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1
Wordeman, L., K. L. McDonald, and W. Z. Cande. "The distribution of cytoplasmic microtubules throughout the cell cycle of the centric diatom Stephanopyxis turris: their role in nuclear migration and positioning the mitotic spindle during cytokinesis." Journal of Cell Biology 102, no. 5 (May 1, 1986): 1688–98. http://dx.doi.org/10.1083/jcb.102.5.1688.
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The cell cycle of the marine centric diatom Stephanopyxis turris consists of a series of spatially and temporally well-ordered events. We have used immunofluorescence microscopy to examine the role of cytoplasmic microtubules in these events. At interphase, microtubules radiate out from the microtubule-organizing center, forming a network around the nucleus and extending much of the length and breadth of the cell. As the cell enters mitosis, this network breaks down and a highly ordered mitotic spindle is formed. Peripheral microtubule bundles radiate out from each spindle pole and swing out and away from the central spindle during anaphase. Treatment of synchronized cells with 2.5 X 10(-8) M Nocodazole reversibly inhibited nuclear migration concurrent with the disappearance of the extensive cytoplasmic microtubule arrays associated with migrating nuclei. Microtubule arrays and mitotic spindles that reformed after the drug was washed out appeared normal. In contrast, cells treated with 5.0 X 10(-8) M Nocodazole were not able to complete nuclear migration after the drug was washed out and the mitotic spindles that formed were multipolar. Normal and multipolar spindles that were displaced toward one end of the cell by the drug treatment had no effect on the plane of division during cytokinesis. The cleavage furrow always bisected the cell regardless of the position of the mitotic spindle, resulting in binucleate/anucleate daughter cells. This suggests that in S. turris, unlike animal cells, the location of the plane of division is cortically determined before mitosis.
2
Whitehead, C. M., and J. B. Rattner. "Expanding the role of HsEg5 within the mitotic and post-mitotic phases of the cell cycle." Journal of Cell Science 111, no. 17 (September 1, 1998): 2551–61. http://dx.doi.org/10.1242/jcs.111.17.2551.
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The BimC family of kinesin like proteins are involved in spindle dynamics in a wide variety of organisms. The human member of this family, HsEg5, has been implicated in centrosome separation during prophase/prometaphase and in the organization of in vitro mitotic asters. HsEg5 displays a complex distribution during mitosis, associating with the centrosomes, spindle microtubules, specific regions of the intracellular bridge and a microtubule bundle that forms in association with the post-mitotic migration of the centrosome. In an effort to determine the function of HsEg5 during late mitotic events and refine its proposed function during early mitotic centrosome separation, we microinjected antibodies specific to HsEg5 into HeLa cells during various stages of mitosis. In the presence of HsEg5 antibodies we find that the microtubule arrays responsible for both pre- and post-mitotic centrosome movement never form. Similarly, the microtubule bundle within the intracellular bridge becomes prematurely altered following karyokinesis resulting in the loss of the microtubule array at either end of the bridge. In addition, some peri-centrosomal material at the spindle poles becomes fragmented and the distribution of the spindle protein NuMA becomes more concentrated at the minus ends of the spindle microtubules. Our study also provides direct evidence that there is a link between post-mitotic centrosome migration and Golgi complex positioning and reformation following mitosis. We conclude that HsEg5 plays a recurrent role in establishing and/or determining the stability of specific microtubule arrays that form during cell division and that this role may encompass the ability of HsEg5 to influence the distribution of other protein components associated with cell division
3
de Saint Phalle, Brigitte, and William Sullivan. "Spindle Assembly and Mitosis without Centrosomes in Parthenogenetic Sciara Embryos." Journal of Cell Biology 141, no. 6 (June 15, 1998): 1383–91. http://dx.doi.org/10.1083/jcb.141.6.1383.
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In Sciara, unfertilized embryos initiate parthenogenetic development without centrosomes. By comparing these embryos with normal fertilized embryos, spindle assembly and other microtubule-based events can be examined in the presence and absence of centrosomes. In both cases, functional mitotic spindles are formed that successfully proceed through anaphase and telophase, forming two daughter nuclei separated by a midbody. The spindles assembled without centrosomes are anastral, and it is likely that their microtubules are nucleated at or near the chromosomes. These spindles undergo anaphase B and successfully segregate sister chromosomes. However, without centrosomes the distance between the daughter nuclei in the next interphase is greatly reduced. This suggests that centrosomes are required to maintain nuclear spacing during the telophase to interphase transition. As in Drosophila, the initial embryonic divisions of Sciara are synchronous and syncytial. The nuclei in fertilized centrosome-bearing embryos maintain an even distribution as they divide and migrate to the cortex. In contrast, as division proceeds in embryos lacking centrosomes, nuclei collide and form large irregularly shaped nuclear clusters. These nuclei are not evenly distributed and never successfully migrate to the cortex. This phenotype is probably a direct result of a failure to form astral microtubules in parthenogenetic embryos lacking centrosomes. These results indicate that the primary function of centrosomes is to provide astral microtubules for proper nuclear spacing and migration during the syncytial divisions. Fertilized Sciara embryos produce a large population of centrosomes not associated with nuclei. These free centrosomes do not form spindles or migrate to the cortex and replicate at a significantly reduced rate. This suggests that the centrosome must maintain a proper association with the nucleus for migration and normal replication to occur.
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Figueroa, Ricardo A., Santhosh Gudise, and Einar Hallberg. "Microtubule-associated nuclear envelope proteins in interphase and mitosis." Biochemical Society Transactions 39, no. 6 (November 21, 2011): 1786–89. http://dx.doi.org/10.1042/bst20110680.
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The LINC (linker of nucleoskeleton and cytoskeleton) complex forms a transcisternal bridge across the NE (nuclear envelope) that connects the cytoskeleton with the nuclear interior. This enables some proteins of the NE to communicate with the centrosome and the microtubule cytoskeleton. The position of the centrosome relative to the NE is of vital importance for many cell functions, such as cell migration and division, and centrosomal dislocation is a frequent phenotype in laminopathic disorders. Also in mitosis, a small group of transmembrane NE proteins associate with microtubules when they concentrate in a specific membrane domain associated with the mitotic spindle. The present review discusses structural and functional aspects of microtubule association with NE proteins and how this association may be maintained over the cell cycle.
5
Barton, R., and K. Gull. "Variation in cytoplasmic microtubule organization and spindle length between the two forms of the dimorphic fungus Candida albicans." Journal of Cell Science 91, no. 2 (October 1, 1988): 211–20. http://dx.doi.org/10.1242/jcs.91.2.211.
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Candida albicans is a dimorphic fungus capable of growing as a budding yeast and as a filamentous hypha. We have used the technique of immunofluorescence to study the changes in the microtubule cytoskeleton during the cell cycle in both growth forms. This approach has revealed the presence of an extensive system of microtubules, including cytoplasmic microtubules and a rod-like intranuclear spindle. We have provided a complete description of the arrangement of cytoplasmic and spindle microtubules at each phase of the yeast cell cycle. A novel and characteristic feature of the yeast phase of Candida is the presence of an array of short microtubules at the neck of the doublet cell. This neck-associated array (NAA), is apparently organized independently of the main microtubule-organizing centre, the spindle pole bodies, at late anaphase. Analysis of microtubule patterns in the hyphal state reveals that the general arrangements of microtubules are similar to those seen in the yeast phase. These patterns suggest a role for the cytoplasmic microtubules in the nuclear migration that occurs during hyphal growth. A major finding is that the mitotic spindle in hyphae is considerably longer (12–20 microns) than the spindle in yeast cells (7–8 microns). This may reflect the role of the hyphal mitotic spindle not only in nuclear division but also in the positioning of the daughter nuclei at the centres of hyphal compartments.
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Mana-Capelli, Sebastian, Janel R. McLean, Chun-Ti Chen, Kathleen L. Gould, and Dannel McCollum. "The kinesin-14 Klp2 is negatively regulated by the SIN for proper spindle elongation and telophase nuclear positioning." Molecular Biology of the Cell 23, no. 23 (December 2012): 4592–600. http://dx.doi.org/10.1091/mbc.e12-07-0532.
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In Schizosaccharomyces pombe, a late mitotic kinase pathway called the septation initiation network (SIN) triggers cytokinesis. Here we show that the SIN is also involved in regulating anaphase spindle elongation and telophase nuclear positioning via inhibition of Klp2, a minus end–directed kinesin-14. Klp2 is known to localize to microtubules (MTs) and have roles in interphase nuclear positioning, mitotic chromosome alignment, and nuclear migration during karyogamy (nuclear fusion during mating). We observe SIN-dependent disappearance of Klp2 from MTs in anaphase, and we find that this is mediated by direct phosphorylation of Klp2 by the SIN kinase Sid2, which abrogates loading of Klp2 onto MTs by inhibiting its interaction with Mal3 (EB1 homologue). Disruption of Klp2 MT localization is required for efficient anaphase spindle elongation. Furthermore, when cytokinesis is delayed, SIN inhibition of Klp2 acts in concert with microtubules emanating from the equatorial microtubule-organizing center to position the nuclei away from the cell division site. These results reveal novel functions of the SIN in regulating the MT cytoskeleton and suggest that the SIN may have broader functions in regulating cellular organization in late mitosis than previously realized.
7
Sutradhar, Sabyasachi, Vikas Yadav, Shreyas Sridhar, Lakshmi Sreekumar, Dibyendu Bhattacharyya, Santanu Kumar Ghosh, Raja Paul, and Kaustuv Sanyal. "A comprehensive model to predict mitotic division in budding yeasts." Molecular Biology of the Cell 26, no. 22 (November 5, 2015): 3954–65. http://dx.doi.org/10.1091/mbc.e15-04-0236.
