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PubMed1978

Analysis of the distribution of spindle microtubules in the diatom Fragilaria.

The spindle of the colonial diatom Fragilaria contains two distinct sets of spindle microtubules (MTs): (a) MTs comprising the central spindle, which is composed of two half-spindles interdigitated to form a region of "overlap"; (b) MTs which radiate laterally from the poles. The central spindles from 28 cells are reconstructed by tracking each MT of the central spindle through consecutive serial sections. Because the colonies of Fragilaria are flat ribbons of contiguous cells (clones), it is possible, by using single ribbons of cells, to compare reconstructed spindles at different mitotic stages with minimal intercellular variability. From these reconstructions we have determined: (a) the changes in distribution of MTs along the spindle during mitosis; (b) the change in the total number of MTs during mitosis; (c) the length of each MT (measured by the number of sections each traverses) at different mitotic stages; (d) the frequency of different classes of MTs (i.e., free, continuous, etc.); (e) the spatial arrangement of MTs from opposite poles in the overlap; (f) the approximate number of MTs, separate from the central spindle, which radiate from each spindle pole. From longitudinal sections of the central spindle, the lengths of the whole spindle, half-spindle, and overlap were measured from 80 cells at different mitotic stages. Numerous sources of error may create inaccuracies in these measurements; these problems are discussed. The central spindle at prophase consists predominantly of continuous MTs (pole to pole). Between late prophase and prometaphase, spindle length increases, and the spindle is transformed into two half-spindles (mainly polar MTs) interdigitated to form the overlap. At late anaphase-telophase, the overlap decreases concurrent with spindle elongation. Our interpretation is that the MTs of the central spindle slide past one another at both late prophase and late anaphase. These changes in MT distribution have the effect of elongating the spindle and are not involved in the poleward movement of the chromosomes. Some aspects of tracking spindle MTs, the interaction of MTs in the overlap, formation of the prophase spindle, and our interpretation of rearrangements of MTs, are discussed.

Cell Cycle↗

PubMed1979

Chromosome movement and spindle birefringence in locally heated cells: interaction versus local control.

A microheater was used to produce a temperature gradient within the mitotic spindle of living cells. The slope of the temperature gradient was estimated from thermal conductivity calculations and confirmed by measurements of spindle birefringence and by experiments on striated muscle. When the microheater was placed at one spindle pole or at one group of kinetochores, the gradient was steep enough to cause a large difference in birefringence between the two half-spindles, but the velocity of chromosome movement in anaphase was nearly the same in the warmer and cooler half-spindles. When the heater was shifted from the pole toward the interzone, the average velocity of chromosome movement increased approximately two-fold but was, again, nearly uniform in the two half-spindles. The rate of spindle elongation was especially sensitive to the site of heating, increasing ten-fold when the heater was shifted from the pole to the interzone. Regardless of heater position, the rate of chromosome movement was determined largely by the temperature of the coolest spindle region--chromosomes in the warmer half-spindle moved more slowly than expected from estimates of the temperature in that region. Since the microheater produces a substantial temperature gradient within the spindle, the near uniformity of chromosome velocity in both half-spindles must be due to some biological property of the spindle. Two very different explanations for the results are considered the most likely. According to one explanation, the near uniformity of velocity in both half-spindles is determined by the structure of the interpolar spindle, while changes in velocity involve force producers located both in the half-spindles and in the interzone. On the other explanation, the velocity is nearly the same in both half-spindles because the force producers are located exclusively in the interzone (Margolis et al., 1978).

Anaphase↗

PubMed1975

Barbiturate spindle activity in functionally corresponding thalamic and cortical somato-sensory areas in the cat.

