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Density of muscle spindles in the jaw muscles of the Japanese flying squirrel and the guinea pig.

The number of muscle spindles and their distribution pattern in the jaw muscles were examined in serial sections of the heads of the Japanese flying squirrel and the guinea pig, and were compared with each other. The Japanese flying squirrel had 127 to 177 spindles, the length of which ranged from 140 microns to 1400 microns; 74 to 99 in the masseter, 39 to 47 in the temporalis, 11 to 28 in the zygomaticomandibularis, zero to one in the maxillomandibularis and two to three in the medial pterygoid muscles. The guinea pig had 322 to 346 spindles, the length of which ranged from 140 microns to 2800 microns; 116 to 125 in the masseter, 15 to 16 in the temporalis, 98 to 107 in the zygomaticomandibularis, 89 to 94 in the maxillomandibularis, and three to five in the medial pterygoid muscles. The lateral pterygoid, mylohyoid and anterior digastric muscles were devoid of spindles, as was the case in the other rodent species examined.

Animals↗

[Responses of the muscle spindles of tenotomized and hypertrophied muscles to stretch and vibration].

Responses of primary and secondary endings of muscle spindles were studied after tenotomy and hypertrophy in the cat soleus muscles. Under constant load (100 g) background activity of primary endings in a control muscle was 17 +/- 1.5 imp/s. It did not change in hypertrophic muscle and increased after tenotomy to 26 +/- 1.5 imp/s. Responses of secondary endings did not change under the same conditions. The frequency of a response from primary and secondary endings of tenotomized and hypertrophic muscles were higher during dynamic and static phase of stretching as compared to response of normal muscle spindles. Primary endings of tenotomized and hypertrophic muscles were able to repeat vibration up to 200 Hz, while secondary endings responded only with some increase in the discharge frequency.

Animals↗

Effects of fine-grained diet on annulospiral endings of muscle spindles in the masseter muscle in developing and adult mice.

In the masticatory muscles, neuromuscular spindles have a very important role in controlling the jaw movement since they act as stretch receptors in skeletal muscles. The continuous intake of fine-grained diet which is easily chewable leads to degeneration of the sensory endings of Ia fibers in many muscle spindles of the mouse masseter muscle in only 120 d after birth.

Animals↗

Effects of hypodynamia-hypokinesia on the muscle spindle discharges of rat soleus muscle.

The aim of this study was to determine whether Ia and II fiber discharges of soleus muscle spindles were modified after a 14-day period of hypodynamia (absence of weight bearing) and hypokinesia (reduction of motor activity). Fifty-one and 38 afferent fibers were studied, respectively, in control and hypodynamia-hypokinesia (HH) groups. Under deep anesthesia (pentobarbital, 30 mg/kg), a L3-L6 laminectomy was performed. Unitary potentials from the L5 dorsal root were recorded in response to ramp-and-hold stretches applied at two stretch amplitudes (3 and 4 mm) and four stretch velocities (6, 10, 15, and 30 mm/s) and to sinusoidal stretches applied at four stretch amplitudes (0.12, 0.25, 0.5, and 1 mm) and six stretch frequencies (0.5, 1, 2, 3, 6, and 10 Hz). In both animal groups, the Ia fibers showed higher dynamic index values, smaller linear range, and higher vibration sensitivity than the II fibers. They also exhibited a pause in their discharges during the stretch release contrary to II fibers, which displayed no pause in their responses. After HH, our results showed that for both fiber types all parameters measured under ramp-and-hold stretches (except the static sensitivity) were significantly increased and under sinusoidal stretches, the vibration sensitivity increased, and the response amplitude only increased at 0.12-mm stretch amplitude. The linear range of Ia afferents was limited to 0.12 mm, whereas it was unchanged for the II fibers. After HH, the stretches could be better transmitted to the muscle spindles, probably resulting from changes in passive mechanical properties of the soleus.

Animals↗

The response of primary muscle spindle endings to random muscle stretch: a quantitative analysis.

