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Muscle spindle is the key proprioceptor in skeletal muscles and plays important roles in many physiological activities, such as maintaining posture, regulating movement and controlling speed variation. It has significant clinical relevance and is emerging as a promising therapeutic target for the treatment of motor functional impairment and metabolic diseases. In this review, we summarized muscle spindle distribution and the mechanism of mechanical signal transmission, and reviewed the research progress on morphological and structural characteristics of muscle spindles.

The palmaris brevis muscle contains numerous muscle spindles to control changes of the muscle length but is devoid of tendon-associated neuronal elements (e.g. Golgi tendon organs or Ruffini-like corpuscles) controlling changes in muscle strength. Pacinian bodies, frequently seen in the palm of the hand, show no direct association to the muscle bundles. The observed innervation pattern of the palmaris brevis muscle points to a specific type of neuronal regulation, present in skeletal muscles with no skeletal connection., (© 2019 The Authors. Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society.)

We examined the six small lateral rotators of the hip joint, which is one of the most flexible joints and allows kinematically complex motions of the hindlimb, to elucidate the functional differentiation among these muscles and to test the hypothesis that species-specific characteristics in hindlimb use during locomotion are reflected in the muscle spindle density and in other parameters of the deep small hip joint rotators. For these purposes, we estimated the number of muscle spindles of the superior gemellus muscle (SG), inferior gemellus muscle, quadratus femoris muscle, obturator internus muscle (OI), obturator externus muscle, and piriformis muscle in three Japanese macaques and a gibbon, using 30-µm-thick serial sections throughout each muscle length after azan staining. The numbers of muscle spindles per 10,000 muscle fibers were determined to compare inter-muscle variation. The spindle density was highest in the SG and lowest in the OI in the Japanese macaques, suggesting that the SG, which is attached to the tendon of the OI, functions as a kinesiological monitor of the OI. On the other hand, SG the was missing in the gibbon, and the OI in the gibbon contained more spindles than that in the Japanese macaques. This suggests that the SG and the OI fused into one muscle in the gibbon. We postulate that the relative importance of the deep small hip rotator muscles differs between the Japanese macaques and gibbon and that the gibbon's muscles are less differentiated in terms of the spindle density, probably because this brachiating species uses its hindlimbs less frequently., (© 2020 S. Karger AG, Basel.)

Muscle spindles are ubiquitous encapsulated mechanoreceptors found in most mammalian muscles. There are two types of endings, primary and secondary, and both are sensitive to changes in muscle length and velocity, with the primary endings having a greater dynamic sensitivity. Unlike other mechanoreceptors in the somatosensory system, muscle spindles are unique in possessing motor innervation, via γ-motoneurons (fusimotor neurons), that control their sensitivity to stretch. Much of what we know about human muscles spindles comes from studying the behavior of their afferents via intraneural microelectrodes (microneurography) inserted into accessible peripheral nerves. We review the functional properties of human muscle spindles, comparing and contrasting with what we know about the functions of muscle spindles studied in experimental animals. As in the cat, many human muscle spindles possess a background discharge that is related to the degree of muscle stretch, but mean firing rates are much lower (~10 Hz). They can faithfully encode changes in muscle fascicle length in passive conditions, but higher level extraction of information is required by the central nervous system to measure changes in muscle length during muscle contraction. Moreover, although there is some evidence supporting independent control of human muscle spindles via fusimotor neurons, any effects are modest compared with the clearly independent control of fusimotor neurons observed in the cat.

The mimic muscles are usually described as containing no muscle spindles. In the present publication the human platysma was reinvestigated concerning its content of corpuscular sensors. Serial sections through the platysma of seven donors revealed numerous muscle spindles but no Pacini corpuscules. The muscle spindles were located in the cranial two-thirds of the platysma, and were evenly distributed with a tendency to have more spindles in the lateral part of the muscle. Immunohistochemical staining with S46 antibodies revealed a predominance of nuclear bag fibers. The results point to an extended function of the platysma as an afferent center of the lower face mimic muscles., (© 2017 Anatomical Society.)

Published descriptions about the sites of origin of the human medial pterygoid muscle vary and there are few reports on the distribution and density of muscle spindles in this muscle. We aimed to: (1) determine the extent of anatomical variability in the origins and insertions of the superficial and deep heads of the human medial pterygoid muscle and (2) determine the extent of variation in the distribution of spindles in the two heads of the muscle. Thirty-nine human cadaver hemi-heads were dissected and the attachments of the medial pterygoid muscle examined. The whole muscle was removed, weighed, cut into segments and embedded in wax for light microscopy. Sections were stained with Weigert-Van Gieson stain and scanned into digital images. Spindles were manually counted. In seven specimens, the deep head of the muscle arose from the medial surface of the lateral pterygoid plate and the pterygoid fossa. In 28 specimens, the origin extended onto the lateral surface of the medial pterygoid plate. There were abundant muscle spindles in the middle of the muscle, slightly fewer 1 cm toward the insertion, significantly fewer 1 cm toward the origin, and few or no spindles near the origins of the superficial and deep heads or near their insertion. In conclusion, firstly, this study shows that in 72% of the specimens examined, the origin of the medial pterygoid is wider than conventionally described in anatomical textbooks. Secondly, the segmental distribution of muscle spindles is described for the first time. Clin. Anat. 30:1064-1071, 2017. © 2017 Wiley Periodicals, Inc., (© 2017 Wiley Periodicals, Inc.)

The first descriptions of muscle spindles with intrafusal fibres containing striated myofibrils and nervous elements were given approximately 150 years ago. It took, however, another 100 years to establish the presence of two types of intrafusal muscle fibres: nuclear bag and nuclear chain fibres. The present paper highlights primarily the contribution of Robert Banks in fibre typing of intrafusal fibres: the confirmation of the principle of two types of nuclear bag fibres in mammalian spindles and the variation in occurrence of a dense M-band along the fibres. Furthermore, this paper summarizes how studies from the Umeå University group (Laboratory of Muscle Biology in the Department of Integrative Medical Biology) on fibre typing and the structure and composition of M-bands have contributed to the current understanding of muscle spindle complexity in adult humans as well as to muscle spindle development and effects of ageing. The variable molecular composition of the intrafusal sarcomeres with respect to myosin heavy chains and M-band proteins gives new perspectives on the role of the intrafusal myofibrils as stretch-activated sensors influencing tension/stiffness and signalling to nuclei., (© 2015 The Authors Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society.)

