Intrinsic excitability differs between murine hypoglossal and spinal motoneurons.

Motoneurons differ in the behaviors they control and their vulnerability to disease and aging. For example, brain stem motoneurons such as hypoglossal motoneurons (HMs) are involved in licking, suckling, swallowing, respiration, and vocalization. In contrast, spinal motoneurons (SMs) innervating the limbs are involved in postural and locomotor tasks requiring higher loads and lower movement velocities. Surprisingly, the properties of these two motoneuron pools have not been directly compared, even though studies on HMs predominate in the literature compared with SMs, especially for adult animals. Here we used whole cell patch-clamp recording to compare the electrophysiological properties of HMs and SMs in age-matched neonatal mice (P7-P10). Passive membrane properties were remarkably similar in HMs and SMs, and afterhyperpolarization properties did not differ markedly between the two populations. HMs had narrower action potentials (APs) and a faster upstroke on their APs compared with SMs. Furthermore, HMs discharged APs at higher frequencies in response to both step and ramp current injection than SMs. Therefore, while HMs and SMs have similar passive properties, they differ in their response to similar levels of depolarizing current. This suggests that each population possesses differing suites of ion channels that allow them to discharge at rates matched to the different mechanical properties of the muscle fibers that drive their distinct motor functions.

Intrinsic and synaptic homeostatic plasticity in motoneurons from mice with glycine receptor mutations.

Inhibitory synaptic inputs to hypoglossal motoneurons (HMs) are important for modulating excitability in brainstem circuits. Here we ask whether reduced inhibition, as occurs in three murine mutants with distinct naturally occurring mutations in the glycine receptor (GlyR), leads to intrinsic and/or synaptic homeostatic plasticity. Whole cell recordings were obtained from HMs in transverse brainstem slices from wild-type (wt), spasmodic (spd), spastic (spa), and oscillator (ot) mice (C57Bl/6, approximately postnatal day 21). Passive and action potential (AP) properties in spd and ot HMs were similar to wt. In contrast, spa HMs had lower input resistances, more depolarized resting membrane potentials, higher rheobase currents, smaller AP amplitudes, and slower afterhyperpolarization current decay times. The excitability of HMs, assessed by "gain" in injected current/firing-frequency plots, was similar in all strains whereas the incidence of rebound spiking was increased in spd. The difference between recruitment and derecruitment current (i.e., ΔI) for AP discharge during ramp current injection was more negative in spa and ot. GABAA miniature inhibitory postsynaptic current (mIPSC) amplitude was increased in spa and ot but not spd, suggesting diminished glycinergic drive leads to compensatory adjustments in the other major fast inhibitory synaptic transmitter system in these mutants. Overall, our data suggest long-term reduction in glycinergic drive to HMs results in changes in intrinsic and synaptic properties that are consistent with homeostatic plasticity in spa and ot but not in spd. We propose such plasticity is an attempt to stabilize HM output, which succeeds in spa but fails in ot.

Mapping the contribution of single muscles to facial movements in the rhesus macaque.

The rhesus macaque (Macaca mulatta) is the most utilized primate model in the biomedical and psychological sciences. Expressive behavior is of interest to scientists studying these animals, both as a direct variable (modeling neuropsychiatric disease, where expressivity is a primary deficit), as an indirect measure of health and welfare, and also in order to understand the evolution of communication. Here, intramuscular electrical stimulation of facial muscles was conducted in the rhesus macaque in order to document the relative contribution of each muscle to the range of facial movements and to compare the expressive function of homologous muscles in humans, chimpanzees and macaques. Despite published accounts that monkeys possess less differentiated and less complex facial musculature, the majority of muscles previously identified in humans and chimpanzees were stimulated successfully in the rhesus macaque and caused similar appearance changes. These observations suggest that the facial muscular apparatus of the monkey has extensive homology to the human face. The muscles of the human face, therefore, do not represent a significant evolutionary departure from those of a monkey species. Thus, facial expressions can be compared between humans and rhesus macaques at the level of the facial musculature, facilitating the systematic investigation of comparative facial communication.

Motor unit physiology: some unresolved issues.

