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PubMed1986

The innervation of tandem muscle spindles in the cat neck.

Patterns of innervation were examined in tandem muscle spindles teased from silver-stained muscles of the cat neck. Each tandem spindle was composed of two or more encapsulated receptors linked in series by a shared bag2 fiber. In most tandem spindles, two different types of encapsulation were identified according to differences in their intrafusal fiber content. One type, the b1b2c unit, contained typical bag1, bag2, and chain fibers and was structurally similar to single spindles described in other cat muscles. Each b1b2c unit contained a single primary sensory ending and 1-6 secondary endings. Fusimotor innervation was supplied by many axons. Some fusimotor axons ended in trail ramifications on bag2 and chain fibers, others ended in plates on the bag1 or long chain fiber. The other type of tandem encapsulation, the b2c unit, had only bag2 and chain fibers in its intrafusal fiber bundle. The b2c unit was usually supplied by only one sensory axon that ended on the nucleated part of the intrafusal fiber bundle. This single ending had a more variable terminal morphology than the primary ending in b1b2c units. A few b2c units (3/49) were also supplied by a secondary ending. The fusimotor innervation of the b2c unit was relatively simple. A single pole of the b2c unit was usually supplied by only one to three axons, all ending in trail ramifications. No plate endings were found in b2c units. These morphological specializations suggest that b1b2c and b2c units in tandem spindles differ in both their transductive and fusimotor mechanisms. Thus, the tandem spindle is a specialized structure that may provide additional proprioceptive information beyond that available from single muscle spindles.

Animals↗

PubMed1991

A 210 kDa nuclear matrix protein is a functional part of the mitotic spindle; a microinjection study using SPN monoclonal antibodies.

Six monoclonal antibodies identify a 210 kDa polypeptide which shows a cell cycle specific redistribution from the nucleus to the mitotic spindle. In interphase cells this polypeptide was localized in the nucleus and behaved during differential cell extraction as a component of the nuclear matrix. It accumulated in the centrosome region at prophase, in the pole regions of the mitotic spindle at metaphase and in crescents at the poles in anaphase, and reassociated with the nuclei as they reformed in telophase. Due to its staining pattern we call the protein the Spindle Pole-Nucleus (SPN) antigen. The localization of SPN antigen during mitosis was dependent on the integrity of the spindle since treatment of cells with nocodazole resulted in the dispersal of SPN antigen into many small foci which acted as microtubule organizing centres when the drug was removed. The SPN antigen was present in nuclei and mitotic spindles of all human and mammalian cell lines and tissues so far tested. When microinjected into the cytoplasm or nuclei of HeLa cells, one antibody caused a block in mitosis. Total cell number remained constant or decreased slightly after 24 h. At this time, about half the cells were arrested in a prometaphase-like state and revealed aberrant spindles. Many other cells were multinucleate. These results show that the SPN antigen is a protein associated with mitotic spindle microtubules which has to function correctly for the cell to complete mitosis.

Antibodies, Monoclonal↗

PubMed1984

A comparison of spindle concentrations in large and small muscles acting in parallel combinations.

A small short muscle frequently acts across a joint in parallel with a vastly larger and longer muscle; therefore it should play a minimal role in the mechanical control of that joint. This study provides evidence suggesting that the small member of such a "parallel muscle combination" (PMC) may serve an important sensory feedback role. The spindle densities of large and small members of PMCs in man and the dog were determined and compared. Epaxial PMCs controlling canine intervertebral joints were dissected and tissue samples were embedded in paraffin, sectioned transversely to the muscles' long axis and, stained with hematoxylin-periodic acid-Schiff (PAS). Representative tissue sections were projected on to stereological grids and the percentage volume of spindles was determined. Data existing in the literature were used to ascertain spindle densities in human PMCs controlling joints in the cervico-occipital region and the extremities. The spindle density for each muscle in a group of PMCs controlling a particular motion was listed, and the mean spindle densities were determined for both the large and the small members of the group. Student's unpaired t test was used to determine the significance of the differences between mean spindle densities. Linear regression was calculated and the data were plotted graphically. In all PMCs examined, the spindle density of the small muscles was significantly higher than that of their large counterparts. It is therefore proposed that the small muscles of PMCs may function as "kinesiological monitors" generating important proprioceptive feedback to the central nervous system.

