Motor-unit recruitment in self-reinnervated muscle

1. Recruitment order of motor units in self-reinnervated medial gastrocnemius (MG) muscles was studied in decerebrate cats 16 mo after surgical reunion of the cut MG nerve. Pairs of MG motor units were isolated by dual microelectrode penetration of ventral roots to measure their recruitment sequence during cutaneous reflexes in relation to their physiological properties. 2. Physiological properties of reconstituted motor units appeared normal, as expected. Also normal were the relationships among these properties: twitch and tetanic tension tended to increase with axonal conduction velocity and decrease with twitch contraction time. A small fraction of motor units (10/116) in reinnervated muscles produced either no measurable tension or unusually large amounts of tension compared with controls. This was the only distinct feature of the sample of reconstituted units. 3. In muscles reinnervated after nerve section, stretch was notably ineffective in eliciting reflex contraction of MG muscles or their constituent motor units (only 5/116 units). Incomplete recovery from nerve section was probably the cause of this impairment, because stretch reflexes were readily evoked in adjacent untreated muscles and in one reinnervated MG muscle that was studied 16 mo after nerve crush. In contrast with the ineffectiveness of muscle stretch, sural nerve stimulation succeeded in recruiting 49/116 units, a proportion fairly typical of normal MG muscles. 4. The contractions of the first unit recruited in cutaneous reflexes tended to be slower and less forceful than those of the other unit in a pair. By these measures, recruitment obeyed the size principle. This recruitment order with respect to unit contractile properties was not significantly different (P > 0.05) between untreated and reinnervated muscles but was significantly (P < 0.005) different from random order in both groups. The same recruitment pattern was observed for pairs of motor units sampled from the muscle reinnervated after nerve crush, whether units were recruited by muscle stretch or sural nerve stimulation. 5. The usual tendency for motor units with slower conduction velocity (CV) to be recruited in sural nerve reflexes before those with faster CV was not strong in reinnervated muscles. After nerve section the proportion of units exhibiting the usual recruitment pattern was not significantly different (P > 0.05) from a random pattern for CV. 6. The central finding is that the normal recruitment patterns recover from nerve injury in a muscle that is reinnervated by its original nerve. By contrast, stretch reflexes do not recover well from nerve section, and this deficiency may contribute to motor disability.     

Fast-to-slow conversion following chronic low-frequency activation of medial gastrocnemius muscle in cats. II. Motoneuron properties

Chronic stimulation (for 2-3 mo) of the medial gastrocnemius (MG) muscle nerve by indwelling electrodes renders the normally heterogeneous MG muscle mechanically and histochemically slow (type SO). We tested the hypothesis that motoneurons of MG muscle thus made type SO by chronic stimulation would also convert to slow phenotype. Properties of all single muscle units became homogeneously type SO (slowly contracting, nonfatiguing, nonsagging contraction during tetanic activation). Motoneuron electrical properties were also modified in the direction of type S, fatigue-resistant motor units. Two separate populations were identified (on the basis of afterhyperpolarization, rheobase, and input resistance) that likely correspond to motoneurons that had been fast (type F) or type S before stimulation. Type F motoneurons, although modified by chronic stimulation, were not converted to the type S phenotype, despite apparent complete conversion of their muscle units to the slow oxidative type (type SO). Muscle units of the former type F motor units were faster and/or more powerful than those of the former type S motor units, indicating some intrinsic regulation of motor unit properties. Experiments in which chronic stimulation was applied to the MG nerve cross-regenerated into skin yielded changes in motoneuron properties similar to those above, suggesting that muscle was not essential for the effects observed. Modulation of group Ia excitatory postsynaptic potential (EPSP) amplitude during high-frequency trains, which in normal MG motoneurons can be either positive or negative, was negative in 48 of 49 chronically stimulated motoneurons. Negative modulation is characteristic of EPSPs in motoneurons of most fatigue-resistant motor units. The general hypothesis of a periphery-to-motoneuron retrograde mechanism was supported, although the degree of control exerted by the periphery may vary: natural type SO muscle appears especially competent to modify motoneuron properties. We speculate that activity-dependent regulation of the neurotrophin-(NT) 4/5 in muscle plays an important role in controlling muscle and motoneuron properties.     

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.     

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 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.     

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.     

