same neuromuscular compartment) are activated homogeneously with activity peaks in the same "best" direction(s). In the present study, the best directions of SMUs in human biceps (both heads) and deltoid (anterior, medial, and posterior portions) were determined by measuring the firing rate and threshold force of units for recruitment during isometric force ramps in many different directions. For all muscles studied, neighboring motor units could have significantly different best directions, suggesting that each muscle receives multiple directional commands. Furthermore, 17% of the units sampled clearly had a second-best direction, consistent with a convergence of different directional commands onto the same motoneuron. The best directions of the units changed gradually with location in the muscle. Best directions did not cluster into separate groups, thus, not supporting the existence of clearly distinguished neuromuscular compartments. Instead, the results reveal a more gradually distributed activation of the biceps and deltoid motoneuron pools. A model is proposed in which the central control mechanism optimizes the fulfillment of the continuously changing directional force requirements of a movement by gradually recruiting and derecruiting those units ideally suited for the production of the required force vector at any given time.     

Heteronymous H-reflex in temporal muscle motor units

An electric stimulation of the masseteric nerve elicits a heteronymous H-reflex in the temporal muscle. The characteristics of this reflex response were investigated by analysis of the firing probability changes of single motor units. Eleven healthy subjects participated in the experiments. The heteronymous H-reflex of the temporal muscle was electrically elicited by stimulation of the masseteric nerve at 120% of the intensity needed for the maximal masseteric M-wave. From 8 to 24 motor units were sampled from the temporal muscle of each subject. Peri-stimulus time histograms of motor unit recordings were built with a 0.5-ms bin width. The mean firing probability was calculated for the 20 ms preceding the stimulus. The firing probability was considered increased when it exceeded the mean by 3 standard deviations. Of 104 sampled motor units, 40 motor units showed a significant increase of the firing probability, which lasted 1 ms or less in 29 of them. In 12 out of 16 motor units, a significant increase of firing probability also persisted at a lower stimulation intensity (120% of the threshold needed to elicit a masseteric M wave). These data indicate that: (1) some temporal muscle motor units are modulated by afferents from the masseter muscle, (2) the heteronymous H-reflex has a monosynaptic component, and (3) there might be a more complex than just monosynaptic organization serving the heteronymous temporal H-reflex. For the latter conclusion regarding synaptic wiring, however, PSTH studies like the present one can offer only indirect evidence, and this question could be better studied in animals.     

Organization of single motor units in feline sartorius

1. We depleted single motor units in feline sartorius muscles of glycogen by stimulating their motoneurons intracellularly. We mapped the intramuscular distribution of depleted fibers by inspecting histological cross-sections throughout the length of sartorius. 2. We selected ten depleted motor units for detailed study and quantitative analysis. Nine motor units were located in the anterior head of sartorius. One was located in a muscle whose distal half appeared to have been damaged some time before the acute experiment. A single motor unit was located in the medial head of sartorius. 3. Five motor units were composed of fast-twitch glycolytic (FG) muscle fibers, two of fast-twitch oxidative glycolytic (FOG) muscle fibers, and three of slow-twitch oxidative (SO) muscle fibers. Estimates of the numbers of depleted fibers in motor units of anterior sartorius indicated that FG motor units were larger (mean 566 fibers) than FOG and SO motor units (SO mean 190, FOG mean 156 fibers). The SO motor unit in the damaged muscle had 550 fibers. One motor unit depleted in the medial head of sartorius had 270 fibers with FG profiles. 4. Muscle fibers belonging to each anterior motor unit were never distributed throughout the whole cross-section of anterior sartorius at any proximodistal level. Furthermore, fibers were distributed nonuniformly along the proximodistal axis of the muscle. In most muscles at least a few depleted fibers were found at all proximodistal levels. However, in one normal muscle and the damaged muscle, depleted fibers were confined to the proximal end. 5. The fibers in the medial motor unit were confined to a strip that did not extend across the whole cross-section of the muscle head. Fibers within this strip were scattered quite evenly from origin to insertion. This medial FG motor unit occupied a smaller territory and contained fewer fibers than anterior motor units of the same histochemical type. 6. These results show that sartorius motor units are not distributed uniformly in the mediolateral plane; those in anterior sartorius were distributed asymmetrically in the proximodistal axis as well. This finding has important functional implications for the way in which we model force development and transmission in sartorius and other long muscles.     

Measures of "fastness": force profiles of twitches and partly fused contractions in rat medial gastrocnemius and tibialis anterior muscle units

Recordings of isometric force were obtained for twitches and (sub)maximal tetani of gastrocnemius medialis (MG) and tibialis anterior (TA) muscle units in female Wistar rats. We assessed the relationships between unit properties that have all been associated with "speed": (1) the relative degree of peak force attained during repetitive activation at 40 Hz (P40/Pmax), (2) the relative degree of final twitch fusion during the same test burst (Fus-end), and (3) various measures of the time-course of single twitches, including twitch time-to-peak and a parameter referred to as "initial fusion ratio" (Fus-in; relative decline from peak force at 25 ms from twitch onset). The various measures of twitch time-course were significantly correlated to each other with correlation coefficients varying over a fairly wide range (0.35-0.64 for MG; 0.50-0.80 for TA). Twitch time-course was also significantly correlated with Fus-end during the 40-Hz repetitive activation; the highest correlation coefficient (0.69 for MG, 0.80 for TA) was obtained for Fus-in, which was also numerically similar to Fus-end. Thus, the degree of fusion indeed seemed to be largely dependent upon aspects of twitch time-course. However, the relative degree of force mobilization obtained in the same contractions elicited by stimulation at 40 Hz was not consistently better correlated with Fus-end than with measures of single twitch time-course. Furthermore, in fast-twitch units having the same twitch time-to-peak, the force mobilization elicited by stimulation at 40 Hz (P40/Pmax) was the same for MG and TA, while the degree of fusion was significantly smaller for TA than for MG units. The results demonstrate the complexity of the concept of isometric "speed" and underline the need for using several speed indicators in parallel in studies concerning the differentiation of muscle (unit) properties.