Animals↗

PubMed1976

Fine structure of the avian muscle spindle capsule.

The capsule of the muscle spindle from the anterior and posterior latissimus dorsi muscles of the adult domestic chicken has been studied with the electron microscope. As in other species, two distinct portions of the spindle capsule are distinguished: an outer capsule and an inner capsule. The outer capsule is structurally similar to and continuous with the perineural epithelium. Outer capsule cells are noted by the abundance of pinocytotic vesicles and a network of 6-7-nm microfilaments. The disposition of these microfilaments is circumferential with respect to the longitudinal axis of the spindle. It is proposed that they may provide a contractile mechanism for the capsule which may be related to the over-all functioning of the spindle during movements of the muscle. The inner capsule is composed of a contiguous network of cells possessing long cytoplasmic processes which envelop the intrafusal fibers and their nerve endings in sensory equatorial regions of the spindle. These cells may elaborate the fibrillar and amorphous extracellular material found in periaxial spindle space. They also possess modified cilia with a "9+0" microtubular pattern. It is suggested that these cilia may behave as sensory transducers, relating fluid changes in the periaxial space to the intrafusal fiber nerve endings. Capillaries of the non-fenestrated variety commonly traverse the outer and inner portions of the capsule and are usually completely surrounded by tenuous overlapping processes of inner capsule cells. These findings suggest that the spindle capsule plays a role as a metabolically-active diffusion barrier to the entrance of substances from the external milieu.

Animals↗

PubMed1995

Neomyogenesis in neonatally de-efferented and postnatally denervated rat muscle spindles.

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.

Animals↗

PubMed1992

Calbindin D-28k-immunoreactivity in rat muscle spindles during postnatal maturation and after denervation.

Calbindin D-28k-immunoreactivity has been demonstrated in some of the intrafusal muscle fibres and in the capsule of adult rat muscle spindles. In this study, the immunocytochemical localization of calbindin D-28k in the muscle spindles of triceps surae muscle was studied during postnatal maturation and after denervation. In young rats calbindin D-28k-immunoreactivity was seen in a few intrafusal fibres, first at the age of 4 days. At the 7th day, three calbindin D-28k-immunoreactive fibres and one unlabelled fibre were seen in most muscle spindles, as in adult rats. The spindle capsule and perineurial sheath of nerves were first seen to exhibit calbindin D-28k immunoreactivity at the age of 14 days, and thereafter the localization of calbinding D-28k-like immunoreactivity was similar to that in adult rats. After denervation, calbindin D-28k-immunoreactivity remained in intrafusal muscle fibres and the spindle capsule for a long period. After two months of denervation, calbindin D-28k immunoreactivity could still be seen in the spindle capsule, but the intrafusal fibres were not labelled. The innervation is known to have trophic effects on the intrafusal fibres. The present findings suggest that the expression of calbindin D-28k-immunoreactivity in maturating muscle spindles may be induced by the developing innervation. The decrease of calbindin D-28k-immunoreactivity in intrafusal fibres after denervation may be due to the loss of trophic factors released by the nerves.

Animals↗

PubMed1990

Increase of elastic fibres in muscle spindles of rats following single or repeated denervation with or without reinnervation.

