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Structure, distribution and innervation of muscle spindles in avian fast and slow skeletal muscle.

Muscle spindles in 2 synergistic avian skeletal muscles, the anterior (ALD) and posterior (PLD) latissimus dorsi, were studied by light and electron microscopy to determine whether morphological or quantitative differences existed between these sensory receptors. Differences were found in the density, distribution and location of muscle spindles in the 2 muscles. They also differed with respect to the morphology of their capsules and intracapsular components. The slow ALD possessed muscle spindles which were evenly distributed throughout the muscle, whereas in the fast PLD they were mainly concentrated around the single nerve entry point into the muscle. The muscle spindle index (number of spindles per gram wet muscle weight) in the ALD was more than double that of its fast-twitch PLD counterpart (130.5+/-2.0 vs 55.4+/-2.0 respectively, n = 6). The number of intrafusal fibres per spindle ranged from 1 to 8 in the ALD and 2 to 9 in the PLD, and their diameters varied from 5.0 to 16.0 microm and 4.5 to 18.5 microm, respectively. Large diameter intrafusal fibres were more frequently encountered in spindles of the PLD. Unique to the ALD was the presence of monofibre muscle spindles (12.7% of total spindles observed in ALD) which contained a solitary intrafusal fibre. In muscle spindles of both the ALD and PLD, sensory nerve endings terminated in a spiral fashion on the intrafusal fibres in their equatorial regions. Motor innervation was restricted to either juxtaequatorial or polar regions of the intrafusal fibres. Outer capsule components were extensive in polar and juxtaequatorial regions of ALD spindles, whereas inner capsule cells of PLD spindles were more numerous in juxtaequatorial and equatorial regions. Overall, muscle spindles of the PLD exhibited greater complexity with respect to the number of intrafusal fibres per spindle, range of intrafusal fibre diameters and development of their inner capsules. It is postulated that the differences in muscle spindle density and structure observed in this study reflect the function of the muscles in which they reside.

Animals↗

Modification by lidocaine of the discharges of primary endings of muscle spindles in cat tenuissimus muscle.

The effects of lidocaine (1--20 microgram/ml) on afferent discharge patterns of primary endings of muscle spindles in cat tenuissimus muscle were investigated. Discharge from the endings, recorded in Ia afferent axons, was evoked by ramp stretch of the muscle, stimulation of single static or dynamic fusimotor axons or by a combination of stretch and fusimotor stimulation. Spontaneous discharge of the endings at the initial length of the muscles was reduced by 2--5 microgram/ml and abolished by 10--15 microgram/ml lidocaine. The static but not the dynamic discharge elicited by muscle stretch was blocked by concentrations of 10--15 microgram/ml. The same concentrations abolished static and dynamic fusimotor influences on primary ending discharge. However, in one experiment where the spindle was microscopically observed, fusimotor stimulation still resulted in contraction of the intrafusal muscle even when fusimotor stimulation failed to elicit changes in discharge response patterns of the primary endings. These findings indicate that lidocaine interferes with the encoding mechanism prior to block of impulse conduction in either the fusimotor or afferent axons.

Animals↗

Dynamic and static stretch responses in muscle spindle receptors in fatigued muscle.

Dynamic and static stretch responses in muscle spindles were investigated in fatigued muscle to determine if acute adaptations do occur in receptor discharge as has been shown after contractions of short duration. Fatigue to 60-50% maximum tetanic tension was induced in the isolated gastrocnemius muscle in 16 cats by sustained, 7 X threshold electrical stimulation (100 Hz) of the cut L7 ventral root and S1 ventral root. Group Ia and II afferent fiber responses to slow ramp stretches (5 mm X s-1) and vibration (100 Hz) applied to the Achilles tendon were monitored before and immediately after muscle tetany to fatigue. Changes in firing characteristics were similar when results from faster (25-30 mm X s-1) ramp stretches were contrasted. During muscle fatigue, decreases in response latency to displacement and increases in resting discharge, mean frequency during stretch, and frequency of firing to vibration were predominant in both afferent fiber types. Static responses were generally lower, indicating a decrease in position sensitivity. Resting muscle force and passive peak muscle stiffness were consistently higher following contraction. The sum effects of these proprioceptive afferent and mechanical muscle responses would be to increase muscle stiffness and thus resist yield in muscle length to perturbations at lower muscle forces. The magnitude of these adaptations in proprioceptive discharge appears dependent on intrafusal muscle fiber activation.