(1) Spontaneous barbiturate spindles were recorded from a locus in the thalamic nucleus ventralis postero-lateralis (VPL) and the corresponding projection area in the somato-sensory cortex (SI) in the cat. (2) Type I spindles32 recorded from such sites in the thalamus and cortex, regarded as being functionally on line, were cross-correlated. The analysis gave high negative cross-correlation coefficients, indicating that a considerable degree of wave synchrony existed between the spindles. Maximal wave synchrony was present in the first and middle part of the spindle. (3) The thalamo-cortical spindle wave synchrony was reduced by a small displacement of the cortical electrode within the somato-sensory cortex. Starting with a very lightly anaesthetized animal, small amounts of barbiturate did not influence the size of the cortical area which had spindles in synchrony with the thalamic spindles. (4) Spindles recorded from a VPL locus and from its cortical projection site started simultaneously. When the thalamic and cortical electrodes were positioned functionally off line this spindle coincidence decreased. The intervals between the spindle onsets increased with increasing off line positions of the electrodes. Those few thalamic and cortical spindles which occurred simultaneously showed no, or only weak, spindle wave synchrony. (5) These data suggest a point-to-point topographical relation between thalamic and cortical loci engaged in barbiturate spindle activity. (6) Type I and type II spindles32 differed with respect to spindle characteristics (amplitude, duration, incidence) and also with respect to corresponding thalamic rhythmic activity. These spindle types are, therefore, probably produced by different mechanisms. We suggest that type II spindles represent a purely cortical process initiated by type I spindles.

Animals↗

PubMed1975

Barbiturate spindle activity in the thalamic lateral ventro-posterior nucleus and the second somato-sensory area of the cortex.

(1) Barbiturate spindles recorded from the second somato-sensory cortical area (SII) were similar to spindles in the primary somato-sensory area (SI) both with respect to incidence, duration of each spindle and per cent spindle time. The spindle wave amplitude was smaller in SII. The highest spindle wave amplitude was observed in the anterior part of SII which receives input from nucleus ventralis postero-lateralis (VPL). No spindle activity was observed in the posterior part of SII which receives input from the posterior nuclear group (PO) of the thalamus. (2) Barbiturate spindles recorded from a locus in VPL and its projection area in SII were cross-correlated. The analysis resulted in high cross-correlation factors, indicating that a considerable degree of spindle wave synchrony existed between the spindles. This wave synchrony was reduced by moving the cortical electrode a short distance. (3) Cortical spindles recorded from corresponding sites in SI and SII were cross-correlated, and gave a high cross-correlation coefficient. This synchrony was markedly reduced if one of the electrodes was moved a few millimetres away from the optimal point. (4) Spindles started simultaneously in corresponding sites of SI and SII. A high degree of coincidence was found also between spindles in a VPL locus and the corresponding projection site in SII. Local anaesthesia applied to or total removal of SI failed to influence the spindle activity in SII and vice versa. Similarly, the SI-SII synchrony survived a deep incision cutting all connections between the two areas. (6) It is suggested that spindles in corresponding sites of SII and SI have a common thalamic pacemaker which probably projects to both areas by axonal branching.

Anesthesia, General↗

PubMed1992

Role of astral microtubules and actin in spindle orientation and migration in the budding yeast, Saccharomyces cerevisiae.

In the yeast Saccharomyces cerevisiae, before the onset of anaphase, the spindle apparatus is always positioned with one spindle pole at, or through, the neck between the mother cell and the growing bud. This spindle orientation enables proper chromosome segregation to occur during anaphase, allowing one replicated genome to be segregated into the bud and the other to remain in the mother cell. In this study, we synchronized a population of cells before the onset of anaphase such that > 90% of the cells in the population had spindles with the correct orientation, and then disrupted specific cytoskeletal elements using temperature-sensitive mutations. Disruption of either the astral microtubules or actin function resulted in improper spindle orientation in approximately 40-50% of the cells. When cells with disrupted astral microtubules or actin function entered into anaphase, there was a 100-200-fold increase in the frequency of binucleated cell bodies. Thus, the maintenance of proper spindle orientation by these cytoskeletal elements was essential for proper chromosome segregation. These data are consistent with the model that proper spindle orientation is maintained by directly or indirectly tethering the astral microtubules to the actin cytoskeleton. After nuclear migration, but before anaphase, bulk chromosome movement occurs within the nucleus apparently because the chromosomes are attached to a mobile spindle. The frequency and magnitude of bulk chromosome movement is greatly diminished by disruption of the astral microtubules but not by disruption of the nonkinetochore spindle microtubules. These results suggest that astral microtubules are not only important for spindle orientation before anaphase, but they also mediate force on the spindle, generating spindle displacement and in turn chromosome movement. Potential roles for this force in spindle assembly and orientation are discussed.

Actins↗

PubMed1976

The innervation of muscle spindles in the snake, Elaphe quadrivirgata.

