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determines the division plane in animal cells is perhaps the most enduring mystery in cytokinesis and a matter of intense debate. There exist three hypotheses of furrow induction. The first is the astral stimulation hypothesis, which postulates that astral microtubules from the spindle poles carry a furrow-inducing signal to the cell cortex, where signals from two poles are somehow focused into a ring at the spindle. A second possibility, called the central spindle hypothesis, is that the cleavage furrow is induced by a positive stimulus that originates in the central spindle equator. The central spindle may contribute to the specification of the division plane by promoting concentration and activation of the small GTPase RhoA at the equatorial cortex. A third hypothesis is the astral relaxation hypothesis. It postulates that active actin-myosin bundles are distributed throughout the cell cortex, and inhibition of their contraction near the spindle poles results in a gradient of contractile activity that is highest at the midpoint between poles. In other words, astral microtubules generate a negative signal that increases cortical relaxation close to the poles. Genetic and laser-micromanipulation studies in C. elegans embryos have shown that the spindle sends two redundant signals to the cell cortex, one originating from the central spindle, and a second signal deriving from the spindle aster, suggesting the involvement of multiple mechanisms combined in the positioning of the cleavage furrow. The predominance of one particular signal varies between cell types and organisms. And the multitude and partial redundancy of signals may be required to make the system robust and to increase spatial precision.
505:(adjacent to the cell membrane). Rho protein family (RhoA protein in mammalian cells) is a key regulator of contractile ring formation and contraction in animal cells. The RhoA pathway promotes assembly of the actin-myosin ring by two main effectors. First, RhoA stimulates nucleation of unbranched actin filaments by activation of Diaphanous-related formins. This local generation of new actin filaments is important for the contractile ring formation. This actin filament formation process also requires a protein called profilin, which binds to actin monomers and helps load them onto the filament end. Second, RhoA promotes myosin II activation by the kinase ROCK, which activates myosin II directly by phosphorylation of the myosin light chain and also inhibits myosin phosphatase by phosphorylation of the phosphatase-targeting subunit MYPT. Besides actin and myosin II, the contractile ring contains the scaffolding protein anillin. Anillin binds to actin, myosin, RhoA, and CYK-4, and thereby links the equatorial cortex with the signals from the central spindle. It also contributes to the linkage of the actin-myosin ring to the plasma membrane. Additionally, anillin generates contractile forces by rectifying thermal fluctuations. Another protein, septin, has also been speculated to serve as a structural scaffold on which the cytokinesis apparatus is organized. Following its assembly, contraction of the actin-myosin ring leads to ingression of the attached plasma membrane, which partitions the cytoplasm into two domains of emerging sister cells. The force for the contractile processes is generated by movements along actin by the motor protein myosin II. Myosin II uses the free energy released when
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transition leads to dephosphorylating of inhibitory sites on multiple central spindle components. First of all, the removal of a CDK1 phosphorylation from a subunit of the CPC (the chromosomal passenger complex) allows its translocalization to the central spindle from the centromeres, where it is located during metaphase. Besides being a structural component of the central spindle itself, CPC also plays a role in the phosphoregulation of other central spindle components, including PRC1 (microtubule-bundling protein required for cytokinesis 1) and MKLP1 (a kinesin motor protein). Originally inhibited by CDK1-mediated phosphorylation, PRC1 is now able to form a homodimer that selectively binds to the interface between antiparallel microtubules, facilitating spatial organization of the microtubules of the central spindle. MKLP1, together with the Rho-family GTPase activating protein CYK-4 (also termed MgcRacGAP), forms the centralspindlin complex. Centralspindlin binds to the central spindle as higher-order clusters. The centralspindlin cluster formation is promoted by phosphorylation of MLKP1 by Aurora B, a component of CPC. In short, the self-assembly of central spindle is initiated through the phosphoregulation of multiple central spindle components by the decline of CDK1 activity, either directly or indirectly, at the metaphase-anaphase transition. The central spindle may have multiple functions in cytokinesis including the control of cleavage furrow positioning, the delivery of membrane vesicles to the cleavage furrow, and the formation of the midbody structure that is required for the final steps of division.
685:
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RhoA after furrow ingression. Actin disassembly is further controlled by the GTPase Rab35 and its effector, the phosphatidylinositol-4,5-bisphosphate 5-phosphatase OCRL. The final step of abscission is controlled by the recruitment and polymerization of the endosomal sorting complex required for transport III (ESCRT-III), which serves to physically constrict and separate the plasma membrane of the two adjoined daughter cells.
741:. Since Myosins are recruited to the medial region, the contractile forces acting on the cortex resemble a 'purse string' constriction pulling inwards. This leads to the inward constriction. The plasma membrane by virtue of its close association with the cortex via crosslinker proteins To the constriction of the cleavage furrow, the total surface area should be increased by supplying the plasma membrane via exocytosis.
