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Crossed and uncrossed central effects of muscle spindle afferents from the lateral pterygoid muscle of the guinea pig.

Physiological evidence is presented for the presence of stretch reflexes in the lateral pterygoid (Pt) muscle of the guinea pig. The central reflex effects of excitation of Pt stretch reflex afferents were also investigated. Passive lateral jaw displacement, which resulted in stretch of the Pt muscle on the side of jaw movement and stretch of the zygomatico-mandibularis (Z) muscle on the side contralateral to the movement, evoked increased EMG activity in these muscles. Stimulation of the trigeminal mesencephalic nucleus (mes V) evoked monosynaptic reflexes in both the Pt and Z nerves. Tonic stretch of the Pt muscle facilitated the monosynaptic reflex in the Pt nerve evoked by stimulation of mes V. Tonic vibration of the Pt muscle facilitated the mes V evoked monosynaptic reflex in the nerves to the ipsilateral Pt and contralateral Z muscles. Conversely, tonic vibration of the Z muscle facilitated the monosynaptic reflex evolved by mes V stimulation in the contralateral Pt and ipsilateral Z nerve. The results support the view that muscle spindles exist in the Pt and Z muscles and that there is a monosynaptic stretch reflex for both the Pt and Z muscles with cell bodies located in the mes V nucleus. It was also shown that the ipsilateral Pt muscle and the contralateral Z muscle act as synergists in the production of lateral jaw movements and that the organization of the stretch reflexes originating from the Pt and Z muscles support their synergistic action.

Animals↗

Muscle spindles in the human anterior digastric muscle.

The occurrence of muscle spindles in the anterior digastric muscle in man was investigated and the fibre calibre spectrum of the corresponding nerve was determined. After removal at autopsy from five individuals of both sexes (aged 23--73), the muscles were stained with Weigert's iron hematoxylin-van Gieson stain and the nerves according to the Alzheimer - Mann - Häggqvist method. Altogether 12 spindles were found in five out of ten muscles. Only muscles from one individual were devoid of spindles. This sparsity or absence was supported by analyses of fibre calibre spectra. The small number of spindles and the fact that they do not occur in all muscles or in all individuals, suggest that they are not an essential source of sensory information.

Adult↗

Superfluousness of motor innervation for the formation of muscle spindles in neonatal rats.

Muscle spindles form de novo in reinnervated muscles of neonatal rats treated with nerve growth factor. Whether the spindles can also form in muscle reinnervated only by afferents was investigated by removing the lumbosacral segment of the spinal cord immediately after crushing the nerve to the medial gastrocnemius muscle at birth, and administering nerve growth factor for 10 days afterwards. As predicted, the medial gastrocnemius muscles were reinnervated by afferents, but not efferents. No motor endplates were visible on any muscle fibers, and extrafusal fibers were atrophied. The reinnervated muscles contained spindle-like encapsulations of one to four fibers at 5, 7, 9 and 30 days after the nerve crush. The number of spindles as well as encapsulated fibers exceeded that of normal medial gastrocnemius muscles. The encapsulated fibers resembled typical intrafusal fibers. They had normal sensory-muscle contacts, but no motor endings. The fibers displayed equatorial clusters of myonuclei and expressed the spindle-specific slow-tonic myosin heavy chain isoform at postnatal day 30. Thus, efferents are not essential for the formation and differentiation of muscle spindles in reinnervated muscles of neonatal rats.

Animals↗

Responses of Ia afferent axons from muscle spindles lacking a bag1 intrafusal muscle fibre.

The Ia afferent axon innervating the muscle spindle is characterised by the dynamic nature of its response to stretch. Traditionally this response has been attributed to the properties of the bag1 intrafusal muscle fibre, a view supported recently by reports of stretch-activation of the bag1 fibre. Some spindle capsules lack a bag1 fibre and the Ia afferent axon terminates on the bag2 and chain fibres only. Such axons may be identified by their lack of an enhanced dynamic response to infusions of succinylcholine. In these experiments the responses to passive stretch of these axons have been investigated and found to be indistinguishable from those of the main Ia population. This implies that the bag1 fibre is not essential to the generation of the dynamic response of primary endings.

Afferent Pathways↗

Microvasculature of rabbit muscle spindles.

The microvasculature of the muscle spindle and its relationship to the microcirculation of teniussimus extrafusal muscle is described. Muscle spindles lie in close proximity and parallel to the central artery, vein, and nerve. The arterioles to spindle capillaries are third to fourth order branches of the central artery, whereas most arterioles to extrafusal capillaries are sixth to eighth order. Two or three capillaries enter each spindle. At least one entry consistently was encountered in the equatorial area near the sensory endings. Branches of intrafusal capillaries run longitudinally, anastomose with each other, and cradle the sensory zone in a longitudinal capillary loop. Capillaries in muscle spindles are larger than those in extrafusal muscle. These characteristic features are presumed to enhance the capability of these capillaries to provide sufficient circulation to the spindle, particularly to the region of the sensory endings.

Animals↗

Muscle spindle supply to the human jaw muscle.