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High-fidelity chromosome segregation during cell division depends on a series of concerted interdependent interactions. Using a systems biology approach, we built a robust minimal computational model to comprehend mitotic events in dividing budding yeasts of two major phyla: Ascomycota and Basidiomycota. This model accurately reproduces experimental observations related to spindle alignment, nuclear migration, and microtubule (MT) dynamics during cell division in these yeasts. The model converges to the conclusion that biased nucleation of cytoplasmic microtubules (cMTs) is essential for directional nuclear migration. Two distinct pathways, based on the population of cMTs and cortical dyneins, differentiate nuclear migration and spindle orientation in these two phyla. In addition, the model accurately predicts the contribution of specific classes of MTs in chromosome segregation. Thus we present a model that offers a wider applicability to simulate the effects of perturbation of an event on the concerted process of the mitotic cell division.
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Renzaglia, Karen Sue, Angel R. Maden, Jeffrey G. Duckett, and Dean P. Whittier. "Monoplastidy in spermatogenesis of Lycopodium obscurum." Canadian Journal of Botany 72, no. 10 (October 1, 1994): 1436–44. http://dx.doi.org/10.1139/b94-177.
Full textAbstract:
Unlike Lycopodium laterale, which is polyplastidic during spermatogenesis, Lycopodium obscurum exhibits monoplastidy beginning in the early proliferative stages of antheridial development. Previous cell generations are polyplastidic and plastid fusion involving connective cylinders establishes the monoplastidic condition. Plastid and nuclear divisions are coordinated in L. obscurum with the plastids positioned at opposite poles prior to spindle development. Unlike monoplastidic cell divisions with morphogenetic plastid migration and polarity in other lycophytes, mosses, and hornworts, however, the spindles in L. obscurum do not originate from the plastid envelopes but from endoplasmic reticulum positioned against the plastid. In the final divisions, spindle microtubules emanate from structurally defined microtubule organizing centers that develop between the plastids and nucleus. Preceding the appearance of centrioles in the spermatid mother cell, the centrosomes comprise electron-dense granular matrices with associated vesicles and endoplasmic reticulum. Among archegoniate microtubule organizing centers, the discrete acentriolar centrosomes of Lycopodium most closely resemble the microtubule organizing centers in moss spore development and the polar organizer of liverwort mitosis. Key words: annulate lamellae, centrosome, Lycopodium, microtubule organizing center, monoplastidy, plastid dividing ring.
9
Brown, R. C., and B. E. Lemmon. "Morphogenetic plastid migration and spindle dynamics in simple land plants." Proceedings, annual meeting, Electron Microscopy Society of America 47 (August 6, 1989): 764–65. http://dx.doi.org/10.1017/s0424820100155797.
Full textAbstract:
In many of the simple land plants mitosis and/or meiosis occurs in cells that contain a single plastid. Preparation for division in these monoplastidic cells is especially obvious because of predictive migration and division of the single plastid. The plastids serve as foci for organization of spindle microtubules (Mts) resulting in infallible coordination of plastid and nuclear division. Thus unlike most plant cells where a distinct MTOC is difficult to distinguish, monoplastidic cells provide a system in which the material responsible for nucleating Mts is closely associated with the plastid envelope and can be followed throughout the cell cycle. The intimate association of plastids with spindle poles has suggested the concept of plastid polarity which holds that the behavior of plastids parallels that of animal centrosomes. This concept is supported by recent studies using correlated methods of modern botanical microscopy.Mitosis was studied in the basal meristem of hornwort sporophytes and in root tips of Isoetes and Selaginella.
10
Gholkar, Ankur A., Keith Cheung, Kevin J. Williams, Yu-Chen Lo, Shadia A. Hamideh, Chelsea Nnebe, Cindy Khuu, Steven J. Bensinger, and Jorge Z. Torres. "Fatostatin Inhibits Cancer Cell Proliferation by Affecting Mitotic Microtubule Spindle Assembly and Cell Division." Journal of Biological Chemistry 291, no. 33 (July 4, 2016): 17001–8. http://dx.doi.org/10.1074/jbc.c116.737346.
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The sterol regulatory element-binding protein (SREBP) transcription factors have become attractive targets for pharmacological inhibition in the treatment of metabolic diseases and cancer. SREBPs are critical for the production and metabolism of lipids and cholesterol, which are essential for cellular homeostasis and cell proliferation. Fatostatin was recently discovered as a specific inhibitor of SREBP cleavage-activating protein (SCAP), which is required for SREBP activation. Fatostatin possesses antitumor properties including the inhibition of cancer cell proliferation, invasion, and migration, and it arrests cancer cells in G2/M phase. Although Fatostatin has been viewed as an antitumor agent due to its inhibition of SREBP and its effect on lipid metabolism, we show that Fatostatin's anticancer properties can also be attributed to its inhibition of cell division. We analyzed the effect of SREBP activity inhibitors including Fatostatin, PF-429242, and Betulin on the cell cycle and determined that only Fatostatin possessed antimitotic properties. Fatostatin inhibited tubulin polymerization, arrested cells in mitosis, activated the spindle assembly checkpoint, and triggered mitotic catastrophe and reduced cell viability. Thus Fatostatin's ability to inhibit SREBP activity and cell division could prove beneficial in treating aggressive types of cancers such as glioblastomas that have elevated lipid metabolism and fast proliferation rates and often develop resistance to current anticancer therapies.
11
Smith, Joshua J., J. Sebastian Yakisich, Geoffrey M. Kapler, Eric S. Cole, and Daniel P. Romero. "A β-Tubulin Mutation Selectively Uncouples Nuclear Division and Cytokinesis in Tetrahymena thermophila." Eukaryotic Cell 3, no. 5 (October 2004): 1217–26. http://dx.doi.org/10.1128/ec.3.5.1217-1226.2004.
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ABSTRACT The ciliated protozoan Tetrahymena thermophila contains two distinct nuclei within a single cell—the mitotic micronucleus and the amitotic macronucleus. Although microtubules are required for proper division of both nuclei, macronuclear chromosomes lack centromeres and the role of microtubules in macronuclear division has not been established. Here we describe nuclear division defects in cells expressing a mutant β-tubulin allele that confers hypersensitivity to the microtubule-stabilizing drug paclitaxel. Macronuclear division is profoundly affected by the btu1-1 (K350M) mutation, producing cells with widely variable DNA contents, including cells that lack macronuclei entirely. Protein expressed by the btu1-1 allele is dominant over wild-type protein expressed by the BTU2 locus. Normal macronuclear division is restored when the btu1-1 allele is inactivated by targeted disruption or expressed as a truncated protein. Immunofluorescence studies reveal elongated microtubular structures that surround macronuclei that fail to migrate to the cleavage furrows. In contrast, other cytoplasmic microtubule-dependent processes, such as cytokinesis, cortical patterning, and oral apparatus assembly, appear to be unaffected in the mutant. Micronuclear division is also perturbed in the K350M mutant, producing nuclei with elongated early-anaphase spindle configurations that persist well after the initiation of cytokinesis. The K350M mutation affects tubulin dynamics, as the macronuclear division defect is exacerbated by three treatments that promote microtubule polymerization: (i) elevated temperatures, (ii) sublethal concentrations of paclitaxel, and (iii) high concentrations of dimethyl sulfoxide. Inhibition of phosphatidylinositol 3-kinase (PI 3-kinase) with 3-methyladenine or wortmannin also induces amacronucleate cell formation in a btu1-1-dependent manner. Conversely, the myosin light chain kinase inhibitor ML-7 has no effect on nuclear division in the btu1-1 mutant strain. These findings provide new insights into microtubule dynamics and link the evolutionarily conserved PI 3-kinase signaling pathway to nuclear migration and/or division in Tetrahymena.
12
McEwen, B. F., C. E. Hsieh, R. M. Barnard, and C. L. Rieder. "Using High Pressure Freezing and Freeze Substitution to Investigate Kinetochore Ultrastructure in Vetebrate Somatic Cells." Microscopy and Microanalysis 5, S2 (August 1999): 436–37. http://dx.doi.org/10.1017/s1431927600015506.
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In vertebrate somatic cells, sister kinetochores attach chromosomes to the mitotic spindle by capturing microtubules (Mts) emanating from opposite spindle poles. Kinetochores are also required for chromosome alignment at the spindle equator, stabilizing kinetochore Mts, cell cycle control of anaphase onset, and poleward migration of sister chromatids during anaphase. Thus kinetochores play a critical role in the distribution of genetic information to daughter cells during cell division. Understanding the molecular mechanisms behind kinetochore function requires knowing how the molecular components are arranged relative to each other, and relative to the kinetochore Mts. Currently, however, our knowledge of kinetochore composition is incomplete, and until recently our knowledge of kinetochore ultrastructure was limited to the familiar trilamellar model derived from conventionally fixed and dehydrated specimens(Figures 1a&b).Highly fibrous structures, such as the kinetochore, are particularly vulnerable to distortions caused by the chemical fixation and dehydration methods used in conventional specimen preparations. Such distortions can be largely avoided by using high pressure freezing and freeze substitution (HPF/FS).
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Gratao, A. A., A. Beck, M. Reichenbach, H. D. Reichenbach, E. Wolf, and F. A. Habermann. "104 3-D VISUALIZATION OF THE FIRST CLEAVAGE AND ABERRATIONS OF BOVINE ZYGOTES BY CONFOCAL LASER SCANNING MICROSCOPY." Reproduction, Fertility and Development 25, no. 1 (2013): 199. http://dx.doi.org/10.1071/rdv25n1ab104.
