Spinal stretch reflexes of triceps surae in active and passive movements [proceedings].
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Biomedical subjects
Publications and source records attributed to J Noth.
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1. Single fusimotor fibres were stimulated repetitively to test their action on the responsiveness of muscle spindle primary endings in the cat soleus to sinusoidal stretching of both large and small amplitude. Frequencies of 0.06-4 Hz were used at amplitudes from 10 mum to 3 mm.2. The response was assessed by fitting a sinusoid to the cycle histogram of the afferent firing throughout the course of the cycle; this linear approximation measures the fundamental of the response and ignores any harmonics. The sine was allowed to project to negative values and any empty bins in the histogram were ignored when fitting.3. With small amplitudes of stretching the histograms were reasonably sinusoidal, but with large amplitudes they showed appreciable distortion of the wave form for the passive ending and during dynamic fusimotor stimulation. Non-linearity of response manifested itself also, with increasing amplitude of stretching, by an increase in the phase advance of the response, by increasing r.m.s. deviation of the histogram points from the fitted sine and (for dynamic stimulation) by an increase in the mean value of the fitted sine.4. With increasing amplitude the response modulation ceased to increase proportionately with the stimulus, so that the sensitivity of the ending to a large stretch (defined as afferent modulation/stretch amplitude) was appreciably less than for a small stretch. This effect was most pronounced for the passive ending.5. Whatever the amplitude of movement the modulation during static stimulation was less than that for the passive or during dynamic stimulation. For small amplitudes the response during dynamic stimulation was less than that of the passive, but for large amplitudes the response during dynamic stimulation was always the greater. At some intermediate cross-over amplitude the two responses were the same size, though still differing slightly in other respects. The value of the cross-over amplitude was usually about 200 mum at 1 Hz, and increased on lowering the frequency. Thus dynamic fusimotor action does not uniformly produce either an increase or a decrease in the sensitivity of the ending in relation to the passive.6. Bode plots, for each amplitude, of sensitivity and phase against frequency suggested that(a) under all conditions the ending is relatively insensitive to frequency in the range studied, for the slope of the log-log sensitivity lines was only 0.15-0.2 (3.5-6 db/decade);(b) the mechanism which makes for non-linearity is not particularly frequency sensitive;(c) static fusimotor stimulation does not change the frequency sensitivity of the ending;(d) dynamic fusimotor stimulation very slightly increases the frequency sensitivity of the ending for large amplitudes.In reaching these conclusions more attention was paid to the slope of the sensitivity lines than to the values of phase.7. It appears that the major effect of fusimotor action, whether static or dynamic, is to regulate the sensitivity of the primary ending to stretching for all amplitudes of movement (i.e. gain) rather than to control the relative values of its sensitivity to length and to velocity (i.e. crudely, the damping in a feed-back loop).
1. A pair of fusimotor fibres, one static and the other dynamic, were stimulated simultaneously to test their combined action on the response of muscle spindle primary endings in the cat soleus to sinusoidal stretching. A frequency of 1 Hz was chiefly used, with a wide range of amplitudes (10 micronm-2 mm). The response of the ending was assessed from the parameters of the sine fitted to its firing averaged throughout the course of the cycle; this was felt useful even though the responses to the larger stretches showed certain non-linear features. 2. With small stretches (up to about 50 micronm amplitude) static action dominated, and the modulation of firing during conbined stimulation was little or no larger than that found during the static stimulation on its own, and much smaller than that found during the static stimulation on its own, and much smaller than that found during the dynamic stimulation. The phase of the response was, however, much the same for all three conditions. 3. With larger stretches the modulation with combined stimulation was intermediate between the values found on stimulating either fusimotor fibre on its own; the dynamic contribution increased progressively with the amplitude of stretching. 4. With larger stretches the phase of the response during combined stimulation was appreciably closer to that for static action than to that for dynamic action. But the differences between the various conditions were small (below 20 degrees) and seem attributable to various distortions of the response wave from away from a true sinusoid, rather than betokening a difference in the ratio of velocity to length sensitivity under the various conditions. This view was supported by the effects on phase of grading the rate of stimulation of one fusimotor fibre while holding that of the other constant. 5. Detailed comparison of the cycle histograms obtained under different conditions showed an interestingly asymmetrical pattern of summation and occlusion of the effects of the two kinds of fusimotor fibre. At the peak of the response to a large stretch static action summed with dynamic action, which was here the stronger, so that at this phase of the cycle the firing was greater with the combined stimulation than with either fibre on its own. But, in the trough of the response to the same stretch static action occluded any dynamic action, which was now the weaker, so that at this phase of the cycle the firing with combined stimulation was virtually the same as that with static stimulation on its own. With a small stretch, static action normally occluded dynamic action throughout the cycle; this is in line with the firing during static action now usually being greater than that during dynamic action for all phases of the cycle.