Histological study of the human jaw muscle revealed that the temporal muscle displayed 342 muscle spindles, 208 in the horizontal and 134 in the vertical portion; the masseter muscle contained 114 spindles, 91 in the superficial and 23 in the profound portion; the medial pterygoid muscle had 59; and the lateral pterygoid muscle contained 6, four in the upper head and two in the lower head. These data suggest that the extensive mobility of the temporomandibular joint, and the maintenance of mandibular posture during mastication and speech are strongly influenced by proprioceptive mechanisms.

Fetus↗

Third-order reverse correlation analysis of muscle spindle primary afferent fiber responses to random muscle stretch.

The response of primary muscle spindle afferent fibers to muscle stretch is nonlinear. Now spindle responses (trains of action potentials) to band-limited Gaussian white noise length perturbations of the gastrocnemius muscles (input signal) are described in cats. The input noise upper cutoff frequency was clearly above the frequency range of physiological length changes in cat hindleg muscles. The input-output relation was analyzed by means of peri-spike averages (PSAs), which could be shown to correspond to the kernels of Wiener's white noise approach to systems identification. The present approach (the reverse correlation analysis) was applied up to the third order. An experiment consisted of two recordings: one (the source recording) to determine PSAs and the other (the test recording) to provide an input signal for predicting responses. The predictions of different orders were compared with the actual neuronal response (the observation) of the test recording. Four different approximation procedures were developed to adapt prediction and observation and to determine weighting factors for the predictions of different orders. The approximations also yielded the value of the power density P of the input noise signal: at a variety of stimulus parameters, P from approximations had the same magnitude as P determined directly from the input signal amplitude spectrum. The prediction of a sequence of action potentials improved the higher the order of components. 37 of 42 action potentials of a test recording (the observation) could be confidently predicted from PSAs or kernels. Compared with the size of the linear first-order prediction curve, the relative sizes of the second and third-order prediction curves were: 1.0:0.47:0.26.

Action Potentials↗

An exceptionally high density of muscle spindles in a slow-tonic pigeon muscle.

Histochemical and histological observations on the tiny wing muscle, M. coracotriceps, of the pigeon revealed a remarkably high density of muscle spindles (14,582 +/- 2,302/g of muscle)--approximately 15 times the highest densities hitherto reported for any muscle. Furthermore, all of the extrafusal fibers of this muscle were of the slow-tonic variety. This unique muscle probably functions as a mechanoreceptor extremely sensitive to changes in its own length.

Animals↗

Muscle spindles in nonhuman primate extrinsic auricular muscles.

Muscle spindles were sought in peri-auricular muscles of several primate species (rhesus monkey, woolly monkey, and baboon). Transverse sections cut at 10 mu and stained primarily by a silver impregnation technique were examined using light microscopy. Spindles were identified on the basis of standard criteria. Posterior and/or superior auricular muscles of each species were found to contain spindles. At least some muscles innervated by facial nerve have classical spindles as component structures.

Animals↗

Coupling between single muscle spindle afferent and EMG in human wrist extensor muscles: physiological evidence of skeletofusimotor (beta) innervation.

OBJECTIVES: This study was conducted to find physiological evidence of skeletofusimotor innervation in man. METHODS: Discharges of 38 single muscle spindle afferents from m. extensor carpi radialis were recorded from 9 subjects during steady isometric contractions of wrist extension. Correlation between these afferents and rectified surface EMG was investigated by estimating cumulant density function. RESULTS: In the cumulant density estimate between spindle afferent and EMG, a positive EMG peak was obtained prior to afferent firing between -30 and -10 ms in 15 afferents (39%). CONCLUSIONS: The present finding indicates coupling between spindle afferents and extrafusal activity and suggests the widespread skeletofusimotor innervation in human muscle spindle.

Adult↗

Reflex influences on muscle spindle activity in relaxed human leg muscles.

The study was designed to determine whether low-threshold cutaneous and muscle afferents from the foot reflexly activate gamma-motoneurons innervating relaxed muscles of the leg. In 15 experiments multiunit recordings were made from 21 nerve fascicles innervating triceps surae or tibialis anterior. In a further nine experiments the activity of 19 identified single muscle spindle afferents was recorded, 13 from triceps surae, 5 from tibialis anterior, and 1 from extensor digitorum longus. Trains of electrical stimuli (5 stimuli, 300 Hz) were delivered to the sural nerve at the ankle (intensity, twice sensory threshold) and the posterior tibial nerve at the ankle (intensity, 1.1 times motor threshold for the small muscles of the foot). In addition, a tap on the appropriate tendon at varying times after the stimuli was used to assess the dynamic responsiveness of the afferents under study. The conditioning electrical stimuli did not change the discharge of single spindle afferents. Recordings of rectified and averaged multiunit activity also revealed no change in the overall level of background neural activity following the electrical stimuli. The afferent responses to tendon taps did not differ significantly whether or not they were preceded by stimulation of the sural or posterior tibial nerves. These results suggest that low-threshold afferents from the foot do not produce significant activation of fusimotor neurons in relaxed leg muscles, at least as judged by their ability to alter the discharge of muscle spindle afferents. As there may be no effective background activity in fusimotor neurons innervating relaxed human muscles, it is possible that these inputs from the foot could influence the fusimotor system during voluntary contractions when the fusimotor neurons have been brought to firing threshold. In one subject trains of stimuli were delivered to the posterior tibial nerve at painful levels (30 times motor threshold). They produced an acceleration of the discharge of a spindle in soleus at a latency of approximately 125 ms, in advance of detectable activity in skeletomotor neurons and before an increase in muscle length was noted. It presumably resulted from activation of gamma-motoneurons innervating soleus by small myelinated afferents (A-delta range).