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A high proportion of bovine oocytes fertilized in vitro fail to develop beyond the first 4 cleavage cycles. The first mitotic division of the zygote and proper segregation of chromosomes and cytoplasmic components seems to be a particularly delicate task. Notably, zygotes cleaving with a delay of only a few hours seem to have a very low chance of developing to the blastocyst stage. But what exactly goes wrong, how often, and why? To answer such questions we have to visualize in greater detail basic structures and processes such as the sperm aster, DNA replication, migration and apposition of the 2 pronuclei, synchronous chromosome condensation and breakdown of the nuclear envelopes, assembly of the first mitotic spindle and chromosome congression, anaphase, and cytokinesis. Oocytes fertilized in vitro were fixed at different time points around the first cleavage and stained for DNA, Ser10-phosphorylated histone H3, microtubules, and microfilaments. Zygotes were imaged in toto by recording confocal serial sections at 1-µm intervals using a 40× objective (NA = 1.3). Details were recorded with high spatial sampling densities (pixel size 50 × 50 nm, z-step size of 200 nm) close to the Nyquist criterion and restored by maximum likelihood estimation deconvolution using the real point spread function. We present a series of 3-D confocal images captured at different stages of the first cleavage. The images reveal new insights into the formation, structure, and function of the first mitotic spindle and the occurrence of spindle aberrations, irregular chromosome segregation, and abnormal cytokinesis. The microscopic findings guide us to candidate proteins for localization analyses and functional studies based on 3-D fluorescence live-cell imaging of zygotes and early embryos. This work is supported by the Deutsche Forschungsgemeinschaft (DFG FOR 1041).
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Tanaka, K., and T. Kanbe. "Mitosis in the fission yeast Schizosaccharomyces pombe as revealed by freeze-substitution electron microscopy." Journal of Cell Science 80, no. 1 (February 1, 1986): 253–68. http://dx.doi.org/10.1242/jcs.80.1.253.
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Nuclear division in Schizosaccharomyces pombe has been studied in transmission electron micrographs of sections of cells fixed by a method of freeze-substitution. We have found cytoplasmic microtubules in the vicinity of the spindle pole bodies and two kinds of microtubules, short discontinuous ones and long, parallel ones in the intranuclear mitotic spindle. For most of the time taken by nuclear division the spindle pole bodies face each other squarely across the nuclear space but early in mitosis they briefly appear twisted out of alignment with each other, thereby imparting a sigmoidal shape to the bundle of spindle microtubules extending between them. This configuration is interpreted as indicating active participation of the spindle in the initial elongation of the dividing nucleus. It is proposed that mitosis is accompanied by the shortening of chromosomal microtubules simultaneously with the elongation of the central pole-to-pole bundle of microtubules of the intranuclear spindle. Daughter nuclei are separated by the sliding apart of interdigitating microtubules of the spindle at telophase. Some of the latter bear dense knobs at their ends.
15
Bouguenina, Habib, Danièle Salaun, Aurélie Mangon, Leslie Muller, Emilie Baudelet, Luc Camoin, Taro Tachibana, et al. "EB1-binding–myomegalin protein complex promotes centrosomal microtubules functions." Proceedings of the National Academy of Sciences 114, no. 50 (November 21, 2017): E10687—E10696. http://dx.doi.org/10.1073/pnas.1705682114.
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Control of microtubule dynamics underlies several fundamental processes such as cell polarity, cell division, and cell motility. To gain insights into the mechanisms that control microtubule dynamics during cell motility, we investigated the interactome of the microtubule plus-end–binding protein end-binding 1 (EB1). Via molecular mapping and cross-linking mass spectrometry we identified and characterized a large complex associating a specific isoform of myomegalin termed “SMYLE” (for short myomegalin-like EB1 binding protein), the PKA scaffolding protein AKAP9, and the pericentrosomal protein CDK5RAP2. SMYLE was associated through an evolutionarily conserved N-terminal domain with AKAP9, which in turn was anchored at the centrosome via CDK5RAP2. SMYLE connected the pericentrosomal complex to the microtubule-nucleating complex (γ-TuRC) via Galectin-3–binding protein. SMYLE associated with nascent centrosomal microtubules to promote microtubule assembly and acetylation. Disruption of SMYLE interaction with EB1 or AKAP9 prevented microtubule nucleation and their stabilization at the leading edge of migrating cells. In addition, SMYLE depletion led to defective astral microtubules and abnormal orientation of the mitotic spindle and triggered G1 cell-cycle arrest, which might be due to defective centrosome integrity. As a consequence, SMYLE loss of function had a profound impact on tumor cell motility and proliferation, suggesting that SMYLE might be an important player in tumor progression.
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TERRY, R. S., A. M. DUNN, and J. E. SMITH. "Segregation of a microsporidian parasite during host cell mitosis." Parasitology 118, no. 1 (January 1999): 43–48. http://dx.doi.org/10.1017/s0031182098003540.
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We investigated the segregation of an intracellular microsporidian parasite during host cell division. A time-course experiment was carried out to examine the distribution of parasites relative to host chromosomal DNA via light and electron microscopy. Fluorescent light microscopy and EM studies showed that the parasite lay in the perinuclear zone of the host cell during interphase and segregated to daughter cells at mitosis. At metaphase, the parasite was frequently closely associated with host microtubules and mitochondria. Electron-dense bridges were observed between the parasites and the host microtubules and also between host mitochondria and microtubules. The study suggests that both the parasite and the host cell organelles segregate in association with spindle microtubules.
17
Berlin, V., C. A. Styles, and G. R. Fink. "BIK1, a protein required for microtubule function during mating and mitosis in Saccharomyces cerevisiae, colocalizes with tubulin." Journal of Cell Biology 111, no. 6 (December 1, 1990): 2573–86. http://dx.doi.org/10.1083/jcb.111.6.2573.
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BIK1 function is required for nuclear fusion, chromosome disjunction, and nuclear segregation during mitosis. The BIK1 protein colocalizes with tubulin to the spindle pole body and mitotic spindle. Synthetic lethality observed in double mutant strains containing a mutation in the BIK1 gene and in the gene for alpha- or beta-tubulin is consistent with a physical interaction between BIK1 and tubulin. Furthermore, over- or underexpression of BIK1 causes aberrant microtubule assembly and function, bik1 null mutants are viable but contain very short or undetectable cytoplasmic microtubules. Spindle formation often occurs strictly within the mother cell, probably accounting for the many multinucleate and anucleate bik1 cells. Elevated levels of chromosome loss in bik1 cells are indicative of defective spindle function. Nuclear fusion is blocked in bik1 x bik1 zygotes, which have truncated cytoplasmic microtubules. Cells overexpressing BIK1 initially have abnormally short or nonexistent spindle microtubules and long cytoplasmic microtubules. Subsequently, cells lose all microtubule structures, coincident with the arrest of division. Based on these results, we propose that BIK1 is required stoichiometrically for the formation or stabilization of microtubules during mitosis and for spindle pole body fusion during conjugation.
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Hagan, I. M., and J. S. Hyams. "The use of cell division cycle mutants to investigate the control of microtubule distribution in the fission yeast Schizosaccharomyces pombe." Journal of Cell Science 89, no. 3 (March 1, 1988): 343–57. http://dx.doi.org/10.1242/jcs.89.3.343.
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We have characterized the changes in microtubule organization that occur through the cell division cycle of the fission yeast Schizosaccharomyces pombe by indirect immunofluorescence microscopy. During interphase, groups of cytoplasmic microtubules, independent of the spindle pole body (SPB), form an array extending between the cell tips. These microtubules are involved in positioning the nucleus at the cell equator and in the establishment of cell polarity. At mitosis, the interphase array disappears and is replaced by an intranuclear spindle extending between the now duplicated SPBs. Elongation of the spindle sees the appearance of astral microtubules emanating from the cytoplasmic face of the SPBs. These persist until the end of anaphase whereupon the spindle microtubules depolymerize and two microtubule organizing centres (MTOCs) at the cell equator re-establish the interphase array. We have used the unique properties of various cell division cycle mutants to investigate further the function of these different microtubule arrays and their temporal and positional control.
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Friedman, D. B., H. A. Sundberg, E. Y. Huang, and T. N. Davis. "The 110-kD spindle pole body component of Saccharomyces cerevisiae is a phosphoprotein that is modified in a cell cycle-dependent manner." Journal of Cell Biology 132, no. 5 (March 1, 1996): 903–14. http://dx.doi.org/10.1083/jcb.132.5.903.
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Spc110p (Nuf1p) is an essential component of the yeast microtubule organizing center, or spindle pole body (SPB). Asynchronous wild-type cultures contain two electrophoretically distinct isoforms of Spc110p as detected by Western blot analysis, suggesting that Spc110p is modified in vivo. Both isoforms incorporate 32Pi in vivo, suggesting that Spc110p is post-translationally modified by phosphorylation. The slower-migrating 120-kD Spc110p isoform after incubation is converted to the faster-migrating 112-kD isoform after incubation with protein phosphatase PP2A, and specific PP2A inhibitors block this conversion. Thus, additional phosphorylation of Spc110p at serine and/or threonine residues gives rise to the slower-migrating 120-kD isoform. The 120-kD isoform predominates in cells arrested in mitosis by the addition of nocodazole. However, the 120-kD isoform is not detectable in cells grown to stationary phase (G0) or in cells arrested in G1 by the addition of alpha-factor. Temperature-sensitive cell division cycle (cdc) mutations demonstrate that the presence of the 120-kD isoform correlates with mitotic spindle formation but not with SPB duplication. In a synchronous wild-type population, the additional serine/threonine phosphorylation that gives rise to the 120-kD isoform appears as cells are forming the mitotic spindle and diminishes as cells enter anaphase. None of several sequences similar to the consensus for phosphorylation by the Cdc28p (cdc2p34) kinase is important for these mitosis-specific phosphorylations or for function. Carboxy-terminal Spc110p truncations lacking the calmodulin binding site can support growth and are also phosphorylated in a cell cycle-specific manner. Further truncation of the Spc110p carboxy terminus results in mutant proteins that are unable to support growth and now migrate as single species. Collectively, these results provide the first evidence of a structural component of the SPB that is phosphorylated during spindle formation and dephosphorylated as cells enter anaphase.