28:
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425:. The process can be divided to the following distinct steps: anaphase spindle reorganization, division plane specification, actin-myosin ring assembly and contraction, and abscission. Faithful partitioning of the genome to emerging daughter cells is ensured through the tight temporal coordination of the above individual events by molecular signaling pathways.
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is largely responsible for the disassembly of microtubule bundles inside the intercellular bridge. Complete cortical constriction also requires removal of the underlying cytoskeletal structures. Actin filament disassembly during late cytokinesis depends on the PKCε–14-3-3 complex, which inactivates
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The process of abscission physically cleaves the midbody into two. Abscission proceeds by removal of cytoskeletal structures from the cytokinetic bridge, constriction of the cell cortex, and plasma membrane fission. The intercellular bridge is filled with dense bundles of antiparallel microtubules
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cells a cleavage furrow is observed to form and ingress, but then regress before cytokinesis is complete). The process of mitotic spindle reorganization and central spindle formation is caused by the decline of CDK1 activity during anaphase. The decline of CDK1 activity at the metaphase-anaphase
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that drives the cleavage process, during which cell membrane and wall grow inward, which eventually pinches the mother cell in two. The key components of this ring are the filamentous protein actin and the motor protein myosin II. The contractile ring assembles equatorially (in the middle of the
1838:
Behnam
Rashidieh; Belal Shohayeb; Amanda Louise Bain; Patrick R. J. Fortuna; Debottam Sinha; Andrew Burgess; Richard Mills; Rachael C. Adams; J. Alejandro Lopez; Peter Blumbergs; John Finnie; Murugan Kalimutho; Michael Piper; James Edward Hudson; Dominic C. H. Ng; Kum Kum Khanna (October 2021).
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The second step of animal cell cytokinesis involves division plane specification and cytokinetic furrow formation. Precise positioning of the division plane between the two masses of segregated chromosomes is essential to prevent chromosome loss. Meanwhile, the mechanism by which the spindle
215:, usually multiple, chromosomes of eukaryote. Accordingly, bacteria construct no mitotic spindle in cell division. Also, duplication of prokaryotic DNA takes place during the actual separation of chromosomes; in mitosis, duplication takes place during the
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Theoretical models show that symmetric constriction requires both lateral stabilization and constriction forces. Reduction of external pressure and of surface tension (by membrane trafficking) reduce the required stabilization and constriction forces.
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In some cases, a cell may divide its genetic material and grow in size, but fail to undergo cytokinesis. This results in larger cells with more than one nucleus. Usually this is an unwanted aberration and can be a sign of cancerous cells.
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Plant cytokinesis differs from animal cytokinesis, partly because of the rigidity of plant cell walls. Instead of plant cells forming a cleavage furrow such as develops between animal daughter cells, a dividing structure known as the
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Baluska F, Liners F, Hlavacka A, Schlicht M, Van Cutsem P, McCurdy DW, Menzel D (October 2005). "Cell wall pectins and xyloglucans are internalized into dividing root cells and accumulate within cell plates during cytokinesis".
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portion of normal proliferative cell divisions. To achieve this, many components of the cytokinesis machinery are highly regulated to ensure that they are able to perform a particular function at only a particular stage of the
620:; (2) trafficking of vesicles to the division plane and their fusion to generate a tubular-vesicular network; (3) continued fusion of membrane tubules and their transformation into membrane sheets upon the deposition of
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number and complement are maintained from one generation to the next and that, except in special cases, the daughter cells will be functional copies of the parent cell. After the completion of the
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associated with its absence varies from one species to the next (for example, certain
Drosophila cell types are incapable of forming a cleavage furrow without the central spindle, whereas in both
716:, which is polymerized directly at the cell plate by callose synthases. As the cell plate continues to mature and fuses with the parental plasma membrane, the callose is slowly replaced with
540:(composed of electron-dense, proteinaceous material) is formed, where the actin-myosin ring has reached a diameter of about 1–2 μm. Most animal cell types remain connected by an intercellular
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to the phragmoplast midzone. These vesicles contain lipids, proteins and carbohydrates needed for the formation of a new cell boundary. Electron tomographic studies have identified the
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Virginia
Andrade et al. Caveolae promote successful abscission by controlling intercellular bridge tension during cytokinesis. Sci. Adv.8 (2022). DOI:10.1126/sciadv.abm5095
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Betts, J Gordon; Desaix, Peter; Johnson, Eddie; Johnson, Jody E; Korol, Oksana; Kruse, Dean; Poe, Brandon; Wise, James; Womble, Mark D; Young, Kelly A (June 8, 2023).
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that derive from the central spindle. These microtubules overlap at the midbody, which is generally thought to be a targeting platform for the abscission machinery.