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About 2/3 of the gamma efferents isolated from the medial gastrocnemius nerve were inhibited by longitudinal high-frequency vibration applied to the tendons of the non-contracting pretibial flexors (decerebrate cats). The inhibition appeared at 15-25 mum amplitude of vibration and increased up to maximum at nearly 100 mum. Increasing the frequency of vibration from 100 to 300 Hz increased the inhibition. The reflex effects elicited by muscle vibration corresponded well in incidence and magnitude with those evoked by tetanization of the deep peroneal nerve at group I stimulus strength. The reflex disappeared when the nerve supply of the vibrated muscles was cut. The sensitivity of some pretibial proprioceptors to vibration was also tested. It is concluded that primary spindle endings of the pretibial flexors inhibit the extensor gamma motoneurons. Some findings hint at a spinal pathway involving Ia inhibitory interneurons. In addition, an inhibitory action of pretibial group II afferents, probably secondary spindle endings, on extensor gamma efferents was demonstrated. The described fusimotor inhibition by antagonistic muscle spindle afferents is a further example of alpha-gamma-linkage.
1. High frequency vibration was applied to the tendon of the non-contracting triceps surae muscle while recording the background discharges of single gamma fibres only small nerve bundles were cut, leaving most of the nerve supply to the triceps intact. 2. 22% out of a total of sixty-three gamma efferents were tonically inhibited by vibration. The inhibition appeared between 25 and 50mum peak-to-peak amplitude of vibration and increased to a plateau for amplitudes of about 100mum. The dependence of the tonic vibration reflex of alpha-efferents on the amplitude of vibration was found to be similar. Increasing the frequency of vibration from 150 to 300 Hz increased the degree of inhibition. 3. 33% of the fusimotor neurones investigated responded to muscle vibration with an increase in discharge rate. The threshold amplitudes of this reflex ranged from 20 to 50mum. Some features of the reflex, in particular the parallel post-vibratory facilitation found in alpha and gamma efferents, pointed to a polysynaptic pathway organized in an alpha-gamma linkage. 4. All gamma efferents inhibited by vibration showed inhibitory responses to antidromic stimulation of the parent ventral root, and most of them were inhibited by ramp stretch of the triceps. The gamma motoneurones facilitated by vibration, however, were excited by muscle stretch and were less susceptible to antidromic inhibition, some lacking it completely. 5. Cutting the nerves to triceps abolished the inhibitory as well as the excitatory responses of gamma efferents to muscle vibration. Both fusimotor reflexes were preserved after spinal section and subsequent administration of L-DOPA. 6. It is concluded that both of the fusimotor reflex effects of vibration are caused by excitation of primary spindle endings within the triceps. The inhibition of fusimotor neurones is thought to be mediated by Renshaw cells activated during vibration. The significance of positive feed-back on to gamma motoneurones as a result of autogenetic facilitation by Ia afferents is discussed in connexion with stability in the stretch reflex loop.
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The effect of hypertonic solutions on the tension of isolated twitch muscle fibers of the frog has been investigated. Increased tonicity up to about 1.7 times normal (1.7 T) caused a very small, graded, maintained tension increase. Above about 1.7 T a large, transient contracture response was superimposed on the small tension. The contracture response was graded with tonicity and reached a maximum at 2.5 T of 108 +/- 25 mN.mm(2) a third of the maximum tetanic tension in isotonic solution. Contracture tension developed with a delay which decreased with increased tonicity. The contracture threshold was lower and the delay shorter in small fibers than in large. Contractures were obtained equally well in depolarized as in polarized fibers. They were completely suppressed by 0.1-0.5 mM tetracaine. The possible mechanism responsible for the tension-inducing effect of hypertonic solutions is discussed in terms of the close similarity between the properties of these contractures and those caused by caffeine, and it is suggested that the effect is due to a release of calcium from internal stores.
Resting tension and short-range elastic properties of isolated twitch muscle fibers of the frog have been studied while bathed by solutions of different tonicities. Resting tension in isotonic solution at 2.3-microm sarcomere spacing averaged 0.46 mN.mm(-2) and was proportional to the fiber cross-section area. Hypertonic solutions, containing 0.1-0.5 mM tetracaine to block contracture tension, caused a small sustained tension increase, which was proportional to the fiber cross-section area and which reached 0.9 mN.mm(-2) at two times normal tonicity (2T). Further increases in tonicity caused little increase in tension. Hypotonic solutions decreased tension. Thus, tension at 2.3 microm is a continuous, direct function of tonicity. The dependence of tension on tonicity lessened at greater sarcomere lengths. At 3.2 microm either a very small rise or, in some fibers, a fall in tension resulted from an increase in tonicity. Hypertonic solutions also decreased the tension of extended sarcolemma preparations. In constant-speed stretch experiments the elastic modulus, calculated from the initial part of the stretch response, rose steeply with tonicity over the whole range investigated (1-2.5T). The results show that tension and stiffness of the short-range elastic component do not increase in parallel in hypertonic solutions.
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