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The responses of muscle spindles to small, slow movements in passive muscle and during fusimotor activity.

We have previously shown that movement detection thresholds at the human elbow joint were less than a degree of joint rotation in the passive limb but were higher if they were measured while subjects co-contracted elbow muscles [A.K. Wise, J.E. Gregory, U. Proske, J. Physiol., 508 (1998) 325-330]. Here we report observations on the responses of muscle spindles of the soleus muscle of the anaesthetised cat to determine their ability to signal small length changes in the passive muscle and during a contraction, under conditions resembling those of the human experiments. After appropriate conditioning of the muscle to control for history effects, primary endings of muscle spindles showed thresholds to ramp stretch at 20 micrometers s-1 of between less than 5 micrometers and 15 micrometers, which translates to 0.05 degrees -0.15 degrees of human elbow joint rotation. Thresholds were much higher following conditioning to introduce slack in the muscle. Since during a voluntary contraction there is likely to be alpha:gamma co-activation, responses of spindles were also recorded during slow stretches (100 micrometers at 20 micrometers s-1) during static fusimotor stimulation, dynamic fusimotor stimulation, combined fusimotor stimulation and fusimotor plus skeletomotor stimulation. Invariably, responses to passive stretch were larger than during motor stimulation. It is concluded that spindles are sensitive enough to signal fractions of a degree of elbow joint rotation and that the rise in threshold observed during a voluntary contraction may be accounted for by the actions of fusimotor and skeletomotor axons on spindle stretch responses.

Animals↗

Influence from stretch-sensitive receptors in the collateral ligaments of the knee joint on the gamma-muscle-spindle systems of flexor and extensor muscles.

In lightly alpha-chloralose anaesthetized cats, activity was recorded simultaneously from 2-4 primary muscle spindle afferents from triceps surae and/or posterior biceps and semitendinosus muscles. Increased tension in medial or lateral collateral ligament was found to evoke changes in the responses of the primary spindle afferents to sinusoidal stretching of their parent muscles. It is concluded that these changes were due to reflex actions of stretch/tension-sensitive receptors in the collateral ligaments on fusimotor neurones. Thus, receptor afferents from the collateral ligaments may contribute to the regulation of the activity in the gamma-motor-muscle-spindle system, and thereby also be of importance for the control of muscular stiffness and knee joint stability.

Animals↗

A quantitative study of muscle spindles and tendon organs in some intrinsic muscles of the hand in the bonnet monkey (Macaca radiata).

The number and density of muscle spindles and tendon organs have been determined in the following intrinsic muscles of the hand of bonnet monkeys: I lumbrical, II lumbrical, abductor digiti minimi, adductor pollicis, and I dorsal interosseous. All these muscles were found to be very rich in muscle spindles (17.6 to 42.31 per gram wet weight) but relatively poor in tendon organs (0.606 to 10.06 per gram wet weight). The lumbricals have very few tendon organs. The possible functional significance of these findings has been discussed.

Animals↗

[The absence of support loading on the rat m. soleus leads to an increase in the number of intrafusal muscle fibres in muscle spindles].

The ultrastructure of muscle spindles (incapsulated mechanoreceptors of stretch of extrafusal muscle fibres) of m. soleus in adult Wistar rats after repeated unloading of support on hind limbs with preservation of support loading on fore limbs has been studied by transmissing electron microscopy. It was shown that, along with muscle spindles with the ordinary number of intrafusal muscle fibres (four), m. soleus contains spindles with an increased number of intrafusal fibers (five to six). It was assumed that the increase in the number of intrafusal muscle fibers is due to the proliferation of their satellite cells.

Animals↗

Muscle spindles in the mylohyoid muscle of rats.

The mylohyoid muscle has several functions in relation to respiration, deglutition, and phonation, but these functions are not fully understood. The interaction of the mylohyoid nerve and muscle in 25 rats was studied by neurophysiologic and histologic methods. Stretching of the muscle elicited electrical responses from the branch of the mylohyoid nerve innervating the mylohyoid muscle, and stretch-sensitive receptors were demonstrated histologically in the mylohyoid muscle. This study indicates that the mylohyoid muscle plays an active role in functions such as swallowing, breathing, and phonation.