165:, which in most species die without function, though they do take on various special functions in other species. Another form of mitosis occurs in tissues such as
700:. The order in which different cell wall components are deposited has been determined largely by immuno-electron microscopy. The first components to arrive are
207:, but because of differences between prokaryotic and eukaryotic cell structures and functions, the mechanisms differ. For instance, a bacterial cell has a
1041:
Kucera, Ondrej; Siahaan, Valerie; Janda, Daniel; Dijkstra, Sietske H; Pilatova, Eliska; Zatecka, Eva; Diez, Stefan; Braun, Marcus; Lansky, Zdenek (2021).
1098:
1665:"Cross-linking Proteins with Distinct Properties Organize Different Assemblies of Actin Filaments" - Molecular Biology of the Cell, 4th Ed, 2002: Cell
600:, cytokinesis in plant cells is significantly different from that in animal cells, Rather than forming a contractile ring, plant cells construct a
1940:"Animal Cytokinesis: from parts list to mechanism" by Eggert, U.S., Mitchison, T.J., Field, C.M. (2006), Annual Review of Cell Biology 75, 543-66
433:
Animal cell cytokinesis starts with the stabilization of microtubules and reorganization of the mitotic spindle to form the central spindle. The
2019:
1319:"Three-dimensional analysis of syncytial-type cell plates during endosperm cellularization visualized by high resolution electron tomography"
1190:"Polo-like kinase 1 triggers the initiation of cytokinesis in human cells by promoting recruitment of the RhoGEF Ect2 to the central spindle"
1368:"Electron tomographic analysis of somatic cell plate formation in meristematic cells of Arabidopsis preserved by high-pressure freezing"
573:. Cytokinesis happens only after APC binds with CDC20. This allows for the separation of chromosomes and myosin to work simultaneously.
2311:
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724:(a glue-like layer containing pectin) develops from the cell plate, serving to bind the cell walls of adjoining cells together.
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for up to several hours until they are split by an actin-independent process termed abscission, the last step of cytokinesis.
441:) forms when non-kinetochore microtubule fibers are bundled between the spindle poles. A number of different species including
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886:
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Staehelin LA, Moore I (1995). "The Plant Golgi
Apparatus: Structure, Functional Organization and Trafficking Mechanisms".
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Mishima M, Pavicic V, Grüneberg U, Nigg EA, Glotzer M (August 2004). "Cell cycle regulation of central spindle assembly".
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These tubules then widen and fuse laterally with each other, eventually forming a planar, fenestrated sheet . As the
2012:
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Cytokinesis must be temporally controlled to ensure that it occurs only after sister chromatids separate during the
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as the source of these vesicles, but other studies have suggested that they contain endocytosed material as well.
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Esau's Plant
Anatomy: Meristems, Cells, and Tissues of the Plant Body: Their Structure, Function, and Development
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673:, often in an asymmetrical fashion, thus completing cytokinesis. The remaining fenestrae contain strands of
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causes this cleavage furrow to ingress (move inwards), a striking process that is clearly visible through a
2005:
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Particular functions demand various deviations from the process of symmetrical cytokinesis; for example in
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CEP55 is a mitotic phosphoprotein that plays a key role in cytokinesis, the final stage of cell division.
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carried by the secretory vesicles that fuse to form the cell plate. The next component to be added is
1270:"Cytokinesis in tobacco BY-2 and root tip cells: a new model of cell plate formation in higher plants"
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cell divides into two daughter cells. Cytoplasmic division begins during or after the late stages of
1950:
More description and nice images of cell division in plants, with a focus on fluorescence microscopy
941:
Battaglia, Emilio (2009). Caryoneme alternative to chromosome and a new caryological nomenclature.
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1841:"Cep55 regulation of PI3K/Akt signaling is required for neocortical development and ciliogenesis"
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807: – Process in which chromosomes are replicated and separated into two new identical nuclei
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require the central spindle in order to efficiently undergo cytokinesis, although the specific
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2195:
1461:"Endocytosis of cell surface material mediates cell plate formation during plant cytokinesis"
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Dhonukshe P, Baluska F, Schlicht M, Hlavacka A, Samaj J, Friml J, Gadella TW (January 2006).
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is hydrolyzed to move along these actin filaments, constricting the cell membrane to form a
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1233:(December 2000). "Cytokinesis in flowering plants: more than one way to divide a cell".
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783: – Final stage of a cell division for eukaryotic cells both in mitosis and meiosis
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1957:"Cytokinesis in Prokaryotes and Eukaryotes: Common Principles and Different Solutions"
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Beltrán-Heredia, Elena; Monroy, Francisco; Cao-García, Francisco J. (2019-11-21).
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Fededa JP, Gerlich DW (May 2012). "Molecular control of animal cell cytokinesis".
819: – Network of filamentous proteins that forms the internal framework of cells
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Charles E. Allen (July 1901). "On the Origin and Nature of the Middle
Lamella".
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Otegui MS, Mastronarde DN, Kang BH, Bednarek SY, Staehelin LA (September 2001).