Muscle spindles in the lower lumbrical muscles of rats were studied by transmission electron microscopy following denervation with or without reinnervation. The number and total area of elastic fibres per muscle spindle increased at 3-12 months following various experimental procedures: (1) denervation and reinnervation after a single crush lesion to the sciatic nerve; (2) reinnervation after four-fold repeated crush injuries; and (3) transection and suture of the nerve. The increased number of oxytalan and elaunin fibres, the precursors of mature elastic fibres, within these muscle spindles provided further evidence for their numerical and dimensional increase. An attachment site of elastic fibres at the spindle pole was identified at the inner cells of the outer spindle capsule. The processes of these cells embraced terminating elastic fibres tightly. Attachment of elastic fibres to intrafusal muscle fibres was less conspicuous since they were not similarly embraced but were rather indistinctly, though closely, associated with the basal lamina along longitudinal surface indentations of intrafusal muscle fibres. It is concluded from this series of experiments that muscle spindles, as dynamic mechanoreceptors, maintain their elastic properties even under pathological conditions. The increase of elastic fibres following denervation and reinnervation represents an obviously meaningful reaction that may compensate for loss of tonic properties of muscle spindles without causing stiffness.

Animals↗

PubMed1998

Trichlorfon exposure, spindle aberrations and nondisjunction in mammalian oocytes.

Consumption of trichlorfon-poisoned fish by women in a small Hungarian village has been associated with trisomy resulting from an error of meiosis II in oogenesis. We therefore examined mouse oocytes exposed for 3 h during fertilization to 50 microg/ml trichlorfon. Spindle morphology was not visibly altered by the pesticide. Chromosomes segregated normally at anaphase II with no induction of aneuploidy. However, formation of a spindle was disturbed in many oocytes resuming meiosis I in the presence of trichlorfon. In spite of the spindle aberrations and the failure of bivalents to align properly at the equator, oocytes did not become meiotically arrested but progressed to metaphase II. At this stage, spindles were highly abnormal, and chromosomes were often totally unaligned, unattached or dispersed on the elongated and disorganized spindle. By causing spindle aberrations and influencing chromosome congression, trichlorfon appears, therefore, to predispose mammalian oocytes to random chromosome segregation, especially when they undergo a first division and develop to metaphase II during exposure. This is the first case in which environmentally induced human trisomy can be correlated with spindle aberrations induced by chemical exposure. Our observations suggest that oocytes may not possess a checkpoint sensing displacement of chromosomes from the equator at meiosis I and may therefore be prone to nondisjunction.

Anaphase↗

PubMed1996

A numerical simulation of muscle spindle ensemble encoding during planar movement of the human arm: correlation sensitivity and parameter dependence.

We extend the analysis developed in the preceding paper in which we correlated kinematic parameters of planar movements of the human arm made by subjects moving to a visual target with numerical estimates of the ensemble encoding of muscle spindles within some of the muscles of this limb. Three possible models for the inclusion of noise in the calculations of the ensemble encodings are considered: (i) random errors in the angular coordinates from which muscle fascicle, and hence spindle length are calculated, (ii) variability of spindle discharge rates, and (iii) variability in the calculation of the ensemble encoding. In each case the correlations between kinematic variables of the movements and the resultant ensemble encodings decrease as the contribution of the noise term to the calculation of the encodings increases. Subject to the constraint that the magnitude of the noise term remains within physiologically realistic limits, however, the observed correlations persist at statistically significant levels. We also investigate the dependence of the observed correlations on the choice of model parameters, namely (i) the absolute and relative contributions made by simulated spindle primary and secondary afferents to the ensemble encoding, (ii) the inclusion of explicit length-related terms in the model of muscle spindle discharge, and (iii) the fractional power of velocity experienced by the model spindles during movement. The resulting correlations are approximately independent of both the fractional power of velocity and absolute firing levels of both the primary and secondary afferents of the spindle model. The inclusion of explicit length-dependent terms in the model does result in differences in the observed correlation coefficients. In this case, however, the magnitudes of the differences are small. On the basis of these findings we conclude that the correlations between kinematic variables of movement and the associated ensemble encodings are robust with regard to both the choice of model parameters and noise inherent at all stages of the transduction and processing of proprioceptive information. The findings of the present study provide further evidence, therefore, to support the hypothesis that motor structures capable of deriving such an ensemble encoding would be provided with information regarding ongoing movements in both intrinsic (body-centered) and extrinsic (Cartesian) coordinate systems.

Animals↗

PubMed2004

Gamma-tubulin localization changes from discrete polar organizers to anastral spindles and phragmoplasts in mitosis of Marchantia polymorpha L.