The muscle spindle has been well studied in mammals, birds, and some reptiles, but little is known about its structure and function in amphibians. Several physiological and behavioral studies in frogs and toads suggest that the submentalis muscle has an important role in jaw-tongue coordination during feeding through sensory feedback mechanisms. Evidence suggests this mechanism is generated through muscle spindles, or spindle-like structures. However, the presence of these has not been investigated. Using charged fluorescent molecules that are taken up by endocytosis at activated synapses, numerous muscle spindles were identified in the submentalis muscle of the marine toad, Bufo marinus. These were further verified with brightfield and differential interference contrast microscopy. The results provide anatomical evidence for muscle spindles in the submentalis muscle and suggest a mechanism for the submentalis muscle's role in jaw-tongue coordination in B. marinus., (2010. © 2010 Wiley-Liss, Inc.)

Objective: To investigate the characteristics in the topographic distribution of the muscle spindles in human masticatory muscles., Methods: The muscle spindles of the masticatory muscles on both sides of 3 human corpses fixed in formalin for more than 1 year were observed histologically and analyzed quantitatively using an image analysis system., Results: The mean muscle spindle density was 32.11/g in the masseter, significantly higher in the deep muscular fibers (70.76/g) with high proportion of spindle complexes than in the superficial layers of the muscle belly (P<0.05). The mean spindle density in the temporalis was 5.44/g, higher in the anterior portion (7.44/g) than in the posterior portion (3.78/g, P<0.01). The medial pterygoid had a mean spindle density of 4.43/g., Conclusion: These results indicate a heterogeneous distribution of muscle spindles in the masticatory muscles with different morphologies or in different portions of the same muscle, depending on muscle fiber type and muscular architecture.

A frequency domain approach and a time domain approach have been combined in an investigation of the behaviour of the primary and secondary endings of an isolated muscle spindle in response to the activity of two static fusimotor axons when the parent muscle is held at a fixed length and when it is subjected to random length changes. The frequency domain analysis has an associated error process which provides a measure of how well the input processes can be used to predict the output processes and is also used to specify how the interactions between the recorded processes contribute to this error. Without assuming stationarity of the input, the time domain approach uses a sequence of probability models of increasing complexity in which the number of input processes to the model is progressively increased. This feature of the time domain approach was used to identify a preferred direction of interaction between the processes underlying the generation of the activity of the primary and secondary endings. In the presence of fusimotor activity and dynamic length changes imposed on the muscle, it was shown that the activity of the primary and secondary endings carried different information about the effects of the inputs imposed on the muscle spindle. The results presented in this work emphasise that the analysis of the behaviour of complex systems benefits from a combination of frequency and time domain methods.

The encapsulated sensory endings of mammalian skeletal muscles are all mechanoreceptors. At the most basic functional level they serve as length sensors (muscle spindle primary and secondary endings), tension sensors (tendon organs), and pressure or vibration sensors (lamellated corpuscles). At a higher functional level, the differing roles of individual muscles in, for example, postural adjustment and locomotion might be expected to be reflected in characteristic complements of the various end-organs, their sensory endings and afferent nerve fibres. This has previously been demonstrated with regard to the number of muscle-spindle capsules; however, information on the other types of end-organ, as well as the complements of primary and secondary endings of the spindles themselves, is sporadic and inconclusive regarding their comparative provision in different muscles. Our general conclusion that muscle-specific variability in the provision of encapsulated sensory endings does exist demonstrates the necessity for the acquisition of more data of this type if we are to understand the underlying adaptive relationships between motor control and the structure and function of skeletal muscle. The present quantitative and comparative analysis of encapsulated muscle afferents is based on teased, silver-impregnated preparations. We begin with a statistical analysis of the number and distribution of muscle-spindle afferents in hind-limb muscles of the cat, particularly tenuissimus. We show that: (i) taking account of the necessity for at least one primary ending to be present, muscles differ significantly in the mean number of additional afferents per spindle capsule; (ii) the frequency of occurrence of spindles with different sensory complements is consistent with a stochastic, rather than deterministic, developmental process; and (iii) notwithstanding the previous finding, there is a differential distribution of spindles intramuscularly such that the more complex ones tend to be located closer to the main divisions of the nerve. Next, based on a sample of tendon organs from several hind-foot muscles of the cat, we demonstrate the existence in at least a large proportion of tendon organs of a structural substrate to account for multiple spike-initiation sites and pacemaker switching, namely the distribution of sensory terminals supplied by the different first-order branches of the Ib afferent to separate, parallel, tendinous compartments of individual tendon organs. We then show that the numbers of spindles, tendon organs and paciniform corpuscles vary independently in a sample of (mainly) hind-foot muscles of the cat. Grouping muscles by anatomical region in the cat indicated the existence of a gradual proximo-distal decline in the overall average size of the afferent complement of muscle spindles from axial through hind limb to intrinsic foot muscles, but with considerable muscle-specific variability. Finally, we present some comparative data on muscle-spindle afferent complements of rat, rabbit and guinea pig, one particularly notable feature being the high incidence of multiple primary endings in the rat.

Age-related physiological and morphological changes of muscle spindles were examined in rats (male Fischer 344/DuCrj: young, 4-13 months; middle-aged, 20-22 months; old, 28-31 months). Single afferent discharges of the muscle spindles in gastrocnemius muscles were recorded from a finely split dorsal root during ramp-and-hold (amplitude, 2.0 mm; velocity, 2-20 mm s(-1)) or sinusoidal stretch (amplitude, 0.05-1.0 mm; frequency, 0.5-2 Hz). Respective conduction velocities (CVs) were then measured. After electrophysiological experimentation, the muscles were dissected. The silver-impregnated muscle spindles were teased and then analysed using a light microscope. The CV and dynamic response to ramp-and-hold stretch of many endings were widely overlapped in old rats because of the decreased CV and dynamic response of primary endings. Many units in old rats showed slowing of discharge during the release phase under ramp-and-hold stretch and continuous discharge under sinusoidal stretch, similarly to secondary endings in young and middle-aged rats. Morphological studies revealed that primary endings of aged rat muscle spindles were less spiral or non-spiral in appearance, but secondary endings appeared unchanged. These results suggest first that primary muscle spindles in old rats are indistinguishable from secondary endings when determined solely by previously used physiological criteria. Secondly, these physiological results reflect drastic age-related morphological changes in spindle primary endings.