Electric Stimulation↗

High incidence of multiple-bag fiber muscle spindles in the articularis humeri muscle of the horse.

The articularis humeri (AH) muscle of the horse is a small muscle composed of histochemically identified type I and IIA extrafusal fibers and a large number of muscle spindles. A total of 150 complete spindles with both spindle poles available were examined in serial transverse sections. On the basis of myosin ATPase-staining reactions after alkaline and acid preincubations, four types of intrafusal fibers, namely, bag1, bag2, "mixed" bag, and chain fibers, were identified. A high proportion of the spindle population (62.6%) consisted of multiple-bag spindles containing three or more (up to six) bag fibers. Also one-bag-fiber spindles were observed. The one-bag-fiber spindles containing a bag2 fiber could be traced into tandem linkages. "Mixed" bag intrafusal fibers, differing in their ATPase staining profile at the two poles, were found in spindles containing also at least one bag1 and one bag2 fiber. An unusually long extracapsular tract (up to 5,500 microns) of the bag intrafusal fibers was observed.

Animals↗

An investigation into the site of termination of static gamma fibres within muscle spindles of the cat peroneus longus muscle.

1. The distribution of static fusimotor fibres to intrafusal muscle fibres of cat peroneus longus muscle spindles was investigated using the glycogen-depletion technique of Edström & Kugelberg (1968). Single static gamma fibres were stimulation intermittently at high rates for 3 hr with the blood supply occluded for some of this time. Subsequently the portion of muscle containing the activated spindles was fixed, sectioned and stained for glycogen with the periodic acid-Schiff (PAS) method. 2. Ten static axons caused depletion in eleven spindles. In five of these the only glycogen-depleted fibres were nuclear chain fibres. In the other six spindles one nuclear bag fibre was depleted in addition to chain fibres and this was always the larger of the two within the spindle. 3. These results on a medium-sized hind limb muscle are compared with findings concerning the distribution of static gamma fibre axons previously investigated only in very small muscles. The results agree in showing that nearly all static gamma fibres innervate nuclear chain fibres but that in 50-75% of the times in which static gamma fibres innervate spindles the distribution is to bag fibres as well as to chain fibres. The interpretation to be put upon this is uncertain. One possibility with which the results from peroneus longus are consistent is that the bag fibres which are usually innervated by static axons are the 'intermediate' bag fibres whose ultrastructure has recently been shown to resemble that of chain fibres.

Animals↗

The fine structure of muscle spindles in the lumbrical muscles of the rat.

Lumbrical muscles of young rats were fixed with OsO(4) and embedded in methacrylate for electron microscopy. The spindle capsule was found to be continuous with and similar in structure to the sheath of Henle surrounding the nerves supplying the spindle. The capsule consists of several closely applied concentric cytoplasmic sheets. Each sheet is about 1,000 A thick and has no fenestrations. Many caveolae and vesicles in the cytoplasm suggest active transport through the sheets. The periaxial space fluid contains much solid material. It is suggested that the capsule and periaxial space regulate internal chemical environment. The interfibrillar structures are less evident in the polar regions of intrafusal fibres than in extrafusal fibres. Simple motor end-plates occur on the polar regions of intrafusal fibres. In the myotube region of the intrafusal fibre a peripheral zone of myofibrils surrounds a cytoplasmic core containing nuclei, mitochondria, Golgi bodies, reticulum, and a few lipid-like granules. Naked sensory endings lie on the myotube "in parallel" with the underlying myofilaments. Naked processes of the primary sensory ending deeply indent the muscle plasma membrane and the underlying wisps of myofilament in the nuclear bag region. The plasma membranes of sensory nerve ending and intrafusal muscle fibre are about 200 A apart.

Animals↗

Studies on the site of termination of static and dynamic fusimotor fibres within muscle spindles of the tenuissimus muscle of the cat.

1. The site of termination of static and dynamic fusimotor fibres has been mapped by finding which intrafusal muscle fibres have been depleted of glycogen as a result of tetanic stimulation of single gamma fibres. Long periods of stimulation coupled with occlusion of the blood supply were necessary to cause glycogen depletion.2. In cat tenuissimus muscle, dynamic gamma motor fibres always activated bag intrafusal muscle fibres, and occasionally chain fibres. Static gamma fibres always activated chain fibres and frequently activated bag fibres as well.3. It is argued that these results can be fitted into the hypothesis of the mechanism of internal functioning of the spindle originally proposed by Jansen & Matthews (1962). It is also pointed out that the results raise problems concerning the mechanism of development of the spindle motor innervation.

Action Potentials↗