Unlike the astral mitotic spindle that is organized at discrete centriolar centrosomes, the spindle of land plants is typically anastral and its origin has remained obscure. Gamma tubulin (gamma-tubulin), an important component of the centrosome, has been demonstrated at microtubule-nucleating sites in plant cells. Mitotic spindles of certain hepatics are initiated at distinct acentriolar polar organizers (POs) that appear de novo at the onset of mitosis. Data on the relationship of gamma-tubulin to POs and to microtubule arrays throughout the cell cycle were collected from rapidly dividing cells of Marchantia polymorpha (Bryophyta) that were triple-stained for gamma-tubulin, microtubules, and nuclei. POs at opposite ends of the elongated nucleus in early prophase stain brightly for gamma-tubulin and astral microtubules emanating from them initiate the spindle. As the spindle develops, however, the gamma-tubulin becomes dispersed from the highly concentrated spherical form of the POs to more diffusely organized cups at tips of the fusiform nucleus. By the end of prophase, all astral microtubules have disappeared and the gamma-tubulin is located in several minipoles along the now broad polar regions of the spindle. At metaphase, gamma-tubulin extends into the spindle itself. By telophase, the gamma-tubulin has migrated from distal to proximal surfaces of the sister nuclei and extends into the phragmoplast. Upon completion of cytokinesis, gamma-tubulin appears diminished and surrounds the nuclear envelopes. These data show that gamma-tubulin is only briefly concentrated in the PO, migrates in a cell-cycle-specific manner, and is consistently present at all putative sites of microtubule nucleation.

Cell Nucleus↗

PubMed1987

Sharing of sensory terminals between the dynamic bag1 and static bag2 fibers in the rat muscle spindle.

The nuclear bag1 intrafusal fiber mediates the dynamic (velocity) sensitivity, whereas the nuclear bag2 and nuclear chain fibers mediate the static (length) sensitivity of muscle spindles to stretch. The pattern of branching of primary and secondary afferents, the distribution of their terminals to the 3 types of intrafusal fibers, and the incidence of sensory cross-terminals were determined by reconstruction of 4 spindles from serial 1-micron and ultrathin transverse sections of rat extensor digitorum longus muscles. A single primary afferent supplied each spindle, and secondary afferents innervated intrafusal fibers in 3 spindles. Only static intrafusal fibers shared cross-terminals of the secondary afferents. In contrast, the dynamic bag1 and static bag2 fibers of each spindle shared at least one terminal of the primary afferent. Cross-terminals shared by the dynamic bag1 and static bag2 fiber parallel the presence of fusimotor (gamma) axons which coinnervate these types of intrafusal fibers in muscle spindles of rats. Consequently, the greater degree of overlap of elements comprising the dynamic and static systems of spindles of the rat relative to that of the cat reduces the probability of generating a purely dynamic or purely static response to an applied stretch.

Animals↗

PubMed1989

Spindle-pole organization during early mouse development.

Spindle-pole organization during early mouse development was examined using a variety of immunological reagents that recognize centrosomal components. Spindle poles of unfertilized eggs and blastocysts were found to react positively with two antisera (centrin and NRS-01), whereas poles of activated eggs and early cleavage-stage embryos were negative when treated with the same sera. In contrast, a third antiserum (5051) showed positive spindle-pole staining throughout the preimplantation stages of development. Two monoclonal antibodies (MPM-1 and MPM-2) that are known to react with mitotic phosphoproteins were also used in this study. Both antibodies stained the cytoplasm of mitotic cells with extremely high intensity. In addition, MPM-2 was found to stain spindle poles. These results suggest that organizational changes in the spindle pole are occurring during early mouse development. Embryos homozygous for a recessive lethal mutation known as oligosyndactyly (Os) were also treated with the reagents described above. This mutation results in a metaphase arrest at the blastocyst stage with intact spindles being present. Spindle poles were observed in Os homozygous mutants stained with centrin, NRS-01, and 5051. However, when Os mutants were stained with the MPM monoclonal antibodies, about half of the mitotic cells completely lacked the dramatic cytoplasmic staining. This observation is in contrast to that observed for wild-type embryos, where greater than 95% of mitotic cells showed positive cytoplasmic staining.