We have tested our hypothesis suggesting (i) that for the reliable determination and counting of muscle spindles (Msp) at the light microscopy level in extraocular muscles (EOM), analysis of the spindle specific myosin heavy chain (MyHC) immunoreactivity of intrafusal fibers, especially after staining with anti-slow-tonic MyHC antibodies, is the most convenient tool, (ii) that the number of Msp determined by the slow-tonic MyHC immunoreactivity of intrafusal fibers in EOM is much lower than that based on histological examination and (iii) that the previously reported numbers of Msp based on histological examination of EOM could be overestimated. In order to determine the number and distribution of Msp and to analyze the MyHC isoform immunoreactivity of intrafusal fibers in porcine EOM, paraffin sections of three 9-month-old pig medial (MR) and lateral rectus (LR), levator palpebrae (LP) and retractor bulbi (RB) muscles were stained histologically or using specific monoclonal antibodies (mAbs) against MyHC isoforms. Msp in recti and LP muscles studied by immunocytochemistry contained nuclear bag (NB) fiber(s) reacting with mAbs against slow-tonic, slow-twitch, alpha-cardiac and neonatal MyHCs, but not with the mAb against fast-twitch MyHC, which, on the contrary, stained nuclear chain (NC) fibers. Based on determination of spindle specific slow-tonic MyHC isoform immunoreactivity we have found 72 Msp in the MR and 68 Msp in the LR and 12 Msp in LP muscles, which was only 62, 55 and 32% of the Msp total counts according to histological examination, respectively. In the RB muscle, we have even found only 15 spindle-like-structures composed of encapsulated thin muscle fibers, which possessed only a reaction with anti-fast-twitch MyHC mAb, but lacked slow-tonic, slow-twitch or alpha-cardiac MyHCs immunoreactivity. Our analysis of porcine EOM confirmed the above suggestions, demonstrating, for the first time in the pig, the presence of "false Msp" mimicking encapsulated muscle fibers on histological sections that lack spindle specific MyHC immunoreactivity. In analogy with other muscles we suggest that "false Msp" are not innervated by sensory axons and therefore do not contribute to the physiological sensation of the muscle length changes. Our results thus show that the reliable identification of functionally effective Msp in EOM must involve immunohistochemical analysis of spindle specific MyHC isoforms of intrafusal fibers, as "false" spindles appearing on histologically stained sections as encapsulated muscle fibers could be regarded as "true" Msp and thus increase the spindle number counts in earlier studies.

Objective: To determine the morphologic and the morphometrical features of spindles in biopsies of patients with different types of muscular dystrophy and investigate the possible involvement of the spindle in the pathologic process of these diseases., Study Design: The following variables were studied in biopsy specimens from 10 patients with Duchenne or Becker dystrophy, 9 with limb-girdle dystrophy, 3 with congenital dystrophy and 3 with facioscapulohumeral dystrophy: diameter and area of spindle; thickness of the capsule; number, diameter and area of intrafusal fibers; and number and area of nuclei., Results: The statistical evaluation of the data showed significant differences regarding the thickness of the capsule, which was greater in patients than in controls, while the diameter and the area of the fibers were all smaller in patients than in controls. The area of nuclei of fibers was increased; this was a common feature for all types of muscular dystrophy., Conclusion: These findings indicate that the spindle possibly participates in the pathologic process of different types of muscular dystrophies.

An allometric analysis of the number of muscle spindles in relation to muscle mass in mammalian (mouse, rat, guinea-pig, cat, human) skeletal muscles is presented. It is shown that the trend to increasing number as muscle mass increases follows an isometric (length) relationship between species, whereas within a species, at least for the only essentially complete sample (human), the number of spindles scales, on average, with the square root rather than the cube root of muscle mass. An attempt is made to reconcile these apparently discrepant relationships. Use of the widely accepted spindle density (number of spindles g(-1) of muscle) as a measure of relative abundance of spindles in different muscles is shown to be grossly misleading. It is replaced with the residuals of the linear regression of ln spindle number against ln muscle mass. Significant differences in relative spindle abundance as measured by residuals were found between regional groups of muscles: the greatest abundance is in axial muscles, including those concerned with head position, whereas the least is in muscles of the shoulder girdle. No differences were found between large and small muscles operating in parallel, or between antigravity and non-antigravity muscles. For proximal vs. distal muscles, spindles were significantly less abundant in the hand than the arm, but there was no difference between the foot and the leg.

Objective: To study the morphometric characteristics of the human muscle spindle in normal muscle and to investigate the influence of aging., Study Design: The following variables were studied in 72 spindles: area and diameter of the spindle; thickness of the capsule; number, area and diameter of fibers; and number and area of nuclei., Results: In deltoid and extensor digitorum brevis muscles, a reduction in the diameter of the spindle as a function of age was found, while no statistically significant change in the variables as a function of age was observed in the quadriceps femoris and biceps muscles. In the deltoid, a reduction in the number of fibers and an increase in their diameter were also observed., Conclusion: These findings could prove useful in the study of the spindle in relation to disease.

The distribution pattern of muscle spindles in the skeletal musculature has been reviewed in a large number of muscles (using the literature data especially from cat and man), and the relation of spindle content to muscle mass was quantitatively examined in 36 cat and 140 human muscles. In both species, the number of spindles increases with increasing muscle mass in a power law fashion of the form y=bx+a, whereby y denotes the logarithm of spindle content within a muscle, and x is the logarithm of muscle mass. For the cat, slope b and intercept a were estimated as 0.39 and 1.53, and for man as 0.48 and 1.33, respectively. The results show that the spindle content of a muscle may be related to its mass, confirming a similar analysis made previously by Banks and Stacey (Mechano receptors, Plenum Press, New York, 1988, pp. 263-269) in a different data set. With regard to the histological profile of muscle fibers, (as it is already well documented by many groups) muscle spindles tend to be located in deeper muscle regions where oxidative fibers predominate, and are far scarcer in superficial and flat muscle regions where glycolytic fibers predominate. These discrete muscle regions differ also in the properties of the vessel tree supplying them, for which the term oxidative and glycolytic "angiotype" has been used. The results from these three aspects of analysis (relation to muscle mass, relation to muscle regions with high oxidative index and relation to muscle regions with dense vascular supply) were combined with histological findings showing that spindles may be in systematic anatomical contact to intramuscular vessels. Based on these data a hypothesis is proposed according to which, both the number and intramuscular placement of muscle spindles are related to the oxidative angiotype supplying the muscle territories rich in oxidative fibers. The hypothesis is discussed.