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Raemaekers, Tim, Katharina Ribbeck, Joël Beaudouin, Wim Annaert, Mark Van Camp, Ingrid Stockmans, Nico Smets, Roger Bouillon, Jan Ellenberg, and Geert Carmeliet. "NuSAP, a novel microtubule-associated protein involved in mitotic spindle organization." Journal of Cell Biology 162, no. 6 (September 8, 2003): 1017–29. http://dx.doi.org/10.1083/jcb.200302129.
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Here, we report on the identification of nucleolar spindle–associated protein (NuSAP), a novel 55-kD vertebrate protein with selective expression in proliferating cells. Its mRNA and protein levels peak at the transition of G2 to mitosis and abruptly decline after cell division. Microscopic analysis of both fixed and live mammalian cells showed that NuSAP is primarily nucleolar in interphase, and localizes prominently to central spindle microtubules during mitosis. Direct interaction of NuSAP with microtubules was demonstrated in vitro. Overexpression of NuSAP caused profound bundling of cytoplasmic microtubules in interphase cells, and this relied on a COOH-terminal microtubule-binding domain. In contrast, depletion of NuSAP by RNA interference resulted in aberrant mitotic spindles, defective chromosome segregation, and cytokinesis. In addition, many NuSAP-depleted interphase cells had deformed nuclei. Both overexpression and knockdown of NuSAP impaired cell proliferation. These results suggest a crucial role for NuSAP in spindle microtubule organization.
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McHugh, Toni, Agata A. Gluszek, and Julie P. I. Welburn. "Microtubule end tethering of a processive kinesin-8 motor Kif18b is required for spindle positioning." Journal of Cell Biology 217, no. 7 (April 16, 2018): 2403–16. http://dx.doi.org/10.1083/jcb.201705209.
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Mitotic spindle positioning specifies the plane of cell division during anaphase. Spindle orientation and positioning are therefore critical to ensure symmetric division in mitosis and asymmetric division during development. The control of astral microtubule length plays an essential role in positioning the spindle. In this study, using gene knockout, we show that the kinesin-8 Kif18b controls microtubule length to center the mitotic spindle at metaphase. Using in vitro reconstitution, we reveal that Kif18b is a highly processive plus end–directed motor that uses a C-terminal nonmotor microtubule-binding region to accumulate at growing microtubule plus ends. This region is regulated by phosphorylation to spatially control Kif18b accumulation at plus ends and is essential for Kif18b-dependent spindle positioning and regulation of microtubule length. Finally, we demonstrate that Kif18b shortens microtubules by increasing the catastrophe rate of dynamic microtubules. Overall, our work reveals that Kif18b uses its motile properties to reach microtubule ends, where it regulates astral microtubule length to ensure spindle centering.
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Rizzelli, Francesca, Maria Grazia Malabarba, Sara Sigismund, and Marina Mapelli. "The crosstalk between microtubules, actin and membranes shapes cell division." Open Biology 10, no. 3 (March 2020): 190314. http://dx.doi.org/10.1098/rsob.190314.
Full textAbstract:
Mitotic progression is orchestrated by morphological and mechanical changes promoted by the coordinated activities of the microtubule (MT) cytoskeleton, the actin cytoskeleton and the plasma membrane (PM). MTs assemble the mitotic spindle, which assists sister chromatid separation, and contact the rigid and tensile actomyosin cortex rounded-up underneath the PM. Here, we highlight the dynamic crosstalk between MTs, actin and cell membranes during mitosis, and discuss the molecular connections between them. We also summarize recent views on how MT traction forces, the actomyosin cortex and membrane trafficking contribute to spindle positioning in isolated cells in culture and in epithelial sheets. Finally, we describe the emerging role of membrane trafficking in synchronizing actomyosin tension and cell shape changes with cell–substrate adhesion, cell–cell contacts and extracellular signalling events regulating proliferation.
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Sawicki, Mark P., Ankur A. Gholkar, and Jorge Z. Torres. "Menin Associates With the Mitotic Spindle and Is Important for Cell Division." Endocrinology 160, no. 8 (June 18, 2019): 1926–36. http://dx.doi.org/10.1210/en.2019-00274.
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Abstract Menin is the protein mutated in patients with multiple endocrine neoplasia type 1 (MEN1) syndrome and their corresponding sporadic tumor counterparts. We have found that menin functions in promoting proper cell division. Here, we show that menin localizes to the mitotic spindle poles and the mitotic spindle during early mitosis and to the intercellular bridge microtubules during cytokinesis in HeLa cells. In our study, menin depletion led to defects in spindle assembly and chromosome congression during early mitosis, lagging chromosomes during anaphase, defective cytokinesis, multinucleated interphase cells, and cell death. In addition, pharmacological inhibition of the menin-MLL1 interaction also led to similar cell division defects. These results indicate that menin and the menin-MLL1 interaction are important for proper cell division. These results highlight a function for menin in cell division and aid our understanding of how mutation and misregulation of menin promotes tumorigenesis.
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TRAAS, J. A., S. BURGAIN, and R. DUMAS DE VAULX. "The Organization of the Cytoskeleton During Meiosis in Eggplant (Solanum Melongena (L.)): Microtubules and F-Actin are Both Necessary for Coordinated Meiotic Division." Journal of Cell Science 92, no. 4 (April 1, 1989): 541–50. http://dx.doi.org/10.1242/jcs.92.4.541.
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Address for reprints Because two division planes form at right angles, male meiosis in higher plants provides striking examples of both division control and spatial programming. To investigate these processes we have stained microtubules and actin filaments during male meiosis in the eggplant. Our results indicate the following. (1) That microtubules and their nucleation sites are involved in the establishment of polarity; this is supported by our observation that the drug CIPC affects spindle polarity.(2) That actin microfilaments are involved in spindle formation and integrity, but not in the establishment of polarity: cytochalasin B and D affect the organization of the spindle microtubules, but not their polarized distribution.(3) That microtubules radiating from the daughter nuclei at the cell poles during interkinesis probably establish the future division plane by concentrating actin in that plane (cf. the proposed role of asters in positioning the contractile ring in animal cells).(4) That this concentration of F-actin in the division plane may be involved in preparing the cytoplasm for cytokinesis and in memorizing the division plane (much as the preprophase band observed in polarized tissues does).(5) That phragmoplast formation is a two-step process. No phragmoplast forms after metaphase I, but a four-way phragmoplast forms after metaphase II, indicating that mitosis and cytokinesis are not obligatorily coupled. These studies demonstrate that actin and microtubules are jointly involved in the spatial coordination of the division process.
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Vanneste, David, Vanessa Ferreira, and Isabelle Vernos. "Chromokinesins: localization-dependent functions and regulation during cell division." Biochemical Society Transactions 39, no. 5 (September 21, 2011): 1154–60. http://dx.doi.org/10.1042/bst0391154.
Full textAbstract:
The bipolar spindle is a highly dynamic structure that assembles transiently around the chromosomes and provides the mechanical support and the forces required for chromosome segregation. Spindle assembly and chromosome movements rely on the regulation of microtubule dynamics and a fine balance of forces exerted by various molecular motors. Chromosomes are themselves central players in spindle assembly. They generate a RanGTP gradient that triggers microtubule nucleation and stabilization locally and they interact dynamically with the microtubules through motors targeted to the chromatin. We have previously identified and characterized two of these so-called chromokinesins: Xkid (kinesin 10) and Xklp1 (kinesin 4). More recently, we found that Hklp2/kif15 (kinesin 12) is targeted to the chromosomes through an interaction with Ki-67 in human cells and is therefore a novel chromokinesin. Hklp2 also associates with the microtubules specifically during mitosis, in a TPX2 (targeting protein for Xklp2)-dependent manner. We have shown that Hklp2 participates in spindle pole separation and in the maintenance of spindle bipolarity in metaphase. To better understand the function of Hklp2, we have performed a detailed domain analysis. Interestingly, from its positioning on the chromosome arms, Hklp2 seems to restrict spindle pole separation. In the present review, we summarize the current knowledge of the function and regulation of the different kinesins associated with chromosome arms during cell division, including Hklp2 as a novel member of this so-called chromokinesin family.
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Bobinnec, Y., M. Fukuda, and E. Nishida. "Identification and characterization of Caenorhabditis elegans gamma-tubulin in dividing cells and differentiated tissues." Journal of Cell Science 113, no. 21 (November 1, 2000): 3747–59. http://dx.doi.org/10.1242/jcs.113.21.3747.