Animals↗

The muscle spindles in slow and twitch skeletal muscle of the lizard.

1. Responses from stretch receptors, identified as muscle spindles, were recorded in filaments of the nerve supplying a twitch muscle, semimembranosus, and a slow muscle, semitendinosus in the lizard Tiliqua.2. While recording afferent discharges in one filament of the motor nerve, several adjacent filaments were each in turn stimulated repetitively until one was encountered which on stimulation produced a powerful increase in spindle firing. Such an effect of the motor stimulus was interpreted as resulting from intrafusal contraction. Any interference with spindle firing patterns from extrafusal contraction produced by the motor stimulation was removed by differentially blocking the contraction with the drug curare.3. Discharge patterns of spindles in response to a slow stretch of the muscle were compared with the response to the same stretch, but during repetitive stimulation of the motor nerve filament which produced an intrafusal contraction.4. At the initial length, the firing rate of spindles in the twitch muscle was greatly increased by the motor tetanus. There was little further increase in the response during and following stretch of the muscle.5. While the spindles in the slow muscle were only moderately excited by the motor tetanus at the initial length of the muscle, a large increase was recorded during the dynamic component of the stretch. At the new length, the steady-state firing continued at a rate well above that for the initial length.6. The effect of the motor tetanus on the response to stretch of muscle spindles in the slow muscle could be mimicked by adding succinyl choline (5 mug/ml.) to the perfusion solution. Spindles in the twitch muscle did not show a sustained sensitivity to the drug.7. It is suggested that while the different effects of motor stimulation on the responses to stretch of spindles in slow and twitch muscle can be explained by propositions based on the sliding filament theory of contraction, the sustained elevation, at the new length, of firing frequencies of spindles in slow muscle might require an additional explanation.

Action Potentials↗

Idiopathic focal dystonia: a disorder of muscle spindle afferent processing?

As muscle spindles are involved in the sensation of position and movement of the body, we tested their involvement in the pathophysiology of idiopathic focal dystonia. Twenty patients with torticollis, nine with writer's cramp, two with blepharospasm and 16 healthy control subjects participated. In the first task, the blindfolded subject matched the position of the passively moved forearm with the opposite forearm when the elbow joint was slowly flexed. In a second matching task, passive movement was replaced by stimulation of one biceps tendon with a 50-Hz vibratory stimulus (a selective stimulus for muscle spindle Ia afferents). In normal individuals, this stimulus produces flexion of the vibrated arm around the elbow joint. Movement in both arms was recorded electronically. In experiments without vibratory stimuli, dystonic subjects showed normal movement of the tracking arm during attempts to match the position of the passively moved arm and no difference between the arms in the initial and final steady state positions. In experiments using vibratory stimuli, vibration of biceps tendons in normal subjects elicited flexion of the stimulated arm at the elbow and a matching movement of the opposite arm. In patients with dystonia, there was a similar flexion response to the vibratory stimulus in the stimulated arm but movement of the tracking arm was reduced. Taken together, these experiments suggest that there is abnormal perception of motion, but not position, in dystonic subjects. Dystonic subjects showed bilateral abnormalities of perception of the tonic vibration reflex which were remote from the clinically affected site. These findings are discussed in relationship to the role of muscle spindle Ia afferents in focal dystonia.

Adult↗

Is muscle spindle proprioceptive function spared in muscular dystrophies? A muscle tendon vibration study.

Muscular dystrophies (MDs) are characterized by the degeneration of skeletal muscle fibers. The aim of the present study was to determine whether the intrafusal fibers of muscle spindles are also affected in MD. The functional integrity of muscle spindles was tested by analyzing their involvement in the perception of body segment movements and in the control of posture. Twenty MD patients (4 with dystrophinopathy, 5 with myotonic dystrophies, 5 with fascioscapulohumeral MD, and 6 with limb-girdle dystrophies) and 10 healthy subjects participated in the study. The MD patients perceived passive movements and experienced illusory movements similar to those perceived by healthy subjects in terms of their direction and velocity. Vibratory stimulation applied to the neck and ankle muscle tendons induced postural responses in MD patients with spatial and temporal characteristics similar to those produced by healthy subjects. These results suggest that the proprioceptive function of muscle spindles is spared in muscular dystrophies.