The purpose of this review was to examine three issues that limit our understanding of motor unit physiology: (1) the range and distribution of the innervation ratios in a muscle; (2) the association between discharge rate and force; and (3) the variation in motor unit activity across contractions that differ in speed and type. We suggest that if more data were available on these issues, the understanding of neuromuscular function would be enhanced substantially, especially with regard to plasticity in the motor neuron pool, adequacy of the neural drive to muscle, and flexibility of activation patterns across various types of contractions. Current data are limited and these limitations influence our ability to interpret adaptations in muscle function in health and disease.

Discharge behaviour of single motor units during maximal voluntary contractions of a human toe extensor.

1. While it is known that the average firing rate of a population of motoneurones declines with time during a maximal voluntary contraction, at least for many muscles, it is not known how the firing patterns of individual motoneurones adapt with fatigue. To address this issue we used tungsten microelectrodes to record spike trains (mean +/- s.e.m., 183 +/- 27 spikes per train; range, 100-782 spikes) from 26 single motor units in extensor hallucis longus during sustained (60-180 s) maximal dorsiflexions of the big toe in seven human subjects. 2. Long spike trains were recorded from 13 units during the first 30 s of a maximal voluntary contraction (mean train duration, 9.6 +/- 1.2 s; range, 3.6-21.9 s) and from 13 units after 30 s (mean train duration, 16.6 +/- 3.7 s; range, 7.1-58.1 s). Maximal isometric force generated by the big toe declined to 78.3 +/- 6.3 % of its control level by 60-90 s and to 39.5 +/- 1.4 % of control by 120-150 s. Despite this substantial fatigue, mean firing rates did not change significantly over time, declining only slightly from 15.8 +/- 0.7 Hz in the first 30 s to 14.0 +/- 0.5 Hz by 60-90 s and 13.6 +/- 0.3 Hz by 120-150 s. 3. To assess fatigue-related adaptation in discharge frequency and variability of individual motor units, each spike train was divided into 2-15 equal segments containing at least 50 interspike intervals. Discharge variability was measured from the coefficient of variation (s.d. /mean) in the interspike intervals, with the s.d. being calculated using a floating mean of 19 consecutive intervals. Adaptation was computed as the average change in firing rate or variability that would occur for each 1 s of activity. There were no systematic changes in either firing rate or variability with time. 4. We conclude that single motoneurones supplying the extensor hallucis longus, a muscle comprised primarily of slow twitch muscle units, show little adaptation in firing with fatigue, suggesting that a progressive reduction in firing rate is not an invariable consequence of the fatigue associated with sustained maximal voluntary contractions.

Cessation of human motor unit discharge during sustained maximal voluntary contraction.

The purpose of this study was to determine whether cessation of motor unit discharge contributes to fatigue in human subjects. Multiple fine-wire and tungsten microelectrodes were inserted into the extensor digitorum or extensor indicis muscles of the forearm in an attempt to record the activity of the same motor unit from different locations within either muscle while subjects maintained a maximal voluntary contraction of the finger extensors until force dropped by approximately 50%. The activities of 13 motor units were followed for extended periods during the fatigue task. Of these, six appeared to cease discharging prior to the end of the task, which could not be attributed to electrode movement. These findings suggest that some motor neurons may not be able to discharge continuously in the presence of sustained volitional synaptic drive or that excitatory drive may diminish during maximal voluntary effort.

Force-frequency and fatigue properties of motor units in muscles that control digits of the human hand.

Modulation of motor unit activation rate is a fundamental process by which the mammalian nervous system encodes muscle force. To identify how rate coding of force may change as a consequence of fatigue, intraneural microstimulation of motor axons was used to elicit twitch and force-frequency responses before and after 2 min of intermittent stimulation (40-Hz train for 330 ms, 1 train/s) in single motor units of human long finger flexor muscles and intrinsic hand muscles. Before fatigue, two groups of units could be distinguished based on the stimulus frequency needed to elicit half-maximal force; group 1 (n = 8) required 9.1 +/- 0.5 Hz (means +/- SD), and group 2 (n = 5) required 15.5 +/- 1.1 Hz. Twitch contraction times were significantly different between these two groups (group 1 = 66. 5 ms; group 2 = 45.9 ms). Overall 18% of the units were fatigue resistant [fatigue index (FI) > 0.75], 64% had intermediate fatigue sensitivity (0.25

Simulation of motor unit recruitment and microvascular unit perfusion: spatial considerations.