In cats the stretch responses of deafferented Ia afferent fibers from gastrocnemius muscle spindles were measured. The response properties were derived by correlating the action potentials to random mechanical stimuli applied to the gastrocnemius muscle. The muscle stretch consisted of band-limited, normally distributed random changes in muscle length. The upper cut-off frequency of the noise stimuli fc (i.e. the -3 dB point of the amplitude spectrum) was varied systematically between 1.5 and 600 Hz and the amplitude of the noise stimuli sigma (i.e. the standard deviation of the distribution of muscle displacements) between 1.5 and 1200 microns. The neuronal responses were analysed by averaging the stimulus signals centered on the Ia action potentials (the "peri-spike average", PSA). The cross-spectrum between stimulus and neuronal response was determined by computing the Fourier expansion of the PSA. Immediately before the action potentials the PSA decreased below mean muscle length and increased thereafter rapidly. As a rule, the action potentials were elicited close to the point of maximum stretch velocity. The cross-spectra revealed a monotonic increase in gain with stimulus frequency, corresponding to a power function with an average exponent of 0.86. The PSA-shape obtained at stimulus cut-off frequencies between 1.5 and 260 Hz remained fairly independent of fc when the abscissa was normalized with 1/fc. In the cross-spectra a maximum gain was always reached near the actual fc-value for upper cut-off frequencies up to 260 Hz. Above 260 Hz maximum gain remained independently of fc at about 260 Hz. The peak-to-peak displacement A of the PSA increased linearly with the logarithm of fc. The shapes of the PSAs and the cross-spectra did not depend on the noise amplitude sigma in the range above 20 microns. A was linearly related to sigma (about 1.6 to 2.4 . sigma for sigma greater than 10 microns and 75 Hz less than fc less than 450 Hz). A unifying mathematical description of the responses, valid for a wide range of stretch parameters, was obtained. We demonstrated that the application of one short run of random muscle stretch of an intermediate noise amplitude and medium cut-off frequency provides substantial information on the spindle stretch response and is recommended for investigations in which stable recordings cannot be obtained for a sufficient length of time.

Action Potentials↗

Distribution, density, and structure of muscle spindles in the vastus intermedius and the peroneus longus muscles of sheep.

Muscle spindles are not always distributed more in postural muscles with many slow-twitch-oxidative (SO) myofibers than in locomotory muscles with few SO myofibers. The purpose of present study was to examine the distribution, density, and structure of muscle spindles in the vastus intermedius muscle: an antigravity muscle and the peroneus longus muscle: a locomotory muscle in the sheep. Muscle spindles were reconstructed from serial sections at 300 microns intervals throughout the muscles. Myofiber types were classified into SO, fast-twitch-oxidative-glycolytic, and fast-twitch-glycolytic myofibers by differences in histochemical reactivity. No significant difference in the density of muscle spindles (DMS) existed between the vastus intermedius (DMS: 5.3) and peroneus longus (DMS: 5.7) muscles. The muscle spindles were distributed more in the distal portion than in the proximal portion of the vastus intermedius muscle. The muscle spindles were distributed in the proximal and middle portion but hardly in the distal portion of the peroneus longus muscle. Muscle spindles were classified into simple, tandem, and compound muscle spindles. Most of the muscle spindles were the simple type. The differences in size of the muscle spindle and numbers of the intrafusal myofibers were not significant between the two muscles. The results show that the density and structure of the muscle spindles do not differ between the postural and locomotory muscles in the sheep.

Animals↗

Occurrence and distribution of muscle spindles in masticatory and suprahyoid muscles of the rat.

The occurrence and distribution of muscle spindles was studied in histochemically and conventionally stained serial cross sections of 6-week-old and adult rat masticatory and suprahyoid muscles. Spindles were present in moderate to large numbers in jaw closers, but they were absent in jaw openers and two of four muscles of an accessory suprahyoid group. In jaw closers, 67% or more of the total spindle population was concentrated relatively distant from the temporomandibular joint, in muscle portions which contained large numbers of extrafusal fibers reacting strongly for oxidative enzymes. Because of their location, spindles in these portions should be stretched more and, subsequently, should respond with a greater afferent discharge at any given muscle length than spindles situated nearer to the joint. Spindles in jaw closers, especially the medial pterygoid and deep masseter, often occurred in clusters and complex forms near the terminal branching of intramuscular nerve trunks. No such concentrations were seen in the two muscles of the accessory suprahyoid group that had spindles. The association in jaw closers of spindles with extrafusal fibers high in oxidative enzyme activity is consistent with the view that spindles are the sensory component of a reflex system that recruits these fibers for finely-graded contractions in response to small internal length-changes of the muscle (Botterman et al., '78); however, in jaw openers and two muscles of the accessory suprahyoid group, the absence of spindles, coupled with the presence of large populations of extrafusal fibers high in oxidative enzyme activity, is not easily reconciled with this concept.

