A flagellum (/fləˈdʒɛləm/; pl.: flagella) (Latin for 'whip' or 'scourge') is a hair-like appendage that protrudes from certain plant and animal sperm cells, from fungal spores (zoospores), and from a wide range of microorganisms to provide motility.[1][2][3][4] Many protists with flagella are known as flagellates.
A microorganism may have from one to many flagella. A gram-negative bacterium Helicobacter pylori, for example, uses its flagella to propel itself through the stomach to reach the mucous lining where it may colonise the epithelium and potentially cause gastritis, and ulcers – a risk factor for stomach cancer.[5] In some swarming bacteria, the flagellum can also function as a sensory organelle, being sensitive to wetness outside the cell.[6]
Across the three domains of Bacteria, Archaea, and Eukaryota, the flagellum has a different structure, protein composition, and mechanism of propulsion but shares the same function of providing motility. The Latin word flagellum means "whip" to describe its lash-like swimming motion. The flagellum in archaea is called the archaellum to note its difference from the bacterial flagellum.[7][8]
Eukaryotic flagella and cilia are identical in structure but have different lengths and functions.[9] Prokaryotic fimbriae and pili are smaller, and thinner appendages, with different functions. Cilia are attached to the surface of flagella and are used to swim or move fluid from one region to another. [10]
The three types of flagella are bacterial, archaeal, and eukaryotic.
The flagella in eukaryotes have dynein and microtubules that move with a bending mechanism. Bacteria and archaea do not have dynein or microtubules in their flagella, and they move using a rotary mechanism.[12]
Other differences among these three types are:
The bacterial flagellum is made up of protein subunits of flagellin.[12] Its shape is a 20-nanometer-thick hollow tube. It is helical and has a sharp bend just outside the outer membrane; this "hook" allows the axis of the helix to point directly away from the cell. A shaft runs between the hook and the basal body, passing through protein rings in the cell's membrane that act as bearings. Gram-positive organisms have two of these basal body rings, one in the peptidoglycan layer and one in the plasma membrane. Gram-negative organisms have four such rings: the L ring associates with the lipopolysaccharides, the P ring associates with peptidoglycan layer, the M ring is embedded in the plasma membrane, and the S ring is directly attached to the cytoplasm. The filament ends with a capping protein.[21][22]
The flagellar filament is the long, helical screw that propels the bacterium when rotated by the motor, through the hook. In most bacteria that have been studied, including the gram-negative Escherichia coli, Salmonella typhimurium, Caulobacter crescentus, and Vibrio alginolyticus, the filament is made up of 11 protofilaments approximately parallel to the filament axis. Each protofilament is a series of tandem protein chains. However, Campylobacter jejuni has seven protofilaments.[23]
The basal body has several traits in common with some types of secretory pores, such as the hollow, rod-like "plug" in their centers extending out through the plasma membrane. The similarities between bacterial flagella and bacterial secretory system structures and proteins provide scientific evidence supporting the theory that bacterial flagella evolved from the type-three secretion system (TTSS).
The atomic structure of both bacterial flagella as well as the TTSS injectisome have been elucidated in great detail, especially with the development of cryo-electron microscopy. The best understood parts are the parts between the inner and outer membrane, that is, the scaffolding rings of the inner membrane (IM), the scaffolding pairs of the outer membrane (OM), and the rod/needle (injectisome) or rod/hook (flagellum) sections.[24]
The bacterial flagellum is driven by a rotary engine (Mot complex) made up of protein, located at the flagellum's anchor point on the inner cell membrane. The engine is powered by proton-motive force, i.e., by the flow of protons (hydrogen ions) across the bacterial cell membrane due to a concentration gradient set up by the cell's metabolism (Vibrio species have two kinds of flagella, lateral and polar, and some are driven by a sodium ion pump rather than a proton pump[26]). The rotor transports protons across the membrane, and is turned in the process. The rotor alone can operate at 6,000 to 100,000 rpm,[27] but with the flagellar filament attached usually only reaches 200 to 1000 rpm. The direction of rotation can be changed by the flagellar motor switch almost instantaneously, caused by a slight change in the position of a protein, FliG, in the rotor.[28] The flagellum is highly energy efficient and uses very little energy.[29] [unreliable source?] The torque is transferred from the MotAB to the torque helix on FliG's D5 domain and with the increase in the requirement of the torque or speed more MotAB are employed.[25] Because the flagellar motor has no on-off switch, the protein epsE is used as a mechanical clutch to disengage the motor from the rotor, thus stopping the flagellum and allowing the bacterium to remain in one place.[30]
The cylindrical shape of flagella is suited to locomotion of microscopic organisms; these organisms operate at a low Reynolds number, where the viscosity of the surrounding water is much more important than its mass or inertia.[31]
The rotational speed of flagella varies in response to the intensity of the proton-motive force, thereby permitting certain forms of speed control, and also permitting some types of bacteria to attain remarkable speeds in proportion to their size; some achieve roughly 60 cell lengths per second. At such a speed, a bacterium would take about 245 days to cover 1 km; although that may seem slow, the perspective changes when the concept of scale is introduced. In comparison to macroscopic life forms, it is very fast indeed when expressed in terms of number o