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R K Powers

Publications and source records attributed to R K Powers.

At least 37 records · Page 2Linked to original sources

Spike frequency adaptation studied in hypoglossal motoneurons of the rat.

1. We studied spike frequency adaptation of motoneuron discharge in the rat hypoglossal nucleus using a brain stem slice preparation. The characteristics of adaptation in response to long (60 s) injected current steps were qualitatively similar to those observed previously in cat hindlimb motoneurons. The discharge rate typically exhibited a rapid initial decline, characterized by a linear frequency-time relation, followed by a gradual exponential decline that continued for the duration of current injection. However, a more systematic, quantitative analysis of the data revealed that there were often three distinct phases of the adaptation rather than two. 2. The three phases of adaptation (initial, early, and late) were present in at least one 60-s trial of repetitive firing in all but a small number of motoneurons. Initial adaptation was limited to the first few spikes except in a few trials (7%) in which there was no initial adaptation. The time course of the subsequent decline in rate could be adequately described by a single-exponential function in about half of the trials (48%). In the remaining trials this subsequent decline in frequency was better described as the sum of two exponential functions: an early phase, lasting < 2 s, and a late phase, which lasted for the duration of the discharge period. 3. The magnitude of initial adaptation was correlated with the initial firing frequency (i.e., the reciprocal of the 1st interspike interval). The magnitudes of the early and late phases of adaptation were correlated with the firing frequency reached at the end of initial adaptation. Neither the magnitudes nor the time courses of the three phases were correlated with other membrane properties such as input resistance, rheobase, or repetitive firing threshold. 4. The slope of the frequency-current (f-I) curve was steeper in the initial phase (first 2-5 spikes) than in either the early (< 2 s) or late (> 2 s) phases of adaptation as previously reported by other investigators. In the absence of early adaptation, a steady state for the f-I slope was reached by 0.7-1 s, the time typically reported in studies of repetitive discharge. However, when early adaptation was present (50% of the trials), a steady-state value for the f-I slope was not reached until the cell had discharged for > 1 s. 5. To characterize the time course of firing rate recovery from the adaptive processes, the current was turned off for periods of < or = 10 s during the course of a 60-s trial.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Effective synaptic current and motoneuron firing rate modulation.

1. We used a modified voltage-clamp technique to measure the steady-state effective synaptic currents (I(N)) produced by activating four different input systems to cat hindlimb motoneurons: Ia afferent fibers, Ia-inhibitory interneurons, Renshaw interneurons, and contralateral rubrospinal neurons. In the same motoneurons, we measured the slope of the firing rate-injected current (f-I) relation in the primary range. We then reactivated these synaptic inputs during steady, repetitive firing to assess their effects on motoneuron discharge rate. 2. Our measurements of I(N) were derived from recordings made near the resting membrane potential, whereas the effects of the synaptic inputs on repetitive discharge were evaluated at more depolarized membrane potentials. Thus we adjusted the I(N) values for these changes in driving force based on estimates of the synaptic reversal potential and the mean membrane potential during repetitive discharge. 3. We found that changes in the steady-state discharge rate of a motoneuron produced by these synaptic inputs could be reasonably well predicted by the product of the estimated value of I(N) during repetitive firing and the slope of the motoneuron's f-I relation. Although there was a high correlation between predicted and observed changes in firing rate for our entire sample of motoneurons (r = 0.93; P < 0.001), the slope of the relation between predicted and observed firing rate modulation was significantly greater than 1. 4. The systematic difference between predicted and observed firing rate modulation observed in the overall sample was primarily due to the fact that our predictions underestimated the changes in firing rate produced by Ia excitation and Ia inhibition.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Intrinsic properties of motoneurons. Implications for muscle fatigue.