Animals↗

Morphological variability and specializations in bovine extraocular muscle spindles.

Bovine extraocular muscles were examined to determine whether the structure of their muscle spindles was notably different from those commonly encountered in mammalian limb muscles. Extraocular muscle spindles on the whole were shorter, and intrafusal fiber counts/spindle were more variable than in somatic muscles. No pronounced nuclear bags were seen in intrafusal fibers. Based on cross-sectional areas, intrafusal fibers in extraocular muscles could be loosely categorized as small or large types. Small fibers expressed more neonatal/fast myosin heavy chain and less embryonic myosin heavy chain than large fibers. When incubated for myosin ATPase, about 70% of the large fibers and 15% of the small fibers in spindles presented profiles that were characteristic of type I extrafusal fibers, and not of nuclear bag or nuclear chain fibers. The ratio of number of small intrafusal fibers to number of large intrafusal fibers in extraocular spindles was on average greater than the ratio of nuclear chain fibers to nuclear bag fibers that is typical for limb spindles of rodents and cats. Structural modifications at muscle spindle sensory regions, extrafusal-like fibers and intrafusal-like fibers with few equatorial nuclei and many myofibrils, may produce distinct afferent signals that are appropriate for sensorimotor integration in the specialized extraocular muscles.

Animals↗

Fusimotor-induced changes in muscle spindle outflow and responsiveness in muscle fatigue in decerebrate cats.

Changes in discharge rate and responsiveness of muscle spindle afferents from triceps surae muscles were studied during long-lasting fatigue isometric contractions of either medial gastrocnemius or lateral gastrocnemius and soleus muscles in decerebrate cats. The rest of the hind limb was either denervated or its innervation was preserved. In denervated preparations a long-lasting post-contraction increase in discharge rate developed in the majority of primary (15 of 18) and in all (20) secondary afferents. This increase was abolished, while the decrease during muscle contraction was enhanced after application of procaine to the corresponding muscle nerve, to block either the small-diameter afferents from the contracting muscle or the fusimotor axons to the spindle of origin of the afferent recorded. In innervated preparations the long-lasting increase was replaced in the majority of primary endings (14 of 22) by a sharp burst at the end of muscle contraction, while in secondary afferents it was either absent or shorter-lasting than in denervated preparations. Changes in responsiveness to sinusoidal muscle length changes, indicating influences of both static and dynamic fusimotor neurons, were, however, similar in innervated and denervated preparations. The results obtained provide evidence that changes in muscle spindle outflow and responsiveness are elicited by the reflex increase in fusimotor activity developing in response to the fatigue-induced afferent discharges from the contracting muscle. Concomitant afferent inflow of another origin to fusimotor neurons affects the changes in spindle outflow, but not in responsiveness. In this way an appropriate increase in support to skeletomotor activity as well as in information on the fatigued muscle of higher motor centres, initiated by the fatigue itself, could be achieved through the gamma loop.

Animals↗

The sensory and motor innervation of muscle spindles in cat tail dorsolateral muscles.