Muscle fiber activity is the principal stimulus for increasing capillary perfusion during exercise. The control elements of perfusion, i.e., microvascular units (MVUs), supply clusters of muscle fibers, whereas the control elements of contraction, i.e., motor units, are composed of fibers widely scattered throughout muscle. The purpose of this study was to examine how the discordant spatial domains of MVUs and motor units could influence the proportion of open capillaries (designated as perfusion) throughout a muscle cross section. A computer model simulated the locations of perfused MVUs in response to the activation of up to 100 motor units in a muscle with 40,000 fibers and a cross-sectional area of 100 mm2. The simulation increased contraction intensity by progressive recruitment of motor units. For each step of motor unit recruitment, the percentage of active fibers and the number of perfused MVUs were determined for several conditions: 1) motor unit fibers widely dispersed and motor unit territories randomly located (which approximates healthy human muscle), 2) regionalized motor unit territories, 3) reversed recruitment order of motor units, 4) densely clustered motor unit fibers, and 5) increased size but decreased number of motor units. The simulations indicated that the widespread dispersion of motor unit fibers facilitates complete capillary (MVU) perfusion of muscle at low levels of activity. The efficacy by which muscle fiber activity induced perfusion was reduced 7- to 14-fold under conditions that decreased the dispersion of active fibers, increased the size of motor units, or reversed the sequence of motor unit recruitment. Such conditions are similar to those that arise in neuromuscular disorders, with aging, or during electrical stimulation of muscle, respectively.

Contractile properties of single motor units in human toe extensors assessed by intraneural motor axon stimulation.

1. Single motor axons innervating human toe extensor muscles were selectively stimulated through a tungsten microelectrode inserted percutaneously into the peroneal nerve. Twitch and tetanic forces were measured from a strain gauge over the proximal phalanx of the toe generating the greatest force. Twitch data were obtained from 19 single motor units in nine subjects: 8 motor units supplied extensor hallucis longus (EHL), 5 motor units supplied extensor digitorum longus (EDL), and 6 motor units supplied extensor digitorum brevis (EDB). Unpotentiated twitch forces ranged from 6.3 to 78.1 mN (20.0 +/- 4.0 mN, mean +/- SE), with the distribution highly skewed toward small forces. Twitch contraction and half-relaxation times were 74.8 +/- 3.9 and 78.6 +/- 6.0 ms, respectively. Compared with motor units in human thenar muscles, those in human toe extensor muscles were stronger but slower. However, as in thenar motor units, twitch force and contraction time were not related. 2. Force-frequency relationships were determined for 13 units (5 EDL, 5 EHL, 3 EDB) by stimulating each unit with short trains (1.0-5.0 s) of constant frequency (2-100 Hz). Peak force was related to stimulus frequency in a sigmoid fashion. The steep region of the curve extended from 5.5 +/- 0.7 (SE) Hz to 16.3 +/- 1.1 Hz for all units, and the stimulus frequency required to generate half-maximal force (9.6 +/- 0.6 Hz) was close to the center of the steep range. This frequency, which was inversely related to twitch contraction time, was lower than the frequency required to develop half-maximal force of human thenar motor units (12 +/- 4 Hz, mean +/- SD). The slopes of the regression lines relating force to frequency, computed over the steep range for each unit, were also lower for the toe extensors (3.7 +/- 0.7 mN/Hz) than for the thenar muscles (6 +/- 1 mN/Hz). 3. Maximal tetanic forces ranged from 29.9 to 188.1 mN (89.0 +/- 16.5 mN, mean +/- SE), and were generated at stimulus frequencies from 15 to 100 Hz (median 50 Hz). The stimulation frequency required for fused tetani (absence of noticeable force fluctuation) was generally less than that required for maximum tetanic force. The mean twitch-tetanus ratio, calculated for unpotentiated twitches, was 0.22 +/- 0.02 (range 0.15-0.41). This ratio was higher than for human thenar motor units (0.14 +/- 0.06, mean +/- SE). After twitch potentiation of 10 units, the mean twitch-tetanus ratio increased to 0.28 +/- 0.04. 4. The effects of preceding each stimulus train with a short interstimulus interval (10 ms) on force production at each frequency were examined in nine motor units. Peak forces at the onset of each contraction were higher when such an "initial doublet" preceded stimulus trains of < or = 20 Hz, but the mean force at the end of each stimulus train was not significantly affected at any frequency. 5. Eight units were stimulated with a train that increased in frequency continuously from 2 to 80 Hz, and then decreased symmetrically. This pattern resulted in peak forces that were higher on the descending limb of the stimulus train, the force-frequency relationship tracing a hysteresis loop. Hysteresis was exhibited because damping in the neuromuscular system causes the mechanical output of muscle to lag behind neural input. Thus, in non-steady-state conditions (as in most forms of natural activity), somewhat higher firing rates may be required to attain a particular level of force; once attained, force output will be transiently unresponsive to diminution of firing rate. 6. We conclude that there are differences in the contractile properties of single motor units in human toe extensor muscles (involved in posture and locomotion) and thenar muscles (involved in prehension and manipulation). Twitch-tetanus ratios were greater for motor units in the toe extensors, and this property accounted for the lower force sensitivity of these units to increases in frequency. (ABSTRACT TRUNCATED)