This study identified the anatomical and close functional relationship between the transverse lingual and superior pharyngeal constrictor muscle. Two en bloc samples (including the tongue and mid-pharyngeal wall) and four whole tongues were obtained from adult human cadavers. We found that fibers of the superior pharyngeal constrictor muscle connected with fibers of the transverse lingual muscle, forming a ring of muscle at the base of the tongue. The average diameters of the transverse muscle fibers increased in size gradually as they approached the base of the tongue. Distribution of the muscle spindles in the transverse lingual muscle and the genioglossus muscle also increased as they reached posteriorly near the base of the tongue. These findings suggest that a ring of muscle formed by the postero-inferior portion of the transverse lingual muscle and the superior pharyngeal constrictor may be largely responsible for the retrusive movement of the tongue and the constrictive movement of the pharyngeal cavity as an antagonist of the genioglossus muscle., (Copyright 2004 Wiley-Liss, Inc.)

Intrinsic laryngeal muscle investigations, especially those of the interarytenoid (IA) muscle, have been primarily teleologically based. We determined IA muscle anatomy and histochemical and immunohistochemical classification of extrafusal and intrafusal (muscle spindle) fibers in 5 patients. Extrafusal fibers were oxidative type I and glycolytic types IIA and IIX. Intrafusal fibers of muscle spindles were identified by the presence of tonic and neonatal myosin. The results demonstrate that the IA muscle has a phenotype similar to that of limb skeletal muscle. Myosin coexpression, the absence of intrafusal fibers, and fiber type grouping were unusual features found previously in the thyroarytenoid and posterior cricoarytenoid muscles, but they were not present in the IA muscle. These findings lead to the conclusion that the IA muscle has functional significance beyond its assumed importance in maintaining vocal fold position during phonation. The presence of spindles demonstrates differences in motor control as compared to the thyroarytenoid and posterior cricoarytenoid muscles. Further, extrafusal fiber characteristics implicate IA muscle involvement in muscle tension dysphonia and adductor spasmodic dysphonia. Given the unique physiologic characteristics of the human IA muscle, further research into the role of the IA muscle in voice disorders is warranted.

In the present study muscle spindles (MSps) and Golgi tendon organs (GTOs) in bovine extraocular muscles (EOMs) were analyzed in detail. The innervation pattern of these proprioceptors was investigated with transmission electron microscope and confocal laser scanning microscope after double-fluorescent labeling. Three-dimensional (3D) reconstructions were performed of GTOs. Muscle spindles. MSps are numerous, each containing two nuclear bag fibers and up to eight nuclear chain fibers. In the equatorial region and paraequatorial region thin axons enwrapping the intrafusal muscle fibers form numerous nerve contacts on the muscle fiber surface. Double staining of such nerve terminals with synaptophysin and alpha-bungarotoxin and their fine structural features confirm their sensory nature. In the encapsulated part of the polar region neuromuscular contacts have structural features of motor nerve terminals and stain positively with alpha-bungarotoxin. Golgi tendon organs. GTOs are numerous in bovine EOMs. Each GTO contains collagen bundles but frequently also intracapsular muscle fibers. Intracapsular muscle fibers either terminate inside the GTO in collagen bundles or pass through the proprioceptor. GTOs are richly supplied with sensory nerve terminals which intermingle with the collagen bundles. Nerve terminals on intracapsular muscle fibers exhibit fine structural characteristics of motor nerve terminals and are alpha-bungarotoxin positive. The 3D images of GTOs show the detailed spatial arrangement of the GTO tissue components. These new insights in the complex and specific morphology of MSps and GTOs in bovine EOMs indicate that we deal with highly developed proprioceptors. These are supposed to provide important information for EOM innervation.

Most studies concerning the structure and function of muscle spindles have utilized the hind limbs of experimental animals. However, little is known about muscle spindles of the back muscles. The purpose of this study was to investigate the sensory innervation of muscle spindles of the paravertebral muscle in the rat. The subjects were 10 normal male rats. The longissimus muscles were isolated and frozen in cooled isopentane (-160 degrees C), and serial transverse sections were made with a cryostat. Histochemical preparations were then made using nicotinamide adenine dinucleotide tetrazolium reductase (NADH-TR) stain and modified Gomori-trichrome stain. The muscle spindles in each segment were identified microscopically by observing the equatorial and polar regions. NADH-TR staining was employed to distinguish nuclear bag1, nuclear bag2, and nuclear chain intrafusal muscle fibers. A total of 20 spindle poles were surveyed. The mean polar length of intrafusal fibers as well as that of each region (A, B, and C) were measured. NADH-TR staining also demonstrated the terminal sites of sensory fibers along intrafusal fibers. All spindle poles surveyed were innervated by secondary sensory fibers in addition to primary sensory fibers. Eight spindle poles were intermediate type muscle spindles that were innervated by one primary sensory fiber and one secondary sensory fiber. Twelve spindle poles were complex type muscle spindles that were innervated by one primary sensory fiber and multiple secondary sensory fibers. The mean length of the A region was 223.1+/-37.9 microm (n=8) for intermediate type spindles and 493.8+/-157.0 microm (n=12) for complex type spindles. The length of the A region was significantly longer in the complex type spindles than in the intermediate type spindles (p<0.001). The results suggest that the innervations of secondary sensory fibers were well developed in the longissimus muscle spindles in the rat. The morphological features of muscle spindles of the longissimus muscle may represent the structural basis for qualitatively different afferent discharges that relate to the characteristic types of locomotion served by paravertebral muscles.

Objective: To study changes in structure and innervation of soleus muscle spindle in rats induced by simulated weightlessness., Method: Weightlessness was simulated by tail suspension in female rats. The technique of Fawosky's Staining was used to detect the morphology of nerve endings of soleus muscle spindle in 7 d, 14 d, 21 d tail-suspended and control rats., Result: The intrafusal muscle fibers were regular and nerve endings were intact and clear in the soleus spindles of control rats. No obvious change was observed in muscle spindles of 7 d tail-suspended rats. Intrafusal muscle fibers became irregular, with rough and vague nerve endings in 14 d tail-suspended rats. The distinct retrograde change of muscle spindles and nerve endings were observed in 21 d tail-suspended rats. Degeneration and rupture were found in nerve endings, which made it unclear and pale in staining., Conclusion: Simulated weightlessness could induce time-related changes in structure and innervation of soleus muscle spindle in rats.