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matures, large amounts of membrane material are removed via clathrin-mediated
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reorganize and disappear into a new cytoskeleton as the cell cycle returns to
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1691:"Regulation of the Total Cell Surface Area in Dividing Dictyostelium Cells"
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between plant daughter cells. It divides the cell into two daughter cells.
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Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2008-06-18).
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components; (4) recycling of excess membrane and other material from the
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1799:"Cep55 stabilization is required for normal execution of cytokinesis"
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839:"Polar Bodies - more a lack of understanding than a lack of respect"
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Seguí-Simarro JM, Austin JR, White EA, Staehelin LA (April 2004).
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Animal cell cytokinesis begins shortly after the onset of sister
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1043:"Anillin propels myosin-independent constriction of actin rings"
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Eventually, the edges of the cell plate fuse with the parental
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1742:"Mechanical conditions for stable symmetric cell constriction"
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passing through them, and are thought to be the precursors of
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Annual Review of Plant
Physiology and Plant Molecular Biology
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The process of
Cytokinesis in a plant cell and an animal cell
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begins within the lumen of the narrow tubules of the young
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For the use of this term in plant and animal shedding, see
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1502:"Polarized cytokinesis in vacuolate cells of Arabidopsis"
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Samuels AL, Giddings TH, Staehelin LA (September 1995).
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Petronczki M, Glotzer M, Kraut N, Peters JM (May 2007).
1910:. Houston: OpenStax CNX. 3.5 Cell Growth and Division.
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van der Horst A, Simmons J, Khanna KK (November 2009).
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Pages displaying short descriptions of redirect targets
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in animals the ovum takes almost all the cytoplasm and
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forms in the cytoplasm and grows into a new, doubled
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Cytokinetic furrow ingression is powered by Type II
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774: – Number of sets of chromosomes of a cell
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142:and cytokinesis, each daughter cell enters the
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837:Schmerler Samuel, Wessel Gary (January 2011).
616:that guides and supports the formation of the
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130:into the cytoplasm of the separating daughter
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1689:Tanaka M, Fujimoto K, Yumura S (April 2020).
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161:. This leaves very little for the resulting
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219:before mitosis begins, though the daughter
2020:
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1935:The Molecular Requirements for Cytokinesis
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488:Actin-myosin ring assembly and contraction
2312:Cellular apoptosis susceptibility protein
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604:in the middle of the cell. The stages of
536:The cytokinetic furrow ingresses until a
173:; it omits cytokinesis, thereby yielding
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26:
18:
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1937:by M. Glotzer (2005), Science 307, 1735
347:("motion, movement"). It was coined by
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1030:. New Science Press. pp. 157–173.
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646:is assembled from the remnants of the
608:formation include (1) creation of the
2001:
1500:Cutler SR, Ehrhardt DW (March 2002).
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223:don't separate completely before the
126:partitions and transports duplicated
102:and part of mitosis during which the
1668:. Garland Science. pp. 1006–.
1573:10.1146/annurev.pp.46.060195.001401
636:; and (5) fusion with the parental
576:After cytokinesis, non-kinetochore
13:
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1637:Evert RF, Eichorn S (2006-09-18).
199:Cytokinesis largely resembles the
14:
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551:The microtubule severing protein
34:undergoing cytokinesis, with the
16:Part of the cell division process
1895: This article incorporates
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795: – Stage of a cell division
749:Proteins involved in cytokinesis
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429:Anaphase spindle reorganization
1973:10.1128/MMBR.65.2.319-333.2001
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921:Merriam-Webster.com Dictionary
908:
879:
830:
801: – Stage of cell division
396:
371:, a hollow), Latin derivative
1:
1247:10.1016/s1369-5266(00)00119-9
823:
591:
524:
498:actin-myosin contractile ring
1858:10.1371/journal.pgen.1009334
1506:Proc. Natl. Acad. Sci. U.S.A
1478:10.1016/j.devcel.2005.11.015
1207:10.1016/j.devcel.2007.03.013
692:The construction of the new
624:, followed by deposition of
479:Division plane specification
366:
354:Origin of this term is from
7:
2322:Maturation promoting factor
1766:10.1103/PhysRevE.100.052408
765:
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231:Etymology and pronunciation
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2463:Postreplication checkpoint
1067:10.1038/s41467-021-24474-1
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134:. It thereby ensures that
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1961:Microbiol. Mol. Biol. Rev
1641:. John Wiley & Sons.
1430:10.1007/s00709-005-0095-5
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596:Due to the presence of a
381:
122:. During cytokinesis the
1955:Nanninga N (June 2001).
