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M D Binder

Publications and source records attributed to M D Binder.

53 records · Page 3Linked to original sources

Recruitment order of motoneurons in stretch reflexes is highly correlated with their axonal conduction velocity.

Motor units of soleus and medial gastrocnemius (MG) muscles were studied in pairs during stretch reflexes in the decerebrate cat to determine the relation between their recruitment orders and axonal conduction velocities. In 97% of soleus pairs, the motor unit with the lower axonal conduction velocity was recruited first. Since the soleus is a homogeneous muscle in the cat, differences in motor-unit type are, therefore, not a sine qua non for orderly recruitment nor is recruitment random within homogeneous populations of motor units, as recently proposed (28). In the medial gastrocnemius, a heterogeneous muscle, the same high correlation (97%) between recruitment sequence and conduction velocity was observed. Thus, the factors that determine recruitment order in heterogeneous muscles are as closely correlated with axonal diameter as they are in homogeneous muscles. Comparison of axonal conduction velocities in our sample of MG units with those in three samples of type-identified MG units studied by other investigators also suggests that motor-unit type is not the critical factor controlling the sequence of activation in heterogeneous muscles. It is concluded that the combined effects of all presynaptic and postsynaptic factors that determine susceptibility to discharge in motoneurons during stretch reflexes are strictly correlated with their axonal conduction velocities, as predicted by the size principle.

Animals↗

A commentary on muscle unit properties in cat hindlimb muscles.

A broad survey of muscle unit properties in 14 muscles of the cat hind limb is presented which emphasizes some general features of unit properties in mammalian muscles. A more detailed analysis of muscle unit properties in three muscles of the posterior compartment of the lower leg is then presented using Burke's tetrapartite (FF, FI or F (Int.), FR, and S) unit classification scheme. Our data on the properties of motor units in cat tibialis posterior (TP) have been compared to those generated by Burke and colleagues on units in flexor digitorum longus (FDL) and medial gastrocnemius (MG). In all three muscles, twitch contraction time was distinctly slower for type S units and specific tension outputs were substantially greater for type FF units than for type S units. The innervation ratios of type FR units were slightly lower than for type S units but the specific tension of the FR units was closer to FF units than to type S units. The FF units controlled 70-74% of the cumulative force output of each muscles, indicating a substantial capacity for powerful rapid contractions of all three of these muscles despite their differences in "size," action, and force generation. Distinctive features of the three muscles included differences in the unit types' force producing capabilities and in the relative representation of "nonfatigable" type FR and S units in each muscle. In particular, TP is endowed with some unusually powerful type FF units and a high percentage (42%) of type S units. In contrast, FDL has units that develop relatively little force and an unusually high representation (56%) of type FR units. The possible relationships between these muscle features and their presumed role in posture and locomotion is discussed.

Animals↗

Topographic organization of monosynaptic reflexes in the cat spinal cord.

The topographic organization of monosynaptic reflexes in the cat spinal cord has been studied by comparing the amplitude of reflex discharges recorded from ventral roots consequent to stimulation of dorsal roots entering the cord at different spinal segments. The results indicate that up to 80% of the potentiated monosynaptic reflex discharge recorded from a ventral root can be attributed to afferent input entering the spinal cord at the same segmental level. Moreover, within the same segment, afferents with a more rostral cord entry level exert a stronger synaptic effect on the more rostral portion of the corresponding ventral root.

Afferent Pathways↗

Motor unit--muscle spindle interactions in active muscles of decerebrate cats.

Single muscle spindle afferent and motor unit EMG spike trains have been recorded simultaneously during periods of spontaneous motor activity in triceps surae muscles of decerebrate cats. The approximate time course and magnitude of the motor unit contractions were extracted from the whole muscle force record by spike-triggered averaging, and the functional interactions between motor unit contractions and spindle discharge were assessed by cross-correlating their respective spike trains. We have found that both spindle group Ia and II afferents are responsive to the contractions of single motor units in the presence of spontaneous motor activity, being strongly coupled to the activity of some motor units and indifferent to the contractions of others. Moreover, the cross-correlation analysis revealed modulation of a single motor unit's discharge pattern by the input of a single Ia afferent.

Action Potentials↗

Responses of Ia and spindle group II afferents to single motor-unit contractions.

