CH391L/S12/Locomotion

Locomotion

Flagella

Structure of Escherichia coli flagellum [1]

The bacterial flagellum is the most common and thoroughly studied prokaryotic motility structure. It resembles a spinning propeller-like structure that is used for swimming in aqueous environments and in some organisms enables swarming across solid surfaces. The flagellum is a very complex organelle consisting of over 20 proteins (flg, flh, fli, flj variants) and as many as 30 proteins assisting in regulation and assembly. Each Escherichia coli or Salmonella cell typically has 6-8 structures. The export system for assembly of the structure represents a classical Type III secretion system (T3SS). The main structure consists of 3 main substructures: the basal body, which anchors the structure in the cell membrane and contains the motor; the filament which acts as the propeller; and the hook, a joint which connects the basal body and filament. Rotation of the filament to generate movement is driven by the proton motive force, whereby H⁺ atoms crossing the cell membrane interact with the motor proteins (MotA, MotB), inducing a conformational change that turns the rotor. This rotation can reach speeds of 18,000rpm and propel the cell 25-35uM per second.

Chemotaxis

Regulation of flagellar rotation

The rotation of the flagellum and the direction of movement is often regulated by sensory stimuli, allowing the cell to migrate towards attractive signals. In E.coli this is achieved through a signal transduction system that controls the phosphorylation state of the response regulator protein CheY. In the absence CheY-P the flagellum rotates CWW in a "run" state. The presence of CheY-P, induces a switch to CW rotation resulting in "tumbling". When an attractant binds to a receptor it initiates a conformational change and downstream cascade that leads to suppression of CheA which can no longer phosphorylate CheY to CheY-P and the cells remain in the "run" state leading to migration towards the signal.

Gas Vesicles

Conical gas vesicles in Serratia sp[2]

Some aquatic bacteria use hollow gas-filled vesicles to provide buoyancy and enable them regulate their position in the water column. Prototrophic bacteria may use the vesicles to find regions with appropriate light intensity[3], similarly aearobic bacteria may use them to float to oxygenated surface waters[4] . Gas vesicles are commonly observed and studied in aquatic cyanobacteria, but have recently been discovered in Serratia sp., an enterobacterium [2]. Typically 10-14 gvp genes are involved in vesicle formation. It has been shown that gas vesicles from Anabaena are permeable to H₂, N₂, 0₂, C0₂, CO, CH₄, and Ar [5].

Twitching Motility

Mattick 2002[6, 7]

Many species of bacteria including Pseudomonas aeruginosa,Neisseria gonorrhoeae and Myxoccocus xanthus use a Type IV pilus (T4P) system for motility. Cell propulsion by T4P involves pilus extension, attachment, and then pilus retraction. This process results in a jerky pattern of movement which hence the name "Twitching motility". Cells move at rates of 0.05 - 1 uM per second, close proximity to other cells is usually required for efficient movement. The process of pilus extension and retraction involves the ATP-dependent assembly or disassembly of PilA monomers in the pilus fiber.

Kearns 2010

Gliding Motility

Focal adhesions/motor complexes in Myxococcus xanthus[8]

Slime production in Myxococcus xanthus[8]

References

<biblio>

1. Jarrell2008 pmid=18461074
2. Ramsey2011 pmid=21873216
3. Walsby1971 pmid=4337701
4. Mattick2002 pmid=12142488
5. Damerval1991 pmid=1904525
6. Zussman2011 pmid=21910630
7. Beard2002 pmid=12167531