1908:Anatomy & Physiology
1708:10.3389/fcell.2020.00238
813: – Biology of cells
710:arabinogalactan proteins
235:The word "cytokinesis" (
23:Cytokinesis illustration
898:Oxford University Press
2445:Cell cycle checkpoints
1527:10.1073/pnas.052712299
1286:10.1083/jcb.130.6.1345
1235:Curr. Opin. Plant Biol
1026:Morgan, David (2007).
689:
410:
374:
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38:being clearly visible.
24:
2472:Other cellular phases
2196:CDK-activating kinase
1816:10.4161/cc.8.22.10047
1335:10.1105/tpc.13.9.2033
1047:Nature Communications
894:UK English Dictionary
757:Clinical significance
687:
675:endoplasmic reticulum
404:
95:) is the part of the
30:
22:
1899:available under the
1099:"Cytokinetic bridge"
349:Charles Otis Whitman
1758:2019PhRvE.100e2408B
1695:Front Cell Dev Biol
1518:2002PNAS...99.2812C
1159:10.1038/nature02767
1151:2004Natur.430..908M
1059:2021NatCo..12.4595K
492:At the cytokinesis
209:Circular chromosome
2458:Spindle checkpoint
2263:P53 p63 p73 family
1384:10.1105/tpc.017749
690:
560:Timing cytokinesis
542:cytokinetic bridge
417:separation in the
411:
379:(cellular), Greek
40:
25:
2505:
2504:
2501:
2500:
2453:Restriction point
1917:978-1-947172-04-3
1746:Physical Review E
1675:978-0-8153-3218-3
1648:978-0-470-04737-8
1596:Botanical Gazette
1109:on 28 August 2019
924:. Merriam-Webster
855:10.1002/mrd.21266
538:midbody structure
124:spindle apparatus
2535:
2352:
2351:
2022:
2015:
2008:
1999:
1998:
1994:
1984:
1944:Campbell Biology
1922:
1921:
1894:
1887:
1881:
1880:
1870:
1860:
1851:(10): e1009334.
1835:
1829:
1828:
1818:
1794:
1788:
1787:
1777:
1737:
1731:
1730:
1720:
1710:
1686:
1680:
1679:
1659:
1653:
1652:
1634:
1628:
1627:
1591:
1585:
1584:
1556:
1550:
1549:
1539:
1529:
1497:
1491:
1490:
1480:
1456:
1450:
1449:
1412:
1406:
1405:
1395:
1363:
1357:
1356:
1346:
1314:
1308:
1307:
1297:
1265:
1259:
1258:
1226:
1220:
1219:
1209:
1185:
1179:
1178:
1145:(7002): 908–13.
1134:
1128:
1125:
1119:
1118:
1116:
1114:
1105:. Archived from
1103:proteinatlas.org
1095:
1089:
1088:
1078:
1038:
1032:
1031:
1023:
1010:
1009:
991:
963:
950:
939:
933:
932:
930:
929:
912:
906:
905:
900:. Archived from
883:
877:
876:
866:
834:
777:
519:light microscope
391:
384:
383:
377:
369:
362:
361:
302:
301:
298:
297:
294:
289:
288:
285:
280:
279:
276:
273:
270:
267:
264:
261:
258:
255:
252:
249:
246:
243:
112:nuclear division
94:
93:
90:
89:
86:
83:
80:
77:
74:
71:
68:
65:
62:
59:
56:
53:
2543:
2542:
2538:
2537:
2536:
2534:
2533:
2532:
2508:
2507:
2506:
2497:
2487:
2467:
2439:
2386:
2381:
2367:
2347:
2341:
2285:
2257:
2200:
2107:
2034:
2026:
1946:(2010), 580-582
1931:
1929:Further reading
1926:
1925:
1918:
1888:
1884:
1836:
1832:
1795:
1791:
1738:
1734:
1687:
1683:
1676:
1660:
1656:
1649:
1635:
1631:
1592:
1588:
1557:
1553:
1498:
1494:
1457:
1453:
1424:(3–4): 141–55.