1. The responses of deefferented Ia and spindle group II afferents to electrically activated twitch contractions of randomly selected motor units of the cat tibialis posterior muscle have been studied. Each afferent was paired with from 8 to 20 of the muscle's 60 motor units, and each afferent-motor unit interaction was recorded to two muscle lengths. 2. Cross-correlation histograms were compiled for each afferent-motor unit interaction studied as well as the average twitch tension produced by the motor unit. A numerical "coupling index" was computed for the histogram distributions to quantitate the extent of mechanical coupling between the receptor and the single motor units. 3. Qualitatively, no consistent differences were noted in the responses of Ia and spindle group II afferents to single motor-unit contractions. However, Ia afferents were responsive to a higher percentage of motor units with which they were tested (89%) and, on the average, displayed a significantly larger magnitude of response (mean coupling index, 0.72 +/- 0.04 SE) than the spindle group II afferents (66% of motor units; mean coupling index, 0.51 +/- 0.03). 4. The extent to which a motor-unit contraction altered the discharge pattern of a spindle afferent was not strictly related to the amount of force generated by the unit, nor to its contraction time. 5. Muscle length exerted a strong influence on both the qualitative and quantitative features of many of the motor unit-muscle receptor interactions. 6. These results suggest that the degree of "mechanical coupling" between a receptor and a motor unit is largely dependent on anatomical arrangements and reinforce the possibility that muscle receptors generate a "sensory partitioning" of the motor-unit population within a muscle.

Animals↗

Tetrapartite classification of motor units of cat tibialis posterior.

The results of this study and its precedents suggest that the tetrapartite classification scheme might have universal applicability to at least the muscle units of cat hindlimb muscles and perhaps any mammalian muscle in which fiber typing reveals the presence of FG, FI, FOG, and type SO fibers. A possible exception to this generalization involves a small (n = 18) but thoroughly examined sample of muscle units from the first deep lumbrical muscle of the cat's foot, which led Kernell et al. (30) to conclude that the FF, FR, and S classification scheme was not directly applicable to that muscle. However, histochemical fiber typing is not yet available for that muscle. Furthermore, more extensive sampling, use of a different stimulation regime in the fatigue test, and a more detailed analysis of the sag property might well reveal that the tetrapartite classification scheme is indeed appropriate for units of cat foot muscles.

Animals↗

Speed-force relations in the motor units of the cat tibialis posterior muscle.

The neuro-mechanical properties and speed-force relations of 81 cat tibialis posterior motor units have been studied. Statistically significant correlations were found between alpha-axonal conduction velocity (CV) and average twitch tension (log10 TwT; r = 0.459, p less than 0.001), between CV and twitch contraction time (CT; r = -0.395, p less than 0.001) and between CT and log10 TwT (r = -0.277, p less than 0.02). The present correlations for the intermediate-sized tibialis posterior are stronger than those previously reported for large muscles such as soleus, medial gastrocnemius, plantaris and flexor hallucis longus. However, they are considerably weaker than those reported for the much smaller lumbrical muscles of the cat's foot. These findings support the contention that the spinal mechanisms governing an orderly recruitment of motor units according to the size of their muscle units must be more complex for large than for small muscles, at least in the cat hindlimb.

Animals↗

The response of Golgi tendon organs to single motor unit contractions.

1. Cross-correlation analysis has been used to quantify the responses of cat soleus tendon organs to repetitive twitch contractions of: (a) different motor units within the muscle, (b) single motor units at different muscle lengths, and (c) single motor units when the pulse-train pattern of stimulation delivered to the motor unit axon was altered. 2. Ib afferents were observed which responded to each of several hundred successive motor unit twitches with identical numbers of spikes and with relatively invariant latencies. 3. The present results show that tendon organs are sensitive to subtle alterations in motor unit twitch wave form and amplitude, and that this sensitivity is reflected in the precise timings of their afferent discharge. 4. Examination of these tendon organ responses indicates that the forces produced by single motor units couples to the receptor capsule are well above threshold. Calculations based on these results, and earlier soleus motor unit and muscle fibre data, suggest that the absolute force threshold for tendon organs may be as little as 4 mg, which is less than the estimated minimum twitch force generated by individual soleus muscle fibres. 5. Considering the number of tendon organs in a muscle, and the likelihood that every motor unit is connected with at least one receptor, the sensitivity of tendon organs ensures that every twitch of every motor unit will be reflected in the population of afferent signals projecting to the spinal cord.

Action Potentials↗

Correlation analysis of muscle spindle responses to single motor unit contractions.

1. Cross-correlation techniques have been used to study the responses of muscle spindle afferents from the soleus muscle of the cat to twitch contractions of single motor units. 2. Cross-correlograms (post-stimulus time histograms) were used to give the frequency of occurrence of a receptor spike at various times following the initiation of a motor unit contraction together with a display of the average twitch tension wave form. 3. The cross-correlograms revealed that the contraction of a single motor unit can be an effective stimulus to a spindle receptor and may induce afferent firing pattern alterations similar to those observed with whole muscle contraction. 4. The cross-correlograms also revealed quantitative differences in the response of a receptor to contraction of different motor units and to contraction of the same motor unit at different lengths. These differences reflect subtle changes in receptor deformation developed by the twitch of a motor unit under different conditions and by the twitches of different motor units. The results are consistent with anatomical data on the number and distribution of motor units and receptor organs in cat soleus. 5. These findings emphasize that rather than simply acting as generalized force or length sensors for the muscle as a whole, each receptor's spike train carries information about the state of a particular set of motor units.