Study Design: Tissue blocks comprising muscle and bone from C5 to C7 segments were harvested at autopsy from 16 individuals ranging in age from 4 to 77 years. The prevertebral longus colli and postvertebral multifidus muscle pairs from one side in each individual were randomly selected for this study of muscle spindles., Objectives: To determine muscle spindle distribution, morphology, and density for the longus colli and multifidus in caudal segments of the human cervical spine, and to assess whether changes are evident from infancy to old age., Summary of Background Data: Age-related changes to the osteoligamentous framework of the cervical spine have been well documented. Postural modification accompanies these structural alterations, but there have been limited attempts to document whether muscle sustains a comparable level of morphologic alteration. Previous studies have examined muscle spindles in the neck muscles of various animal models and in a variety of isolated human muscles. However, most of these studies incurred bias through sampling and methodologic assumptions., Methods: The longus colli and multifidus were resected between C5 and C7, and between left and right pairs selected randomly for spindle analysis. These vertebral segments were selected deliberately because they form the apex of the cervical lordosis and the site at which the greatest age-related modification occurs. The tissue was processed in paraffin, sectioned, and then stained by Masson's trichrome. Spindle characteristics were examined using light microscopy and analyzed by unbiased stereologic methods. A one-sample paired t test was used to ascertain whether the differences in spindle density between the two muscles were statistically significant., Results: The longus colli has a high density of muscle spindles, which appear clustered and concentrated anterolaterally, away from the vertebral body. The multifidus has a low density of muscle spindles, which are found predominantly as single units concentrated closely to the vertebral lamina. No change in spindle distribution, morphology, and density were observed with age., Conclusions: The current study examined spindle characteristics for an intrinsic neck muscle pair whose coactivation contributes to segmental stability of the cervical spine. The distribution and morphology of muscle spindles differ between the longus colli and the multifidus. In addition, these muscles have significant differences in terms of mean spindle density. Spindle characteristics represent one of many factors that govern proprioceptive regulation in skeletal muscle, and in neck muscles, the central connectivity of these receptors remains undefined. Therefore, although there are anatomic differences between the neck flexor and extensor, the functional implications of these differences are not clear. It is also of interest that spindle characteristics remain unchanged in these intrinsic muscles whose underlying segments are subject to age-related osteoligamentous changes.

A model of the muscle spindle was developed based on its anatomical structure. The model contains three intrafusal fibers (bag1, bag2, and chain), two efferents (dynamic gamma efferent to the bag1 fiber and static gamma efferent to bag2 and chain fibers), and two afferents [primary (Ia) and secondary (II)]. As in the real muscle spindle, the spindle model, under the modulation of gamma efferents, responds to the extrafusal muscle fiber length. Both outputs (Ia and II afferents) of the model were compared extensively with published data, under both sinusoidal stretch (with different stretch amplitudes and frequencies) and ramp and hold stretch (with different stretch amplitudes and velocities) in three different fusimotor activation conditions (dynamic gamma stimulation, static gamma stimulation, and without gamma stimulation). Model Ia afferent responses fit the published data well with active gamma input, but less well in the passive state. Model II afferent responses also fit the published data, although less quantitative data were available for comparison. The model correctly predicted the fractional power dependence of the primary and secondary ending responses on stretch velocity. The current model provides a powerful tool for simulation studies of neuromusculoskeletal systems, and demonstrates the feasibility of using a structural approach to model complex neurophysiological systems.

The capacity of muscle spindles in adult human extraocular muscles (EOM) to provide effective proprioception was questioned on the grounds of their peculiar morphology. Their appearance could be attributable to ageing and to test this possibility examples of infant muscle spindles have been examined. Forty encapsulated structures from five extraocular muscles removed post mortem from 4 infant patients aged 6 days, 5, 23 and 30 months were examined by means of light microscopy using serial transverse sections. Seven of them were identified as false spindles. The remaining 33 structures, identified as spindles, contained a total of 175 intrafusal fibres varying from 2 to 12 (mean: 6) in each. 130 of these fibres (74.3%) were of nuclear chain type. Unequivocal evidence of bag fibres was not found. Spindles lacked or had a limited equatorial expansion, and the inner capsule was incomplete and irregularly shaped. 45 (25.7%) of the intrafusal fibres had extrafusal features with large diameters, peripherally placed nuclei, no equatorial modification and without associated sensory nerve terminals. Serial sections revealed that a majority of the nuclear chain fibres were interrupted, fragmented or terminated abruptly, and most spindles contained at least one incomplete fibre. These observations show that the atypical features observed in adult human EOM spindles are also present in infants and are therefore not attributable to ageing., (Copyright 2001 S. Karger AG, Basel)

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.

Although the middle ear muscles have been described for the first time more than four hundred years ago their role in modulation and transmission of sound is not yet fully understood. Surprisingly very little is known about proprioceptors in these muscles, especially in man, although this seems to be the key to the understanding of their various functions. Therefore, the question for proprioceptive sensory organs in these muscles is still relevant. The tensor tympani and stapedius muscles of four women who had donated their bodies to our institute were taken. Complete serial sections of these muscles were made which were either impregnated with silver, stained with ferric oxide for acidic polysaccharides or incubated with antibodies against S-100 protein. Thereby four to eight (mean five) muscle spindles distributed along the whole muscle could be detected in the tensor tympani muscles. These spindles contain one to three intrafusal muscle fibres and their length ranges from 140 to 4270 microm (mean 1492.8 microm). Furthermore, in three stapedius muscles one to two (mean 1.7) muscle spindles were found. They were from 350 to 500 microm (mean 482 microm) long and contained only one intrafusal muscle fiber. Regarding the diameter of intrafusal muscle fibers in both, the tensor tympani as well as the stapedius muscle, no difference to extrafusal muscle fibers of these muscles could be detected. The structure of these spindles differs considerably from those found in skeletal muscles. The morphological findings presented strongly suggest that muscle spindles occur regularly in both middle ear muscles. The results presented herein are consistent with clinical findings obtained from electromyographic studies and may help to elucidate all functions the middle ear muscles might serve in man.