Motor-unit behavior in humans during fatiguing arm movements

1. The activity of 40 triceps brachii motor units was recorded from the dominant arms of 9 healthy adult volunteers (age 27.8 +/- 4.4 yr, mean +/- SD) during a fatigue task that included both isometric and anisometric contractions. The fatigue task lasted 8.3 min and consisted of 50 extension and 50 flexion movements of the elbow. Each movement (40 degrees in 0.8s) was separated by an isometric contraction. A constant load resisting extension of 17.7 +/- 3.0% of maximal voluntary contractions (MVC) was applied throughout the task. This paradigm enabled the direct contrast of motor-unit discharge behavior during the different types of fatiguing contractions. 2. Motor-unit behavior was examined to determine the relative contribution of two mechanisms for optimizing force production under fatiguing conditions: recruitment of motor units and modulation of motor-unit discharge following recruitment. Threshold torques for motor-unit recruitment thresholds were determined by ramp-and-hold isometric contractions. Motor-unit discharge was evaluated during the fatigue task by contrasting the number of motor-unit potentials (spikes) per contraction for concentric eccentric, and isometric contractions. 3. The fatigue task resulted in a 30 +/- 12% decline in the mean MVC of elbow extension. Recruitment of nine new motor units (23%) was evident during the fatiguing extension movements, often within five to seven movements (i.e., within 25-35 s). Each newly recruited motor unit had the largest recruitment threshold torque in that experiment. 4. Analysis of the motor units that were active from the beginning of the fatigue task revealed that the mean number of motor-unit spikes per contraction increased, or remained constant as fatigue ensued, yet for the majority of motor units it increased or remained constant. None of the newly recruited motor units demonstrated decreased number of mean spikes per contraction after recruitment. Further, concurrently active motor units displayed different discharge behavior in two-thirds of the subjects. It is proposed that if the neural drive to the muscle is distributed uniformly upon the motoneuron pool, peripheral feedback from the exercising muscle may modulate specific motoneuron discharge levels during fatigue.     

Extraocular motor unit and whole-muscle responses in the lateral rectus muscle of the squirrel monkey

Because primate studies provide data for the current experimental models of the human oculomotor system, we investigated the relationship of lateral rectus muscle motoneuron firing to muscle unit contractile characteristics in the squirrel monkey. Also examined was the correlation of whole-muscle contractile force with the degree of evoked eye displacement. A force transducer was used to record lateral rectus whole-muscle or muscle unit contraction in response to abducens whole-nerve stimulation or stimulation of single abducens motoneurons or axons. Horizontal eye displacement was recorded using a magnetic search coil. (1) Motor units could be categorized based on contraction speed (fusion frequency) and fatigue. (2) The kt value (change in motoneuronal firing necessary to increase motor unit force by 1.0 mg) of the units correlated with maximum tetanic tension. (3) There was some tendency for maximum tetanic tension of this unit population to separate into three groups. (4) At a constant frequency of 100 Hz, 95% of the motor units demonstrated significantly different force levels dependent on immediately previous stimulation history (hysteresis). (5) A mean force change of 0.32 gm/ degrees and a mean frequency change of 4.7 Hz/ degrees of eye displacement were observed in response to whole-nerve stimulation. These quantitative data provide the first contractile measures of primate extraocular motor units. Models of eye movement dynamics may need to consider the nonlinear transformations observed between stimulation rate and muscle tension as well as the probability that as few as two to three motor units can deviate the eye 1 degrees.     

Average but not continuous speed match between motoneurons and muscle units of rat tibialis anterior