The following is a brief review of the intrinsic properties of motoneurons that contribute to their recruitment and rate modulation. Our emphasis is on properties that may either accelerate or delay the onset of muscular fatigue. In general, intrinsic motoneuron properties are regulated in a way that minimizes energy expenditure. The correlation of recruitment threshold with motoneuron type ensures that the most fatigable motor units are reserved for the most forceful contractions. The variation in minimum firing rates arising from variations in AHP characteristics ensures that motoneurons begin to fire at rates that are matched to the force producing characteristics of their muscle units. Further, it is possible that spike-frequency adaptation contributes to optimization of the tension (force)-firing frequency (T-f) transform of individual motor units.

Animals↗

M/RMS: an EMG method for quantifying upper motoneuron and functional weakness.

A new method for quantifying upper motoneuron (UMN) weakness using standard EMG equipment is presented and compared to the twitch occlusion method. Control subjects and patients with stroke, spinal cord injury (SCI), and peripheral nerve injury (PNI) were evaluated. Force, EMG, and twitch force from nerve electrical stimulation were recorded during isometric contractions of the biceps and triceps. Ratios of the elicited CMAP amplitude (M)/voluntary-root-mean-square EMG activity (RMS) and of the voluntary contraction twitch force (Ti)/the resting twitch force (Tmax) were the two measures of UMN weakness studied. Both ratios are linearly correlated with force for controls, log M/RMS (r = 0.96) and Ti/Tmax (r = 0.86). Log M/RMS ratio was abnormally high (> mean + 2 SD; i.e., > 1.09) for weak muscles affected by stroke and incomplete SCI, but was normal in muscles weakened by PNI. An elevated M/RMS ratio, may aid in quantification of deficient supraspinal activation from UMN or functional weakness.

Adult↗

Electrodiagnosis of upper limb weakness in acute quadriplegia.

Clinical and neuropathologic observations after cervical spinal cord injury suggest varying involvement of gray and white matter. The resulting upper limb weakness may reflect varying degrees of upper motoneuron (UMN) and/or lower motoneuron (LMN) involvement. This study uses electrophysiologic measures, including compound muscle action potential (M response) amplitude, root mean square (RMS) of the surface electromyographic activity during voluntary muscle contractions and the firing rate of motor units, to distinguish UMN and LMN weakness in upper extremities after acute quadriplegia. M response amplitude did not correlate with strength; many muscles had large M responses given their strength. These muscles manifest: (1) high M/RMS ratios (ratio of electrically elicited to voluntarily recruited electromyographic activity) and (2) slow firing rates of single motor units during maximal isometric contractions. For muscles with normal M amplitudes, M/RMS ratio correlates inversely with strength. For muscles with normal M/RMS ratios, M amplitude correlates positively with strength. Cluster analysis was used to distinguish UMN, LMN or Mixed types of weakness. Distinguishing these different types of weakness in acute quadriplegia may allow individualized rehabilitation for the type of weakness present.

Action Potentials↗

How different afferent inputs control motoneuron discharge and the output of the motoneuron pool.

In theory, there are at least two distinct mechanisms by which afferent inputs could alter motoneuron discharge and shape the output of a motoneuron pool: either by delivering synaptic current to the motoneurons' somata ('classic' synaptic transduction); or by altering the motoneurons' voltage-sensitive conductances (neuromodulation). Recent work has confirmed the operation of both of these mechanisms. It has been shown that the effect of a 'classic' synaptic input on motoneuron firing rate is predicted by the product of the effective synaptic current and the slope of the motoneuron's frequency-current relation. It has also been shown that neuromodulators can alter both the slope of a motoneuron's frequency-current relation and its threshold for repetitive firing. It is argued here, however, that when two or more sources of synaptic input are activated concurrently, the distinction between these two mechanisms is blurred. Computer simulations of motoneuron and motor pool behavior have proved extremely useful in understanding these processes.

Animals↗

A variable-threshold motoneuron model that incorporates time- and voltage-dependent potassium and calcium conductances.