Full textAbstract:
gamma-Tubulin is an essential component of the microtubule-nucleation machinery and therefore plays a crucial role during mitosis. To gain further insights into the function of this protein in the events that take place during embryogenesis and differentiation, we carried out detailed studies on gamma-tubulin during all the developmental stages of Caenorhabditis elegans. We identified the gamma-tubulin gene from this organism and analyzed the localization of the protein by both immunofluorescence and GFP reporter construct. We show that gamma-tubulin association with the centrosome is highly dynamic in mitotic cells, being massively recruited at prophase and released at anatelophase. This accumulation in mitotic centrosomes is dramatic during the first embryonic divisions. We provide the first description of the morphological changes at the centrosome level during the orientation of the mitotic spindle and the flattening of the posterior aster. Loss of function of the gamma-tubulin gene by RNAi induces a strong polyploidization of mitotic germ cells and embryos, but does not affect meiosis and pronuclear migration. In addition, we demonstrate the prominent redistribution of gamma-tubulin in adults at basal bodies of amphid and phasmid neurons, and at the apical membrane of polarized intestinal cells.
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Sherwood, Racquel Kim, and Richard J. Bennett. "Microtubule Motor Protein Kar3 Is Required for Normal Mitotic Division and Morphogenesis in Candida albicans." Eukaryotic Cell 7, no. 9 (June 27, 2008): 1460–74. http://dx.doi.org/10.1128/ec.00138-08.
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ABSTRACT The kinesin-related protein Kar3 is a minus end-directed molecular motor that plays a multifunctional role in microtubule-directed nuclear movement. Previously, it was shown that Candida albicans Kar3p is critical for nuclear fusion during mating as kar3 mutants were defective in karyogamy. In this study, we confirm that Kar3p is required for nuclear congression in mating but that neither Kar3p nor the dynein motor protein Dyn1p is required for nuclear migration in the mating projection prior to cell fusion. In addition, we show that C. albicans Kar3p plays an important role in the cell and colony morphology of mitotically dividing cells, as evidenced by diminished filamentation of kar3 cells on Spider medium and an increased tendency of mutant cells to form pseudohyphal cells in liquid culture. Loss of Kar3p also led to defects in nuclear division, causing cells to grow slowly and exhibit reduced viability compared to wild-type cells. Slow growth was due, at least in part, to delayed cell cycle progression, as cells lacking Kar3p accumulated in anaphase of the cell cycle. Consistent with a role in mitotic division, Kar3 protein was shown to localize to the spindle pole bodies. Finally, kar3 cells exhibited unstable or aberrant mitotic spindles, a finding that accounts for the delay in cell cycle progression and decreased viability of these cells. We suggest that the altered morphology of kar3 cells is a direct consequence of the delay in anaphase, and this leads to increased polarized growth and pseudohypha formation.
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Granger, C., and R. Cyr. "Use of abnormal preprophase bands to decipher division plane determination." Journal of Cell Science 114, no. 3 (February 1, 2001): 599–607. http://dx.doi.org/10.1242/jcs.114.3.599.
Full textAbstract:
Many premitotic plant cells possess a cortical preprophase band of microtubules and actin filaments that encircles the nucleus. In vacuolated cells, the preprophase band is visibly connected to the nucleus by a cytoplasmic raft of actin filaments and microtubules termed the phragmosome. Typically, the location of the preprophase band and phragmosome corresponds to, and thus is thought to influence, the location of the cell division plane. To better understand the function of the preprophase band and phragmosome in orienting division, we used a green fluorescent protein-based microtubule reporter protein to observe mitosis in living tobacco bright yellow 2 cells possessing unusual preprophase bands. Observations of mitosis in these unusual cells support the involvement of the preprophase band/phragmosome in properly positioning the preprophase nucleus, influencing spindle orientation such that the cytokinetic phragmoplast initially grows in an appropriate direction, and delineating a region in the cell cortex that attracts microtubules and directs later stages of phragmoplast growth. Thus, the preprophase band/phragmosome appears to perform several interrelated functions to orient the division plane. However, functional information associated with the preprophase band is not always used or needed and there appears to be an age or distance-dependent character to the information. Cells treated with the anti-actin drug, latrunculin B, are still able to position the preprophase nucleus suggesting that microtubules may play a dominant role in premitotic positioning. Furthermore, in treated cells, spindle location and phragmoplast insertion are frequently abnormal suggesting that actin plays a significant role in nuclear anchoring and phragmoplast guidance. Thus, the microtubule and actin components of the preprophase band/phragmosome execute complementary activities to ensure proper orientation of the division plane.
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Meyer, Régis E., Aaron R. Tipton, Rebecca LaVictoire, Gary J. Gorbsky, and Dean S. Dawson. "Mps1 promotes poleward chromosome movements in meiotic prometaphase." Molecular Biology of the Cell 32, no. 10 (May 1, 2021): 1020–32. http://dx.doi.org/10.1091/mbc.e20-08-0525-t.
Full textAbstract:
Mps1 is a kinase that regulates several steps in mitosis and meiosis. Mps1 is essential for the spindle checkpoint and helps stabilize attachment of kinetochores to microtubules. Here we show that following microtubule attachment, Mps1 promotes microtubule depolymerization to trigger migration of the chromosome toward the spindle pole.
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Bulgheresi, Silvia, Elke Kleiner, and Juergen A. Knoblich. "Inscuteable-dependent apical localization of the microtubule-binding protein Cornetto suggests a role in asymmetric cell division." Journal of Cell Science 114, no. 20 (October 15, 2001): 3655–62. http://dx.doi.org/10.1242/jcs.114.20.3655.
Full textAbstract:
Drosophila neuroblasts divide asymmetrically along the apical-basal axis. The Inscuteable protein localizes to the apical cell cortex in neuroblasts from interphase to metaphase, but disappears in anaphase. Inscuteable is required for correct spindle orientation and for asymmetric localization of cell fate determinants to the opposite (basal) cell cortex. Here, we show that Inscuteable also directs asymmetric protein localization to the apical cell cortex during later stages of mitosis. In a two-hybrid screen for Inscuteable-binding proteins, we have identified the coiled-coil protein Cornetto, which shows a highly unusual subcellular distribution in neuroblasts. Although the protein is uniformly distributed in the cytoplasm during metaphase, it concentrates apically in anaphase and forms an apical crescent during telophase in an inscuteable-dependent manner. Upon overexpression, Cornetto localizes to astral microtubules and microtubule spin-down experiments demonstrate that Cornetto is a microtubule-binding protein. After disruption of the actin cytoskeleton, Cornetto localizes with microtubules throughout the cell cycle and decorates the mitotic spindle during metaphase. Our results reveal a novel pattern of asymmetric protein localization in Drosophila neuroblasts and are consistent with a function of Cornetto in anchoring the mitotic spindle during late phases of mitosis, even though our cornetto mutant analysis suggests that this function might be obscured by genetic redundancy.
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Brachat, Arndt, John V. Kilmartin, Achim Wach, and Peter Philippsen. "Saccharomyces cerevisiaeCells with Defective Spindle Pole Body Outer Plaques Accomplish Nuclear Migration via Half-Bridge–organized Microtubules." Molecular Biology of the Cell 9, no. 5 (May 1998): 977–91. http://dx.doi.org/10.1091/mbc.9.5.977.
Full textAbstract:
Cnm67p, a novel yeast protein, localizes to the microtubule organizing center, the spindle pole body (SPB). Deletion ofCNM67 (YNL225c) frequently results in spindle misorientation and impaired nuclear migration, leading to the generation of bi- and multinucleated cells (40%). Electron microscopy indicated that CNM67 is required for proper formation of the SPB outer plaque, a structure that nucleates cytoplasmic (astral) microtubules. Interestingly, cytoplasmic microtubules that are essential for spindle orientation and nuclear migration are still present in cnm67Δ1 cells that lack a detectable outer plaque. These microtubules are attached to the SPB half- bridge throughout the cell cycle. This interaction presumably allows for low-efficiency nuclear migration and thus provides a rescue mechanism in the absence of a functional outer plaque. AlthoughCNM67 is not strictly required for mitosis, it is essential for sporulation. Time-lapse microscopy ofcnm67Δ1 cells with green fluorescent protein (GFP)-labeled nuclei indicated that CNM67 is dispensable for nuclear migration (congression) and nuclear fusion during conjugation. This is in agreement with previous data, indicating that cytoplasmic microtubules are organized by the half-bridge during mating.
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Giansanti, M. G., M. Gatti, and S. Bonaccorsi. "The role of centrosomes and astral microtubules during asymmetric division of Drosophila neuroblasts." Development 128, no. 7 (April 1, 2001): 1137–45. http://dx.doi.org/10.1242/dev.128.7.1137.
Full textAbstract:
Drosophila neuroblasts are stem cells that divide asymmetrically to produce another large neuroblast and a smaller ganglion mother cell (GMC). During neuroblast division, several cell fate determinants, such as Miranda, Prospero and Numb, are preferentially segregated into the GMC, ensuring its correct developmental fate. The accurate segregation of these determinants relies on proper orientation of the mitotic spindle within the dividing neuroblast, and on the correct positioning of the cleavage plane. In this study we have analyzed the role of centrosomes and astral microtubules in neuroblast spindle orientation and cytokinesis. We examined neuroblast division in asterless (asl) mutants, which, although devoid of functional centrosomes and astral microtubules, form well-focused anastral spindles that undergo anaphase and telophase. We show that asl neuroblasts assemble a normal cytokinetic ring around the central spindle midzone and undergo unequal cytokinesis. Thus, astral microtubules are not required for either signaling or positioning cytokinesis in Drosophila neuroblasts. Our results indicate that the cleavage plane is dictated by the positioning of the central spindle midzone within the cell, and suggest a model on how the central spindle attains an asymmetric position during neuroblast mitosis. We have also analyzed the localization of Miranda during mitotic division of asl neuroblasts. This protein accumulates in morphologically regular cortical crescents but these crescents are mislocalized with respect to the spindle orientation. This suggests that astral microtubules mediate proper spindle rotation during neuroblast division.