Adaptation, Physiological↗

Fine structure of nerve processes containing basic fibroblast growth factor in muscle spindles of the rat masseter muscle.

An antiserum against basic fibroblast growth factor (bFGF) was characterized by immunoblot analysis and used to investigate the fine structure of bFGF-containing processes in muscle spindles of the rat masseter muscle. The bFGF antiserum recognized purified bFGF and bFGF-like materials with the same molecular weight as bFGF in crude homogenate of the brainstem from which bFGF fibers in the masseter muscle spindles presumably originate. Immunoelectron micrographs demonstrated that both proprioceptive and motor nerve endings in contact with intrafusal fibers contain bFGF. These findings suggest that bFGF of central (possibly trigeminal mesencephalic and motor nucleus) origin is transported into the muscle spindles subserving mechanoreception.

Animals↗

Existence of a muscle spindle on the posterior cricoarytenoid muscle of the guinea pig.

We examined a muscle spindle in the posterior cricoarytenoid muscle of the adult guinea pig. The spindle capsule contained three intrafusal muscle fibres. At the equatorial region, three intrafusal fibres did not form a typical aggregation of nuclei such as bags and chains observed in common mammalian muscle spindles. Two intrafusal fibres of the three were innervated by sensory endings which branched and arranged irregularly. These sensory endings lay on the surface of intrafusal fibres in shallow grooves or deeply penetrated into the fibre. The third intrafusal fibre, lacking sensory innervation, showed a similar fine structure to extrafusal fibres. These findings suggest that the muscle spindle in the posterior cricoarytenoid muscle of the guinea pig is strikingly different in the structure of intrafusal fibres and in the arrangement of sensory endings from that in common mammalian skeletal muscles. The occurrence and role of these different structures were discussed.

Animals↗

Ultrastructural abnormalities of muscle spindles in the rat masseter muscle with malocclusion-induced damage.

Human temporomandibular disorders due to disturbed occlusal mechanics are characterized by sensory, motor and autonomic symptoms, possibly related to muscle overwork and fatigue. Our previous study in rats with experimentally-induced malocclusion due to unilateral molar cusp amputation showed that the ipsilateral masseter muscles undergo morphological and biochemical changes consistent with muscle hypercontraction and ischemia. In the present study, the masseter muscle spindles of the same malocclusion-bearing rats were examined by electron microscopy. Sham-operated rats were used as controls. In the treated rats, clear-cut alterations of the muscle spindles were observed 26 days after surgery, when the extrafusal muscle showed the more severe damage. The fusal alterations affected predominantly capsular cells, intrafusal muscle fibers and sensory nerve endings. These results suggest that in the malocclusion-bearing rats, an abnormal reflex regulation of the motor activity of the masticatory muscles may take place. They also allow us to hypothesize that muscle spindle alterations might be involved in the pathogenesis of human temporomandibular disorders.

Animals↗

Reflex activation of muscle spindles in human pretibial muscles during standing.

1. Experiments were performed in standing subjects to determine whether low-threshold cutaneous and muscle afferents from mechanoreceptors in the human foot reflexly influence fusimotor neurons innervating pretibial flexor muscles. Recordings were made from 30 identified muscle-spindle afferents, four tendon-organ afferents, and one alpha-motor axon innervating the pretibial flexor muscles. The subjects stood without support or vision on a force platform while trains of electrical stimuli (5 stimuli, 300 Hz) were delivered at nonpainful intensities to the sural nerve or to the posterior tibial nerve at the ankle. 2. Seventeen of the 30 spindle endings had no background discharge, and none was activated by the sural or posterior tibial stimuli. Five silent afferents were given a background discharge by sustained pressure on the relevant tendon, but with two the discharge was dominated by a tremor rhythm obscuring any reflex response to the stimuli. Based on peristimulus time histograms (PSTHs), the sural stimuli then produced increases in discharge of two of the remaining three endings at latencies of 84 and 90 ms. These effects could not be explained by muscle stretch and are presumed to have been fusimotor mediated. 3. When the subjects stood freely without support or vision, 13 muscle-spindle endings had a background discharge, but with three endings tremor developed at the ankle and dominated the spindle discharge. Sural stimuli affected the discharge of five of nine endings unaffected by tremor. With three of these endings, there were changes in discharge that could be explained by muscle stretch.(ABSTRACT TRUNCATED AT 250 WORDS)

Adult↗

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↗

Effect of diazepam on the gamma motor system indicated by the responses of the muscle spindle of the triceps surae muscle of the decerebrate cat to the muscle stretch.

