5.

3 MreB and Cell Morphology

Just as specific proteins direct cell division in prokaryotes, other
specific proteins direct cell shape. Interestingly, these shapedetermining
proteins show significant homology to key cytoskeletal
proteins in eukaryotic cells. Like eukaryotes, prokaryotes
also contain a cell cytoskeleton, and one that is both dynamic and
multifaceted.
Cell Shape and MreB
The major shape-determining factor in Bacteria is a protein
called MreB. MreB forms a simple cytoskeleton in Bacteria and in
a few species of Archaea. MreB forms a helix of filaments around
the inside of the cell, just below the cytoplasmic membrane (Figure
5.5). The MreB cytoskeleton presumably defines cell shape
by recruiting other proteins that function in cell wall growth to
group into a specific pattern. Inactivation of the gene encoding
MreB in rod-shaped bacteria causes the cells to become coccusshaped.
Moreover, most naturally coccoid bacteria lack the MreB
gene and thus do not make MreB. This indicates that the default
morphology for a bacterium is most likely the sphere. Variations
in the arrangement of MreB filaments in cells of nonspherical
bacteria are probably responsible for the different common morphologies
of prokaryotic cells ( Figure 2.11).
How does MreB define a cells shape? The helical structures
formed by MreB (Figure 5.5a) are not static, but instead can
rotate within the cytoplasm of a growing cell. Newly synthesized
peptidoglycan (Section 5.4) is associated with the MreB helices
at points where the helices contact the cytoplasmic membrane
(Figure 5.5a). It is thought that MreB localizes the synthesis of
new cell wall to specific locations along the long axis of a rodshaped
cell during growth. This allows new cell wall to form at
several points along the cell rather than from a single location at
the FtsZ site outward, as in spherical bacteria (see Figure 5.3). By
rotating within the cell cylinder and initiating cell wall synthesis
where it contacts the cytoplasmic membrane, MreB directs new
wall synthesis in such a way that a rod-shaped cell elongates only
along its long axis.
Crescentin
Caulobacter crescentus, a vibrio-shaped species of Proteobacteria
( Section 7.12 and 14.21), produces a shape-determining
protein called crescentin in addition to MreB. Copies of crescentin
protein organize into filaments about 10 nm wide that localize
onto the concave face of the curved cell. The arrangement
and localization of crescentin filaments are thought to impart the
characteristic curved morphology to the C. crescentus cell (Figure
5.5c). Caulobacter is an aquatic bacterium that undergoes a life
cycle in which swimming cells, called swarmers, eventually form