1. A "threshold-crossing" motoneuron model was developed to relate recently described biophysical features of cat alpha-motoneurons to motoneuron discharge behavior. This model incorporated three features not included in precedent models: 1) a low-threshold, persistent calcium current; 2) realistic voltage dependencies of the major ionic conductances; and 3) a variable spike threshold. The effects of these additional biophysical features on model behavior were investigated by successively adding them to a fixed threshold model with a single potassium conductance. 2. Fixed-threshold models with either one or two potassium conductances could not produce appropriate discharge behavior. Steady-state frequency-current (F-I) relations were characterized by a continuously increasing slope, unlike the piecewise linear relations observed in real motoneurons. These models also produced unrealistically high discharge rates at the highest levels of "injected" current. 3. The addition of a variable spike threshold, which was made to increase linearly with the magnitude of injected current, could limit maximum discharge rates to more realistic levels. However, steady-state F-I relations still did not exhibit the appropriate shape. 4. The incorporation of a low-threshold calcium current led to a good quantitative agreement between the steady-state F-I relations produced by the model and those obtained in real motoneurons. In addition, the steady-state relation between total membrane current and membrane voltage (I-V relation) of the model was very similar to those measured in real motoneurons. The model's I-V and F-I relations were both very sensitive to the exact form of the steady-state relation between the magnitude of the calcium conductance and membrane voltage. 5. Additional modifications, which included a second calcium conductance and a factor relating spike threshold to membrane voltage, helped to produce more realistic afterhyperpolarizations and first-interval F-I relations. 6. Bistable discharge behavior could be produced by reducing the slow potassium conductance and increasing the time constants governing the activation and deactivation of the low-threshold calcium conductance. 7. The final model thus reproduces a wide range of motoneuron behaviors including subthreshold rectification, piecewise linear first interval and steady-state F-I relations, and, with appropriate modifications, bistable discharge behavior. Nonetheless, by simplifying the representation of fast spike conductances as well as the kinetics of the other ionic conductances, the model remains simple enough to be incorporated into a larger neural network.

Animals↗

Distribution of rubrospinal synaptic input to cat triceps surae motoneurons.

1. We evoked steady-state synaptic potentials in triceps surae motoneurons of the cat by stimulating the hindlimb projection area of the contralateral magnocellular red nucleus at 200 Hz. We measured the effective synaptic currents (IN) underlying the synaptic potentials using a modified voltage-clamp technique. We also determined the effect of the rubrospinal input on the discharge rate of some of the motoneurons by inducing repetitive discharge with long injected current pulses during which the red nucleus stimulation was repeated. 2. At motoneuron resting potential, the distribution of IN from the red nucleus within the triceps surae pools was qualitatively similar to the distribution of synaptic potentials: 86% of the putative type F motoneurons received a net depolarizing IN from the red nucleus stimulation, whereas only 38% of the putative type S units did so. The mean values of IN were significantly different in the two groups [+4.1 +/- 5.0 nA (SD) for putative type F and -1.6 +/- 3.1 nA for putative type S]. 3. However, when the values of IN at threshold for repetitive firing were estimated, the distribution of IN from the red nucleus was quite different. At threshold, all of the putative type S units received hyperpolarizing IN but so did nearly half of the putative type F units. 4. As would be expected from the wide range of IN at threshold (-20 to +12 nA), the red nucleus input produced dramatically different effects on the discharge of different motoneurons.(ABSTRACT TRUNCATED AT 250 WORDS)

Action Potentials↗

Effective synaptic current can be estimated from measurements of neuronal discharge.