The morphology and distribution of sensory endings in snake (Elaphe quadrivirgata) muscle spindles was studied in silver preparations. The sensory axon supplying long-capsule spindles often bifurcates before or after penetrating the capsule and runs for some distance along the intrafusal fibre. The sensory axon supplying short-capsule spindles penetrates the capsule at a sharp angle, and, without ramification, terminates abruptly on the intrafusal fibre. The sensory terminal is composed of terminal bulbs and small links. In long-capsule spindles the sensory ending is arranged longitudinally in one or two rows. The sensory ending in short-capsule spindles ramifies extensively, covering densely the intrafusal fibre. The area occupied by sensory bulbs was estimated to be slightly higher in short-capsule (15-17?) THAN IN LONG-CAPSULE SPINDLES (12-13). The functional significance of these findings is discussed. Motor innervation of muscle spindles in the snake Elaphe quadrivirgata was studied using AchE staining for light microscopy in conjection with electron microscopy. In the polar region of either type of spindle, the majority of the motor endings are of grape type. In long-capsule spindles plate endings may occur in the spindle pole as well as in the capsular region. Motor endings in the capsular region are mostly of plate type. Plate endings rarely occur in short-capsule spindles. Identified single motor endings were studied by electron microscopy. Intrafusal grape endings are characterized by a smooth post-synaptic membrane. Intrafusal plate endings in the polar region are characterized by junctional folds or gutterings; plate endings in the capsular region show less developed gutterings or indentations. The distribution of intrafusal motor endings was examined along the length of single intrafusal fibres. The long-capsule spindle often receives asymmetrical motor innervation around the capsular region, whereas motor endings in the short-capsule spindle distribute more symmetrically.

Acetylcholinesterase↗

PubMed2025

Zea mays Meiotic Spindle Ultrastructure Reveals Kinetochore-Microtubule Interface and Embedded Membrane Components.

UNLABELLED: Introduction: Spindles are microtubules-based machines whose primary function is to accurately segregate chromosomes in both mitotic and meiotic cell division. The structure of spindles is critical for their function; errors in morphology or attachment to chromosomes lead to aneuploidy, potentially resulting in disease, infertility, and lethality. Electron microscopy studies have yielded fine-detail spindle ultrastructures in many plant and animal species, but no studies have investigated the spindle of Zea mays, a critical crop, and cytogenetic model system. METHODS: Here we use electron tomography (ET), reconstruction, and modeling to obtain three-dimensional, nanometer-resolution of the Z. mays meiotic spindle. Structures such as microtubules, kinetochores, vesicles, membrane channels, and nuclear envelope were modeled through a partial spindle reconstruction, and confirmed using immunostaining and live fluorescence microscopy. RESULTS: ET revealed that maize spindles contain 8-18 kinetochore microtubules (kMTs) per kinetochore, which are approximately 776 nm in diameter and 316 nm in depth. Small ∼37 nm vesicles were identified, as well as larger (∼5 µm long, 800 nm wide) membrane structures with channels that allow spindle microtubules to pass through. These membrane channels stain positively for the ER-marker protein disulfide isomerase. Imaging of prophase meiotic cells revealed a cross-hatch microtubule arrangement in the perinuclear ring on the external surface of the nuclear envelope, which also contained type II nuclear grooves with transnuclear microtubules passing from the nucleus to the cytoplasm. CONCLUSIONS: Z. mays meiotic spindles are similar to animal counterparts with a comparable number of kMTs and pre-spindle transnuclear microtubules but also plant-specific features such as Golgi-derived vesicles to assist cell plate formation, internal ER membrane channels, and a perinuclear microtubule ring that aids spindle assembly. Maize kinetochores have an electron-diffuse ball in cup morphology that is comparable in size to Drosophila kinetochores and larger than mammalian kinetochores. .

Zea mays↗

PubMed1976

Spindle wave synchrony in the somatosensory cortex of the cat.