Animals↗

PubMed1992

Mitotic apparatus formation and cleavage induction by micromanipulation of the nucleus and centrosome: the centrosome forms a spindle together with only the chromosomes at a short distance.

We micromanipulated the nucleus and centrosomes in the zygote of the starfish, Asterina pectinifera, in order to investigate their roles in mitotic apparatus formation and cleavage induction. The zygote cleaved without spindle formation when its nucleus was removed. When one or two centrosomes were transplanted, they formed asters in the recipient cell, which cleaved into three or four blastomeres so that each blastomere might contain one centrosome or aster. When one centrosome was removed, a half-spindle formed in the manipulated cell, which did not cleave until the other centrosome was duplicated. When both centrosomes were removed, no microtubular structures such as the spindle and the aster appeared in the manipulated cell, which failed to cleave. These results indicate that two centrosomes or more in the cell induce cleavage with or without the nucleus and that one centrosome or less does not induce cleavage. It is also concluded that the centrosome(s) together with the nucleus forms a half-spindle or bipolar spindle. However, from the experiments of nucleus transplantation and displacement, spindle formation is found to depend on the distance between chromosomes and centrosomes. The half-spindle formed when the distance from the centrosome to the chromosomes was shorter than 22 microns; on the other hand, when the distance was longer than 22 microns, the nucleus remained apart from the aster, which means that the functional range of the astral microtubule's ability to engage chromosomes was 22 microns from the centrosome.

Animals↗

PubMed1994

Signal transmission from motor axons to group Ia muscle spindle afferents: frequency responses and second-order non-linearities.

Spinal recurrent inhibition via Renshaw cells and proprioceptive feedback via skeletal muscle and muscle spindle afferents have been hypothesized to constitute a compound feedback system [Windhorst (1989) Afferent Control of Posture and Locomotion; Windhorst (1993) Robots and Biological Systems--Towards a New Bionics]. To assess their detailed functions, it is necessary to know their dynamic characteristics. Previously we have extensively described the properties of signal transmission from motor axons to Renshaw cells using random motor axon stimulation and data analysis methods based thereupon. Using the same methods, we here compare these properties, in the cat, with those between motor axons and group Ia muscle spindle afferents in terms of frequency responses and nonlinear features. The frequency responses depend on the mean rate (carrier rate) of activation of motor axons and on the strength of coupling between motor units and spindles. In general, they are those of a second-order low-pass system with a cut-off at fairly low frequencies. This contrasts with the dynamics of motor axon-Renshaw cell couplings which are those of a much broader band-pass with its peak in the range of c. 2-15 Hz [Christakos (1987) Neuroscience 23, 613-623]. The second-order non-linearities in motor unit-muscle spindle signal lines are much more diverse than those in motor axon-Renshaw cell couplings. Although the average strength of response declines with mean stimulus rate in both subsystems, there is no systematic relationship between the amount of non-linearity and the average response in the former, whilst there is in the latter. The qualitative appearance of motor unit-muscle spindle non-linearities was complicated as was the average response to motor unit twitches. Thus, whilst Renshaw cells appear to dynamically reflect motor output rather faithfully, muscle spindles seem to signal local muscle fibre length changes and their dynamics. This would be consistent with the hypothesis that the two feedback pathways monitor different state variables determining the production of muscle force: neural input and length and its changes. Specifically, the dynamic properties of both subsystems may combine favourably to decrease the risk of instability (tremor) in the motoneuron-muscle spindle loop.

Animals↗

PubMed2006

Morphological and morphometrical study of human muscle spindles in Werdnig-Hoffmann disease (infantile spinal muscular atrophy type I).