In muscle spindles of the cat, independent control of dynamic and static components of the response of the primary sensory ending to stretch is provided by separate motor inputs to the various kinds of intrafusal muscle fibre: dynamic axons (gamma or beta) to the bag1 fibres and static axons to the bag2 (typically gamma only) and chain (gamma or beta) fibres. Nonlinear summation of separately evoked effects during combined stimulation of dynamic and static motor axons appears to be due to mutual resetting by antidromic invasion of separate encoding sites, leading to partial occlusion of the momentarily lesser response by the greater. The encoding sites are thought to be located within the primary ending's preterminal branches which from first-order level are normally segregated to the bag1 fibre and to the bag2 and chain fibres. Here we describe the analysis of a special case that arose in a histophysiological study which had shown that the degree of occlusion was related to the minimum number of nodes between the putative encoding sites. Three-dimensional reconstruction of the primary ending revealed that the terminals of one chain fibre were derived entirely from the first-order branch that supplied the bag1 fibre, including one terminal that was shared directly with the bag1 (sensory cross-terminal). The other first-order branch supplied the bag2 and remaining chain fibres as normal. The degree of occlusion seen during simultaneous stimulation of a dynamic beta axon and a static gamma axon indicated that the encoding sites were separated by both first-order branches. Schematic reconstruction of the motor innervation revealed that the static gamma axon was most unlikely to have supplied the chain fibre which shared sensory terminals with the bag1, but that these fibres also shared a motor input with histological characteristics of beta type. Ramp-frequency stimulation of the dynamic beta axon at constant length evoked a driving effect which persisted after fatiguing the extrafusal component and was therefore explicable on the basis of the observed pattern of motor innervation, though the identity of the axon could not be conclusively proved. Individually, instances of shared sensory terminals and motor input of bag1 and chain fibres are rare in the cat; their combination in a single spindle with correlated physiology is described here for the first time. The observation is considered in relation to the importance of dynamic and static segregation in motor control, since it may imply that there is a lower limit to the degree of segregation that the developmental programme can provide.

A reconstruction was made of the trajectory of primary spindle afferents from the intercostal muscles in the spinal cord of the cat. Intraaxonal recordings were performed from the primary spindle afferents that were identified by their response to lung inflation and stimulus threshold to activate the action potentials. The afferents were stained by using intraaxonal injection of horseradish peroxidase (HRP). Results were obtained mainly from internal intercostal Ia fibers, which entered the spinal cord and bifurcated into ascending and descending branches. The ascending branches could be traced up to 10.7 mm, and the descending branches could be traced up to 7.3 mm. The ascending branches extended to the next segment. Collaterals ranging from one to six were given off from these branches. The distances between adjacent collaterals ranged from 0.9 mm to 4.7 mm. Each collateral had similar morphological characteristics. The collaterals entered the dorsal horn and ran toward lamina IX through the medial half of the gray matter. Fine branches and boutons were given off in laminae V, VII, VIII, and IX. The aggregations of these branches were found in lamina VII, mainly in the region of Clarke's column and in the ventral and ventrolateral regions thereof and in lamina IX, mainly in the nucleus lateromedialis. Most terminals did not contact the somata of target neurons in all laminae in which terminals were found. However, a few terminals were found to contact large neurons in lamina IX. In addition to these aggregates, there were some terminals scattered throughout the ventral horn. Thus, it was concluded that single intercostal Ia afferents project to the region of Clarke's column, to the intercostal motor nucleus, and to the intermediate regions.

The South American opossum, Monodelphis domestica, is very immature at birth, and we wished to assess its potential for studies of jaw muscle development. Given the lack of prior information about any Monodelphis fiber types or spindles, our study aimed to identify for the first time fiber types in both adult and neonatal muscles and the location of spindles in the jaw muscles. Fiber types were identified in frozen sections of adult and 6-day-old jaw and limb muscles by using myosin ATPase and metabolic enzyme histochemistry and by immunostaining for myosin isoforms. The distribution of fiber types and muscle spindles throughout the jaw-closer muscles was identified by immunostaining of sections of methacarnoy-fixed, wax-embedded heads. Most muscles contained one slow (type I) and two fast fiber types (equivalent to types IIA and IIX), which were similar to those in eutherian muscle, and an additional (non-IIB) fast type. In jaw-closer muscles, the main extrafusal fiber type was IIM (characteristic of these muscles in some eutherians), and almost all spindles were concentrated in four restricted areas: one in masseter and three in temporalis. Six-day neonatal muscles were very immature, but future spindle-rich areas were revealed by immunostaining and corresponded in position to the adult areas. Extrafusal and spindle fiber types in Monodelphis share many similarities with eutherian mammalian muscle. This finding, along with the immaturity of myosin isoform expression observed 6 days postnatally, indicates that Monodelphis could provide a valuable model for studying early developmental events in the jaw-closer muscles and their spindles.

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.

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.

Background: Increasing importance is attributed to the proprioceptive innervation of extraocular muscles for the development of binocular vision. In the literature, data on proprioceptors in human extraocular muscles are rare and inconsistent. Therefore a detailed morphological analysis of spindles in human extraocular muscles appeared indicated., Material and Methods: Complete serial sections of all extraocular muscles of 3 human individuals, 67, 72, and 83 years of age, were alternately impregnated with silver, stained following Mowry and immunohistochemically stained for S100 or PGP 9.5. Distal parts of extraocular muscles from multiorgan donors, 2, 17, and 34 years of age, were prepared for electron microscopy., Results: Spindles contained 1 to 18 intrafusal muscle fibers, 55% of which were nuclear chain fibers, 2% nuclear bag fibers, and 43% fibers exhibiting the same morphological features as extrafusal fibers. The latter had been previously described as "anomalous fibers" (Ruskell). This intrafusal type exclusively occurs in human extraocular muscle spindles. Ultrastructural analysis revealed morphologically normal sensory terminals on all 3 types of intrafusal fibers. Findings in a two-year-old individual were similar to those in aged individuals., Conclusions: Spindles in human extraocular muscles are specifically structured. Their special morphological features are also present during the development of binocular vision. Spindles are supposed to play a role in the (fine) control of eye movements.

The paper presents the present state of research on the most important receptors of joints and muscles with regard to occurrence, structure, and function. Proprioception relies on the nervous supply of the tissues in the knee joint, and on the surrounding muscles and tendons; it is the basis of coordination.

The structure-function relationships of the macaque extensor digitorum and hallucis brevis muscle (EDB and EHB), the single intrinsic and four-tendoned muscle of the dorsum of the foot, were investigated in the macaque after preparation of samples by two plastination methods, using gelatin and epoxy resin. These preparative methods clearly revealed the muscle fibers, spindles and myelinated axons in the EDB and EHB. In the more medially located two bellies (the medial two bellies of the EDB), the mean relative proportion of red fibers was larger than in the more laterally located two bellies (the lateral belly of the EDB, and the EHB), while those of intermediate and white fibers were smaller than in the more laterally located two bellies. (The difference was significant between some pairs of bellies.) In the more medially located two bellies, the mean cross-sectional area of each type of fiber, namely, the white, intermediate and red fibers, tended to be smaller than in the more laterally located two bellies. (There were significant differences between many pairs of bellies.) The total number of muscle spindles within an EDB and EHB was 15, and larger numbers of muscle spindles were distributed to the medially located two bellies compared to the laterally located two bellies. (There were significant differences between many pairs of bellies.) These results suggest that the more medially located two bellies of the EDB and EHB are adapted to the relatively precise movements of toes, while the more laterally located two bellies are adapted to the prompt extension and abduction that are related to the arboreal habitat of the macaques.