1413:
1409:
1364:
1360:
1315:
1311:
1266:
1262:
1227:
1223:
1186:
1182:
1135:
1131:
1126:
1122:
1112:
1110:
1097:
1096:
1092:
1039:
1035:
1024:
1013:
980:10.1038/ncb2482
964:
953:
940:
936:
927:
925:
914:
913:
909:
885:
884:
880:
835:
831:
826:
775:
768:
759:
751:
735:
730:
671:plasma membrane
656:Golgi apparatus
648:mitotic spindle
594:
562:
534:
527:
511:cleavage furrow
494:cleavage furrow
490:
481:
450:D. melanogaster
439:spindle midzone
435:central spindle
431:
409:and cytokinesis
399:
331:Classical Latin
305:combining forms
291:
282:
240:
236:
233:
171:skeletal muscle
50:
46:
36:cleavage furrow
17:
12:
11:
5:
2541:
2531:
2530:
2525:
2520:
2503:
2502:
2499:
2498:
2496:
2495:
2490:
2485:
2481:
2475:
2473:
2469:
2468:
2466:
2465:
2460:
2455:
2449:
2447:
2441:
2440:
2438:
2437:
2432:
2426:
2421:
2416:
2411:
2406:
2396:
2394:
2388:
2387:
2385:
2384:
2379:
2375:
2370:
2365:
2360:
2358:
2349:
2343:
2342:
2340:
2339:
2329:
2324:
2319:
2314:
2309:
2304:
2299:
2293:
2291:
2287:
2286:
2284:
2283:
2278:
2273:
2267:
2265:
2259:
2258:
2256:
2255:
2237:
2210:
2208:
2202:
2201:
2199:
2198:
2193:
2188:
2183:
2178:
2173:
2168:
2163:
2158:
2153:
2148:
2143:
2138:
2133:
2128:
2123:
2117:
2115:
2109:
2108:
2106:
2105:
2091:
2073:
2059:
2044:
2042:
2036:
2035:
2025:
2024:
2017:
2010:
2002:
1996:
1995:
1952:
1947:
1941:
1938:
1930:
1927:
1924:
1923:
1916:
1882:
1830:
1809:(22): 3742–9.
1789:
1732:
1681:
1674:
1654:
1647:
1629:
1608:10.1086/328131
1586:
1567:(1): 261–288.
1551:
1492:
1451:
1407:
1358:
1329:(9): 2033–51.
1309:
1280:(6): 1345–57.
1260:
1241:(6): 493–502.
1221:
1180:
1129:
1120:
1090:
1033:
1028:The Cell Cycle
1011:
968:Nat. Cell Biol
951:
945:62 (4): 1–80.
934:
907:
904:on 2020-03-22.
878:
843:Mol Reprod Dev
828:
827:
825:
822:
821:
820:
814:
808:
802:
796:
790:
784:
778:
767:
764:
758:
755:
750:
747:
734:
731:
729:
726:
722:middle lamella
706:hemicelluloses
612:, an array of
593:
590:
561:
558:
526:
523:
489:
486:
480:
477:
472:tissue culture
430:
427:
398:
395:
337:, reflecting "
232:
229:
205:binary fission
15:
9:
6:
4:
3:
2:
2540:
2529:
2526:
2524:
2521:
2519:
2516:
2515:
2513:
2494:
2491:
2489:
2482:
2480:
2477:
2476:
2474:
2470:
2464:
2461:
2459:
2456:
2454:
2451:
2450:
2448:
2446:
2442:
2436:
2433:
2430:
2427:
2425:
2422:
2420:
2417:
2415:
2412:
2410:
2407:
2405:
2401:
2398:
2397:
2395:
2393:
2389:
2383:
2376:
2374:
2371:
2369:
2362:
2361:
2359:
2357:
2353:
2350:
2344:
2337:
2333:
2330:
2328:
2325:
2323:
2320:
2318:
2315:
2313:
2310:
2308:
2305:
2303:
2300:
2298:
2295:
2294:
2292:
2288:
2282:
2279:
2277:
2274:
2272:
2269:
2268:
2266:
2264:
2260:
2253:
2249:
2245:
2241:
2238:
2235:
2231:
2227:
2223:
2219:
2215:
2212:
2211:
2209:
2207:
2206:CDK inhibitor
2203:
2197:
2194:
2192:
2189:
2187:
2184:
2182:
2179:
2177:
2174:
2172:
2169:
2167:
2164:
2162:
2159:
2157:
2154:
2152:
2149:
2147:
2144:
2142:
2139:
2137:
2134:
2132:
2129:
2127:
2124:
2122:
2119:
2118:
2116:
2114:
2110:
2103:
2099:
2095:
2092:
2089:
2085:
2081:
2077:
2074:
2071:
2067:
2063:
2060:
2057:
2053:
2049:
2046:
2045:
2043:
2041:
2037:
2033:
2030:
2023:
2018:
2016:
2011:
2009:
2004:
2003:
2000:
1992:
1988:
1983:
1978:
1974:
1970:
1967:(2): 319–33.
1966:
1962:
1958:
1953:
1951:
1948:
1945:
1942:
1939:
1936:
1933:
1932:
1919:
1913:
1909:
1904:
1902:
1898:
1893:
1886:
1878:
1874:
1869:
1864:
1859:
1854:
1850:
1846:
1845:PLOS Genetics
1842:
1834:
1826:
1822:
1817:
1812:
1808:
1804:
1800:
1793:
1785:
1781:
1776:
1771:
1767:
1763:
1759:
1755:
1752:(5): 052408.
1751:
1747:
1743:
1736:
1728:
1724:
1719:
1714:
1709:
1704:
1700:
1696:
1692:
1685:
1677:
1671:
1667:
1666:
1658:
1650:
1644:
1640:
1633:
1625:
1621:
1617:
1613:
1609:
1605:
1601:
1597:
1590:
1582:
1578:
1574:
1570:
1566:
1562:
1555:
1547:
1543:
1538:
1533:
1528:
1523:
1519:
1515:
1512:(5): 2812–7.