1. The basic question of how motoneurons transform synaptic inputs into spike train outputs remains unresolved, despite detailed knowledge of their morphology, electrophysiology, and synaptic connectivity. We have approached this problem by making measurements of a synaptic input under steady-state conditions and combining them with quantitative assessments of their effects on the discharge rates of cat spinal motoneurons. 2. We used a modified voltage-clamp technique to measure the steady-state effective synaptic currents (IN) produced by rubrospinal input to cat triceps surae motoneurons. In the same motoneurons we measured the slope of the firing rate-injected current (f-i) relation in the primary range. We then reactivated the rubrospinal input during steady, repetitive firing to assess its effect on motoneuron discharge rate. 3. We found that changes in the steady-state discharge rate of a motoneuron produced by this synaptic input could be described simply as the product of the net effective synaptic current measured at the soma and the slope of the motoneuron's f-i relation. This expression essentially redefines synaptic efficacy in terms of a cell's basic input-output function. Further, measurements of effective synaptic current simplify the task of estimating synaptic efficacy, because detailed knowledge of neither the electrotonic architecture of the postsynaptic cell nor of the locations of the presynaptic boutons is required.

Animals↗

Alterations in motoneuron properties induced by acute dorsal spinal hemisection in the decerebrate cat.

Using intracellular recording techniques, we studied the response characteristics of two separate populations of triceps surae motoneurons in unanesthetized decerebrate cats, recorded before and after low thoracic hemisection of the spinal cord. In each preparation, we studied the response properties of one group of motoneurons and the protocol was then repeated for a separate group, immediately following the dorsal hemisection. In each group, we examined both the minimum firing rates of motoneurons during intracellular current injection and a range of cellular properties, including input resistance, rheobase current and afterhyperpolarization time course and magnitude. Although earlier studies from this laboratory have shown substantial reductions in minimum firing rate in reflexively active motoneurons in the hemisected decerebrated preparation, the response of motoneurons to intracellular current injection in the current preparation proved to be quite different. Minimum firing rates were either normal or even somewhat higher in the post-lesion group, while the time course of the afterhyperpolarization was shortened. Moreover, these effects were not evenly distributed across the motoneuron pool. The rate effect was most evident in motoneurons with higher conduction velocity, while the afterhyperpolarization effect occurred predominantly in motoneurons with lower conduction velocity. Neither of these effects could be accounted for by lesion-induced changes in other cellular properties. We conclude that tonically active neurons with descending axons traversing dorsolateral white matter may influence both the discharge characteristics and membrane properties of spinal motoneurons in novel ways, presumably by modifying voltage or calcium activated motoneuronal conductances. The previously described reactions in the firing rate of motoneurons after such lesions appear to be mediated by different means, perhaps by alterations in synaptic input from segmental interneurons.

Action Potentials↗

Effects of low-frequency stimulation on the tension-frequency relations of fast-twitch motor units in the cat.

1. Tension-frequency relations were measured in single fast-twitch motor units of the cat flexor digitorum longus (FDL) muscle before and after stimulating each unit with a series of 10-s trains at 20 Hz. The 20-Hz conditioning stimulation produced a combination of potentiating and fatiguing effects, similar to those previously reported to follow higher frequency stimulation of single motor units of the cat and maximal voluntary contractions in man. 2. The conditioning stimulation left three types of after effects: 1) short-term potentiation, 2) a somewhat longer lasting depression of maximal tension, and 3) a delayed depression of low-frequency responses (low-frequency fatigue). 3. The immediate potentiating effect of the conditioning stimulation was most prominent in fatigue-resistant (FR) motor units, whereas depression of maximal tension and low-frequency fatigue were most prominent in fatigue-intermediate (FI) and highly fatigable (FF) motor units. 4. On the basis of our results and those of other investigators, we propose that potentiation, depression of maximal tension, and low-frequency fatigue are independent phenomena, acting at distinct points in the excitation-contraction coupling process. 5. Our results suggest that both potentiation and low-frequency fatigue can result from rather modest amounts of preceding activity. Thus large changes in muscle force production are not unique to maximal contractions but are likely to follow sustained, submaximal contractions as well.

Animals↗

Summation of motor unit tensions in the tibialis posterior muscle of the cat under isometric and nonisometric conditions.