1. Properties of single motoneuron/muscle-unit combinations were determined for tibialis anterior (TA) in rats anesthetized with pentobarbital. The TA observations were systematically compared with those obtained earlier by the use of the same techniques from rat medial gastrocnemius (MG). 2. TA motoneurons were investigated with regard to afterhyperpolarization (AHP; total duration 32-74 ms, amplitude 0.39-4.96 mV) and axonal conduction velocity (41-79 m/s). TA muscle-unit measurements included the time course of the isometric twitch (time-to-peak force 10.8-18.0 ms; total duration 42-92 ms), the maximum tetanic force (22-217 mN), and a measure of fatigue sensitivity (fatigue index 5-100%). The range of twitch and AHP durations ("speed range") was markedly smaller in the present TA material than for MG. 3. The mean duration of the TA motoneuronal AHP (49 +/- 8 ms, mean +/- SD) was close to that of its muscle-unit twitch (56 +/- 12 ms). Thus an "average" speed match existed between TA motoneurons and their muscle fibers. 4. For TA there was no correlation between the time courses of AHP and twitch. Thus there was for TA no "continuous" speed match between the motoneurons and their muscle fibers. 5. For TA twitches or AHPs studied separately, there was a significant correlation between different time course measures. Furthermore, compared with TA units having relatively fast twitches, those with slower twitches tended to show 1) a smaller maximum tetanic force and 2) a greater AHP amplitude. Fatigue-resistant units tended to have slower twitches than fatigue-sensitive ones.(ABSTRACT TRUNCATED AT 250 WORDS)     

1998 ISEK Congress Keynote Lecture: Motor units: how many, how large, what kind? International Society of Electrophysiology and Kinesiology

There are now at least nine methods for motor unit number estimation (MUNE) in living human muscles. All methods are based on the comparison of an average single motor unit potential (or twitch) with the response of the whole muscle. Such estimations have been performed for proximal and distal muscles of the arm and leg in healthy subjects and in patients with various neuromuscular disorders. In healthy subjects there is a loss of motor units which is most evident in distal muscles and after the age of 60 years. Substantial losses of motor units have been measured in patients with ALS, post-polio symptoms, and diabetic peripheral neuropathies. In contrast, normal MUNEs have been found in approximately half of patients with persisting obstetric brachial palsies. The sizes of motor units show considerable variations within the same muscle and also between muscles; very large units are usually present in severe partial denervation. Although many motor unit properties are largely governed by motoneurons, some exhibit less plasticity in humans than in other mammals.     

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.     

123I-MIBG myocardial scintigraphy in diabetic patients: relationship with 201Tl uptake and cardiac autonomic function

The purpose of this study is to investigate the effects of aging on the human stretch reflexes. The EMG and torque responses of the stretch reflex of the wrist flexors were evoked by ramp-and-hold mechanical perturbations. The stretch reflexes were recorded at seven test conditions with different stretch velocity and muscle preload. The test results from young and older healthy adult subjects were compared. In average, the absolute amplitude of the short-latency (20-40 ms) EMG (recorded from flexor carpi radialis) reflex response was significantly lower in the older group. If the data were normalized and expressed in percentage of the maximal voluntary EMG activity, however, this group difference was not significant. There was no change in the reflex gain of the short-latency reflex with aging. For the long-latency (50-90 ms) EMG reflex response, both the normalized amplitude and the reflex gain were significantly enhanced with aging, probably through supraspinal mechanisms. There was no significant difference in the threshold velocity for the evoked EMG reflexive activities between age groups. There were also no changes in the reflexive wrist flexion torque with aging. These results suggested that the number of motor units recruited during the stretch reflex activity declined with aging although the percentage of motor units recruited was not affected by aging. It is concluded that the central regulating mechanisms of the spinal motoneuron excitability are not compromised by aging. The automatic gain compensation phenomenon is also preserved with aging.     

Motor unit recruitment during prolonged isometric contractions

Motor unit recruitment patterns were studied during prolonged isometric contraction using fine wire electrodes. Single motor unit potentials were recorded from the brachial biceps muscle of eight male subjects, during isometric endurance experiments conducted at relative workloads corresponding to 10% and 40% of maximal voluntary contraction (MVC), respectively. The recordings from the 10% MVC experiment demonstrated a characteristic time-dependent recruitment. As the contraction progressed both the mean number of motor unit spikes counted and the mean amplitude of the spikes increased significantly (P < 0.01). This progressive increase in spike activity was the result of a discontinuous process with periods of increasing and decreasing activity. The phenomenon in which newly recruited motor units replace previously active units is termed "motor unit rotation" and appeared to be an important characteristic of motor control during a prolonged low level contraction. In contrast to the 10% MVC experiment, there was no indication of de novo recruitment in the 40% MVC experiment. Near the point of exhaustion a marked change in action potential shape and duration dominated the recordings. These findings demonstrate a conspicuous difference in the patterns of motor unit recruitment during a 10% and a 40% MVC sustained contraction. It is suggested that there is a close relationship between intrinsic muscle properties and central nervous system recruitment strategies which is entirely different in fatiguing high and low level isometric contractions.     