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Lorson, Monique A., H. Robert Horvitz, and Sander van den Heuvel. "LIN-5 Is a Novel Component of the Spindle Apparatus Required for Chromosome Segregation and Cleavage Plane Specification in Caenorhabditis elegans." Journal of Cell Biology 148, no. 1 (January 10, 2000): 73–86. http://dx.doi.org/10.1083/jcb.148.1.73.
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Successful divisions of eukaryotic cells require accurate and coordinated cycles of DNA replication, spindle formation, chromosome segregation, and cytoplasmic cleavage. The Caenorhabditis elegans gene lin-5 is essential for multiple aspects of cell division. Cells in lin-5 null mutants enter mitosis at the normal time and form bipolar spindles, but fail chromosome alignment at the metaphase plate, sister chromatid separation, and cytokinesis. Despite these defects, cells exit from mitosis without delay and progress through subsequent rounds of DNA replication, centrosome duplication, and abortive mitoses. In addition, early embryos that lack lin-5 function show defects in spindle positioning and cleavage plane specification. The lin-5 gene encodes a novel protein with a central coiled-coil domain. This protein localizes to the spindle apparatus in a cell cycle- and microtubule-dependent manner. The LIN-5 protein is located at the centrosomes throughout mitosis, at the kinetochore microtubules in metaphase cells, and at the spindle during meiosis. Our results show that LIN-5 is a novel component of the spindle apparatus required for chromosome and spindle movements, cytoplasmic cleavage, and correct alternation of the S and M phases of the cell cycle.
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Eichenlaub-Ritter, U. "Spatiotemporal control of functional specification and distribution of spindle microtubules with 13, 14 and 15 protofilaments during mitosis in the ciliate Nyctotherus." Journal of Cell Science 76, no. 1 (June 1, 1985): 337–55. http://dx.doi.org/10.1242/jcs.76.1.337.
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The formation of microtubules with more than 13 protofilaments in the ciliate Nyctotherus ovalis Leidy seems to be a highly ordered process. Such microtubules are restricted to the nucleoplasm and, moreover, to certain stages of nuclear division. They assemble during anaphase of micronuclear mitosis and during the elongation phase of macronuclear division. The number of microtubules with more than 13 protofilaments in the micronuclear nucleoplasm increases as anaphase progresses. Furthermore, assembly of microtubules with 14 and 15 protofilaments seems to proceed concomitantly with net disassembly of 13-protofilament microtubules, because the total amount of polymerized tubulin in the interpolar spindle region remains approximately constant between mid anaphase and late telophase. In addition, evidence for spatial control of the distribution of microtubules with different protofilament numbers in the micronuclear stembody has been found. The percentage of microtubules with 13 protofilaments per stembody cross-section is highest at the ends of the stembody, while the percentage of microtubules with either 14 or 15 protofilaments increases as the middle of the stembody is approached. Temporal control of polymerization of microtubules with high protofilament numbers seems to be exerted independently in the two types of nuclei. For example, when the macronucleus starts to elongate it contains microtubules with more than 13 protofilaments but the metaphase micronucleus still possesses only microtubules with 13 protofilaments at this stage. Control of fidelity of protofilament numbers is not lost in the early stages of micronuclear or macronuclear division when cells are exposed to 2H2O or media containing taxol. Even microtubules that reassemble during recovery of metaphase micronuclei from nocodazole-induced microtubule depolymerization, in either the absence or presence of 2H2O and taxol, possess 13 protofilaments. Similarly, if the introduction of microtubules with 14 and 15 protofilaments is inhibited during early micronuclear anaphase and delayed for 60 min by exposure to nocodazole, such microtubules still assemble during telophase when recovery is permitted. Microtubules that have been assembled under normal conditions show differential sensitivity to nocodazole. During metaphase, nocodazole induces disassembly of most microtubules. There is an increase in microtubule stability that coincides with the appearance of microtubules with high protofilament numbers during early anaphase. However, considerable numbers of 13-protofilament microtubules, as well as microtubules with 14 and 15 protofilaments, exhibit such stability during anaphase.(ABSTRACT TRUNCATED AT 400 WORDS)
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Liu, Leyuan, Rui Xie, Chaofeng Yang, and Wallace L. McKeehan. "Dual Function Microtubule- and Mitochondria-Associated Proteins Mediate Mitotic Cell Death." Analytical Cellular Pathology 31, no. 5 (January 1, 2009): 393–405. http://dx.doi.org/10.1155/2009/783817.
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Background: Survival and evolution of aneuploid cells after an asymmetric segregation of chromosomes at mitosis may be the common initiating event and underlying cause of the genetic diversity and adaptability of cancers. We hypothesize that mechanisms exist to detect impending aneuploidy and prevent it before completion of an aberrant mitosis.Methods: The distribution of isoforms of C19ORF5, an interactive partner with mitochondria-associated LRPPRC and tumor suppressor RASSF1A, state of spindle microtubules and mitochondrial aggregation was analyzed in synchronized mitotic cells and cells stalled in mitosis after treatment with paclitaxel.Results: C19ORF5 distributed broadly across the mitotic spindle and reversibly accumulated during reversible mitotic arrest. Prolonged stabilization of microtubules caused an accumulation of a C19ORF5 product with dual MAP and MtAP properties that caused irreversible aggregation of mitochondria and death of mitotic cells.Conclusions: Dual function microtubule-associated (MAP) and mitochondria-associated (MtAP) proteins generated by prolonged mitotic arrest trigger mitochondrial-induced mitotic cell death. This is a potential mechanism to prevent minimal survivable aneuploidy resulting from an aberrant cell division and cancers in general at their earliest common origin.
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Frieders, Elizabeth M., and David J. McLaughlln. "Mitosis in the yeast phase of Agaricostilbum pulcherrimum and its evolutionary significance." Canadian Journal of Botany 74, no. 9 (September 1, 1996): 1392–406. http://dx.doi.org/10.1139/b96-169.
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Agaricostilbum pulcherrimum is an anomaly and is difficult to place systematically. It possesses a yeast phase, and as in most basidiomycetous yeasts, mitosis has not been investigated cytoiogically. Yeast cells of A. pulcherrimum were prepared for immunofluorescence and transmission electron microscopy by a freeze-substitution method. A cladistic analysis of cell cycle characters among A. pulcherrimum and two ascomycetous and two basidiomycetous yeasts, performed with phylogenetic analysis using parsimony, revealed that A. pulcherrimum is basal within these basidiomycetes. Spindle pole bodies are multilayered discs and appear to be intranuclear during early division, similar to meiotic division. Spindle initiation and early elongation occur in the parent, a situation unreported in basidiomycetous yeasts. The site of spindle initiation, the position of the nucleus during division, and the pattern of astral microtubules demonstrate that the mode of nuclear division in A. pulcherrimum is intermediate between those of the investigated ascomycetous and basidiomycetous yeasts. Keywords: basidiomycete, cell cycle, cytoskeleton, immunofluorescence, phylogeny, spindle pole body.
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Uematsu, Keiji, Fumihiko Okumura, Syunsuke Tonogai, Akiko Joo-Okumura, Dawit Hailu Alemayehu, Akihiko Nishikimi, Yoshinori Fukui, Kunio Nakatsukasa, and Takumi Kamura. "ASB7 regulates spindle dynamics and genome integrity by targeting DDA3 for proteasomal degradation." Journal of Cell Biology 215, no. 1 (October 3, 2016): 95–106. http://dx.doi.org/10.1083/jcb.201603062.
Full textAbstract:
Proper dynamic regulation of the spindle is essential for successful cell division. However, the molecular mechanisms that regulate spindle dynamics in mitosis are not fully understood. In this study, we show that Cullin 5–interacting suppressor of cytokine signaling box protein ASB7 ubiquitinates DDA3, a regulator of spindle dynamics, thereby targeting it for proteasomal degradation. The presence of microtubules (MTs) prevented the ASB7–DDA3 interaction, thus stabilizing DDA3. Knockdown of ASB7 decreased MT polymerization and increased the proportion of cells with unaligned chromosomes, and this phenotype was rescued by deletion of DDA3. Collectively, these data indicate that ASB7 plays a crucial role in regulating spindle dynamics and genome integrity by controlling the expression of DDA3.
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Cha, B., L. Cassimeris, and D. L. Gard. "XMAP230 is required for normal spindle assembly in vivo and in vitro." Journal of Cell Science 112, no. 23 (December 1, 1999): 4337–46. http://dx.doi.org/10.1242/jcs.112.23.4337.
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XMAP230 is a high molecular mass microtubule-associated protein isolated from Xenopus oocytes and eggs, and has been recently shown to be a homolog of mammalian MAP4. Confocal immunofluorescence microscopy revealed that XMAP230 is associated with microtubules throughout the cell cycle of early Xenopus embryos. During interphase XMAP230 is associated with the radial arrays of microtubules and midbodies remaining from the previous division. During mitosis, XMAP230 is associated with both astral microtubules and microtubules of the central spindle. Microinjection of affinity-purified anti-XMAP230 antibody into blastomeres severely disrupted the assembly of mitotic spindles during the rapid cleavage cycles of early development. Both monopolar half spindles and bipolar spindles were assembled from XMAP230-depleted extracts in vitro. However, spindles assembled in XMAP230-depleted extracts exhibited a reduction in spindle width, reduced microtubule density, chromosome loss, and reduced acetylation of spindle MTs. Similar defects were observed in the spindles assembled in XMAP230-depleted extracts that had been cycled through interphase. Depletion of XMAP230 had no effect on the pole-to-pole length of spindles, and depletion of XMAP230 from both interphase and M-phase extracts had no effect on the rate of microtubule elongation. From these results, we conclude that XMAP230 plays an important role in normal spindle assembly, primarily by acting to stabilize spindle microtubules, and that the observed defects in spindle assembly may result from enhanced microtubule dynamics in XMAP230-depleted extracts.