Limitations of the surface electromyography technique for estimating motor unit synchronization.

Motor unit synchronization was estimated from the surface electromyograms (EMG) of the first dorsal interosseus muscle of human volunteers by a simplified surface-EMG technique (Milner-Brown et al. 1973, 1975). Single motor units were identified from intramuscular recordings and were used to obtain a spike-triggered average of the surface-EMG. The discharge rate of a reference motor unit was controlled at two levels (high and low), and the effect of motor unit activity on the surface-EMG estimate of synchronization was studied in 56 motor units. The surface-EMG estimate of motor unit synchronization was significantly higher when the reference motor unit discharged at the high rate than when it discharged at the low rate. A regression analysis indicated that the synchronization ratio calculated from the surface EMG was significantly correlated with the level of EMG activity in the muscle. Motor unit synchronization was also estimated from surface-EMG measurements that were derived by computer simulation. The simulation permitted manipulation of motor unit activity (discharge rate and recruitment) with a complete absence of synchrony among the units in the pool. The stimulated surface-EMG index was influenced by an artifact associated with signal rectification, and this effect changed non-monotonically with motor unit activity. Furthermore, the increase in the motor unit activity reduced the signal-to-noise ratio of the spike-triggered surface EMG average, and consequently decreased the sensitivity of the surface-EMG index as an estimate of motor unit synchronization. We conclude that the simplified surface-EMG method (Milner-Brown et al. 1973, 1975) does not provide a useful index of motor unit synchronization due to its inability to accurately distinguish the synchronization from methodological effects related to a rectification artifact and variation in the signal-to-noise ratio.

Motor unit activity during isometric and concentric-eccentric contractions of the human first dorsal interosseus muscle.

1. Motor unit activity was recorded with intramuscular fine wire electrodes during isometric, concentric, and eccentric activity in the human first dorsal interosseus muscle. Twenty-one units from 11 subjects were sampled. 2. During isotonic cycles of shortening and lengthening, 18 of 21 units were recruited during the concentric phase, increased their discharge rates as the concentric movement progressed, then decreased their discharge rate during the eccentric phase, and were derecruited. 3. A different pattern of recruitment was observed in recordings from three units. These units were recruited during the eccentric phase, at a time when other units were decreasing their discharge rate or being derecruited. In two of the units selectively recruited during the eccentric phase, it was possible to determine their isometric thresholds, which were higher than those of units exhibiting the more common pattern of recruitment. 4. For two of the three units exhibiting selective recruitment during eccentric contraction, the unit was recorded simultaneously with different pairs of recording wires separated by 5-10 mm. Each discharge of these units was detected by both electrodes, making it unlikely that movement artifact was responsible for the initiation or cessation of discharge. 5. The recruitment patterns observed suggest that changes in the type or distribution of synaptic inputs to motoneurons during movement can, in some instances, override pre- and postsynaptic factors that shape recruitment order in isometric conditions.

Short-term immobilization has a minimal effect on the strength and fatigability of a human hand muscle.