Fast-to-slow conversion following chronic low-frequency activation of medial gastrocnemius muscle in cats. I. Muscle and motor unit properties

This study of cat medial gastrocnemius (MG) muscle and motor unit (MU) properties tests the hypothesis that the normal ranges of MU contractile force, endurance, and speed are directly associated with the amount of neuromuscular activity normally experienced by each MU. We synchronously activated all MUs in the MG muscle with the same activity (20 Hz in a 50% duty cycle) and asked whether conversion of whole muscle contractile properties is associated with loss of the normal heterogeneity in MU properties. Chronically implanted cuff electrodes on the nerve to MG muscle were used for 24-h/day stimulation and for monitoring progressive changes in contractile force, endurance, and speed by periodic recording of maximal isometric twitch and tetanic contractions under halothane anesthesia. Chronic low-frequency stimulation slowed muscle contractions and made them weaker, and increased muscle endurance. The most rapid and least variable response to stimulation was a decline in force output of the muscle and constituent MUs. Fatigue resistance increased more slowly, whereas the increase in time to peak force varied most widely between animals and occurred with a longer time course than either force or endurance. Changes in contractile force, endurance, and speed of the whole MG muscle accurately reflected changes in the properties of the constituent MUs both in extent and time course. Normally there is a 100-fold range in tetanic force and a 10-fold range in fatigue indexes and twitch time to peak force. After chronic stimulation, the range in these properties was significantly reduced and, even in MU samples from single animals, the range was shown to correspond with the slow (type S) MUs of the normal MG. In no case was the range reduced to less than the type S range. The same results were obtained when the same chronic stimulation pattern of 20 Hz/50% duty cycle was imposed on paralyzed muscles after hemisection and unilateral deafferentation. The findings that the properties of MUs still varied within the normal range of type S MUs and were still heterogeneous despite a decline in the variance in any one property indicate that the neuromuscular activity can account only in part for the wide range of muscle properties. It is concluded that the normal range of properties within MU types reflects an intrinsic regulation of properties in the multinucleated muscle fibers.     

Reversal of the influence of group Ib afferents from plantaris on activity in medial gastrocnemius muscle during locomotor activity

1. Rhythmic locomotor activity was evoked in clonidine-treated acute and chronic spinal cats, and the effect of stimulating group I afferents from the plantaris muscle on the timing and magnitude of bursts in medial gastrocnemius (MG) motoneurons was examined. 2. The locomotor rhythm was entrained when group I afferents in the plantaris nerve were electrically stimulated with trains of stimuli presented at rates above and below the intrinsic frequency of the rhythmic activity. During entrainment at rates higher than the intrinsic frequency, a burst of activity in ipsilateral MG motoneurons was initiated approximately 40 ms after the onset of each stimulus train. At lower rates of entrainment the onset of MG bursts preceded the onset of the stimulus trains, and each stimulus train had an excitatory effect on the MG burst with a latency in the range of 30-50 ms. A similar excitatory effect was observed when the stimulus trains were triggered at a preset delay after the endogenous generation of the MG bursts. 3. The excitatory action of plantaris group I afferents on the MG motoneurons was only seen during periods of locomotor activity. In the absence of rhythmic activity, the same stimulus trains reduced any ongoing tonic activity in MG motoneurons. 4. Vibration of the plantaris muscle to preferentially activate group Ia afferents neither entrained the locomotor rhythm nor increased the magnitude of the MG bursts. 5. We conclude that during locomotor activity, input from group Ib afferents of the plantaris muscle has an excitatory action on the system of interneurons generating the extensor bursts, i.e., on the extensor half-center of the central rhythm generator.(ABSTRACT TRUNCATED AT 400 WORDS)     

The influence of age on the assessment of motor unit activation in a human hand muscle