A study was made of the morphological and morphometrical features of muscle spindles in biopsies of patients with Werdnig-Hoffmann disease (infantile spinal muscular atrophy type I) to investigate the possible involvement of the muscle spindles in the pathological processes of the disease. A total of 57 muscle spindles from 26 cases were studied. The parameters determined were: the diameter and area of spindles, the number, diameter and area of intrafusal fibers, the number and area of nuclei. In addition, the ratio of the area of the intrafusal fibers to the area of nuclei and the ratio of the area of the spindle to the area of the intrafusal fibers were calculated. Statistical evaluation of the data showed significant differences regarding the area of the muscle spindle, the diameter of the intrafusal fibers and the mean area of nuclei of the intrafusal fibers, which were all smaller in patients than in controls (p=0.03, 0.01 and 0.02, respectively), while the thickness of the capsule was greater in patients than in controls (p=0.01). Our results indicate that the muscle spindle participates in the pathological processes of Werdnig-Hoffmann disease.

Adult↗

PubMed2005

Asymmetric spindle positioning.

When a spindle is positioned asymmetrically in a dividing cell, the resulting daughter cells are unequal in size. Asymmetric spindle positioning is driven by regulated forces that can pull or push a spindle. The physical and molecular mechanisms that can position spindles asymmetrically have been studied in several systems, and some themes have begun to emerge from recent research. Recent work in budding yeast has presented a model for how cytoskeletal motors and cortical capture molecules can function in orienting and positioning a spindle. The temporal regulation of microtubule-based pulling forces that move a spindle has been examined in one animal system. Although the spindle positioning force generators have not been identified in most animal systems, the forces have been found to be regulated by both PAR polarity proteins and G-protein signaling pathways in more than one animal system.

Animals↗

PubMed2004

Eg5 causes elongation of meiotic spindles when flux-associated microtubule depolymerization is blocked.

In higher eukaryotes, microtubules (MT) in both halves of the mitotic spindle translocate continuously away from the midzone in a phenomenon called poleward microtubule flux. Because the spindle maintains constant length and microtubule density, this microtubule translocation must somehow be coupled to net MT depolymerization at spindle poles. The molecular mechanisms underlying both flux-associated translocation and flux-associated depolymerization are not well understood, but it can be predicted that blocking pole-based destabilization will increase spindle length, an idea that has not been tested in meiotic spindles. Here, we show that simultaneous addition of two pole-disrupting reagents p50/dynamitin and a truncated version of Xklp2 results in continuous spindle elongation in Xenopus egg extracts, and we quantitatively correlate this elongation rate with the poleward translocation of stabilized microtubules. We further use this system to demonstrate that this poleward translocation requires the activity of the kinesin-related protein Eg5. These results suggest that Eg5 is responsible for flux-associated MT translocation and that dynein and Xklp2 regulate flux-associated microtubule depolymerization at spindle poles.

Animals↗

PubMed2006

Cdc42 activation couples spindle positioning to first polar body formation in oocyte maturation.

During vertebrate egg maturation, cytokinesis initiates after one pole of the bipolar metaphase I spindle attaches to the oocyte cortex, resulting in the formation of a polar body and the mature egg. It is not known what signal couples the spindle pole positioning to polar body formation. We approached this question by drawing an analogy to mitotic exit in budding yeast, as asymmetric spindle attachment to the appropriate cortical region is the common regulatory cue. In budding yeast, the small G protein Cdc42 plays an important role in mitotic exit following the spindle pole attachment . We show here that inhibition of Cdc42 activation blocks polar body formation. The oocytes initiate anaphase but fail to properly form and direct a contractile ring. Endogenous Cdc42 is activated at the spindle pole-cortical contact site immediately prior to polar body formation. The cortical Cdc42 activity zone, which directly overlays the spindle pole, is circumscribed by a cortical RhoA activity zone; the latter defines the cytokinetic contractile furrow . As the RhoA ring contracts during cytokinesis, the Cdc42 zone expands, maintaining its complementary relationship with the RhoA ring. Cdc42 signaling may thus be an evolutionarily conserved mechanism that couples spindle positioning to asymmetric cytokinesis.

Animals↗