Afferent Pathways↗

Functional identification of the input-output transforms of mammalian motoneurones.

We studied the responses of rat hypoglossal and cat lumbar motoneurones to a variety of excitatory and inhibitory injected current transients during repetitive discharge. The amplitudes and time courses of the transients were comparable to those of the synaptic currents underlying postsynaptic potentials (PSPs) recorded in these cells. Poisson trains of these current transients were combined with an additional independent, high frequency random waveform to approximate band-limited white noise. The composite, white noise waveform was then superimposed on long duration suprathreshold current steps. We used the responses of the motoneurones to the white noise stimulus to derive zero-, first- and second-order Wiener kernels, which provide a quantitative description of the relation between injected current and discharge probability. The convolution integral computed for an injected current waveform and the first-order Wiener kernel provides the best linear prediction of the associated peristimulus time histogram (PSTH). This linear model provided good matches to most of the PSTHs compiled between the times of occurrence of individual current transients and motoneurone discharges. However, for the largest amplitude current transients, a significant improvement in the PSTH match was often achieved by expanding the model to include the convolution of the second-order Wiener kernel with the input. The overall transformation of current inputs into firing rate could be approximated by a second-order Wiener Model, i.e., a cascade of a dynamic, linear filter followed by a static non-linearity. At a given mean firing rate, the non-linear component of the motoneurone's response could be described by the square of the linear component multiplied by a constant coefficient. The amplitude of the response of the linear component increased with the average firing rate, whereas the value of the multiplicative coefficient in the nonlinear component decreased. As a result, the overall transform could be predicted from the mean firing rate and the linear impulse response, yielding a relatively simple, general description of the motoneurone's input-output function.

Action Potentials↗

Synaptic integration in spinal motoneurones.

Spinal motoneurones receive thousands of presynaptic excitatory and inhibitory synaptic contacts distributed throughout their dendritic trees. Despite this extensive convergence, there have been very few studies of how synaptic inputs interact in mammalian motoneurones when they are activated concurrently. In the experiments reported here, we measured the effective synaptic currents and the changes in firing rate evoked in cat spinal motoneurones by concurrent repetitive activation of two separate sets of presynaptic neurons. We compared these effects to those predicted by a linear sum of the effects produced by activating each set of presynaptic neurons separately. We generally found that when two inputs were activated concurrently, both the effective synaptic currents and the synaptically-evoked changes in firing rate they produced in motoneurones were generally linear, or slightly less than the linear sum of the effects produced by activating each input alone. The results suggest that the spatial distribution synaptic terminals on the dendritic trees of motoneurones may help isolate synapses from one another, minimizing non-linear interactions.

Animals↗

Multiple mechanisms of spike-frequency adaptation in motoneurones.

Spike-frequency adaptation is the continuous decline in discharge rate in response to a constant stimulus. We have described three distinct phases of adaptation in rat hypoglossal motoneurones: initial, early and late. The initial phase of adaptation is over in one or two intervals, and is primarily due to summation of the calcium-activated potassium conductance underlying the medium duration afterhyperpolarization (mAHP). The biophysical mechanisms underlying the later phases of adaptation are not well understood. Two of the previously-proposed mechanisms for adaptation are an increase in outward current flowing through calcium-activated potassium channels and increasing outward current produced by the electrogenic sodium-potassium pump. We found that neither of these mechanisms are necessary for the expression of the early and late phases of adaptation. The magnitude of the initial phase of adaptation was reduced when the calcium in the external solution was replaced with manganese, but the magnitudes of the early and late phases were consistently increased under these conditions. Partial blockade of the sodium-potassium pump with ouabain had no significant effect on any of the three phases of adaptation. Our current working hypothesis is that the magnitude of late adaptation depends upon the interplay between slow inactivation of sodium currents, that tends to decrease discharge rate, and the slow activation or facilitation of a calcium current that tends to increase discharge rate. Adaptation is often associated with a progressive decrease in the peak amplitude and rate of rise of action potentials, and a computer model that incorporated slow inactivation of sodium channels reproduced this phenomenon. However, the time course of adaptation does not always parallel changes in spike shape, indicating that the progressive activation of another inward current might oppose the decline in frequency caused by slow sodium inactivation.

Action Potentials↗