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Yokobayashi, Shihori, Masayuki Yamamoto, and Yoshinori Watanabe. "Cohesins Determine the Attachment Manner of Kinetochores to Spindle Microtubules at Meiosis I in Fission Yeast." Molecular and Cellular Biology 23, no. 11 (June 1, 2003): 3965–73. http://dx.doi.org/10.1128/mcb.23.11.3965-3973.2003.
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ABSTRACT During mitosis, sister kinetochores attach to microtubules that extend to opposite spindle poles (bipolar attachment) and pull the chromatids apart at anaphase (equational segregation). A multisubunit complex called cohesin, including Rad21/Scc1, plays a crucial role in sister chromatid cohesion and equational segregation at mitosis. Meiosis I differs from mitosis in having a reductional pattern of chromosome segregation, in which sister kinetochores are attached to the same spindle (monopolar attachment). During meiosis, Rad21/Scc1 is largely replaced by its meiotic counterpart, Rec8. If Rec8 is inactivated in fission yeast, meiosis I is shifted from reductional to equational division. However, the reason rec8Δ cells undergo equational rather than random division has not been clarified; therefore, it has been unclear whether equational segregation is due to a loss of cohesin in general or to a loss of a specific requirement for Rec8. We report here that the equational segregation at meiosis I depends on substitutive Rad21, which relocates to the centromeres if Rec8 is absent. Moreover, we demonstrate that even if sufficient amounts of Rad21 are transferred to the centromeres at meiosis I, thereby establishing cohesion at the centromeres, rec8Δ cells never recover monopolar attachment but instead secure bipolar attachment. Thus, Rec8 and Rad21 define monopolar and bipolar attachment, respectively, at meiosis I. We conclude that cohesin is a crucial determinant of the attachment manner of kinetochores to the spindle microtubules at meiosis I in fission yeast.
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Reinsch, S., and E. Karsenti. "Orientation of spindle axis and distribution of plasma membrane proteins during cell division in polarized MDCKII cells." Journal of Cell Biology 126, no. 6 (September 15, 1994): 1509–26. http://dx.doi.org/10.1083/jcb.126.6.1509.
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MDCKII cells differentiate into a simple columnar epithelium when grown on a permeable support; the monolayer is polarized for transport and secretion. Individual cells within the monolayer continue to divide at a low rate without disturbing the function of the epithelium as a barrier to solutes. This presents an interesting model for the study of mitosis in a differentiated epithelium which we have investigated by confocal immunofluorescence microscopy. We monitored the distribution of microtubules, centrioles, nucleus, tight junctions, and plasma membrane proteins that are specifically targeted to the apical and basolateral domains. The stable interphase microtubule cytoskeleton was rapidly disassembled at prophase onset and reassembled at cytokinesis. As the interphase microtubules disassembled at prophase, the centrioles moved from their interphase position at the apical membrane to the nucleus and acquired the ability to organize microtubule asters. Orientation of the spindle parallel to the plane of the monolayer occurred between late prophase and metaphase and persisted through cytokinesis. The cleavage furrow formed asymmetrically perpendicular to the plane of the monolayer initiating at the basolateral side and proceeding to the apical domain. The interphase microtubule network reformed after the centrioles migrated from the spindle poles to resume their interphase apical position. Tight junctions (ZO-1), which separate the apical from the basolateral domains, remained assembled throughout all phases of mitosis. E-cadherin and a 58-kD antigen maintained their basolateral plasma membrane distributions, and a 114-kD antigen remained polarized to the apical domain. These proteins were useful for monitoring the changes in shape of the mitotic cells relative to neighboring cells, especially during telophase when the cell shape changes dramatically. We discuss the changes in centriole position during the cell cycle, mechanisms of spindle orientation, and how the maintenance of polarized plasma membrane domains through mitosis may facilitate the rapid reformation of the polarized interphase cytoplasm.
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Victoria, V. W., W. Allen, and D. L. Kropf. "Nuclear rotation and lineage specification in Pelvetia embryos." Development 115, no. 3 (July 1, 1992): 873–83. http://dx.doi.org/10.1242/dev.115.3.873.
Full textAbstract:
The first division of Pelvetia zygotes is an unequal div-ision which produces two cells with distinct developmental fates. The smaller rhizoid cell gives rise to the holdfast of the mature plant, and the larger thallus cell is the pro-genitor of the stipe and fronds. We have investigated the role of the cytoskeleton in determining the orientation of this invariant division. Prior to mitosis, microtubule-organizing centers (MtOCs) associated with the nuclear envelope undergo a precise realignment from transverse to axial with respect to the rhizoid/thallus axis. This is accomplished by a 90 deg rotation of the entire nuclear/MtOC complex. After rotation, each MtOC serves as a spindle pole during mitosis, and subsequently cytokinesis bisects the spindle. Both nocodazole and cytochalasin D cause incorrect alignment of MtOCs, indicating that both microtubules and microfilaments are required for nuclear/MtOC rotation. These inhibitors also result in aberrant orientation of the first division plane. Microtubules visualized by confocal microscopy connect the rotating nucleus to the apical cortex and may provide the force for rotation.
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Skinner, John J., Stacey Wood, James Shorter, S. Walter Englander, and Ben E. Black. "The Mad2 partial unfolding model: regulating mitosis through Mad2 conformational switching." Journal of Cell Biology 183, no. 5 (November 24, 2008): 761–68. http://dx.doi.org/10.1083/jcb.200808122.
Full textAbstract:
The metamorphic Mad2 protein acts as a molecular switch in the checkpoint mechanism that monitors proper chromosome attachment to spindle microtubules during cell division. The remarkably slow spontaneous rate of Mad2 switching between its checkpoint inactive and active forms is catalyzed onto a physiologically relevant time scale by a self–self interaction between its two forms, culminating in a large pool of active Mad2. Recent structural, biochemical, and cell biological advances suggest that the catalyzed conversion of Mad2 requires a major structural rearrangement that transits through a partially unfolded intermediate.
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Jones, Matthew C., Junzhe Zha, and Martin J. Humphries. "Connections between the cell cycle, cell adhesion and the cytoskeleton." Philosophical Transactions of the Royal Society B: Biological Sciences 374, no. 1779 (July 2019): 20180227. http://dx.doi.org/10.1098/rstb.2018.0227.
Full textAbstract:
Cell division, the purpose of which is to enable cell replication, and in particular to distribute complete, accurate copies of genetic material to daughter cells, is essential for the propagation of life. At a morphological level, division not only necessitates duplication of cellular structures, but it also relies on polar segregation of this material followed by physical scission of the parent cell. For these fundamental changes in cell shape and positioning to be achieved, mechanisms are required to link the cell cycle to the modulation of cytoarchitecture. Outside of mitosis, the three main cytoskeletal networks not only endow cells with a physical cytoplasmic skeleton, but they also provide a mechanism for spatio-temporal sensing via integrin-associated adhesion complexes and site-directed delivery of cargoes. During mitosis, some interphase functions are retained, but the architecture of the cytoskeleton changes dramatically, and there is a need to generate a mitotic spindle for chromosome segregation. An economical solution is to re-use existing cytoskeletal molecules: transcellular actin stress fibres remodel to create a rigid cortex and a cytokinetic furrow, while unipolar radial microtubules become the primary components of the bipolar spindle. This remodelling implies the existence of specific mechanisms that link the cell-cycle machinery to the control of adhesion and the cytoskeleton. In this article, we review the intimate three-way connection between microenvironmental sensing, adhesion signalling and cell proliferation, particularly in the contexts of normal growth control and aberrant tumour progression. As the morphological changes that occur during mitosis are ancient, the mechanisms linking the cell cycle to the cytoskeleton/adhesion signalling network are likely to be primordial in nature and we discuss recent advances that have elucidated elements of this link. A particular focus is the connection between CDK1 and cell adhesion. This article is part of a discussion meeting issue ‘Forces in cancer: interdisciplinary approaches in tumour mechanobiology’.
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Lee, Wei-Lih, Jessica R. Oberle, and John A. Cooper. "The role of the lissencephaly protein Pac1 during nuclear migration in budding yeast." Journal of Cell Biology 160, no. 3 (February 3, 2003): 355–64. http://dx.doi.org/10.1083/jcb.200209022.
Full textAbstract:
During mitosis in Saccharomyces cerevisiae, the mitotic spindle moves into the mother–bud neck via dynein-dependent sliding of cytoplasmic microtubules along the cortex of the bud. Here we show that Pac1, the yeast homologue of the human lissencephaly protein LIS1, plays a key role in this process. First, genetic interactions placed Pac1 in the dynein/dynactin pathway. Second, cells lacking Pac1 failed to display microtubule sliding in the bud, resulting in defective mitotic spindle movement and nuclear segregation. Third, Pac1 localized to the plus ends (distal tips) of cytoplasmic microtubules in the bud. This localization did not depend on the dynein heavy chain Dyn1. Moreover, the Pac1 fluorescence intensity at the microtubule end was enhanced in cells lacking dynactin or the cortical attachment molecule Num1. Fourth, dynein heavy chain Dyn1 also localized to the tips of cytoplasmic microtubules in wild-type cells. Dynein localization required Pac1 and, like Pac1, was enhanced in cells lacking the dynactin component Arp1 or the cortical attachment molecule Num1. Our results suggest that Pac1 targets dynein to microtubule tips, which is necessary for sliding of microtubules along the bud cortex. Dynein must remain inactive until microtubule ends interact with the bud cortex, at which time dynein and Pac1 appear to be offloaded from the microtubule to the cortex.