The spatial representation of muscle spindles (Sps) in the small (approximately 0.2 g), simply structured capsularis muscle that crosses anterior to the cat's hip joint was compared with the distribution of the slow oxidative (SO) and few (< 10%) fast oxidative-glycolytic (FOG) fibers of which it is composed to see if their distributions were consistent with a hypothesis that sensory input from Sps influences the incidence of extrafusal fiber types. In frozen sections from 4 muscles, FOG fibers were enumerated along 1-mm strips across the muscle's maximum width, and between the 'superficial' surface and the 'deep' one that contacts the joint. The locations of Sps in complete serial sections of 2 paraffin-embedded muscles, one perfused with the hip joint flexed and the other with it extended, were plotted on an outline of each muscle at its midlength, and their numbers and density in horizontal and sagittal 'strata' determined. In general, the incidence of Sps increased down the superficial-to-deep axis, while FOG fibers became fewer, as is consistent with support of SO status by Sp input. Along the craniocaudal axis, i.e. width, the numbers of FOG fibers rose toward the hip joint, but this was not associated with a monomodal gradient of Sps. In the extended muscle, however, the lengths of the axial bundle and capsular space of Sps in the half of the muscle next to the joint exceeded those in the longer, cranial half, implying that under stretch the input from Sps became higher toward the joint. In the non-extended muscle these lengths did not differ, although the lengths of extrafusal fibers isolated from 2 macerated muscles and normalized according to sarcomere length decreased linearily by approximately 50% along craniocaudal axis. It is explained that if elastic resistance of a Sp's sensory region exceeded that of an equivalent length of septal tissue in-series, the progressive shift in the ratio of compliances across this trapezoidally-shaped muscle should result in relatively greater lengthening of Sps at the shorter border as the muscle was extended. Levels of discharge conducive to transition of some motor units to the FOG type might be attained. Thus, gradients in the discharge of Sps (but not necessarily incidence) along both transverse and superficial-to-deep axes may be consistent with Sp sensory input influencing the distribution of at least these types of motor units.

The review provides data on the pre- and postnatal ontogenesis of muscle spindles. I demonstrated the roles of various sections of the nervous system in the mechanism of development and differentiation of the intrafusal muscle fibres. It provides data on the structure, dynamic and static nuclear bag muscle and nuclear chain fibres, their motoric and sensory innervations. It discussed the functional characteristics of responses from the primary and secondary endings to various mechanical stimulations. The paper discusses the results describing potentials (local and distributing) of intrafusal fibres of all types. It also describes the role of fusimotor innervation in regulation of the sensory activity of muscle spindles.

Superficial lumbrical spindles of the cat show several important differences as compared with the classic picture of the spindle, derived principally from tenuissimus. Thus long chain fibres occur in a high proportion of superficial lumbrical spindles, and there is great individual variation in the number of bag fibres present. This study is especially concerned with the morphology and relationships of the bag1, fibre, which are again highly unusual. Most notably, bag1 fibres were almost always the longest intrafusal fibres, typically exceeding bag2 fibres by about 2 mm in total length. They also commonly exhibited the greatest diameter, especially in the extracapsular region where, uniquely, they normally increased in size on leaving the capsule, often becoming similar to extrafusal fibres in this respect. In a few spindles where there was only a single bag fibre this was more often a bag1 than is the case for similar spindles in tenuissimus, and in tandem-linked spindles the continuous bag fibres were more often bag1 than bag2. These results contribute to growing evidence that different muscles exhibit characteristic features of the structure and innervation of their proprioceptors, which may reflect adaptations to local control requirements.

The objectives of this study were to identify the exact location of spindles in jaw-closer muscles of the cat, to count the total number of spindles and to compare their distribution with the distribution of slow extrafusal fibres. The jaw-closer muscle group with all the skeletal attachments intact was fixed in a modified Carnoy solution, decalcified and processed through to wax. Complete series of sections were cut transverse, sagittal and perpendicular to the anterior temporalis muscle. At regular intervals, serial sections were stained by the Weigert-van Gieson method or immunostained for myosin isoforms. Spindle counts were made only from muscle areas where fibres were cut in transverse section, and the spindles were followed individually. The extrafusal fibres were identified by indirect immunoperoxidase staining with antibodies specific for the slow (type I) and fast (type IIM) isoforms of myosin found in jaw-closer muscles. The mean numbers and locations of spindles found were 13 in medial pterygoid (close to inferior border), 123 (one count only) in the deep anterior portion of the temporalis muscle (between the coronoid process of the mandible and the cranium), 50.5 in a small deep zone of superficial masseter anterior to the temporomandibular joint, and 50 in zygomaticomandibularis (the deepest portion of masseter). Most of the spindles were simple spindles. Spindle complexes (4 or more spindle units fused in parallel) were rare and were found only in zygomaticomandibularis and in masseter. Most parts of the jaw-closer muscles had no spindles and contained only fast fibres. Cosegregation of muscle spindles with slow fibres was found in most parts of this muscle group, but the distribution of spindles was more restricted than that of slow fibres.

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.

The fiber-type composition of postnatal chicken leg muscle spindles with from one to four intrafusal fibers was examined in sections incubated with monoclonal antibodies against fast and slow myosin heavy chains. In monofibral spindles the lone intrafusal fiber was almost always fast. In duofibral spindles usually one slow and one fast fiber were present. Trifibral spindles most often displayed two fast and one slow fiber, whereas quadrofibral receptors characteristically contained two slow and two fast fibers. Earlier results showed that the primary intrafusal myotube in nascent spindles has almost always a fast myosin heavy chain profile and that the proportion of slow myotubes and fibers increases as intrafusal fiber bundles grow in size. Data from postnatal chicken leg muscles collected here suggest that up to the first four fibers this proportional increase can be largely accounted for if consecutive intrafusal fibers arise in a fast-slow-fast-slow sequence. The late recognition during myogenesis of primary intrafusal myotubes and their fast myosin heavy chain profiles warrant exploring if nascent chicken muscles spindles are first seeded by fast fetal myoblasts.