Adult↗

Re-examination of the possible role of Golgi tendon organ and muscle spindle reflexes in proprioceptive neuromuscular facilitation muscle stretching.

Literature concerning the theoretical role of spinal reflex circuits and their sensorimotor signals in proprioceptive neuromuscular facilitation (PNF) muscle stretching techniques was examined. Reviewed data do not support the assertion commonly made in PNF literature that contraction of a stretched muscle prior to further stretch, or contraction of opposing muscles during muscle stretch, produces relaxation of the stretched muscle. Further, following contraction of a stretched muscle, inhibition of the stretch reflex response lasts only 1 s. Studies examined suggested that decreases in the response amplitude of the Hoffmann and muscle stretch reflexes following a contraction of a stretched muscle are not due to the activation of Golgi tendon organs, as commonly purported, but instead may be due to presynaptic inhibition of the muscle spindle sensory signal. The current view on the complex manner by which the spinal cord processes proprioceptive signals was discussed. The ability of acute PNF stretching procedures to often produce a joint range of motion greater than that observed with static stretching must be explained by mechanisms other than the spinal processing of proprioceptive information. Studies reviewed indicate that changes in the ability to tolerate stretch and/or the viscoelastic properties of the stretched muscle, induced by PNF procedures, are possible mechanisms.

Elasticity↗

A comparative study of muscle spindles in slow and fast neonatal muscles of normal and dystrophic mice.

Muscle spindles from the slow-twitch soleus and the fast-twitch extensor digitorum longus (EDL) muscles of genetically dystrophic mice of the dy2J/dy2J strain were compared with age-matched normal animals at neonatal ages of 1-3 weeks according to histochemical, quantitative, and ultrastructural parameters. Intrafusal fibers in both the soleus and EDL exhibited similar regional differences in myosin ATPase activity, and conformed to those noted previously in various adult species. In distal polar regions, all nuclear bag fibers resembled extrafusal fibers of the type 1 variety, whereas in capsular zones they could be divided into two subtypes. Nuclear chain fibers possessed a staining pattern similar to type 2 extrafusal fibers, and in contrast to the bag fibers they exhibited no regional variations. These features were consistently observed in both the normal and dystrophic muscles at all ages. Spindles varied only slightly in their number and distribution in the two types of muscle, and their location followed the neurovascular branching pattern in each. Irrespective of age or genotype, spindles in the soleus were more homogeneously dispersed, but those in the EDL were concentrated along the dorsal aspect of the muscle. No significant differences were noted in the total number of spindles between normal and dystrophic muscles. In addition, no dramatic differences were observed in the muscle spindle index for soleus and EDL. The first obvious disease-related changes were noted in extrafusal fibers of the soleus of 3-week-old mice, and spindles were often located close to these areas of fiber degeneration. Despite alterations in the surrounding tissue, however, spindles appeared morphologically unaltered in dystrophy. These observations indicate that intrafusal fibers of spindles in neonatal mice appear enzymatically and histologically unaffected in incipient stages of progressive muscular dystrophy.

Adenosine Triphosphatases↗

Physiological properties of muscle spindles in dorsal neck muscles of the cat.