1. The tension produced by the combined stimulation of two to four single motor units of the cat tibialis posterior muscle was compared with the algebraic sum of the tensions produced by each individual motor unit. Comparisons were made under isometric conditions and during imposed changes in muscle length. 2. Under isometric conditions, the tension resulting from combined stimulation of units displayed marked nonlinear summation, as previously reported in other cat hindlimb muscles. On average, the measured tension was approximately 20% greater than the algebraic sum of the individual unit tensions. However, small trapezoidal movements imposed on the muscle during stimulation significantly reduced the degree of nonlinear summation both during and after the movement. This effect was seen with imposed movements as small as 50 microns. 3. The degree of nonlinear summation was not dependent on motor unit size or on stimulus frequency. The effect was also unrelated to tendon compliance because the degree of nonlinear summation of motor unit forces was unaffected by the inclusion of different amounts of the external tendon between the muscle and the force transducer. 4. Our results support previous suggestions that the force measured when individual motor units are stimulated under isometric conditions is reduced by friction between the active muscle fibers and adjacent passive fibers. These frictional effects are likely to originate in the connective tissue matrix connecting adjacent muscle fibers. However, because these effects are virtually eliminated by small movements, linear summation of motor unit tensions should occur at low force levels under nonisometric conditions.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Evidence of shared, direct input to motoneurons supplying synergist muscles in humans.

Cross-correlation techniques were used to test for the presence of shared, direct input to motoneurons innervating different synergist elbow flexor muscles in man. Motor unit activity was recorded intramuscularly from two elbow flexor muscles during steady isometric elbow flexion in normal and paretic subjects. To increase the probability of detecting weak synchrony, one of the intramuscular needles was positioned to record multiunit activity. Significant correlogram peaks were obtained in 25/57 runs in normal subjects, and the features of the correlograms were similar to those previously reported based on cross-correlation of two single units within the same muscle. Further, the characteristics of discharge synchrony measured in paretic stroke patients are consistent with other reports on the effects of stroke on synchrony among motoneurons belonging to the same pool, i.e. narrow correlogram peaks were rare in paretic subjects and significant correlogram peaks often had longer than normal durations.

Cerebrovascular Disorders↗

Stretch reflex dynamics in spastic elbow flexor muscles.

Previous studies of stretch reflexes in patients with spastic hypertonia have emphasized the dynamic character of stretch reflex output. In contrast, our own studies of stretch reflex dynamics in spastic elbow flexor muscles of 14 hemiparetic human subjects have shown that stretch-evoked torque displays a relatively weak dependence on stretch velocity, and there is generally no preferential enhancement of dynamic as compared with static reflex output. Moreover, stretch reflex dynamics are broadly similar in voluntarily activated spastic and normal elbow flexor muscles. These findings support our hypothesis that spastic hypertonia results primarily from a decrease in stretch reflex threshold. The strong velocity dependence of stretch-evoked electromyographic activity in initially inactive spastic muscles could be due to a decrease in reflex threshold with increasing stretch velocity, rather than an abnormal velocity-dependent increase in stretch reflex responsiveness.

Adult↗

Quantitative relations between hypertonia and stretch reflex threshold in spastic hemiparesis.

The relative contributions of variations in stretch reflex threshold and total joint stiffness to changes in stretch-evoked torque were assessed in the spastic elbow muscles of 14 hemiparetic spastic subjects. For a given subject, variations in torque, measured after a constant angular deflection, were mediated largely by changes in stretch reflex threshold, rather than by changes in reflex stiffness. Between-subject comparisons were sensitive to stiffness differences between limbs, but reflex thresholds were still broadly correlated with torque magnitude, suggesting that reductions in stretch reflex threshold are uniformly present in spastic muscles. These findings, coupled with the apparent similarity of reflex stiffness estimates in voluntarily activated spastic and normal muscles, suggest that the central disturbance in spasticity is a reduction in the threshold of the stretch reflex, without a significant enhancement of reflex gain.