The sensory and motor innervation of a sample of 202 muscle spindles from the dorsolateral tail muscles of eight cats were studied by silver and gold staining techniques. The range of diameters of Group Ia afferent nerve fibres to primary (P) sensory endings was 2.0-14.0 micron with a peak at 8.5 micron. The range of Group II fibres to secondary (S) sensory endings was 2.0-8.5 micron with a peak at 5.0 micron. In a sample of 234 spindle capsules, 171 (84.2%) were single capsule spindles and 31 (15.8%) were tandem spindles. With respect to the sensory endings of single capsule spindles, 19.7% showed only a P-ending, 32.5% had an additional S-ending and 15.8%, 3.0% and 2.1% had two, three and four S-endings, respectively. In the tandem spindle capsules, 0.8% of the larger proximal capsules showed only a P-ending, 5.1% had an additional S-ending and 4.7%, 2.6% and 0.4% had two, three and four S-endings, respectively. In the 31 tandem spindles, one of which had a triple capsule, there were 14 types of sensory ending combination in the proximal and distal capsules. The most common (44.4%) innervation was by a P-ending only, while 27.0% had an additional S-ending and 17.5%, 9.5% and 1.6% had two, three and four S-endings, respectively. Of the distal capsules, 92.9% showed only a P-ending. There were three types of motor nerve ending, namely the p1-plate, the p2-plate and the trail (tr) ending, in spindle polar regions. Spindle poles without a motor innervation also occurred. There were 16 types of motor ending combination in the two polar regions of muscle spindles. The most common type of combination was the trp2-ending (41.8%), receiving an average of 5.8 fusimotor fibre branches and the next common was the tr-ending only (38.7%), innervated by an average of 3.2 fibres. The least common combination was with the p1-plate ending (9.5%), receiving an average of 6.9 fibres. The mean number of fusimotor fibre branches per spindle pole was 4.2.

Animals↗

Distribution of muscle spindles in a simply structured muscle: integrated total sensory representation.

BACKGROUND: The distribution of muscle spindles (Sps) in a small muscle of simple architecture, the capsularis at the cat's hip joint, was quantified to reveal the patterns of proprioceptive representation in the transverse and sagittal planes as well as to model the effect a local disturbance in muscle length would have on total Sp discharge. METHODS: Locations in serial cross-sections of the 32 and 38 Sps in 2 muscles, 1 perfused with the hip joint flexed and the other extended, were plotted, and their patterns of integrated sensitivity calculated assuming that (1) the discharge rate of a Sp afferent varies linearly with change in length along the Sp's axis, and (2) that within a local disturbance produced by contraction of a motor unit (MU), lengths decrease either linearly or as the square of the distance from its center. RESULTS: The isomeric pattern of "integrated, total Sp representation on cross-section" showed two peaks of sensitivity in the half of the muscle that had been next to the joint capsule, offset by low representation in a small, central area and along the extensive zone bordering the laterally facing "superficial surface." The equivalent radius of an idealized symmetrical MU territory was estimated from distributions of the few fast, oxidative-glycolytic fibers found in two muscles, and the effect of a MU's contraction on net Sp discharge predicted when the unit was positioned at distinctive sites within the pattern. As an index of Ia and II afferent representation in the sagittal plane, the distribution of the nucleated regions of Sps and the summed lengths of segments of Sp axial bundles and capsules, respectively, within successive 1-mm segments of the muscle were graphed. CONCLUSIONS: The longitudinal representation and structure of the muscle are not suited for reflex adjustment of differences in length along the muscle. The isomeric pattern of high relief in the transverse plane suggests that in this approximately 0.2 g muscle, the localization of myotatic reflexes might be accommodated but the need for adjustment in activation of MUs seems minimal. This is because the muscle is not compartmentalized, its fibers extend between the muscle's origin and insertion, their angle of pinnation is low, and greater than 90% are of slow type. The distribution of Sps is consistent with gauging length of the entire muscle and hence angulation at the hip joint.

Animals↗

Topography of muscle spindles and Golgi tendon organs in shoulder muscles of "Monodelphis domestica".

The topography of muscle spindles and Golgi tendon organs in the rotator cuff and surrounding shoulder muscles of a small laboratory marsupial (monodelphis domestica) were studied using light microscopy of serial sections. The shoulder joint of monodelphis has a large degree of freedom of movement allowing this animal to use the upper extremities for a wide range of activities like climbing and manipulating food. Thus, similar to the situation in man the shoulder joint is mainly secured by muscles. Silver stained serial paraffin sections were examined under the light microscope and the distribution of muscle spindles and Golgi tendon organs was reconstructed using three-dimensional image processing. In the two animals examined 113 and 131 muscle spindles respectively were found within the 4 rotator cuff muscles. In addition, 76 and 40 Golgi tendon organs respectively were seen at the musculo-tendinous junctions of these muscles preferentially close to the insertion at the humerus head. Also the surrounding shoulder muscles contain both muscle spindles and Golgi tendon organs in large numbers, but the ratio of Golgi tendon organs per muscle spindle appears to be lower. Number and localization of muscle spindles and Golgi tendon organs suggest, that these receptors are important for both reflex control of shoulder muscle tone as well as monitoring of static position and movement in the shoulder joint.