The experiments were performed on 37 intercollicularly decerebrate cats. The triceps surae muscle was subjected to trapezoidal stretches of 10 mm/s or 2 mm/s over a length of 14 mm. The reflex arc was left partially or fully intact. The effect of diazepam given intravenously on the discharge frequencies of group Ia afferent fibres from muscle spindles in the belly of the triceps surae muscle was studied. Diazepam depressed both the dynamic and the static gamma motor responses. When gamma motor fibres were selectively blocked by procaine a parallel shift of the tension-extension curve was observed. From comparison of the effect of procaine and diazepam on the phasic and static responses of the muscle spindle it was concluded that directly after the injection of diazepam the gamma motor activity was completely abolished at a dose as low as 0.1 mg/kg whereas the alpha motor system was only partly depressed even at higher doses. Differences in the effect of diazepam on the dynamic and static gamma system respectively, could not be found.

Action Potentials↗

Possible fusimotor innervation to frog muscle spindles.

In some muscle spindles located in or near the red area of semitendinosus muscle of the bull frog, a fine motor axon innervates the intracapsular compact zone, without any branches onto the extrafusal muscle fibers. The fine motor axon is 3.9 microns in mean diameter (measured in living axon) and 9.8 m X s-1 in mean conduction velocity at 20 degrees C. A ramp-and-hold stretch during 20 Hz repetitive stimulation of the fine motor axon markedly increased the rate of afferent discharges during static phase. Histochemical studies indicated depletion of glycogen in a compact zone along a small diameter intrafusal muscle fiber of the muscle spindles frozen immediately after tetanic stimulation of the fine motor axon. These results suggest that the fine motor axon is a fusimotor system in the frog muscle spindle.

Action Potentials↗

Immunocytochemical localization of Na, K-ATPase in rat muscle spindles.

In the muscle spindle, one of the major sensory receptors in the vertebrate skeletal muscle, it was demonstrated that stretching caused a conductance increase of the sensory terminal membrane mainly to Na+ (Hunt, Wilkinson and Fukami, 1978 (6)). Since the muscle spindle is a slowly adapting stretch receptor, and even at rest some spindles are active, a vigorous Na, K-pump activity is expected to counteract the incessant inflow of Na+ into the terminal. To test this assumption, rat muscle spindles were examined by immunofluorescence microscopy as well as by the electron microscopic immunogold technique using antibody against rat alpha-subunit of Na, K-ATPase. The results indicate that the sensory ending has the highest density of the enzyme among the other cellular components examined, and that the enzyme density appears to be higher in the plasma membrane of the sensory ending facing the intrafusal muscle fiber (synaptic membrane) than the rest of the membrane (extra-synaptic membrane). The functional significance of the above findings was discussed.

Animals↗

Morphology and distribution of muscle spindles in dorsal muscles of the cat neck.

Silver-stained sections have been examined from the five dorsal neck muscles splenius, biventer, cervicis, complexus, rectus capitis major, and occipitoscapularis. Every serial section was examined for at least one muscle of each type so that a complete description of the spindle distribution and morphology could be made. With the exception of occipitoscapularis (whose prime function is probably in scapula rotation and not in head movement), neck muscles have a remarkably high spindle density. Occipitoscapularis has a spindle density similar to that of hindleg locomotor muscles (13-19 per gram), but splenius has a density of 46-66 per gram, biventer cervicis 74-96 per gram, complexus 71-107 per gram, and rectus capitis major 48-84 per gram. Such high densities have only previously been seen in small muscles whose total spindle population is not large. Because of the relatively large size of some neck muscles, individual muscles with a spindle content of up to 254 spindles have been found.

Animals↗