Bacterial Cell Division

Abstract

Bacterial cell division or cytokinesis is the process in which a bacterial cell is split into two progeny cells, each with a copy of the chromosome. In most bacteria this process in initiated by the formation of the Z ring, a dynamic structure consisting of polymers of FtsZ, a tublin family member. The Z ring recruits additional division proteins to form the septal ring, also called the divisome, which leads to the synthesis of the septum separating the progeny cells. Spatial regulation of Z‐ring formation occurs primarily through negative regulators of FtsZ assembly that are positioned within the cell. The Z ring forms where the concentration of these negative regulators is at a minimum. A variety of regulators and mechanisms for positioning them have been identified in different bacteria. Among the mechanisms are the binding of a regulator to the segregating chromosome or the divisome itself and a self‐organizing system.

Key concepts:

  • Division depends on a cytoskeletal element (Z ring) that functions as a scaffold to recruit all the division proteins.

  • Spatial regulation of Z ring placement involves positioning inhibitors of FtsZ in the cell to prevent FtsZ polymers from coalescing into the ring.

  • Dynamic self‐organization – the ability of proteins to form dynamic structures and patterns fuelled by nucleotide hydrolysis; the Min system produces an oscillatory pattern and FtsZ filaments have the ability to form rings when attached to a lipid bilayer that has a cylindrical shape.

Keywords: FtsZ; septum; cytokinesis; tubulin; Z ring; binary fission

Figure 1.

Diagram of FtsZ. The first 320 residues (numbering from E. coli) comprise the domain involved in polymerization and GTPase activity. The tail of FtsZ extends an additional 63 residues and the extreme end is required for interaction with additional proteins (FtsA and ZipA) involved in tethering FtsZ to the membrane and an inhibitor of Z ring formation (MinC/D).

Figure 2.

Cell division in E. coli. Before the appearance of the Z ring FtsZ is present in dynamic spirals that move about the cell. As DNA replication nears completion the Z ring forms. It consists of FtsZ filaments tethered to the membrane by linkage to FtsA and ZipA. After a lag (up to half the cell division cycle in rapidly growing cells) the complete divisome is assembled and septation begins. The proteins essential for division are depicted. Many more proteins localize to the division site than depicted here and are required for cell wall synthesis, splitting the septal crosswall and invagination of the outer membrane.

Figure 3.

Two examples of spatial regulation of cell division in bacteria. Z ring placement is regulated by negative regulatory systems that prevent FtsZ filaments from coalescing into a ring. These negative regulators are positioned by the nucleoid and septation itself by forming the cell poles. In E. coli the Min system restricts Z ring assembly away from midcell through the placement of MinC at the poles. The inhibitory zone set up by the Min system overlaps that imposed by SlmA bound to the nucleoid (Noc). These inhibitory zones are indicated by the coloured lines above and below the cell. In Caulobacter crescentus a newly formed stalked cell has FtsZ present at the nascent pole formed in the previous division. As the replication origin is duplicated one of the origins, along with the bound ParB and the MipZ gradient it generates (indicated by the orange colour), migrates to the opposite pole releasing FtsZ. FtsZ assembles near midcell where the concentration of MipZ is lowest.

Figure 4.

Min oscillation is a self‐organizing system that prevents Z ring placement near the poles of the cell. In E. coli the Min proteins undergo a coupled oscillation. MinD binds to the membrane in an ATP‐dependent manner and recruits MinC. MinE binds to MinD, displaces MinC and stimulates the ATPase activity of MinD. This action of MinE releases the proteins from the membrane. In models of the oscillation the only requirements are MinE, MinD, ATP and an enclosed system. Z ring assembly in the polar region is prevented by the membrane‐bound MinC which antagonizes FtsZ filaments. Modified with permission of Karger S and Basel AG, from Dajkovic and Lutkenhaus .

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Further Reading

Goley ED, Iniesta AA and Shapiro L (2007) Cell cycle regulation in Caulobacter: location, location, location. Journal of Cell Science 120: 3510–3517.

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Lutkenhaus, Joe(Dec 2009) Bacterial Cell Division. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1002/9780470015902.a0000294.pub2]