The purpose of this study was to determine the association between reduced fatigability typically observed in disused muscle and an improved resistance to the impairment of neuromuscular propagation. Endurance time of an isometric contraction sustained at 35% of maximum voluntary contraction (MVC) force and the fatigue-induced change in the evoked compound muscle action potential (M wave) were measured in the first dorsal interosseus muscle of human subjects before, during, and after 3 (n = 9) or 5 wk (n = 2) of immobilization. The immobilization procedure caused a substantial decline in the chronic electromyographic (EMG) activity (to 4% of control value) of the first dorsal interosseus muscle. Endurance time was found to be significantly correlated to the maintenance of M-wave amplitude during the fatigue task. However, neither of these variables was significantly affected by immobilization. Also, immobilization had no significant effect on the prefatigue values of MVC force and EMG or twitch contraction time or on the postfatigue changes in MVC force and EMG, M wave duration, twitch amplitude, and contraction time. In the unfatigued muscle, immobilization did cause an increase in twitch force (153%) and a decrease in M-wave amplitude (67%). It appears, therefore, that a healthy first dorsal interosseus muscle is generally resistant to adaptation when its use has been reduced for 3-5 wk by immobilization.

The role of the sarcolemma action potential in fatigue.

A prevalent feature of neuromuscular fatigue is a decline in the extracellularly recorded myoelectric signal. One factor that could underlie this change is a decrease in the amplitude of the sarcolemmal action potential. Based on observed reductions in action potential amplitude without effect on force, it has been argued that changes in the action potential during sustained activity would be unlikely to contribute to fatigue. However, those observations were primarily from experiments in which 1) high frequency stimulation may have caused signal cancellation due to action potential overlap; or 2) sustained membrane depolarization may have directly activated excitation-contraction coupling. The relatively low and narrow range of membrane depolarization required for full activation of amphibian and slow-twitch mammalian fibers makes them resistant to incomplete activation if action potentials are depressed during fatigue. Mammalian fast-twitch fibers, on the other hand, require greater depolarization for full activation and also exhibit a greater decrease in action potential amplitude with fatigue. Therefore, it seems probable that fatigue-related decline in action potential amplitude in these fibers leads to incomplete activation and loss of force.

Crosstalk in surface electromyography: Theoretical and practical estimates.

The purpose of this paper is to address four aspects of surface electromyography associated with crosstalk between adjacent recording sites. The first issue that is addressed in the potential crosstalk between electrodes located on muscles with different functions: antagonist pairs, or muscles with one common and one different function (i.e. soleus/peroneus longus or soleus/ gastrocnemius). Practical functional tests are utilized to demonstrate the crosstalk between muscle pairs to be negligible. The second goal is to estimate the depth of pick-up and the crosstalk between myoelectric signals from agonist muscles using a theoretical model. The depth of pick-up was estimated to be 1.8 cm (including a 2 mm layer of skin and fat) using electrodes of 49 mm(2) with bipolar spacing of 2.0 cm. A cross-correlation technique is demonstrated which predicts the common signal (crosstalk) between surface electrodes with electrode-pair spacing of 1 cm around a hypothetical muscle. The predicted crosstalk using cross-correlation measures was 49% at 1 cm electrode-pair spacing dropping to 13% at 2 cm spacing and 4% at 3 cm. The third part compares these predictions with crosstalk measures from experimental recordings taken from electrode pairs spaced 2.5 cm apart around the quadriceps. At 2.5 cm spacing there was 22-24% common signal dropping to between 4-7% at 5 cm and to between 1 and 2% at 7.5 cm. The fourth and last component of this report assesses three methods to decrease the range of pick-up and thereby potential crosstalk: electrodes of smaller surface area, reduced bipolar spacing and mathematical differentiation. All three techniques reduce the common signal by varying amounts; all three techniques combined reduce the predicted crosstalk for the 1.0 cm electrode-pair spacing from 49-10.5%.

Models of recruitment and rate coding organization in motor-unit pools.

