Cytokinesis

Abstract

Cytokinesis is the term given to the division of a cell into two daughter cells. Cytokinesis in all cells requires the resolution of the single lipid bilayer of the parent cell into separate bilayers enclosing the two daughter cells, yet cytokinesis mechanisms differ markedly in different cell types. In bacteria, a ring of FtsZ protein filaments constricts the membrane. FtsZ‐like proteins operate in some Archeal species, but others use a different mechanism. In plants, vesicles are deposited by microtubules in the centre of the cell to create sheets of membrane that grow outwards until they reach and fuse with the cytoplasmic membrane. In animal cells, actin and myosin form a contractile ring, which draws the membrane inwards. Bundled microtubules and vesicle trafficking then complete the separation (abscission). In all cases, cytokinesis closely follows chromosome segregation and constriction occurs between the separating chromosomes to ensure the genetic integrity of the daughter cells.

Key Concepts:

  • To make two cells from one, the DNA needs to be copied, the two copies separated and the cell divided (cytokinesis) so that one DNA copy resides in each daughter cell.

  • The cytoplasmic membrane of the parent cell must resolve into the two cytoplasmic membranes of the daughter cells.

  • Cell division frequently involves the formation of a ring of protein filaments that attaches to the membrane and constricts, drawing the membrane inwards.

  • In many bacterial cells, the main component of this ring is the FtsZ protein. In animal cells, the main components are actin and myosin.

  • In plant cells, microtubules form across the future site of division and deposit membrane vesicles which fuse to form the new membrane.

  • In many eukaryote cells, the mitotic spindle allows the DNA to separate, and then signals the cell to determine the plane of division.

  • In animals, the final separation of daughter cells involves vesicles fusing with the cell membrane.

  • Errors in cytokinesis have serious implications for cancer.

Keywords: cell division; abscission; mitosis; spindle; cytoskeleton; DNA; aneuploidy

Figure 1.

A model for the oscillation of MinCDE localisation to suppress the formation of the FtsZ ring at the poles of an E. coli cell. MinD‐ATP binds to the membrane at one pole of the cell. MinC binds to MinD‐ATP and co‐localises with it, inhibiting FtsZ ring formation. MinE, present as a ring at the edge of the MinD zone, displaces MinC and stimulates the MinD ATPase resulting in release of MinD from the membrane. The ADP bound to the released MinD is then exchanged for ATP, which results in re‐dimerisation, localisation to the membrane at the other pole and recruitment of MinC to that pole. Once the MinE ring reaches the cell pole MinE is released and reassembles at the edge of the new MinD zone at the other end of the cell. Because of this oscillation, MinC alternately inhibits FtsZ polymerisation at the poles, resulting in assembly of the FtsZ ring (orange) in the middle of the long axis of the cell. Adapted with permission from Lutkenhaus J and Sundaramoorthy M (2003) MinD and role of the deviant Walker A motif, dimerisation and membrane binding in oscillation. Molecular Microbiology 48: 295–303. Reprinted with permission from John Wiley & Sons Ltd. (http://onlinelibrary.wiley.com/DOI/10.1046/j.1365‐2958.2003.03427.x/abstract).

Figure 2.

Animal cytokinesis and the contractile ring. Diagrammatic representation of actomyosin contractile rings in animal cells. Attachment of the actin filaments to the plasma membrane via a formin is hypothetical. Movement of myosin‐II towards the barbed (anchored) end of an actin filament it will draw the plasma membrane inwards. Illustrations by Graham Johnson. Reprinted with permission from Pollard ; Elsevier B.V. (http://www.sciencedirect.com/science/journal/09550674).

Figure 3.

Positioning the contractile ring. Centralspindlin complexes move to the midzone along microtubules where they concentrate and activate the RhoGEF through a direct protein–protein interaction. This leads to a cortical ring of activated Rho1 between the separated chromosomes. The activated Rho1 leads to formation and activation of the actomyosin contractile ring (not shown). Reprinted with permission from Saint R and Somers WG (2003) Animal cell division: a fellowship of the double ring? Journal of Cell Science116: 4277–4281. (http://jcs.biologists.org/cgi/pmidlookup?view=long&pmid=14514883) The Company of Biologists.

Figure 4.

Establishing the contractile ring. (a) Possible role for the scaffold protein, Anillin, in establishing the actoymosin contractile ring. Following activation of RhoA by Ect2A/Pbl at the cortical midzone, RhoA and RacGap interact with the Anillin protein, which may recruit actomyosin to the membrane to form the contractile ring. (b) Possible roles of RhoA in cytokinesis Activated RhoA is known to trigger the two key events in formation of actomyosin contractile networks. First, it can bind to and activate Formin proteins, the first step in a cascade of events that leads to actin polymerisation. Secondly, it can bind to and activate protein kinases such as Rho kinase (ROK) and citron kinase (CITK), which can phosphorylate the myosin regulatory light chain (MRLC), causing myosin contraction. Adapted with permission from D'Avino PP (2009) How to scaffold the contractile ring for a safe cytokinesis – lessons from Anillin‐related proteins. Journal of Cell Science122: 1071–1079. (http://jcs.biologists.org/cgi/pmidlookup?view=long&pmid=19339546). The Company of Biologists.

Figure 5.

Abscission Possible role of microtubules and vesicle trafficking in abscission during animal cell cytokinesis. Components are delivered by molecular motors such as KIF13A, which move to the plus ends of the midbody microtubules and, mediated perhaps by association with components of the ESCRT‐III complex, induce abscission. Reprinted from Nezis IP, Sagona AP, Schink KO and Stenmark H (2010) Divide and ProsPer: The emerging role of PtdIns3P in cytokinesis. Trends in Cell Biology20: 642–649 with permission from Elsevier.

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Gregory, Stephen L, and Saint, Robert(May 2011) Cytokinesis. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1002/9780470015902.a0022536]