(1) Spontaneous barbiturate spindles were recorded from the primary and secondary somatosensory cortex. The recordings were concentrated to areas surrounding several reference loci. The recording sites producing the maximal response evoked by stimulation of an exposed nerve in a contralateral limb were used as reference loci. (2) Spindles recorded at various distances from the respective reference loci were cross-correlated to spindles developing simultaneously in the latter. High correlation coefficients, indicating a considerable degree of wave synchrony, were obtained between spindles in the reference locus and spindles recorded a few millimeters from this site. The correlation coefficients decreased with increasing interelectrode distance. A relatively sharp fall in the correlation coefficients was generally found 2-3 mm from the reference locus. Small amounts of sodium pentobarbital, given intravenously at intervals of 5 min, had no effect upon this pattern. (3) The change in the correlation coefficients was followed by a parallel change in the amplitude of the evoked potentials. The iso-correlation lines of spindle wave synchrony and the iso-amplitude lines of the evoked potentials had a similar distribution and extension for each particular reference locus. (4) Lateral spread of spindle waves in the cortex seems to be of minor importance, since a vertical lesion cutting the cortico-cortical fibres did not reduce the wave synchrony of the spindles recorded from either side of the lesion. (5) The majority of the spindles recorded in the close vicinity of a reference locus started simultaneously within +/- 0.1 sec. This pattern changed with increasing distance from the reference locus and 5.6 mm away only a fraction of the spindles started simultaneously. However, within the entire primary somatosensory cortex a small but significant coupling existed between onset of the spindles.

Animals↗

PubMed1979

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

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

Animals↗

PubMed1975

Discharge of spindle afferents from jaw-closing muscles during chewing in alert monkeys.

The discharge of muscle spindle afferents from monkey spindle afferents from monkey jaw-closing muscles was studied during mastication of natural foods by extracellular recording from the fibers or cell bodies of the tract and mesencephalic nucleus of the fifth nerve. In all, 39 muscle afferents were studied. The spindle associated with 18 of the afferents was positively identified by the afferent's response to gentle, localized palpation of either the temporalis or masseter muscle. Discharge patterns were observed during mastication, and in the majority of cases the qualitative passive response characteristics of the spindle afferent were determined. During steady chewing spindle afferent discharge typically paused briefly during the initial rapid upward part of the chewing cycle. Firing generally began as the jaw slowed its upward movement, and firing rates during the slow grinding portion of the upward movement were within the range of 50-80 spikes/s. All spindles exhibited a brisk discharge during the opening movement, typically within the range of 100-150 spikes/s. One-third of the spindle afferents exhibited a brief, high-frequency burst of firing at the very beginning of the opening movement, presumably as a result of stretch applied to a spindle just previously subjects to fusimotor excitation. Although the results of the study make it clear that spindles in jaw-closing muscles are coactived along with the extrafusal muscle fibers, the fusimotor bias does not seem capable of sustaining discharge in the face of rapid shortening of the muscle. Furthermore, the fact that discharge rate during opening, when the jaw-closing motoneurons are quiescent, is much higher than at any part of the closing cycle, when the motoneurons are active, suggests that the muscle spindles cannot provide the primary excitatory drive to the motoneurons.

Acoustic Stimulation↗

PubMed1979

Electron microscopy of the spindle in locally heated cells.

Individual living cells in metaphase were exposed to a steep temperature gradient by placing a microheater near one spindle pole. The cells were then fixed and the spindle was examined by electron microscopy. The structure of the warmer half-spindle differed from the cooler half-spindle in several ways. Kinetochore microtubules were nearly parallel in the warmer half-spindle but were divergent in the cooler. The total length of microtubules in the warmer half-spindle was 52 per cent greater and the number of kinetochore microtubules per kinetochore averaged 16 per cent higher than in the cooler half-spindle. The warmer half-spindle was longer than the cooler. These observations clearly demonstrate a locally enhanced assembly of microtubules in the warmer half-spindle. The electron microscope study makes still clearer the unusual character of chromosome movement in the differentially heated cells: the structure of the warmer half-spindle is hard to distinguish from that in normal cells, yet chromosome movement there is far slower than normal (Nicklas, 1979).

Animals↗

PubMed1975

Responses of muscle spindles in a tortoise.