Adult↗

Disturbances of motor output in a cat hindlimb muscle after acute dorsal spinal hemisection.

Force and electromyogram (EMG) responses of the medial gastrocnemius muscle were assessed during isometric contractions in 8 decerebrate cat preparations, before and after acute dorsal hemisection of the spinal cord at the T12 level. The measures derived included the relation between static force and mean rectified EMG, the EMG amplitude distribution, EMG power spectral density, and force power spectral density. Our findings were that the spinal lesion induced modifications in the shape of the EMG amplitude distribution, a substantial increase in mean rectified EMG per unit force, and increases in EMG spectral power and force spectral power over a broad band of frequencies. In 7/8 preparations, there was disproportionate enhancement of EMG spectral power below 40 Hz, with a commensurate reduction in the EMG mean power frequency (MPF) in 6 of these 7 cases. Recordings of motoneuron discharge from 9 decerebrate preparations taken before and after the spinal hemisection revealed that the lesion-induced changes in EMG and force power spectra were accompanied by lower mean discharge rates, and by a compression of the range of recruitment force. These changes in motoneuron rate and recruitment were probably responsible for the changes in EMG and force measures, especially for the relative increase in low-frequency EMG power. If these acute disturbances of motoneuron rate and recruitment persist in chronic human neurological disorders, they represent an important and largely unrecognized source of muscular weakness and increased fatigability.

Action Potentials↗

Effects of acute dorsal spinal hemisection on motoneuron discharge in the medial gastrocnemius of the decerebrate cat.

1. The discharge of single alpha-motoneuron axons was recorded from small cut filaments of the medial gastrocnemius (MG) muscle nerve in the decerebrated cat preparation before and after a dorsal hemisection of the thoracic spinal cord. The remainder of the MG muscle nerve was left intact, and muscle force and multiunit electromyographic (EMG) activity were recorded along with alpha-motoneuron discharge, while motor output was varied by manual stimulation of the contralateral hindlimb. 2. We recorded activity in 32 motoneurons before and after the spinal lesion, and pre- and postlesion recruitment forces and minimum firing rates were determined for 30 of these. Postlesion decreases in minimum firing rates were observed in 25/30 motoneurons, and decreases in recruitment force were seen in 21/30 motoneurons. The remaining motoneurons, which generally had low presection recruitment forces and minimum rates, exhibited postlesion increases in both parameters (see below). 3. The effects of the spinal lesion on the recruitment force and minimum firing rate of a motoneuron were related to the prelesion values of these parameters; the largest postlesion decreases were seen in motoneurons with the highest prelesion rates and recruitment forces. Spinal lesions thus acted to shift and compress the range of recruitment forces and minimum firing rates, so that after the lesion all motoneurons tended to exhibit discharge behavior typical of that seen only in the lowest threshold motoneurons before the lesion. In addition, motoneurons with low prelesion recruitment forces (less than 1.0 N of active force) generally showed an increase in recruitment force after the lesion, indicating that the lesion may have led to changes in the prelesion recruitment order. Direct evidence of recruitment reversals was obtained in 4/14 experiments where two or more motoneurons were followed pre- and postlesion. 4. The lesion-induced changes in motoneuron discharge characteristics were associated with changes in the relations between muscle force, rectified EMG, and motoneuron rate. Postlesion discharge rates were always significantly lower than the prelesion rates when compared over the same range of EMG levels. This postlesion drop in discharge rates was generally associated with inefficient force production, as evidenced by a significant drop in muscle force for matched EMG levels. 5. The degree of discharge synchrony in MG motoneurons was assessed by calculating a spike-triggered average (STA) between axonal discharge and multiunit rectified EMG. Significant STA peaks were rare before the lesion (4/32 motoneurons) but were quite common after the lesion (29/32 motoneurons).(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials↗