1. Isometric muscle force and the surface electromyogram (EMG) were simulated from a model that predicted recruitment and firing times in a pool of 120 motor units under different levels of excitatory drive. The EMG-force relationships that emerged from simulations using various schedules of recruitment and rate coding were compared with those observed experimentally to determine which of the modeled schemes were plausible representations of the actual organization in motor-unit pools. 2. The model was comprised of three elements: a motoneuron model, a motor-unit force model, and a model of the surface EMG. Input to the neuron model was an excitatory drive function representing the net synaptic input to motoneurons during voluntary muscle contractions. Recruitment thresholds were assigned such that many motoneurons had low thresholds and relatively few neurons had high thresholds. Motoneuron firing rate increased as a linear function of excitatory drive between recruitment threshold and peak firing rate levels. The sequence of discharge times for each motoneuron was simulated as a random renewal process. 3. Motor-unit twitch force was estimated as an impulse response of a critically damped, second-order system. Twitch amplitudes were assigned according to rank in the recruitment order, and twitch contraction times were inversely related to twitch amplitude. Nonlinear force-firing rate behavior was simulated by varying motor-unit force gain as a function of the instantaneous firing rate and the contraction time of the unit. The total force exerted by the muscle was computed as the sum of the motor-unit forces. 4. Motor-unit action potentials were simulated on the basis of estimates of the number and location of motor-unit muscle fibers and the propagation velocity of the fiber action potentials. The number of fibers innervated by each unit was assumed to be directly proportional to the twitch force. The area of muscle encompassing unit fibers was proportional to the number of fibers innervated, and the location of motor-unit territories were randomly assigned within the muscle cross section. Action-potential propagation velocities were estimated from an inverse function of contraction time. The train of discharge times predicted from the motoneuron model determined the occurrence of each motor-unit action potential. The surface EMG was synthesized as the sum of all motor-unit action-potential trains. 5. Two recruitment conditions were tested: narrow (limit of recruitment < 50% maximum excitation) and broad recruitment range conditions (limit of recruitment > 70% maximum excitation).(ABSTRACT TRUNCATED AT 400 WORDS)

Reduced control of motor output in a human hand muscle of elderly subjects during submaximal contractions.

1. The effect of age on the motor output of the first dorsal interosseous muscle of 22 (6 female, 16 male) human subjects was investigated. The purpose of the study was to determine the effect of age on the control of muscle force and the associated changes in the discharge behavior and mechanical properties of single motor units. 2. Each subject performed three tasks requiring isometric abduction of the left index finger: a maximum voluntary contraction (MVC), a constant-force task, and a threshold task. The ability to control force was assessed during the constant-force task by quantifying the variation in isometric force about four submaximal target forces (5, 20, 35, and 50% MVC). The threshold task involved sustaining the discharge of the isolated motor unit at a low, steady rate for approximately 3 min. 3. The discharge behavior and the mechanical properties of single motor units were determined during the threshold task by measuring the interimpulse intervals and the peak amplitude and time to peak of the spike-triggered average force. 4. The data indicated that age had an effect on the variation of force about submaximal target forces (range: 5-50% MVC), and that these force variations, when calculated relative to the target force, were greater at lower force levels in the elderly subjects.(ABSTRACT TRUNCATED AT 250 WORDS)

Impairment of neuromuscular propagation during human fatiguing contractions at submaximal forces.

1. The purpose of the study was to examine the dependence of neuromuscular propagation impairment on the level of isometric force sustained to the endurance limit. The task involved human volunteers sustaining a submaximal abduction force with the index finger by activating the first dorsal interosseous muscle as long as possible. 2. The submaximal force was sustained at one of three levels (20, 35 or 65% of maximum) by increasing motor unit activity, as indicated by the electromyogram (EMG), during the fatiguing contraction. Although the EMG increased during the fatiguing contraction, the EMG was significantly less than maximum at the endurance limit for all subjects (deficit of 19-55% of maximum). This deficit was inversely related to the level of the sustained submaximal force. 3. The maximum voluntary contraction and twitch forces were significantly reduced following the fatiguing contraction. As with the EMG, the degree of force reduction was greatest for the subjects who sustained the low target forces. 4. The fatiguing contraction caused a 12-23% decline in M wave amplitude, a 33-51% increase in M wave duration, and no change in M wave area. The decline in M wave amplitude, which is an index of neuromuscular propagation impairment, was greatest among the subjects who sustained the low target forces. 5. The mean power frequency of the EMG decreased by a similar amount (50-57%) during the fatiguing contraction for all three groups of subjects. 6. A model representing the interaction of processes that enhance and impair force was developed to explain the recovery of twitch force following the sustained contractions at different target forces. 7. We conclude that the fatigue experienced by a subject when force is sustained at a submaximal value does involve an impairment of neuromuscular propagation. This impairment is one factor that limits muscle excitation during a submaximal, fatiguing contraction and contributes to the diminished force capability by the end of the fatigue task.