Responses were recorded from muscle spindles in the extensor digitorum longus muscle of the freshwater tortoise Chelodina longicollis. Most spindles showed no resting discharge unless the muscle was stretched up to a millimetre beyond the minimum body length. The firing rate then increased by 2-5 imp./sec/mm extension. All spindles were relatively insensitive to the rate of change of muscle length. The dynamic index was small and increased in direct proportion to the rate of stretch. Stimulation of selected filaments of the nerve produced a response from the spindle that was attributed to intrafusal contraction. A comparison of the response of the spindle to muscle stretch with the response during simultaneous stretch and stimulation of a motor filament revealed two distinct types of behaviour. In the first, a dynamic effect, motor stimulation greatly increased the response of the spindle to the velocity component of stretch. Other filaments had a static action: an increase in firing of the spindle with the muscle at constant length, but little change in the response to the velocity component of stretch. Stimulation of up to 7 different filaments produced an effect on a spindle. When the motor effects for a given spindle could be clearly characterised, they were all static or all dynamic. Both static and dynamic effects on the one spindle were seen only rarely.

Action Potentials↗

PubMed1976

Experimental manipulation of the amount of tubulin available for assembly into the spindle of dividing sea urchin eggs.

Spindle assembly is studied in the eggs of the sea urchin Lytechinus variegatus by experimentally varying the amount of polymerizable tubulin within the egg. Aliquots of fertilized eggs from the same female are individually pulsed for 1-6 min with 1 X 10(-6) M Colcemid at least 20 min before first nuclear envelope breakdown. This treatment inactivates a portion of the cellular tubulin before the spindle is formed. Upon entering mitosis, treated eggs form functional spindles that are reduced in length and birefringent retardation but not width. With increased exposure to Colcemid, the length and retardation of the metaphase spindles are progressively reduced. Similar results are obtained by pulsing the eggs with Colcemid before fertilization, which demonstrates that the tubulin found in unfertilized sea urchin eggs is later used in spindle formation. Spindles, once assembled, are responsive to increases in the amount of polymerizable tubulin within the cell. Rapid increases in the amount of polymerizable tubulin within a Colcemid-treated cell can be experimentally effected by irradiating the cells with 366-nm light. This treatment photochemically inactivates the Colcemid, thereby freeing the tubulin to polymerize. Upon irradiation, the small prometaphase spindles of Colcemid-treated cells immediately increase in length and retardation. In these irradiated cells, spindle length and retardation increase as much as four times faster than they do during prometaphase for normal spindles. This suggests that the rate of the normal prometaphase increase in retardation and spindle size may be determined by factors other than the maximum rate of tubulin polymerization in the cell.

Animals↗

PubMed1977

A 'late supernormal period' in the recovery of excitability following an action potential in muscle spindle and tendon organ receptors.

1. Discharge patterns have been recorded from five types of stretch receptor; frog muscle spindles, lizard tendon organs, cat soleus tendon organs and primary and secondary endings of cat soleus muscle spindles.2. The fully adapted discharge of each type of receptor is irregular, especially for frog spindles and primary endings of cat spindles as compared with the other three types (the ;regularly firing' receptors). Frog spindles and some cat spindle primary endings would maintain a discharge at very low mean rates (1/sec or less) while the remaining receptors would stop suddenly, as soon as their rate of discharge fell below a critical value characteristic for each individual ending.3. This pattern of discharge suggests that there is a peak in the excitability of ;regularly firing' receptors at a time following a preceding impulse, which corresponds to the intervals between impulses at each particular receptor's slowest rate of maintained firing, and that the excitability subsequently falls again. Primary endings of cat muscle spindles also showed some evidence of such a ;late supernormal period', but frog spindles did not.4. Direct evidence for the ;late supernormal period' was obtained from experiments in which a maintained discharge was restarted by an antidromic action potential in a receptor which had stopped firing, and to which had been applied a stretch just too small to restart the discharge.5. It is shown in an Appendix that a model receptor in which the recovery of excitability following an impulse has a hyperbolic time course, and in which Gaussian distributed noise is superimposed on the generator potential, can have a discharge pattern very closely resembling that of a frog spindle (cf. Buller, 1965).6. After addition of a late supernormal period to the model, its discharge pattern could mimic closely that of a lizard or cat tendon organ, or of a secondary ending of a cat spindle.

Action Potentials↗

PubMed1978

Muscle spindle activity in man during shortening and lengthening contractions.