Two indices of motor unit recruitment, the ramp-force and repetitive-discharge thresholds, were compared in the first dorsal interosseus muscle of ten young and twelve elderly subjects. The purpose was to determine the effect of age on the relationship between the two recruitment thresholds and the spike-triggered average force of motor units. Each subject performed three tasks requiring isometric abduction of the left index finger: a maximum voluntary contraction (MVC), a ramp-and-hold contraction, and a repetitive-discharge task. The elderly subjects used coactivation of the antagonist muscle (second palmar interosseus) more frequently than the young subjects during the ramp-and-hold contraction. Many elderly subjects expressed difficulty with the controlled ramp-down phase of the ramp-and-hold contraction and preferred a coactivation strategy to a derecruitment strategy for this task. There were no differences due to age or gender in the ramp-force thresholds between the various groups. However, the normalized repetitive-discharge threshold was significantly less for the younger subjects and for the male subjects. Nonetheless, the two recruitment thresholds were able to predict the spike-triggered average force with similar success for both the young and the elderly subjects. These data suggest that the recruitment threshold of a motor unit in first dorsal interosseus was characterized equally well by either the ramp-force or repetitive-discharge measurement for both young and elderly subjects but that coactivation was used more frequently by the elderly subjects during the ramp-and-hold task.     

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.     

Lateral rectus whole muscle and motor unit contractile measures with the extraocular muscles intact

Both extracellular and intracellular stimulation of single motoneurons were shown to be similarly effective and consistent in eliciting contractile responses in single lateral rectus muscle motor units. The whole muscle was activated by stimulating the sixth nerve in the brain stem. Both whole muscle and motor unit contractile characteristics, under isometric conditions, were found to remain consistent regardless of whether this extraocular muscle was detached or left attached to the globe. In addition, whole muscle twitch and maximum tetanic tension evoked by sixth nerve stimulation was significantly less than would be predicted by the linear summation of individual motor unit twitch and maximum tetanic tensions.     

Task-dependent facilitation of motor evoked potentials during dynamic and steady muscle contractions

Task-dependent differences in the facilitation of motor evoked potentials (MEPs) following cortex stimulation were studied in a proximal (deltoid) and a distal muscle (abductor digiti minimi; ADM) in 23 healthy subjects during both dynamic and steady contractions of the target muscle under isometric and under nonisometric conditions. In the deltoid, MEP amplitudes were significantly greater if stimulation was performed during dynamic contractions than during steady contractions, despite equal background electromyographic levels just prior to the stimulus. The same task-specific extra facilitation of deltoid MEP amplitudes was also found with magnetic stimulation of the brain stem instead of the cortex in 3 subjects. In the ADM, no such task-dependent extra facilitation of MEPs during dynamic contractions was found. It is concluded that in the deltoid, during dynamic contractions, a greater proportion of the spinal motoneurons is close to depolarization threshold (greater "subliminal fringe") whereas the number of firing motoneurons is similar to that during steady contraction. The lack of task-dependent extra facilitation of MEPs in the ADM is explained by the predominant recruitment principle for force gradation in small hand muscles, which is in contrast to the predominant frequency principle used in proximal muscles.     

Muscle fiber length and moment arm coordination during dorsi- and plantarflexion in the mouse hindlimb

The purpose of this study was to test the hypothesis that muscle fiber length and joint moment arm are combined in such a way that maximum muscle force is produced during locomotion. Plantarflexor (soleus, SOL and medial gastrocnemius, MG) and dorsiflexor (extensor digitorum longus, EDL and tibialis anterior, TA) muscle architecture in the mouse was measured along with their associated moment arms. Fiber length varied significantly between muscles ranging from 5.7 +/- 0.2 mm (MG) to 7.6 +/- 0.2 mm (TA). Plantarflexor moment arms were over twice as large as dorsiflexor moment arms (1.88 +/- 0.06 mm vs. 0.84 +/- 0.03 mm) suggesting a greater muscle length change with joint angle for plantarflexors compared to dorsiflexors. Using a simple muscle-joint model, the active sarcomere length range in these muscle groups was calculated and proved to be quite similar between functional groups. The active range for dorsiflexors was 2.2-2.4 microns, while the active range for plantarflexors was 2.2-2.5 microns, indicating that both muscle groups operate primarily near the plateau of their length tension-relation. Finally, when calculating force produced by muscles during locomotion, the combination of moment arm and fiber length measured in all muscle groups yielded muscle-joint systems that produced near maximal forces at the velocities modeled. These data indicate that fiber length and moment arm appear to be coordinated to yield the greatest possible force production during locomotion.