Estimating the strength of common input to human motoneurons from the cross-correlogram.

1. The relationship between the motor unit discharge pattern (rate and variability) and synchronization of motor unit pairs was studied in the first dorsal interosseus muscle of human subjects. In separate trials of up to 4 min duration, subjects voluntarily controlled the mean discharge rate of an identified motor unit at one of several prescribed rates (range 7.5-17.5 Hz). 2. The effect of discharge rate on the synchronous peak in the cross-correlogram was examined in eighty motor unit pairs from six subjects. Five commonly used synchronization indices were used to quantify synchrony in the cross-correlograms constructed from different discharge-rate trials. For each synchronization index, the apparent magnitude of synchrony increased at lower motor unit discharge rates. The synchronization indices were not equally sensitive to discharge rate; increases in the different indices ranged from 72 to 494% between the highest and lowest discharge rates. 3. A model of the membrane potential trajectory underlying rhythmic motoneuron discharge was used to explain the observed increase in the magnitude of the synchronization indices at lower discharge rates. The essential feature of this model is that the probability of a common-input EPSP causing a synchronous discharge in two motoneurons is independent of discharge rate. This means that the number of synchronous action potentials in excess of chance in any trial depends on the properties of the common-input EPSPs and the duration of the trial, but is not related to motor unit discharge rates. The model also demonstrated that when the excess synchronous counts are normalized to motor unit discharge rate, or baseline counts in the histogram (as in the conventional synchronization indices), the magnitude of the index increases when the motor unit discharge rates are low. 4. The strength of common input to motoneurons could be misinterpreted if conventional synchronization indices are used because of discharge-rate effects. The model was used to derive an index of the strength of common input to motoneurons (CIS) that was independent of motor unit discharge rate. CIS is the frequency of synchronous action potentials in the motor unit pair in excess of those expected due to chance (calculated during periods of tonic discharge in both units). The mean CIS in first dorsal interosseus motor unit pairs ranged from 0.052 to 1.005 extra synchronous action potentials per second across subjects. 5. Discharge variability was correlated with each of the synchronization indices and the CIS.(ABSTRACT TRUNCATED AT 400 WORDS)

Detection of motor unit action potentials with surface electrodes: influence of electrode size and spacing.

A model of the motor unit action potential was developed to investigate the amplitude and frequency spectrum contributions of motor units, located at various depths within muscle, to the surface detected electromyographic (EMG) signal. A dipole representation of the transmembrane current in a three-dimensional muscle volume was used to estimate detected individual muscle fiber action potentials. The effects of anisotropic muscle conductance, innervation zone location, propagation velocity, fiber length, electrode area, and electrode configuration were included in the fiber action potential model. A motor unit action potential was assumed to be the sum of the individual muscle fiber action potentials. A computational procedure, based on the notion of isopotential layers, was developed which substantially reduced the calculation time required to estimate motor unit action potentials. The simulations indicated that: 1) only those motor units with muscle fibers located within 10-12 mm of the electrodes would contribute significant signal energy to the surface EMG, 2) variation in surface area of electrodes has little effect on the detection depth of motor unit action potentials, 3) increased interelectrode spacing moderately increases detection depth, and 4) the frequency content of action potentials decreases steeply with increased electrode-motor unit territory distance.

Ground reaction forces in running: a reexamination.

Ground reaction force (GRF) data were collected on twenty adult males during running stance to establish normative standards to aid in assessment of the gait of atypical runners. Subjects ran between 30 and 40 trials across a Kistler 0.6 X 0.9 m force platform at self-selected speeds ranging from 2.5 to 5.5 m s-1. Best fit polynomials for a given descriptor variable were constructed for each subject and the polynomials were evaluated as a function of running speed. Predicted means and standard deviations (based on the polynomials) were calculated and multivariate analyses of variance were performed. The descriptor variables: impact peak, loading rate, thrust maximum, decay rate, average vertical GRF, change in vertical velocity, braking impulse, propulsive impulse and stance time were determined to be running speed dependent (p less than 0.001). Specific patterns associated with the breaking component of the antero-posterior GRF of heel-strikers included single, double and multiple peaks. Three dimensional graphic displays showed that, despite considerable group variability in medial-lateral GRF-time histories, consistency was evident in the patterns of individuals across speeds. Individual right-left asymmetries were clearly shown in these displays.