1. The responses of forty-one muscle spindle endings, mostly in tibialis anterior, were studied in human subjects during voluntary movements of the ankle joint performed at various speeds against different external loads. 2. During slow shortening contractions, the discharge rates of spindle endings in the contracting muscle accelerated after the appearance of the first e.m.g. potentials but before sufficient force had been generated to move the limb. With some endings, the discharge rate decreased during the shortening movement while the e.m.g. activity was increasing, but it always remained higher than before the onset of contraction. If the speed of the movement was increased fewer spindle discharges were seen during muscle shortening. If the shortening contraction was opposed by an external load, so that greater effort was required to perform the same movement, more discharges were seen and the discharge pattern became less modulated by the change in muscle length. 3. These findings indicate that during shortening contractions the fusimotor system is activated together with the skeletomotor system. However, the fusimotor drive is generally insufficient to maintain a significant spindle discharge unless movement is slow or the muscle is shortening against an external load. 4. During lengthening contractions the spindle responses were greater than to passive stretch of similar amplitude and velocity, suggesting heightened fusimotor outflow. 5. During shortening and lengthening contractions small iregularities in the speed of movement occurred commonly. Unintended acceleration of a shortening movement caused a pause in spindle firing, and unintended acceleration of a lengthening movement caused an increased discharge from spindle endings. These spindle responses were associated with corresponding alterations in the discharge pattern of the voluntarily activated motor units at latencies consistent with the operation of spinal reflex mechanisms. 6. It is suggested that a functional role for the fusimotor activation during slow shortening contractions is to provide spindle endings with a background discharge so that they can detect irregularities in the movement and initiate the appropriate reflex correction.

Action Potentials↗

PubMed1979

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

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

Adaptation, Physiological↗

PubMed1992

Effects of vinblastine, podophyllotoxin and nocodazole on mitotic spindles. Implications for the role of microtubule dynamics in mitosis.

Inhibition of mitosis by many drugs that bind to tubulin has been attributed to depolymerization of microtubules. However, we found previously that low concentrations of vinblastine and vincristine blocked mitosis in HeLa cells with little or no depolymerization of spindle microtubules, and spindles appeared morphologically normal or nearly normal. In the present study, we characterized the effects of vinblastine, podophyllotoxin and nocodazole over broad concentration ranges on mitotic spindle organization in HeLa cells. These three drugs are known to affect the dynamics of microtubule polymerization in vitro and to depolymerize microtubules in cells. We wanted to probe further whether mitotic inhibition by these drugs is brought about by a more subtle effect on the microtubules than net microtubule depolymerization. We compared the effects of vinblastine, podophyllotoxin and nocodazole on the organization of spindle microtubules, chromosomes and centrosomes, and on the total mass of microtubules. Spindle organization was examined by immunofluorescence microscopy, and microtubule polymer mass was assayed on isolated cytoskeletons by a quantitative enzyme-linked immunoadsorbence assay for tubulin. As the drug concentration was increased, the organization of mitotic spindles changed in the same way with all three drugs. The changes were associated with mitotic arrest, but were not necessarily accompanied by net microtubule depolymerization. With podophyllotoxin, mitotic arrest was accompanied by microtubule depolymerization. In contrast, with vinblastine and nocodazole, mitotic arrest occurred in the presence of a full complement of spindle microtubules. All three drugs induced a nearly identical rearrangement of spindle microtubules, an increasingly aberrant organization of metaphase chromosomes, and fragmentation of centrosomes. The data suggest that these anti-mitotic drugs block mitosis primarily by inhibiting the dynamics of spindle microtubules rather than by simply depolymerizing the microtubules.

Cell Cycle↗

PubMed1976

Spindle and kinetochore morphology of Dictyostelium discoideum.

The metaphase spindle of haploid Dictyostelium discoideum (n = 7) is 2 mum long. It consists of some 20 microtubules which seem continuous between the spindle pole bodies and there are about 20 chromosomal microtubules at each end of the spindle. During anaphase the central spindle elongates and the chromosomal microtubules shorten. The spindle length and structure at this stage suggests that lengthening is caused by elongation as well as parallel sliding of the nonchromosomal microtubules. The nuclear envelope remains mostly intact during mitosis, and nuclear separation through medial constriction takes place when the spindle is 6 mum long. Cytokinesis occurs when the spindle is 10 mum long. At that time the kinetochores double in size. During interphase, the spindle pole body separates from the nucleus to a distance of 0.7 mum, and it returns at the onset of the next prophase when it becomes functionally double, thereby starting the formation of a central spindle. When comparing mitosis in the cellular slime molds Polysphondylium violaceum and D. discoideum, several similarities and some differences are apparent.

Cell Division↗