1. Single-fiber recording was used to examine the properties of 107 spindle endings in cat biventer cervicis (BC) and complexus (CM) muscles. Responses of receptors were examined following muscle contraction and ramp and hold stretch. Twenty-two endings in splenius (SP) were also examined, but their responses could not be quantitated because the anatomy of SP prevented the application of appropriate stretches. 2. Conduction velocitites of spindle afferents ranged from 13 to 90 m/s. Endings with primary response patterns usually had faster conduction velocities than secondary endings, but there was overlap in the conduction velocity ranges of the two subgroups. 3. Most neck spindle afferents could be classified as either primary or secondary by a constellation of physiological criteria including dynamic response pattern, dynamic index, and variability of resting discharge frequency. However, 22 of 107 endings from BC and CM had responses with characteristics intermediate between primary and secondary responses. The possible sources of these characteristics are discussed. 4. Despite the similarity in properties between spindles of different neck muscles, the length sensitivities of CM spindles were high compared to those of BC spindles. CM spindles showed length-related modulation of firing frequency over a more restricted range of initial muscle lengths than did BC spindles. 5. Eight Golgi tendon organs (GTO) were identified by their characteristics responses. Conduction velocities obtained for five GTO afferent nerves ranged from 50 to 67 m/s. Recordings were also made from receptros in deep muscles surrounding the vertebrae. These receptors had properties characteristic of muscle spindles.

Adaptation, Physiological↗

Muscle spindles are concentrated in the superior vocalis subcompartment of the human thyroarytenoid muscle.

It is hypothesized that different parts of the thyroarytenoid muscle (TA) are functionally specialized. Specifically, the TA is divided into a lateral muscularis compartment and a medial vocalis compartment. This study examined the distribution of muscle spindles throughout the human TA as an indicator of these functional differences. Histological cross-sections from the anterior, middle, and posterior regions of five human membranous vocal folds were examined for the number and location of muscle spindles. There was an average of 6.1 muscle spindles in sections from each region with no significant variation between the different regions (p < .05). However, in sections from all three regions, the muscle spindles were always found to be concentrated in the superior medial quadrant of the TA (mean 85.9%, p < .01). The inferior medial, superior lateral, and inferior lateral quadrants of the TA contained 11.96%, 2.17%, and 0%, respectively, of the total muscle spindles. Within the superior medial quadrant, most of the muscle spindles were localized in the most superficial part of the muscle. The results of this study demonstrate that the majority of TA muscle spindles are concentrated in its superior medial quadrant, an area we have termed the superior vocalis subcompartment (SC). This finding suggests that the superior vocalis SC is functionally distinct from the remainder of the TA. It is hypothesized that tension in the superior vocalis SC can be controlled independently from the remainder of the TA, and this capability is used to effect the biomechanics of vocal fold vibration during phonation.

Humans↗

[The structure of muscle spindles in the intercostal muscles and rectus abdominis of the human fetus].

The structure and innervation of muscle fibres were studied in external intercostal muscles and the streight muscle of the abdomen in 24--26-week human fetuses. The diameter of most spindles was shown to be within the range of 50--70 mkm, while in the streight muscle of the abdomen it could reach 100 mkm. In the external intercostal muscles the spindles, as a rule, were longer (300-500 mkm) than in the streight muscle of the abdomen (100--300 mkm) which was likely due to special functioning of the spindles in rhythmically working muscles. According to sensory innervation all the spindles may be divided into 3 main types: simple, intermediate and complex. In the external intercostal muscles there occur 3 types of spindles, while in the streight muscle of the abdomen there are spindles of a complex type.

Humans↗

[Effects of three Chinese traditional drugs on afferent discharge from single muscle spindle in toads].

OBJECTIVE: To shed some light on treatment of muscle atrophy through testing the effects of several Chinese traditional drugs that can invigorate blood circulation on the afferent discharge of muscle spindle. METHOD: Single muscle spindle was isolated from sartorius muscle of toad. Afferent discharge from the muscle spindle was recorded using air-gap technique and effects of three Chinese traditional drugs on afferent activity of the muscle spindle were observed. RESULT: Ligustrazine could markedly tingle the spontaneous electrical activity of the muscle spindle, and it also showed a good dose-response relationship, while Salvia Miltiorrhiza and Safflower had no such excitative effect. CONCLUSION: It is known that ligustrazine can expand blood vessels and agitate microcirculation, and this effect on the muscle spindle makes it helpful for persons with muscle atrophy.

Animals↗