Animals↗

The number, distribution and density of muscle spindles in two wing muscles of the domestic duck.

A study by serial longitudinal sections of two dorsal wing muscles of the duck, the extensor pollicis and the extensor digitorum communis, has shown that: (1) The extensor pollicis muscle shows a spindle content range of 19--27, mean 23.2, giving a spindle density of 76.7/g; the extensor digitorum communis muscle shows a spindle content range of 37--59, mean 47.6, and a spindle density of 119.0/g. (2) Spindles in the extensor pollicis muscle are randomly distributed, while those in the extensor digitorum muscle are located primarily in the proximal two thirds of the muscle. (3) The present data have been discussed in relation to previous observations of muscle spindles occurring in some avian and mammalian skeletal muscles.

Animals↗

Muscle spindles in the levator palpebrae superioris muscle of human foetuses.

Studies were performed on the levator palpebrae superioris muscle, isolated from foetuses aged 17 to 30 weeks. The number of muscle spindles varied from 2 in the 17th week to 7 in the 30th week. The length of the muscle spindles ranged from 20 to 500 microm, and the diameter varied from 10 to 70 microm. These observations show that the number of muscle spindles in levator palpebrae superioris muscles is significantly lesser than that in the extraocular muscles.

Afferent Pathways↗

Muscle spindle distribution in the levator veli palatini muscle in the rat.

We previously reported that the levator veli palatini muscle (LVP) in the rat is innervated by the glossopharyngeal nerve. The LVP positioned between the mouth and nasopharynx, has important roles in respiration, swallowing, and speech. Muscle spindles, structures scattered through skeletal muscles, appear to function like miniature strain gauges, sensing the degree of tension in the muscle. Muscle spindles were demonstrated in the rat's LVP in our neurophysiologic and histologic studies. We think the stretch of LVP modulates the rapid movements of the LVP by the proprioceptive component of the muscle spindle. Our results infer it is important to protect the innervation and the muscle spindles of the LVP from damage during any surgical dissection of the soft palate musculature.

Animals↗

Postnatal development of the annulospiral endings of Ia fibers in muscle spindles of the mouse temporal muscle.

The formation of the annulospiral endings of Ia fibers in muscle spindles was investigated in the temporal muscle of developing mice. From 15 to 20 d after birth, there were only a few spindles with "++" sensory endings. With growth of mice, the primary endings of the Ia fiber began to develop its spiral endings. And by the 30th postnatal day (that is, after weaning), almost all of the Ia fibers had completely coiled endings though the formation was still continuing in some spindles at this time. Formation of the annulospiral endings of the Ia fibers of muscle spindles in the temporal muscle was completed earlier than that of fibers in the spindles in the masseter muscle.

Animals↗

Critical period in muscle spindle regeneration in grafts of developing rat muscles.

Extensor digitorum longus (EDL) muscles from rats at various intervals after birth were grafted into EDL muscles of adult recipients. Three to twelve months after the operation, host muscles containing the grafts were removed and examined for the presence of muscle spindles in the graft. The aim of the study was to establish when muscle spindles become capable of regeneration during development. Regenerated muscles grafted during the first week after birth were virtually spindleless. Grafts of muscles transplanted 10 and 15 days postnatally contained only 5-8 muscle spindles on average. In contrast, the regenerated grafts originating from muscles of 24- and 28-day-old rats were spindle-rich as in mature muscle grafts; the number of spindles in the transplanted EDL muscles (25.0 +/- 2.3; mean +/- SE) attained values comparable to free standard autografts of these muscles in adult animals. Thus, the critical period after grafting, which also involves the loss of a vascular supply, is considerably longer than the critical period for muscle-spindle survival after nerve injury. Fifteen days after birth, when muscle spindles still survive denervation, only a few regenerated spindles were present in the individual muscle regenerates. We assume that the low resistance of immature spindle capsules to ischaemia accounts for their massive degeneration and abortive spindle regeneration in grafts from 10- to 15-day-old rats.

Adenosine Triphosphatases↗