1511:
1507:
1503:
1496:
1488:
1484:
1479:
1474:
1471:(1): 137–50.
1470:
1466:
1462:
1455:
1447:
1443:
1439:
1435:
1431:
1427:
1423:
1419:
1411:
1403:
1399:
1394:
1389:
1385:
1381:
1378:(4): 836–56.
1377:
1373:
1369:
1362:
1354:
1350:
1345:
1340:
1336:
1332:
1328:
1324:
1320:
1313:
1305:
1301:
1296:
1291:
1287:
1283:
1279:
1275:
1271:
1264:
1256:
1252:
1248:
1244:
1240:
1236:
1232:
1225:
1217:
1213:
1208:
1203:
1200:(5): 713–25.
1199:
1195:
1191:
1184:
1176:
1172:
1168:
1164:
1160:
1156:
1152:
1148:
1144:
1140:
1133:
1124:
1108:
1104:
1100:
1094:
1086:
1082:
1077:
1072:
1068:
1064:
1060:
1056:
1052:
1048:
1044:
1037:
1029:
1022:
1020:
1018:
1016:
1007:
1003:
999:
995:
990:
985:
981:
977:
973:
969:
962:
960:
958:
956:
948:
944:
938:
923:
922:
917:
916:"cytokinesis"
911:
903:
899:
895:
893:
888:
887:"cytokinesis"
882:
874:
870:
865:
860:
856:
852:
848:
844:
840:
833:
829:
818:
815:
812:
809:
806:
803:
800:
797:
794:
791:
788:
785:
782:
779:
773:
770:
769:
763:
754:
746:
742:
740:
739:Myosin ATPase
725:
723:
719:
715:
711:
707:
703:
699:
695:
686:
682:
680:
679:plasmodesmata
676:
672:
668:
664:
659:
657:
653:
649:
645:
640:
639:
635:
631:
627:
623:
619:
615:
611:
607:
603:
599:
589:
587:
583:
579:
574:
572:
567:
557:
554:
549:
545:
543:
539:
532:
522:
520:
516:
512:
508:
504:
501:cell) at the
499:
495:
485:
476:
473:
469:
466:
462:
458:
457:
452:
451:
446:
445:
440:
436:
426:
424:
420:
416:
408:
403:
394:
393:, movement).
392:
390:
378:
376:
370:
368:
357:
352:
350:
346:
345:
340:
336:
335:Ancient Greek
332:
328:
324:
323:
318:
317:
312:
311:
306:
300:
228:
226:
222:
218:
214:
210:
206:
202:
197:
195:
191:
185:
183:
182:
176:
175:multinucleate
172:
168:
164:
160:
156:
151:
149:
145:
141:
137:
133:
129:
125:
121:
117:
113:
109:
105:
101:
98:
97:cell division
92:
44:
37:
33:
29:
21:
2434:
2414:Prometaphase
1964:
1960:
1943:
1934:
1907:
1889:
1885:
1848:
1844:
1833:
1806:
1802:
1792:
1775:10261/205685
1749:
1745:
1735:
1698:
1694:
1684:
1664:
1657:
1638:
1632:
1599:
1595:
1589:
1564:
1560:
1554:
1509:
1505:
1495:
1468:
1464:
1454:
1421:
1417:
1410:
1375:
1371:
1361:
1326:
1322:
1312:
1277:
1274:J. Cell Biol
1273:
1263:
1238:
1234:
1231:Staehelin LA
1224:
1197:
1193:
1183:
1142:
1138:
1132:
1123:
1111:. Retrieved
1107:the original
1102:
1093:
1050:
1046:
1036:
1027:
974:(5): 440–7.
971:
967:
942:
937:
926:. Retrieved
919:
910:
902:the original
890:
881:
846:
842:
832:
817:Cytoskeleton
760:
752:
743:
736:
733:Animal cells
691:
660:
644:phragmoplast
641:
614:microtubules
610:phragmoplast
595:
578:microtubules
575:
563:
550:
546:
541:
535:
513:. Continued
496:, it is the
491:
482:
464:
454:
448:
442:
438:
432:
412:
405:Animal cell
386:
372:
364:
353:
342:
320:
314:
308:
234:
198:
186:
178:
163:polar bodies
152:
106:of a single
42:
41:
2435:Cytokinesis
2404:Preprophase
2348:checkpoints
1602:(1): 1–34.
1418:Protoplasma
1053:(1): 4595.
989:11336/20338
811:Cell theory
667:endocytosis
503:cell cortex
397:Animal cell
203:process of
201:prokaryotic
43:Cytokinesis
2518:Cell cycle
2512:Categories
2356:Interphase
2346:Phases and
2029:Cell cycle
1803:Cell Cycle
1372:Plant Cell
1323:Plant Cell
1229:Otegui M,
943:Caryologia
928:2016-01-21
849:(1): 3–8.