The ultrastructure of muscle spindles de-efferented by the extirpation of the lumbosacral spinal cord at the age of 2 days and subsequently deprived of their sensory innervation by the section of the sciatic nerve at 3-4 weeks of age was studied in serial sections of 2-month-old rat hindlimb muscles. De-efferentation leaves the primary sensory neurons and their peripheral axons intact and capable of inducing the muscle spindle morphogenesis during the critical period of their development. In de-efferented and subsequently denervated muscle spindles, new supernumerary intrafusal muscle profiles (SIPs) appeared in the muscle spindle A region. They were formed in intimate spatial relation with the original intrafusal muscle fibres (IMFs) predominantly from activated satellite cells derived from both nuclear bag (larger diameter) and nuclear chain fibres. SIPs, however, lacked the typical nuclear accumulations, as well as other ultrastructural distinctions present in control IMFs. The majority of differentiated SIPs separated from original IMFs, whereas the less differentiated SIPs were usually closely apposed to the surface of the parent IMFs and both were covered by the common basal lamina. In some spindles, the original IMFs and/or new SIPs at different stages of their differentiation were found together and they formed clusters of variable shape and composition. In the majority of clusters, all profiles seemed to be isolated along their entire length, although in few clusters, occasional cytoplasmic connections of variable length between intrafusal profiles were found. This result is important for the interpretation of the forthcoming study of expression of muscle spindle-specific myosin heavy chain isoforms in denervated SIPs in rat muscle spindles gradually deprived of their motor and sensory innervation.

A quantitative analysis of the motor innervation of intrafusal muscle fibres is described, based on teased silver-impregnated spindles of the tenuissimus muscle of the cat. Included in the analysis are the number and distribution of intrafusal branches of both skeletofusimotor (beta) and purely fusimotor (gamma) axons, and the form of their endings. The number of axonal branches per spindle was found to follow binomial probability distributions, as had previously been shown for the afferent axons. There was a strong correlation between the numbers of gamma intrafusal branches and afferent axons, but none for the intrafusal branches of beta axons. The degree of segregation of gamma input to bag2 and chain fibres was assessed and was found, among other things, to be related to the presence of secondary sensory endings in the same pole. In this and other respects it did not appear to have the properties that would be expected if independent activation of the bag2 and chain fibres were to be functionally important. Morphometric analysis of the motor endings supplied to bag2 or chain fibres by gamma axons revealed some differences between those of intrafusal branches with segregated as opposed to unsegregated distributions, but this cannot be taken as evidence of more than one type of static gamma motoneuron because of the likely contribution of other influential factors such as fibre size. Finally, the relevance of studies on intrafusal motor innervation to the concept of the motor unit and its development are discussed.

To assess the respiratory function of the scalene and sternomastoid muscles in the dog, we studied the effect of graded increases in inspiratory airflow resistance and single-breath airway occlusion on the electrical activity of these muscles in 18 supine anesthetized spontaneously breathing animals. The sternomastoids never showed any activity, and the scalenes showed some inspiratory activity during occlusion in only two animals. The adoption of the prone position and bilateral cervical vagotomy did not affect this pattern. Hypercapnia also did not elicit any sternomastoid activity and induced scalene inspiratory activity during occlusion in only four of nine animals. On microscopic examination, however, both muscles were found to contain large numbers of spindles, suggesting that they have the capacity to respond to stretch. In addition, with increases in inspiratory resistance, both the sternum and ribs were displaced in the caudal direction. As a result, the scalenes demonstrated a gradual inspiratory lengthening and the normal inspiratory lengthening of the sternomastoids was accentuated. Additional studies in three unanesthetized animals showed consistent activity in the scalene and sternomastoid muscles during movements of the trunk and neck but no activity during breathing, including occluded breathing. These observations thus indicate that the alpha-motoneurons of the scalene and sternomastoid muscles in the dog have very small central respiratory drive potentials with respect to their critical firing threshold. In this animal, these muscles do not have a significant respiratory function.

1. By recording, with 2 pairs of wire electrodes, single-fibre action potentials (APs) from lower sacral nerve roots of a brain-dead human and a patient with spinal cord lesion, impulse patterns of afferent APs and impulse trains of oscillatory firing motoneurons could be identified and correlated. 2. Two highly activated secondary muscle spindle afferents increased and decreased their activity at about 0.3 Hz. The duration of the doublet interspike interval of a secondary spindle afferent fibre showed no correlation to the oscillation period of the motoneuron. 3. A continuously oscillatory firing motoneuron innervating the external and sphincter showed more transient breaks with the reduction of the number of phase correlated APs from 2 spindle afferents, indicating a looser oscillation. A transient brake of a 157 msec period alpha 2-oscillation could be correlated to the shift of a interspike interval distribution peak from 150 to 180 msec of the adequate afferent input, which suggests a transient loss of the necessary phase relation. 4. Oscillatory firing alpha 2-motoneurons innervating the external bladder and anal sphincters fired independently according to their phase correlated APs from the urinary bladder stretch receptor and muscle spindle afferents respectively; the bladder motoneuron slightly inhibited the anal motoneuron. 5. Receptors of the afferents and innervation sites of oscillatory firing motoneurons could be located within the urinary tract and the anal canal.

Small samples of the masseter and temporalis muscles of six adult male cats were dissected free, snap-frozen and cryosectioned. Serial sections from areas with muscle spindles were histochemically stained for myofibrillar ATPase after incubation in acid and alkali buffer, and for reactivity to two different myosin isoform-specific antibodies. Differential reactivity to these four stains identified bag1, bag2 and chain intrafusal fibre types. The most typical distribution of fibres within the spindle was 1 bag1, 1 bag2 and 3-4 chains. Although a few spindles were found in the anterior area of the temporalis that attaches to the coronoid process of the mandible and in the anterior area of the superficial masseter, the vast majority were found in the deepest portion of the masseter. Here spindles were characteristically with a number of simple spindles fused together into complex units.

Among the hypotheses regarding fusimotor functions based on earlier animal experiments some are inconsistent, others are in conformity with microneurographic observations in man. The human data provide evidence against the following two theories: (1) the length follow-up servo theory; and (2) the theory that fusimotor neurons can be selectively activated to produce spindle sensitization and stretch reflex reinforcements. The human data support the theory of alpha-gamma coactivation. In particular, in the early phase of isometric voluntary contractions fusimotor-driven afferent spindle activity assists in autogenetic activation of alpha motoneurons and in reciprocal relaxation of antagonists. As muscle fatigue develops, the autogenetic reflex drive via the fusimotor route declines. The fusimotor bias during contraction provides for maintenance of spindle sensitivity to minute perturbations and for load-compensating reflex adjustments to such perturbations. Reflex overcorrections may lead to uncontrollable oscillations of the type seen in enhanced physiological tremor.

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.