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Corbett, Kevin D., and Arshad Desai. "A new piece in the kinetochore jigsaw puzzle." Journal of Cell Biology 206, no. 4 (August 18, 2014): 457–59. http://dx.doi.org/10.1083/jcb.201407048.
Full textAbstract:
In eukaryotic cell division, the kinetochore mediates chromosome attachment to spindle microtubules and acts as a scaffold for signaling pathways, ensuring the accuracy of chromosome segregation. The architecture of the kinetochore underlies its function in mitosis. In this issue, Hornung et al. (2014. J. Cell Biol. http://dx.doi.org/201403081) identify an unexpected linkage between the inner and outer regions of the kinetochore in budding yeast that suggests a new model for the construction of this interface.
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Letort, Gaelle, Isma Bennabi, Serge Dmitrieff, François Nedelec, Marie-Hélène Verlhac, and Marie-Emilie Terret. "A computational model of the early stages of acentriolar meiotic spindle assembly." Molecular Biology of the Cell 30, no. 7 (March 21, 2019): 863–75. http://dx.doi.org/10.1091/mbc.e18-10-0644.
Full textAbstract:
The mitotic spindle is an ensemble of microtubules responsible for the repartition of the chromosomal content between the two daughter cells during division. In metazoans, spindle assembly is a gradual process involving dynamic microtubules and recruitment of numerous associated proteins and motors. During mitosis, centrosomes organize and nucleate the majority of spindle microtubules. In contrast, oocytes lack canonical centrosomes but are still able to form bipolar spindles, starting from an initial ball that self-organizes in several hours. Interfering with early steps of meiotic spindle assembly can lead to erroneous chromosome segregation. Although not fully elucidated, this process is known to rely on antagonistic activities of plus end– and minus end–directed motors. We developed a model of early meiotic spindle assembly in mouse oocytes, including key factors such as microtubule dynamics and chromosome movement. We explored how the balance between plus end– and minus end–directed motors, as well as the influence of microtubule nucleation, impacts spindle morphology. In a refined model, we added spatial regulation of microtubule stability and minus-end clustering. We could reproduce the features of early stages of spindle assembly from 12 different experimental perturbations and predict eight additional perturbations. With its ability to characterize and predict chromosome individualization, this model can help deepen our understanding of spindle assembly.
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Maddox, Paul, E. Chin, A. Mallavarapu, E. Yeh, E. D. Salmon, and K. Bloom. "Microtubule Dynamics from Mating through the First Zygotic Division in the Budding Yeast Saccharomyces cerevisiae." Journal of Cell Biology 144, no. 5 (March 8, 1999): 977–87. http://dx.doi.org/10.1083/jcb.144.5.977.
Full textAbstract:
We have used time-lapse digital imaging microscopy to examine cytoplasmic astral microtubules (Mts) and spindle dynamics during the mating pathway in budding yeast Saccharomyces cerevisiae. Mating begins when two cells of opposite mating type come into proximity. The cells arrest in the G1 phase of the cell cycle and grow a projection towards one another forming a shmoo projection. Imaging of microtubule dynamics with green fluorescent protein (GFP) fusions to dynein or tubulin revealed that the nucleus and spindle pole body (SPB) became oriented and tethered to the shmoo tip by a Mt-dependent search and capture mechanism. Dynamically unstable astral Mts were captured at the shmoo tip forming a bundle of three or four astral Mts. This bundle changed length as the tethered nucleus and SPB oscillated toward and away from the shmoo tip at growth and shortening velocities typical of free plus end astral Mts (∼0.5 μm/min). Fluorescent fiduciary marks in Mt bundles showed that Mt growth and shortening occurred primarily at the shmoo tip, not the SPB. This indicates that Mt plus end assembly/disassembly was coupled to pushing and pulling of the nucleus. Upon cell fusion, a fluorescent bar of Mts was formed between the two shmoo tip bundles, which slowly shortened (0.23 ± 0.07 μm/min) as the two nuclei and their SPBs came together and fused (karyogamy). Bud emergence occurred adjacent to the fused SPB ∼30 min after SPB fusion. During the first mitosis, the SPBs separated as the spindle elongated at a constant velocity (0.75 μm/min) into the zygotic bud. There was no indication of a temporal delay at the 2-μm stage of spindle morphogenesis or a lag in Mt nucleation by replicated SPBs as occurs in vegetative mitosis implying a lack of normal checkpoints. Thus, the shmoo tip appears to be a new model system for studying Mt plus end dynamic attachments and much like higher eukaryotes, the first mitosis after haploid cell fusion in budding yeast may forgo cell cycle checkpoints present in vegetative mitosis.
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Ehler, L. L., J. A. Holmes, and S. K. Dutcher. "Loss of spatial control of the mitotic spindle apparatus in a Chlamydomonas reinhardtii mutant strain lacking basal bodies." Genetics 141, no. 3 (November 1, 1995): 945–60. http://dx.doi.org/10.1093/genetics/141.3.945.
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Abstract The bld2-1 mutation in the green alga Chlamydomonas reinhardtii is the only known mutation that results in the loss of centrioles/basal bodies and the loss of coordination between spindle position and cleavage furrow position during cell division. Based on several different assays, bld2-1 cells lack basal bodies in > 99% of cells. The stereotypical cytoskeletal morphology and precise positioning of the cleavage furrow observed in wild-type cells is disrupted in bld2-1 cells. The positions of the mitotic spindle and of the cleavage furrow are not correlated with respect to each other or with a specific cellular landmark during cell division in bld2-1 cells. Actin has a variable distribution during mitosis in bld2-1 cells, but this aberrant distribution is not correlated with the spindle positioning defect. In both wild-type and bld2-1 cells, the position of the cleavage furrow is coincident with a specialized set of microtubules found in green algae known as the rootlet microtubules. We propose that the rootlet microtubules perform the functions of astral microtubules and that functional centrioles are necessary for the organization of the cytoskeletal superstructure critical for correct spindle and cleavage furrow placement in Chlamydomonas.
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Venuto, Santina, and Giuseppe Merla. "E3 Ubiquitin Ligase TRIM Proteins, Cell Cycle and Mitosis." Cells 8, no. 5 (May 27, 2019): 510. http://dx.doi.org/10.3390/cells8050510.
Full textAbstract:
The cell cycle is a series of events by which cellular components are accurately segregated into daughter cells, principally controlled by the oscillating activities of cyclin-dependent kinases (CDKs) and their co-activators. In eukaryotes, DNA replication is confined to a discrete synthesis phase while chromosome segregation occurs during mitosis. During mitosis, the chromosomes are pulled into each of the two daughter cells by the coordination of spindle microtubules, kinetochores, centromeres, and chromatin. These four functional units tie chromosomes to the microtubules, send signals to the cells when the attachment is completed and the division can proceed, and withstand the force generated by pulling the chromosomes to either daughter cell. Protein ubiquitination is a post-translational modification that plays a central role in cellular homeostasis. E3 ubiquitin ligases mediate the transfer of ubiquitin to substrate proteins determining their fate. One of the largest subfamilies of E3 ubiquitin ligases is the family of the tripartite motif (TRIM) proteins, whose dysregulation is associated with a variety of cellular processes and directly involved in human diseases and cancer. In this review we summarize the current knowledge and emerging concepts about TRIMs and their contribution to the correct regulation of cell cycle, describing how TRIMs control the cell cycle transition phases and their involvement in the different functional units of the mitotic process, along with implications in cancer progression.
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Yao, Changfu, Uttama Rath, Helder Maiato, David Sharp, Jack Girton, Kristen M. Johansen, and Jørgen Johansen. "A nuclear-derived proteinaceous matrix embeds the microtubule spindle apparatus during mitosis." Molecular Biology of the Cell 23, no. 18 (September 15, 2012): 3532–41. http://dx.doi.org/10.1091/mbc.e12-06-0429.
Full textAbstract:
The concept of a spindle matrix has long been proposed. Whether such a structure exists, however, and what its molecular and structural composition are have remained controversial. In this study, using a live-imaging approach in Drosophila syncytial embryos, we demonstrate that nuclear proteins reorganize during mitosis to form a highly dynamic, viscous spindle matrix that embeds the microtubule spindle apparatus, stretching from pole to pole. We show that this “internal” matrix is a distinct structure from the microtubule spindle and from a lamin B–containing spindle envelope. By injection of 2000-kDa dextran, we show that the disassembling nuclear envelope does not present a diffusion barrier. Furthermore, when microtubules are depolymerized with colchicine just before metaphase the spindle matrix contracts and coalesces around the chromosomes, suggesting that microtubules act as “struts” stretching the spindle matrix. In addition, we demonstrate that the spindle matrix protein Megator requires its coiled-coil amino-terminal domain for spindle matrix localization, suggesting that specific interactions between spindle matrix molecules are necessary for them to form a complex confined to the spindle region. The demonstration of an embedding spindle matrix lays the groundwork for a more complete understanding of microtubule dynamics and of the viscoelastic properties of the spindle during cell division.