824:References
698:cell plate
663:cell plate
634:cell plate
628:and other
618:cell plate
606:cell plate
602:cell plate
592:Plant cell
586:cell cycle
584:(see also
582:interphase
571:cell cycle
531:Abscission
525:Abscission
515:hydrolysis
470:and human
465:C. elegans
456:C. elegans
444:H. sapiens
221:chromatids
217:interphase
190:cell plate
159:organelles
148:cell cycle
144:interphase
136:chromosome
128:chromatids
108:eukaryotic
2479:Apoptosis
2429:Telophase
2419:Metaphase
2214:INK4a/ARF
1901:CC BY 4.0
1581:1040-2519
1465:Dev. Cell
1194:Dev. Cell
1113:28 August
799:Metaphase
781:Telophase
718:cellulose
694:cell wall
638:cell wall
630:cell wall
626:cellulose
598:cell wall
461:phenotype
415:chromatid
407:telophase
351:in 1887.
327:Neo-Latin
194:cell wall
181:syncytium
155:oogenesis
140:telophase
104:cytoplasm
2424:Anaphase
2409:Prophase
2032:proteins
1991:11381104
1903:license.
1877:34710087
1825:19855176
1784:31869912
1727:32322581
1624:84936099
1546:11880633
1487:16399085
1446:11881080
1438:16228896
1402:15020749
1353:11549762
1255:11074381
1216:17488623
1167:15282614
1085:34321459
998:22552143
873:21268179
793:Anaphase
787:Prophase
766:See also
652:vesicles
566:anaphase
419:anaphase
225:anaphase
2528:Meiosis
2523:Mitosis
2493:Meiosis
2400:Mitosis
2392:M phase
2373:S phase
2240:cip/kip
1868:8577787
1754:Bibcode
1718:7156592
1701:: 238.
1616:2464904
1514:Bibcode
1304:7559757
1295:2120572
1175:4418281
1147:Bibcode
1076:8319318
1055:Bibcode
1006:3355851
864:3164815
805:Mitosis
772:Diploid
714:callose
702:pectins
622:callose
553:spastin
468:embryos
423:mitosis
389:kínesis
382:κίνησις
344:kinesis
303:) uses
177:cells (
146:of the
120:meiosis
116:mitosis
100:process
32:Ciliate
2332:Cullin
2218:p14arf
2040:Cyclin
1989:
1979:
1914:
1875:
1865:
1823:
1782:
1725:
1715:
1672:
1645:
1622:
1614:
1579:
1544:
1537:122430
1534:
1485:
1444:
1436:
1400:
1393:412860
1390:
1351:
1344:139450
1341:
1302:
1292:
1253:
1214:
1173:
1165:
1139:Nature
1083:
1073:
1004:
996:
892:Lexico
871:
861:
728:Forces
708:, and
341:" and
213:linear
2488:phase
2382:phase
2368:phase
2307:Cdc42
2302:Cdc25
2290:Other
2072:, B3)
1982:99029
1620:S2CID
1612:JSTOR
1442:S2CID
1171:S2CID
1002:S2CID
367:kytos
360:κύτος
356:Greek
329:from
316:kine-
310:cyto-
167:liver
132:cells
2336:CUL7
2297:Cdc2
1987:PMID
1912:ISBN
1897:text
1873:PMID
1821:PMID
1780:PMID
1723:PMID
1670:ISBN
1643:ISBN
1577:ISSN
1542:PMID
1483:PMID
1434:PMID
1398:PMID
1349:PMID
1300:PMID
1251:PMID
1212:PMID
1163:PMID
1115:2019
1081:PMID
994:PMID
947:link
869:PMID
642:The
453:and
437:(or
375:cyto
339:cell
333:and
322:-sis
179:see
169:and
118:and
2327:Wee
2317:E2F
2281:p73
2276:p63
2271:p53
2252:p57
2248:p27
2244:p21
2234:p19
2230:p18
2226:p15
2222:p16
2176:11B
2171:11A
2113:CDK
1977:PMC
1969:doi
1863:PMC
1853:doi
1811:doi
1770:hdl
1762:doi
1750:100
1713:PMC
1703:doi
1604:doi
1569:doi
1532:PMC
1522:doi
1473:doi
1426:doi
1422:225
1388:PMC
1380:doi
1339:PMC
1331:doi
1290:PMC
1282:doi
1278:130
1243:doi
1202:doi
1155:doi
1143:430
1071:PMC
1063:doi
984:hdl
976:doi
859:PMC
851:doi
588:).
507:ATP
421:of
307:of
290:-,-
184:).
114:in
2514::
2250:,
2246:,
2232:,
2228:,
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